[the Families and Genera of Vascular Plants 6] K. Kubitzki (Auth.), Professor Dr. Klaus Kubitzki (Eds.) - Flowering Plants · Dicotyledons_ Celastrales, Oxalidales, Rosales, Cornales, Ericales (2004, Springer-Verlag

THE FAMILIES
AND GENERA
OF VASCULAR PLANTS
Edited by K. Kubitzki
Springer-Verlag Berlin Heidelberg GmbH

Volumes published in this series:
Volume I Pteridophytes and Gymnosperms

Edited by K.U. Kramer and P.S. Green (1990)
Date of publication: 28.9.1990
Volume II Flowering Plants. Dicotyledons. Magnoliid, Hamamelid and Caryophyllid Families

Edited by K. Kubitzki, J.G. Rohwer, and V. Bittrich (1993)
Volume III Flowering Plants. Monocotyledons: Lilianae (except Orchidaceae)

Edited by K. Kubitzki (1998)
Volume IV Flowering Plants. Monocotyledons: Alismatanae and Commelinanae (except

Gramineae)
Volume V Flowering Plants. Dicotyledons: Malvales, Capparales and Non-betalain

Caryophyllales
Edited by K. Kubitzki and C. Bayer (2003)
Volume VI Flowering Plants. Dicotyledons: Celastrales, Oxalidales, Rosales, Cornales, Ericales

The Families
and Genera
of Vascular Plants
Edited by K. Kubitzki
VI Flowering Plants · Dicotyledons

Celastrales, Oxalidales, Rosales, Cornales, Ericales
Volume Editor:
K. Kubitzki
With 137 Figures
~Springer
Professor Dr. KLAUS KuBITZKI
Institut fiir Allgemeine Botanik
OhnhorststraBe 18
22609 Hamburg
Germany
ISBN 978-3-642-05714-4
Library of Congress Cataloging-in-Publication Data

Flowering plants, Dicotyledons : Celastrales, Oxalidales, Rosales, Cornales, Ericales I volume editor, K. Kubitzki.
p. em. - (Families and genera of vascular plants; 6)
Includes bibliographical references (p. ).
ISBN 978-3-642-05714-4 ISBN 978-3-662-07257-8 (eBook)
DOI 10.1007/978-3-662-07257-8
1. Dicotyledons - Classification. I. Kubitzki, Klaus, 1933- II. Series.
QK495.A12F56 2004
583'.01'2- dc22 2003058977
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Preface
During recent years the findings of molecular systematics for the first time have provided
a reliable lead for a higher order classification of angiosperms, as reflected in the classi-
fication of the Angiosperm Phylogeny Group. From Volume III of this series onward, these
new concepts have been used as a guideline for treating angiosperm families in a mean-
ingful context within individual volumes of this series. Among the five orders treated in
this volume, Celastrales and Oxalidales represent novel concepts of which at least the
latter makes sense also in morphological terms. The concept of Ericales followed here has
largely been expanded beyond its conventional limits to include parts of former Ebenales,
Lecythidales, Primulales and other orders. Also, the circumscription of many families has
been revised on the basis of recent molecular studies. Among the recircumscribed fami-
lies, Ericacae, Celastraceae, Cunoniaceae and Elaeocarpaceae appear now in revised and
expanded circumscriptions, and the former Primulales families have been profoundly re-
organised. I am most grateful to my authors who have incorporated into their treatments
the most recent results of molecular and cladistic studies. At the same time I, as the editor,
also feel the dilemma of prejudicing those authors who responded at an early stage of the
preparation of these volumes and whose contributions had to wait for publication until
late authors were able to comply. Lamentably, R.C. Carolin's treatments of Stackhousiaceae
and Tremandraceae, submitted in the early 1990s, have been superseded by later devel-
opments, and I am most grateful to Roger Carolin for his understanding and generous
permission to use his treatments in a changed family context.
I would like to direct the attention of the users of this series to the remarkable treat-
ment of Ericaceae in this volume. Its author, P.F. Stevens, is introducing here an interest-
ing kind of "hierarchical" description, which emphasises apomorphic characters and
leads to great clearness and conciseness, the constantly aspired aims of this series.
It would hardly have been possible to complete this volume without the help so gener-
ously offered by many colleagues. Most substantial support has been provided by C. Bayer
and P.F. Stevens who reviewed many contributions and suggested most helpful improve-
ments. A.S. George is thanked for building up a team that contributed Styphelioideae
and thus made possible completion of Ericaceae in the new circumscription; I am also
grateful for his kind assistance in various other matters. Deep thanks for reading and com-
menting on particular contributions are due to R. Archer (Celastraceae), I.K. Ferguson
(Cornaceae), J.W. Kadereit (Introduction), S.R. Manchester (Cornaceae), M. Matthews
(Cunoniaceae), B. Stahl (Symplocaceae), S. Renner (Sarraceniaceae), R. Rutishauser
(Cunoniaceae) and J. Thompson (tremandraceous Elaeocarpaceae). D. Albach and
L. Hufford are thanked for their comments on problems concerning Hydrostachyaceae,
and M. Matthews and P.K. Endress for showing me papers prior to publication.
My thanks go also to the copyright holders of the illustrations who so generously autho-
rised the use of the material included in this volume.
Finally, it is a pleasure to acknowledge the agreeable collaboration with the staff of
Springer-Verlag and Pro Edit, Heidelberg, who kindly responded to all requests I had in
connection with the completion of this volume.
Hamburg, October 2003 K. KUBITZKI

Contents
Introduction to Families Treated in This Volume

K. KUBITZKI ...... . 1
GENERAL REFERENCES 12
Actinidiaceae S. DRESSLER and C. BAYER 14
Balsaminaceae E. FISCHER • 20
Brunelliaceae K. KUBITZKI 26
Celastraceae M.P. SIMMONS . . • . • . • . 29

Cephalotaceae J.G. CONRAN . . . . . . . . . 65
Clethraceae J.V. SCHNEIDER and C. BAYER 69
Connaraceae R.H.M.J. LEMMENS, F.J. BRETELER
and C.C.H. JoNKIND • . • . • . 74
Cornaceae K. KUBITZKI . • . . . • . • . • • . 82
Cunoniaceae J.C. BRADFORD, H.C. FoRTUNE-HOPKINS
and R.W. BARNES 91
Curtisiaceae K. KUBITZKI 112
Cyrillaceae K. KUBITZKI 114
Diapensiaceae P.J. SCOTT 117

Dirachmaceae C. BAYER 122
Ebenaceae B. W ALLNOFER . . . . . . . . . 125

Elaeagnaceae I.V. BARTISH and U. SWENSON 131
Elaeocarpaceae M.J.E. CoonE . • . . • . • . • . 135
Ericaceae P.F. STEVENS, with J. LUTEYN, E.G.H. OLIVER, T.L. BELL,
E.A. BROWN, R.K. CROWDEN, A.S. GEORGE,
G.J. JoRDAN, P. LADD, K. LEMSON, C.B. McLEAN,
Y. MENADUE, J.S. pATE, H.M. STACE,
C.M. WEILLER . . . . . . . . . . . . . . . . . . . 145
Fouquieriaceae K. KUBITZKI 195
Grubbiaceae K. KUBITZKI 199
Hydrangeaceae L. HUFFORD ••.•• 202

Hydrostachyaceae C. ERBAR and P. LEINS 216
Lecythidaceae G.T. PRANCE and S.A. MoRI 221

Lepidobotryaceae K. KUBITZKI • 233
Lissocarpaceae B. W ALLNOFER 236
Loasaceae M. WEIGEND . 239
VIII Contents
Maesaceae B. STAHL and A.A. ANDERBERG 255

Marcgraviaceae S. DRESSLER . . . . . . . . . . 258
Myrsinaceae B. STAHL and A.A. ANDERBERG 266
Napoleonaeaceae G.T. PRANCE . 282
Oxalidaceae A.A. Cocucci 285
Parnassiaceae M.P. SIMMONS 291

Pellicieraceae K. KUBITZKI . 297
Polemoniaceae D.H. WILKEN . 300
Primulaceae A.A. ANDERBERG 313
Rhamnaceae D. MEDAN and C. SCHIRAREND 320

Roridulaceae J.G. CONRAN 339
Rosaceae C. KALKMAN 343
Samolaceae B. STAHL .. 387

Sapotaceae T.D. PENNINGTON 390
Sarraceniaceae K. KUBITZKI . . . 422
Scytopetalaceae O.APPEL . . . . . 426
Sladeniaceae P.F. STEVENS and A.L. WEITZMAN 431
Styracaceae P.W. FRITSCH . . . 434
Symplocaceae H.P. NooTEBOOM 443
Ternstroemiaceae A.L. WEITZMAN, S. DRESSLER and P.F. STEVENS 450

Tetrameristaceae K. KUBITZKI . . . . . . . . . . . . . . . . . . . 461
Theaceae P.F. STEVENS, S. DRESSLER and A.L. WEITZMAN 463
Theophrastaceae B. STAHL . . . . . . . . . 472
Index of Scientific Names 479

List of Contributors
ANDERBERG, A.A. Department of Phanerogamic Botany, Swedish Museum of

Natural History, PO Box 50007, 104 05 Stockholm, Sweden
APPEL, 0. Bredkamp 36e, 22589 Hamburg, Germany
BARNES, R. Department of Primary Industries, Water and Environment,
GPO Box 44, Hobart, Tasmania 7001, Australia
BARTISH, LV. Institute of Botany, Academy of Sciences,
25243 Pruhonice, Czech Republic
BAYER, C. Palmengarten, Siesmayerstr. 61,60323 Frankfurt/M., Germany
BELL, T.L. Department of Botany, University of Western Australia,
Nedlands, Western Australia 6987, Australia
BRADFORD, J.C. Environmental Science and Policy, University of California at
Davis, 2132 Wickson Hall, Davis, CA 95616, USA
BRETELER, F.J. Herbarium Vadense, Landbouwuniversiteit Wageningen,
Postbus 80120,6700 ED Wageningen, The Netherlands
BROWN, E.A. National Herbarium of New South Wales, Royal Botanic
Gardens, Mrs Macquarie's Road, Sydney, New South Wales
2000, Australia
Cocucci, A.A. Instituto Multidisciplinario de Biologia Vegetal,
C. C. 495, 5000 Cordoba, Argentina
CONRAN, J.G. Department of Environmental Biology, University of
Adelaide, SA 5005, Australia
CoonE, M.J.E. Royal Botanic Gardens Kew, Richmond, Surrey TW9 3AS, UK
CROWDEN, R.K. Plant Science, University of Tasmania, GPO Box 252-55,
Hobart, Tasmania 7001
DRESSLER, S. Forschungsinstitut Senckenberg, Senckenberganlage 25,
60325 Frankfurt/Main, Germany
ERBAR, C. Biodiversitat und Pftanzensystematik, Heidelberger Institut
fiir Pftanzenwissenschaften (HIP), Universitat Heidelberg,
lm Neuenheimer Feld 345,69120 Heidelberg, Germany
FISCHER, E. Institut fiir Biologie, Universitat Koblenz-Landau,
Universitatsstr. 1, 56070 Koblenz, Germany
FoRTUNE-HOPKINS, H. Royal Botanic Gardens Kew,
Richmond, Surrey TW9 3AS, UK
FRITSCH, P. W. Department of Botany, California Academy of Sciences,
Golden Gate Park, San Francisco, CA 94118, USA
GEORGE, A.S. 'Four Gables', 18 Barclay Road, Kardinya,
Western Australia 6163, Australia
X List of Contributors
HUFFORD, L. School of Biological Sciences, Washington State University,

Pullman, WA 99164-4236, USA
JoNGKIND, C.C.H. Herbarium Vadense, Landbouwuniversiteit Wageningen,
JoRDAN, G.J. Plant Science, University of Tasmania, GPO Box 252-55,
KALKMAN,C. Deceased, formerly National Herbarium Netherlands,
Leiden branch
KUBITZKI, K. Institut fiir Allgemeine Botanik, UniversiHit Hamburg,
Ohnhorststr. 18,22609 Hamburg, Germany
LADD,P. Department of Environmental Science, Murdoch University,
Murdoch, Western Australia 6150, Australia
LEINS, P. BiodiversiHit und Pflanzensystematik, Universitat Heidelberg,
Im Neuenheimer Feld 345,69120 Heidelberg, Germany
LEMMENS, R.H.M.J. Herbarium Vadense, Landbouwuniversiteit Wageningen,
LEMSON, K. School of Natural Sciences, Edith Cowan University,
100 Joondalup Drive, Western Australia 6027, Australia
LUTEYN, J. The New York Botanical Garden, Bronx, New York 10458, USA
McLEAN, C.B. ICFR, 500 Yarra Boulevard, Richmond, VIC 3121, Australia
MEDAN, D. Catedra de Botanica, Facultad de Agronomfa de la
Universidad de Buenos Aires, Avenida San Martin 4453,
C1417DSQ Buenos Aires, Argentina
MENADUE, Y. Plant Science, University of Tasmania, GPO Box 252-55,
MORI, S.A. The New York Botanical Garden, Bronx, New York 10458, USA
NooTEBOOM, H.P. National Herbarium of the Netherlands, Leiden branch,
Einsteinweg 2, 2333 CC Leiden, The Netherlands
OLIVER, E.G.H. Compton Herbarium, National Botanical Institute,
Kirstenbosch, P. Bag x7, Claremont 7735, South Africa
PATE, J.S. Department of Botany, University of Western Australia,
PENNINGTON, T.D. Royal Botanic Gardens Kew, Richmond,
Surrey TW9 3AS, UK
PRANCE, G.T. The Old Vicarage, Silver Street, Lyme Regis,
Dorset DT7 3HS, UK
SCHIRAREND, C. Botanischer Garten der Universitat Hamburg,
Hesten 10, 22609 Hamburg, Germany
SCHNEIDER, J.V. Forschungsinstitut Senckenberg, Senckenberganlage 25,
60325 Frankfurt/Main, Germany
SCOTT, P.J. Memorial University of Newfoundland, St. John's,
Newfoundland A1 C 5S7, Canada
SIMMONS, M.P. Department of Biology, Colorado State University
Fort Collins, Colorado 80523, USA
List of Contributors XI
STACE,H.M. Department of Botany, University of Western Australia,

STAHL, B. Hogskolan pa Gotland, 621 57 Visby, Sweden
STEVENS, P.F. Missouri Botanical Garden, PO Box 299,
St. Louis Missouri 64507, USA
SWENSON, U. Department of Phanerogamic Botany, Swedish Museum of
Natural History, PO Box 50007, 104 05 Stockholm, Sweden
W ALLNOFER, B. Naturhistorisches Museum Wien, Botanische Abteilung,
Burgring 7, Postfach 417, 1014 Wien, Austria
WEIGEND,M. lnstitut fiir Biologie/Systematische Botanik, Freie Universitiit
Berlin, Altensteinstr. 6, 14195 Berlin, Germany
WEILLER, C.M. Research School for Biological Sciences, Australian National
University, Australian Capital Territory 0200, Australia
WEITZMAN, A.L. Department of Botany, Smithsonian Institution, Washington,
DC, 20560, USA
WILKEN, D.H. Santa Barbara Botanic Garden, 1212 Mission Canyon Road,
Santa Barbara, CA 93105, USA

K. KUBITZKI
1. Celastrales Fay et al. 2000; Soltis et al. 2000; APG II 2003). The
Celastrales clade comprises only three families,
Lepidobotryaceae, Celastraceae s.l. and Parnassi-
1. Herbs with conspicuous tannin sacs in epidermis; leaves
estipulate; flowers hermaphrodite; disk 0; stigmas com-
aceae. The rbcL analysis by Savolainen, Fay et al.
missural; [ovules tenuinucellate, numerous per loculus, (2000) provided strong support for a position
bitegmic or unitegmic; capsules loculicidal]. 2/71, North- of Lepidobotryaceae as sister to the other two
ern Hemisphere and South America Pa~nassiace~e families. Parnassiaceae, comprising Parnassia
- Usually woody without conspicuous tannm sacs m and Lepuropetalum, have often been included in
epidermis; stipules small and caducous or 0; flowers _her-
maphrodite to unisexual; plant~ (andr?)monoe~wus,
Saxifragaceae. There exists, however, _strong
(gyno)dioecious, or polygamous; d1sk +;stigmas cannal morphological (see Simmons on Parnass1aceae,
2 this volume) and molecular evidence for their
2. Leaves unifoliolate with articulated petioles and solitary exclusion from Saxifragaceae. Various analyses of
stipels; flowers in racemes or spikes; ovules 2 per locule, plastid and nuclear genes (Savolainen, Chase
collateral; stylodia 2 or 3, nearly free, or very short; [cap-
sules septicidal; endocarp and exocarp separating; seed et al. 2000; Soltis et al. 1997, 2000) have resolved
arillate; endosperm 0]. 2/2, Africa and Mesoamerica Parnassiaceae as sister to Celastraceae. In a multi-
Lepidobotryaceae gene analysis of Celastraceae (Simmons et al.
- Leaves simple; flowers in thyrsoids or inflorescences 2001b), Parnassia and Lepuropetalum have been
derived there from but rarely racemes or spikes; ovules
1-numerous/locule (alternating if >1); style simple or
resolved as members of an early branching but
rarely branched or cleft into stylodia; [fruit loculicidally weakly supported lineage of that family, in which
and/ or septicidally capsular, schizocarpic, baccate, they are sister to Perrottetia and Mortonia. The
samaroid, or drupaceous; seed arillate or not, winged latter two genera, as well as the early-derived Quet-
or not; endosperm + or 0]. 98/c. 1200, of worldwide zalia, are somewhat anomalous among Celas-
distribution Celastraceae
traceae in lacking an aril in favour of a sarcotesta,
and partly in possessing scalariform vessel pe:fo-
In traditional classifications Celastrales comprised rations. For the time being it seems therefore JUS-
families of usually woody plants with simple tified to retain family status for Parnassiaceae, as
leaves, haplostemonous flowers with a disk and suggested by Simmons in his contribution to this
apotropous ovules which, in view of this character volume. As a result of Simmons' (2001a, 2001b)
combination, formed an utterly heterogeneous analysis, Celastraceae are now re-circumscribed
assemblage. In the classification of Engler and Gilg to comprise the genera Brexia, Canotia,
(1912), for instance, families as divergent as Bu~ Plagiopteron, Siphonodon, Stackhousiaceae and
aceae, Rhamnaceae, Aquifoliaceae and Balsami- Hippocrateaceae, all of which at one time or
naceae were dumped into their Sapindales another had been related to Celastraceae, and all
(= Celastrales ). In later classifications, most of which have now been shown to be nested within
authors gave up this broad circumscription but the that family.
Celastrales of Takhtajan (1987), to give just one
As discussed above, Celastrales and also Celas-
example, comprised 12 families of which nine,
traceae are weakly characterised morphologically,
according to our present knowledge, would have to as is obvious from the four morphological synapo-
be excluded from this order. In the absence of
morphies Simmons et al. (2001b) adduce for the
convincing morphological evidence, only sequ-
latter, all being subject to reversals: stamen plus
ence-based phylogenetic studies have led to the
staminode number equals petal number; filaments
recognition of a monophyletic order Celastrales
inserted at the outer margin of the disk; styles
which represents a clade of eurosids I that is sister
undivided; and presence of 2-4 ovules per locule.
to Oxalidales + Malpighiales, these three being None of these characters are found in all three
sister to all remaining eurosids I (Fagales, Rosales, families, but it may be added that an indumentum
Zygophyllales, Curcurbitales, Fabales; Savolainen,
is weakly developed in all of them.
2 K. Kubitzki
2. Oxalidales details of how the three tremandraceous genera

are linked to that family are still unknown.
Morphologically, both Elaeocarpaceae and
1. Hetero-(tri- and di- )-stylous; rapanone present; ellagic
acid 0 [petals postgenitally fused into a tube above the free Cunoniaceae are difficult to characterise but, inter-
petal bases; ovules almost orthotropous] 2 estingly, racemisation of thyrso-paniculate inflo-
- Homostylous; rapanone 0; ellagic acid present or not 3 rescences seems to have taken place in both.
2. Herbaceous or woody; carpels 5(4), ovarial portions ± Striking similarities in floral structure between
united; fruit capsular or baccate; ovules bitegmic/tenuinu- Cunoniaceae
and Anisophyllaceae have been
cellate. 6/880, widely distributed in temperate and tropical
regions Oxalidaceae emphasised by Matthews and Endress (2002);
- Woody throughout; carpels 5-1, always free; fruit these are interpreted as convergence by molecular
follicular; ovules bitegmic/crassinucellate. 12/110-200, workers (A. Schwarzbach, in litt., Dec. 2002).
pantropical Connaraceae
3. Petals 0; apocarpous 4
- Petals +; syncarpous, if apocarpous then leaves 3-4 per
node or flowers unisexual; [pollen grains exceptionally 3. Rosales
small] 5
4. Herbaceous rosulate perennial with insect-trapping pitch-
ers; flowers always 6-merous, carpels completely plicate; 1. Androecium of several whorls, basically 10 + 5 + 5, but
stylodia with punctate stigmas; seeds exarillate. 1/1, sometimes reduced to a single whorl; [woody or herba-
Western Australia Cephalotaceae ceous; condensed tannins generally present, ellagitannins
- Woody; leaves simple or imparipinnate, stipulate; flowers and cyanogenic glycosides often present; nodes trilacunar;
4-6( -8)-merous; carpels ascidiate; stylodia with decurrent stipules free or adnate to petiole, rarely 0; hypanthium
stigmas; seeds arillate. 1/61, Mexico to Bolivia, Greater usually well-developed; epicalyx (when present), sepals,
Antilles Brunelliaceae petals and stamens inserted on its rim; nectariferous disk
5. Leaves opposite or ternate (spiral in Davisonia), simple or sometimes present, intrastaminal; carpels 1-many, free or
compound; stipules often interpetiolar; filaments exceed- variously connate; seed coat mostly mesotestal-sclerotic ].
ing the petals; anthers opening longitudinally (apically 85/c. 2000, almost cosmopolitan Rosaceae
in Bauera), dorsifixed; disk inter- or intrastaminal; rarely - Androecium of a single 3-6-merous whorl (in Shepherdia
apocarpous; stylodia free. 27/c. 300, Meso- and South 2 whorls) 2
America, S Africa, Madagascar, Australia, Malesia, and 2. Flowers with sepals and petals; syncarpous 3
islands of W Indian Ocean and S Pacific Cunoniaceae - Flowers apetalous 4
- Leaves usually alternate, simple, stipules lateral or 0; petals 3. Sepals not keeled, not hooded; nectary disk 0; flowers
often fringed but all but one of the numerous New World solitary. 1/2, Socotra and Somalia Dirachmaceae
species of Sloanea apetalous; anthers dehiscing with - Sepals often keeled; petals mostly hooded; nectary disk
terminal pores or slits, basifixed; disk extrastaminal; not present; flowers in cymes or fascicles; [woody or herba-
apocarpous; styles nearly always undivided. 12/c. 550, ceous; calyx valvate; stamens alternisepalous ]. 52/925,
widely distributed in tropical and temperate regions, not almost cosmopolitan Rhamnaceae
in Africa Elaeocarpaceae 4. Gynoecium apparently 1-carpellate or apocarpous;
stipules 0; cystoliths 0; nodes unilacunar; ellagic acid
On morphological grounds, Oxalidaceae have present 5
often been included in Geraniales, but molecular - Gynoecium syncarpous/2-carpellate; stipules present; cys-
toliths often present; nodes trilacunar; ellagic acid 0 6
studies have defined Oxalidales as a clade of 5. Indumentum of simple hairs; nectary disk 0; gynoecium
eurosids I that receives strong statistic support in 1( -3)-carpellate/apocarpous; ovules pendulous. 1/1, arid
independent large-scale analyses (e.g. Savolainen, regions of NE Africa and adjacent Arabian Peninsula
Chase et al. 2000; Savolainen, Fay et al. 2000; Soltis Barbeyaceae (see Vol. II of this series)
et al. 2000), and always groups with Malpighiales - Indumentum of stellate-peltate hairs; nectary disk present;
gynoecium apparently 1-carpellate (pseudomonomerous);
and Celastrales in various constellations. Within ovules ascending. 3/30-50, temperate regions of Northern
Oxalidales, relationships are still unclear but Con- Hemisphere, extending to SE Asia and E Australia
naraceae and Oxalidaceae appear closely related Elaeagnaceae
and may be sister to the remaining families. 6. Flowers bisexual or unisexual; embryo straight; hypan-
thium present; cystoliths rare; pollen 4-6-porate; second-
Morphologically, Connaraceae and Oxalidaceae
ary leaf veins usually ending in teeth; [laticifers 0]. 7/c. 48,
are very close; apart from the tristyly and (near) Northern Hemisphere and extending to Africa and South
apocarpy, they have similar seed coats (Corner America Ulmaceae s.str. (see Vol. II of this series)
1976) and agree in sieve element plastids with pro- - Flowers unisexual throughout; embryo curved; hypan-
teinaceous inclusions (Behnke 1982). The molecu- thium 0; cystoliths common; pollen 2-3-porate; secondary
leaf veins ending subterminally 7
lar data also suggest a close relationship among 7. Laticifers 0 (Humulus and Cannabis with secretory canals
Cunoniaceae/Elaeocarpaceae, Cunoniaceae/ in the phloem containing a watery liquid); fruits drupa-
Brunoniaceae, and Brunelliaceae/Cephalotaceae, ceous, solitary; plants woody. 8/130-140, temperate and
but statistic support is not satisfactory in any of tropical regions worldwide
these cases (Bradford and Barnes 2001). Treman- Celtidaceae (incl. Cannabaceae) (see under Ulmaceae
and Cannabaceae, Vol. II of this series)
draceae are nested within Elaeocarpaceae but
Introduction to Families Treated in This Volume 3
- Laticifers present; fruits drupes or achenes, usually densely treatment of these families ten years ago in Vol. II
clustered; plants woody or herbaceous; [condensed of this series. Thus, Celtidaceae may merit family
tannins ± 0] 8
8. Woody to herbaceous with laticifers throughout the plant status independent from Ulmaceae; Cannabaceae
and containing milky latex; stamens straight; ovules may be a derived lineage of Celtidaceae; and
(sub)apically attached; stigmatic branch(es) 2 (1). 37/1100, Cecropiaceae appear nested within Urticaceae, as
worldwide but mainly tropical indicated in the conspectus.
Moraceae (see Vol. II of this series) The Rhamnalean clade is less well characterised
- Herbs, herbaceous vines, or trees; laticifers restricted
to cortex and containing mucilaginous latex; stamens morphologically and its monophyly appears still
straight or inflexed; ovules (sub- )basally attached; stig- problematic, partly due to contradictions between
matic branch 1. 51/1120, widely distributed in tropical and morphological and molecular evidence, and partly
temperate regions of the world due to poor statistic support (Thulin et al. 1998),
Urticaceae (incl. Cecropiaceae) (see Vol. II of this series)
yet the inclusion of Barbeyaceae and Dirach-
maceae in this lineage seems appropriate.
In the past, Rosaceae had often been linked to
Chrysobalanaceae but, starting with the work of
Chase et al. (1993), many molecular phylogenetic
analyses have contributed to shaping Rosales in 4. Cornales and Ericales
the present circumscription, as reflected in the
ordinal classification of the Angiosperm Phy- Families of Cornales
logeny Group (1998, 2003). Rosales are part of the
nitrogen-fixing clade of eurosids I, which other- 1. Submerged herbs with perianthless flowers; primary root
wise includes Fabales, Cucurbitales and Fagales lacking; stylodia 2; endosperm cellular, in mature seed
(Soltis et al. 1995). Both the nitrogen-fixing scanty or none; stipule intrapetiolar; [male flowers of 1 (or
clade and its subclades are all strongly supported 2?) stamens; female flowers bicarpellate ]. 1/22, C and South
as monophyletic in various broad-based gene Africa, Madagascar Hydrostachyaceae
- Terrestrial plants with primary root; flowers usually with
sequence analyses (e.g. Soltis et al. 2000; a perianth (except Davidia); style simple or with style
Savolainen, Chase et al. 2000). branches; stipules none 2
Rosaceae are resolved as the first branch of 2. Ovule 1 per carpel; ovary inferior, with epigynous
Rosales. The family is well characterised by its disk (except Davidia); fruits drupaceous; trichomes not
tuberculate 3
recently discovered, very specific androecial struc-
- Ovules more than 1 per carpel; ovary superior to inferior;
ture; the gynoecium is largely apocarpous, the fruits capsular, rarely baccate; trichomes tuberculate with
carpels often contain more than one ovule, and the basal cell pedestals 5
seeds are (nearly) exalbuminous. The androecium 3. Leaves strongly serrate; fruits 4-seeded; [disk barbate;
of Rosaceae consists of an outer whorl of paired gynoecium with axile bundle supply to ovules; inflores-
cence terminal, thyrso-paniculate]. 1/1, southern Africa
antepetalous stamens and two simple antesepa- Curtisiaceae
lous and antepetalous whorls (10 + 5 + 5; - Leaves entire (Davidia, Camptotheca serrate/dentate);
Lindenhofer and Weber 1999a, 1999b, 2000). fruits 1(-3-5)-seeded 4
Androecia with antepetalous stamen pairs fol- 4. Inflorescences capitate or cone-like, compound of dicha-
lowed by simple stamen whorls are otherwise sia; ovaries of flowers of one partial inflorescence coherent
or connate; anthers 2-sporangiate, dehiscing with valves;
known from families such as Alismataceae and disk barbate; ovary 2-carpellate; fruit without germination
Aristolochiaceae (Leins and Erbar 1985), and in valve. 1/3, South Africa Grubbiaceae
Rosaceae may be plesiomorphic. The remaining Inflorescences thyrso-paniculate; ovaries not coherent or
Rosales families nearly always have only a single connate; anthers 4-sporangiate, dehiscing by longitudinal
stamen whorl. Among these families, two clades, slits; disk glabrous; ovary 1-2(-9)-carpellate; fruit often
with germination valve. 7/110, temperate and tropical
the Urticalean and the Rhamnalean clades, may be regions mostly of Northern Hemisphere Cornaceae
distinguished. In both clades the carpels are 5. Tanniniferous shrubs, vines, rarely herbs; vessel elements
uniovulate and the corolla has been lost in part of with scalariform, rarely simple perforation; stylodia free,
them, and the intron of the mitochondrial nad1 or style solitary, sometimes branched. 17/220, North and
gene is trans-spliced (in contrast to the cis-spliced South America, Eurasia, Pacific islands Hydrangeaceae
Non-tanniniferous coarse herbs, rarely shrubs, vines,
intron in Rosaceae and most other dicots, Qiu et treelets, ferociously armed with scabrid, glochidiate and/ or
al. 1998). stinging hairs; vessel elements usually with simple perfo-
Support for the Urticalean clade is strong both ration; style unbranched; [ovules tenuinucellate, some
in terms of morphological and molecular evidence crassinucellate?, some with terminal haustoria]. 20/330,
mostly American, some in Africa and Polynesia
(Sytsma et al. 2002; Stevens 2003), and may call for Loasaceae
a revision of family limits in comparison with the
4 K. Kubitzki
Families of Ericales 11. Seed coat reduced to 1 cell layer or lacking altogether; style
hollow, fluted; [vessel element perforation scalariform;
endosperm cellular] 12
1. Placenta free, central; [vessel element perforation simple; - Seed coat usually well-developed, several cells thick; style
flowers haplostemonous with sympetalous corolla; usually not hollow (hollow in many Styracaceae) 14
stamens antepetalous; ovules bitegmic, unitegmic in 12. Corolla sympetalous, cylindrical-urceolate/choripetalo~s;
Aegiceras and Cyclamen, tenuinucellate; endosperm
anthers inverting late/ early in floral ontogenesis,
development nuclear] 2 with/without 2(4) appendages; pollen in monads/tetrads;
- Placenta axile or parietal 6 [anthers tetrasporangiate/bisporangiate, dehiscing with
2. Flowers with staminodes alternating with stamens 3 longitudinal slits/pores; endothecium present/reduced;
- Staminodes lacking 4 ovary 5(-1 or -12)-carpellate]. 124/4100, warm-temperate
3. Woody; upper and lower part of anther thec~e containi~g and montane-tropical Ericaceae
meal of calcium oxalate crystals; ovules not Immersed m - Corolla choripetalous, or petals fused at most up to 1/3;
placentae. 6/90; neotropical Theophrastaceae anthers inverting at most in bud, not appendaged; pollen
- Herbaceous; anthers not containing meal of calcium in monads 13
oxalate; ovules immersed in placentae. 1/12, almost 13. Indumentum of simple and fasciculate hairs; pedicels
cosmopolitan . ~amolac~ae
without prophylls; anthers ventrifixed/versatile, inverting
4. Pedicels provided with prophylls; trees with mixed umse- at anthesis, dehiscing by apical, pore-like slits; ovary 3-
riate and multiseriate rays. 1/150, tropical regions of Old locular. 2/95, North and tropical America, Asia
World Maesaceae Clethraceae
- Prophylls lacking; woody or herbaceous, rays only multi- - Glabrous; pedicels with prophylls; anthers dorsifixed, not
seriate, or rayless 5 inverting at anthesis, dehiscing by longitudinal slits; ovary
5. Herbs, shrubs, or trees with secretory cavities in vegetative 2-5-locular, each locule with 1-3 ovules and only 1 seed;
and reproductive organs; flowers with short tube; ovules [sieve element plastids Pcf (the only example within an
immersed in placentae (Ardisiandra excepted). 48/1500, otherwise Ss type order)]. 2/2, North, Central and N South
almost cosmopolitan Myrsinaceae America Cyrillaceae
- Herbs; secretory cavities lacking; flowers with long or 14. Insect-trapping herbs or shrubs; [vessel element perfora-
short tube; ovules not immersed in placentae (Dionysia tion scalariform; inflorescence a botryoid] 15
excepted). 14/600, predominantly north-temperate - Not insect trapping 16
Primulaceae 15. Shrubs; leaves bearing stalked insect-trapping glands;
6. Raphide cells present; [endosperm cellular; vessel ele- stamens 5; anthers curved in bud and swinging upward
ments usually with simple perforation] 7 when touched at anthesis to become erect; endothecium
- Raphide cells lacking . _11 lacking fibrous thickenings. 1/2, Cape Province of South
7. Ovules unitegmic; iridoids present; [shrubs or hanas w_Ith Africa Roridulaceae
spiral, serrate leaves and often rather flat, setose ~airs; - Perennial herbs; leaves ascidiate traps partly filled with
polystemonous or obdiplostemonous; anthers s?metlmes digestive liquid; stamens 10-numerous; anthers basifixed
poricidal; gynoecium with simple style or stylo~Ia_l. ~/360, or versatile; endothecium well-developed. 3/15, North and
America, Asia Achmd~aceae
N South America Sarraceniaceae
- Ovules bitegmic (rarely unitegmic in Balsaminace~e; 16. Low-growing mycotrophic herbs or shrublets, nearly
unknown in Pellicieraceae); iridoids unknown; [with rayless; [functional stamens 5, alternipetalous, inserted in
integumentary tapetum] . 8 floral tube; vessel elements commonly with simple perfo-
8. Fleshy herbs; branched sclereids absent; [flowers vertiCally rations]. 5/13, circumboreal Diapensiaceae
monosymmetric, inverting during growth; sepals 5(3), Herbs, vines, shrubs or trees; rays usually well-developed
the functional abaxial sepal with prominent nectariferous (not so in some Polemoniaceae) 17
spur; fruit an explosive capsule]. 2~850, mostly_ Old World, 17. Pollen 3-4-porate or 4-60-aperturate 18
Africa, esp. Madagascar to mountamous SE Asia - Pollen 3(4, very rarely [Sapotaceae] 5)-colp(or)ate;
Balsaminaceae
WOO~ 19
- Woody; with branched sclereids (lacking in 18. Trees; pollen 3-porate; flowers 4-merous throughout, sym-
Tetrameristaceae) 9 petalous, with an 8-lobed corona in~erted distally _in fl?ral
9. Inflorescences with abaxially ascidiate, nectar-secreting tube; nectary disk absent; style simple; ovary mfenor;
bracts; lianas, epiphytes, often heterophyllous;. pet~ls ovules 2 per carpel. 1/c. 5, tropical South America
lacking glandular pits; ovules many per carpel; [fruits with Lissocarpaceae
numerous small seeds borne on fleshy, coloured placenta]. - Herbs, rarely vines, shrubs; pollen strikingly sculptured,
7/130, Central and South America Marcgraviaceae (4-)6-60-porate or colp(or)ate with deeply immersed
- Inflorescence without ascidiate bracts; erect shrubs and pores or colpi; flowers usually 5-merous with 3-carpell~te
trees; petals with adaxial glandular pits; ovule I/ carpel 10 gynoecium, corona lacking; intrastaminal ne~tary d1sk
10. Mangrove tree with fluted trunk bases; ~rophylls and usually present; style 3-branched; ovary supenor; ovules
sepals petaloid, surpassing the petals; ovary Imperfectly 2- 1-many per carpel; [vessel element perforation simple].
locular; fruit !-seeded, indehiscent; seeds large, exalbu- 18/380, America, some in Eurasia Polemoniaceae
minous. 1/1, Central and N South America 19. Bizarre spinose shrubs or small trees with woody or suc-
Pellicieraceae culent trunks; [vessel elements with simple perforation;
- Trees and shrubs lacking sclereids; trunks not fluted; pro- nodes unilacunar; prophylls present; flowers sympetalous;
phylls and sepals not petaloid; ovary 4-5-locular; fruit a ovules bitegmic]. 1/11, S USA and Mexico
4-5-seeded berry; endosperm copious. 2/2-4, disjunct Fouquieriaceae
between N South America and SE Asia/Malesia - Woody, not succulent, not spiny (except Chinese
Tetrameristaceae Sinojackia) 20
20. Young axis with cortical vascular bundles; stipules often The recognition of the orders Cornales and
present 21 Ericales in the circumscription of the Angiosperm
- Young axis without cortical vascular bundles; stipules
usually (Sapotaceae excepted) absent 23
Phylogeny Group II (2003), as followed in this
21. Seeds albuminous; [vessel element perforation scalari- volume, is the result of numerous studies in molec-
form; petals replaced by pseudocorolla of 6-28 fused sta- ular systematics, starting with Olmstead et al.
minodes; nectary disk absent; seeds often ruminate]. 6/21, (1992) and Chase et al. (1993), based on sequence
West Africa, tropical South America Scytopetalaceae analyses of the rbcL gene, and many subsequent
- Seeds exalbuminous; [vessel element perforation simple
and scalariform] 22 investigations, which often employed several
22. Petals free, 3-6( -18) (lacking in Foetidia); stamens usually plastid, mitochondrial and/or nuclear genes.
connate at base and often forming a strap-like structure More recently, Xiang et al. (2002) have presented
(sometimes nectariferous) arching over the ovary; nectary detailed molecular analyses of Cornales, and
disk absent; fruit sometimes operculate; seeds arillate or Kallersjo et al. (2000) and Anderberg et al. (2002)
not; embryos undifferentiated or with fleshy cotyledons.
17/282, pantropical Lecythidaceae have analysed the phylogenetic relationships of
- Pseudocorolla of 30-35 fused staminodes; flowers fully the Ericales. There is now ample molecular evi-
actinomorphic; nectary disk annular; fruit drupaceous; dence for the monophyly both of Cornales and
seeds exarillate; embryos well-differentiated. 2/10, West Ericales at the base of a larger monophyletic
Africa Napoleonaeaceae
23. Pubescence of stellate or stellate-peltate trichomes; style
group, Asteridae, with Cornales and Ericales sub-
usually hollow; [cork deep-seated; vessel element perfora- sequently being sister to all remaining asterid taxa
tion scalariform; prophylls absent; sympetalous, usually (Albach et al. 2001c and references therein). Since
campanulate; anthers basifixed; nectary disk absent; fruit the decisive criterion for these ordinal concepts
a capsule or drupe; ovules bitegmic or unitegmic; testa is the statistical support in parsimony and/or
vascularized]. 11/160, America, Europe, E and SE Asia,
Malesia Styracaceae
maximum likelihood analyses, there is little
- Pubescence, if present, different; style not hollow 24 wonder that their circumscription is a far cry from
24. Ovules bitegmic 25 that of traditional orders, and particularly the
- Ovules unitegmic 28 Ericales are an inclusive group embodying also
25. Vessel element perforations simple; bark and heartwood the Primulales, parts of the traditional Theales
black; [secretory cells common; cork surficial; prophylls
lacking; sympetalous; anthers basifixed; ovary 2-8-locular, and Ebenales, and fragments of other traditional
superior; ovules 2 per loculus; style usually very short but orders as well. These components of the Ericales
divided in long style branches; fruit a berry; x = 15]. often seem morphologically to have little, if any-
2/500-600, pantropical Ebenaceae thing, in common, and a cladistic analysis based
- Vessel element perforations scalariform; bark and heart-
on morphological and chemical characters by
wood not black 26
26. Inflorescence cymose; sclereids absent; anthers dehiscing Anderberg (1992) failed to discover synapo-
by apical pores or slits, [basifixed; cork surficial; fruit a morphies for members of these two orders.
schizocarp, with persistent columella]. 2/3, SE Asia, East In the pre-molecular era, Cornales and Ericales
Africa Sladeniaceae had been part of what today is considered the
- Inflorescence racemose, or flowers solitary; sclereids
widely present; anther dehiscence by slits; [leaves serrate,
rosid alliance and were not included in the erst-
elongating while still rolled up; accumulating aluminium, while Sympetalae which, by and large, coincide
and condensed and hydrolysable tannins] 27 with the "higher" asterids or euasterids. Whereas
27. Anthers versatile; cork usually deep-seated; pseudopollen in the euasterids character expressions such as
produced from connective; capsule with persistent sympetalous and haplostemonous flowers, 2-
column; embryo straight. 7/125-420, SE Asia, Indomalesia,
America Theaceae carpellate ovaries, unitegmic and tenuinucellate
- Anthers basifixed; cork usually surficial; pseudopollen ovules, cellular endosperm development, simple
absent; fruit a berry or drupe; embryo often curved. vessel perforations and the lack of tannins prevail,
12/340, worldwide, tropical and subtropical regions, few in in the rosids choripetalous and polystemonous or
Africa Ternstroemiaceae (ob )diplostemonous flowers, ovaries with more
28. Vessel element perforation simple; latex present; leaves
usually entire; hairs often T-shaped, unicellular; inflores- than two carpels, bitegmic and crassinucellate
cence fasciculate, rarely a raceme; prophylls lacking; ovules, cellular and/or nuclear endosperm devel-
stamens antepetalous; ovules 1 per locule; testa shiny, opment, scalariform vessel element perforations
smooth, with rough scar. 53/1100, pantropical and presence of condensed and/or hydrolysable
Sapotaceae
- Vessel element perforation scalariform; no latex; leaves
tannins are the rule. In Cornales and Ericales, a
usually serrate; plants glabrous or with multicellular hairs mixture of character expressions of the two major
and stalked glands; inflorescence thyrso-paniculate; pro- groups is to be found which highlights the inter-
phylls present; ovules 2 per locule; seed enclosed in hard mediate position of these orders. Although tradi-
endocarp, testa thin. 1/c. 300, tropical and subtropical tionally Cornales and Ericales were placed
America, S and E/SE Asia, Australia to Fiji
Symplocaceae
distantly from one another in the system, some
6 K. Kubitzki
authors recognised similarities between them. In derived than Cornaceae (Huber 1963). The posi-
a broad-based comparative study, and guided tion of Loasaceae has been rather dubious for
mainly by the Comus-type wood anatomy, Huber a long time, but their close relationship with
(1963) grouped many families from different tra- Hydrangeaceae is now well settled. On the basis of
ditional orders together, among which Cornaceae morphological characters, Takhtajan (1959) and
and Hydrangeaceae and also Styracaceae and Hufford (1992) suggested a relationship between
Symplocaceae were deemed to be particularly these families; subsequent molecular work has
close to one another. He also emphasised the close- confirmed their sister group status. The many
ness of Cornales and Ericales, which he thought similarities between the basal subfamily of
to be perhaps only separable on the absence vs. Hydrangeaceae, Jamesioideae, and Loasaceae (see
presence of terminal endosperm haustoria. A few discussion under ''Affinities" in Loasaceae, this
years later the significance of the monoterpene- volume) are quite convincing in this respect but
derived iridoids was recognised and, building on Loasaceae, in their floral specialisations and pre-
the pioneering work of, amongst others, Hegnauer vailing herbaceousness, significantly accompanied
(1966), Bate-Smith and Swain (1966) and Kooiman by the lack of condensed tannins, have strongly
(1969), Dahlgren (1975, 1983) shaped his sup- diverged from Hydrangeaceae.
erorder Cornanae based on the correlation In contrast, Cornaceae, Curtisiaceae and Grub-
between sympetaly, unitegmic and tenuinucellate biaceae have mainly haplostemonous, often 4-
ovules, cellular endosperm formation, terminal merous epigynous flowers with inferior ovaries
endosperm haustoria and the occurrence of containing solitary ovules, drupaceous fruits and
iridoids. He also stated (Dahlgren et al. 1981) much-reduced seed coats. Eyde (1988) considered
that "[a]n independent origin of several (or even the transition to epigyny with the concomitant
numerous) groups of plants with this combination evolution of a single-seeded fruit chamber and the
of characters is highly unlikely". Later molecular fruit stone as the key innovation of the Cornus-
work (e.g. Xiang et al. 1998) led to the exclusion nyssoid-mastixioid alliance. These traits must
from Cornales of several smaller families now have been introduced into this lineage before the
included in Garryales, Solanales and Apiales. branching-off of the Grubbia/Curtisia alliance,
Although amply showing the iridoid theme, they because Curtisia has retained a 4-seeded fruit with
lack condensed tannins and ellagitannins and an axile bundle supply, whereas in Cornaceae the
also polyandric flowers, but often contain caffeic 1-seeded condition has gradually evolved during
(or chlorogenic) acid, and thus also morphologi- the Tertiary (see "Palaeobotany" in Cornaceae, this
cally/chemically fit the asterid pattern. volume). Yet wood anatomically, Cornus and allies
The most detailed molecular analyses of the appear to be most archaic; among Cornaceae and
Cornales (Soltis et al. 2000; Albach et al. 2001c; Hydrangeaceae, both fossil and extant, a link as
Xiang et al. 2002) recover a monophyletic group direct as between Hydrangeaceae and Loasaceae
comprising several well supported clades, yet can not be recognised.
leave us uncertain as to their interrelationships. In molecular studies Hydrostachys has been
The following clades can be recognised: 1. Grub- found nearly consistently a member of Cornales,
biaceae + Curtisiaceae, 2. Cornus + Alangium, 3. albeit in unstable positions, moving around in
mastixioids + nyssoids (2 and 3 here combined Hydrangeaceae, more rarely in Loasaceae, and
in the family Cornaceae, see "Phylogeny" under sometimes even outside the Cornales, obviously
Cornaceae, this volume), 4. Hydrangeaceae + due to long-branch attraction. In the analyses
Loasaceae, and 5. Hydrostachys, the latter placed of Xiang et al. (2002), measures were taken to
either at the very basis of the Cornales clade or on minimise the influence of long branches, and
a long branch in different positions nested in Hydrostachys switched into a position at the base
Hydrangeaceae or more rarely Loasaceae. The res- of the cornalean clade.
olution of basal relationships among these clades The original ordinal concept of Ericales com-
remains unknown. prised little more than Ericaceae and their satellite
From the morphological point of view, one families today included in that family. Later, Dia-
would see Hydrangeaceae in a basal position pensiaceae, Clethraceae, Cyrillaceae, Actinidiaceae
within Cornales, at least according to their floral and Grubbiaceae were added, but the strong
and fruit morphology: they have often hemiapoc- expansion of the group, as it stands today, became
arpous gynoecia with free stylodia and many- necessary through insights from molecular
seeded capsular fruits, all perhaps plesiomorphic; studies. These included the work of Morton et al.
wood anatomically, they are somewhat more (1996, 1997), Kallersjo et al. (2000) and Albach et
Balsaminaceae
Pellicieraceae
Tetrameristaceae
Marcgraviaceae
Fouquieriaceae
Polemoniaceae
- Sladeniaceae
Ternstroemiaceae
Theaceae
Ebenaceae
I Lissocarpaceae
Symplocaceae
Maesaceae
Theophrastraceae
Samolaceae
Primulaceae
Myrsinaceae
Styracaceae
Diapensiaceae
Napoleonaeaceae
I
Scytopetalaceae
Lecythidaceae
Sapotaceae
Actinidiaceae
Roridulaceae
Sarraceniaceae
Clethraceae
Cyrillaceae
I Ericaceae
al. (2001c), and finally the broad-based five-gene Fig. 1. Parsimony jackknife tree of Ericales families, based on
analysis of Anderberg et al. (2002), whose lead I an analysis of sequences from the five genes atbP, ndhF, rbcL,
atpl, and matK. Redrawn, with permission, from Anderberg
am following here. The tree in Fig. 1 shows two et a!. (2002); Scytopetalaceae added after Morton et a!. (1997)
clades: Balsaminaceae, Pellicieraceae, Tetrameris-
taceae with Marcgraviaceae; and Fouquieriaceae
with Polemoniaceae subsequently sister to the
remaining families, which form a polytomy of ovules, cellular endosperm formation, and
eight clades, six of which comprise only one or two micropylar endosperm haustoria (embryological
families. data are known only for Marcgraviaceae and
The relationship among the four families of the Balsaminaceae ). Marcgraviaceae and Pellicieraceae
basal clade has first been recognised by Morton et are Neotropical, Tetrameristaceae are disjunct
al. {1996). Although strongly supported, they are between northern South America and South-East
dissimilar yet have in common the possession Asia, and Balsaminaceae are widespread but
of raphides, simple vessel perforations, bitegmic mainly in the Old World. The families of the fol-
8 K. Kubitzki
lowing clades have bitegmic or unitegmic ovules, accumulation of aluminium. van den Oever et al.
the bitegmic having mostly nuclear endosperm, (1981) therefore raise the question whether Sym-
the unitegmic usually with cellular, just as would plocaceae could be closely related to Ternstroemi-
be expected (see Dahlgren 1991; Albach et al. aceae, which would merit further consideration.
2001a). Previously suggested phylogenetic links between
The second clade, consisting of Fouquieriaceae Symplocaceae and Sapotaceae (Morton et al.1996)
and Polemoniaceae, is basal to all remaining are not supported by more recent analyses.
taxa of the Ericales and has been confirmed As expected, the five families of (ex-)Primulales
by several molecular studies. Nevertheless, boot- are firmly grouped together, with Maesaceae
strap support for this clade is only moderate sister to the other families, and Samolaceae
(Anderberg et al. 2002), and Fouquieriaceae and and Theophrastaceae both basal to Primulaceae
Polemoniaceae differ in the number of integu- and Myrsinaceae (Anderberg and Stahl 1995).
ments, the mode of endosperm development, and Morphological and molecular analyses have
the presence of a chalazal endosperm haustorium shown Primulaceae and Myrsinaceae in their
and iridioids. Nash (1903) had already pointed to traditional circumscription to be profoundly
a certain similarity between Gilia and Fouquieria, interdigitated, and a complete reorganisation has
which both inhabit the same region, and Hufford become inevitable, with Maesa and Samolus ele-
(1992) focused on the spiny long-shoots of vated to family rank (see Maesaceae, Samolaceae,
Fouquieria and the polemoniaceous Acanthogilia; this volume).
these are, however, morphologically quite distinct In molecular analyses, Diapensiaceae have
and the relationship between the two may not be shifted from their traditional position close to
as close. For a possible relationship between Pole- Ericales into a position sister to Styracaceae, for
moniaceae and the Primulales, which is not sup- which strong support is available. This alignment
ported by the five-gene analysis, see Albach et al. was first recognised by Morton et al. (1996), who
(2001c: 181). discussed the morphological traits common to
Sladeniaceae and Ternstroemiaceae (the ex- both families; diplostemony may be basic in them.
Theaceae-Ternstroemioideae) form a further However, profound differences between the two
clade, again modestly supported. As far as is should not be overlooked and include gynoecium
known, they have bitegmic ovules and nuclear position, seed coat anatomy and, of course, the
endosperm but differ in various traits. Theaceae s. suite of traits in Diapensiaceae that are related
str. are on a separate clade of the molecular tree; to their pronounced mycotrophy. Also, the seeds
this is reflected by their morphological characters differ strongly: exotestal-theoid in Diapensiaceae,
in which they differ from Ternstroemiaceae, which exotestal-mesotestal, not theoid in Styracaceae.
include versatile anthers with the stamens some- The unitegmic ovules are parallelisms, as Styrax is
times in five antepetalous groups, branched styles bitegmic.
and fruit structure. The phylogenetic relationships among Lecythi-
In the past Ebenaceae and Lissocarpaceae have daceae and their relatives have been explored on
always been grouped together; they form another the basis of morphological and molecular data by
clade in the tree of the five-gene-analysis, which Morton et al. (1996). They found a weak associa-
has high bootstrap support. However, the suggestion of these families with Sapotaceae, which has
tion by Berry et al. (2001) to combine both fami- been confirmed, although with moderate support,
lies into one is not followed here, as both differ in by the five-gene analysis of Anderberg et al. (2002).
several important characters (see Conspectus). This is surprising, because the closest relatives
Symplocaceae are well characterised by repro- of Sapotaceae had been expected to share their
ductive and vegetative traits and are unique in antepetalous stamens, but this sapotaceaous trait
having antesepalous stamen fascicles; increase has no counterpart in the invariably polyandric
in stamen number in other Ericales families lecythidaceous families. Sapotaceae, which are
(Sapotaceae, Styracaceae, Ebenaceae, Theaceae, highly autapomorphic (see Conspectus), agree
Actinidiaceae) is from antepetalous stamens. The with Lecythidaceae and relatives in having mostly
reputedly primitive wood characters of Symplocos, trilacunar nodes and sometimes stipulate leaves,
such as solitary vessels with scalariform perfora- rare character states in the Ericales.
tion plates and mainly scalariform to opposite Scytopetalaceae, differing from Lecythidaceae,
wall pitting, diffuse parenchyma, fibre tracheids i.a. in albuminous seeds, are treated here as a
and heterogeneous rays of two size classes, are separate family; the combined morphological/
also found in Ternstroemiaceae, together with the molecular topology of Morton et al. (1996) there-
fore requires treating Napoleonaea (with Crater- Fig. 2. Ericoid seed coats. A Philadelphus coronarius
anthus) at the same rank. It is interesting - and (Hydrangeaceae). B Souroubea sympetala (Marcgraviaceae). C
Eurya emarginata (Ternstroemiaceae). D Symplocos rigidis-
also somewhat disturbing - that in the basal sima (Symplocaceae). E Roridula gorgonias (Roridulaceae). F
branches of their lecythidalean clade (see Figs. 4 Corema album (Ericaceae). (Huber 1991)
and 5 in Morton et al. 1997) petals have been lost
and replaced by a staminodial corolla substitute
(Napoleonaeaceae and Scytopetalaceae), whereas otestal "theoid" (Nandi et al. 1998) or "ericoid"
in Foetidia within Lecythidaceae, this loss has seed coat (Huber 1991) is well-developed in
occurred again without having been repaired for. core Ericales (Stevens 1971), but also known from
The largest clade within the Ericales is formed other (expanded) Ericales (Fig. 2), and even
by the core Ericales, comprising Roridulaceae, Sar- Hydrangeaceae and Loasaceae (Netolitzky 1926;
raceniaceae, and Actinidiaceae, and Clethraceae, Huber 1991).
Cyrillaceae, and Ericaceae. All have unitegmic This brief survey of the distribution of some
ovules, cellular endosperm, and terminal endo- anatomical, embryological and chemical charac-
sperm haustoria; all except for Actinidiaceae have ters in Cornales and Ericales calls for a com-
scalariform vessel perforation, and iridoids are parative evaluation. This has been attempted in a
present, Cyrillaceae and Clethraceae excepted. cladistic context by several earlier authors such as
Actinidiaceae, Roridulaceae and Sarraceniaceae Hufford (1992) and Albach et al. (2001a) but it
share the possession of a hypostase, which other- is hampered by the lack of resolution of the
wise is rare in the Ericales (Anderberg et al. 2002). basal branches in the analyses available for these
The absence of a fibrous endothecium in most groups. Thus, at present only some general
Ericaceae, Roridula and Heliamphora is obviously comments are possible.
due to the peculiarities of pollen release and may In Cornaceae, the reproductive specialisation
be autapomorphic in each group. Ericaceae, Cyril- (inferior ovaries with one-seeded drupes) con-
laceae, and Clethraceae are held together by traits trasts with the "primitive" traits of the xylem,
such as the reduced seed coat and the ovary cavity whereas in Hydrangeaceae, the situation is inverse.
extending into the "hollow" style, and (except for This reminds us that wood anatomists currently
Cyrillaceae) inversion of the anthers. This is a par- emphasise that xylem structure is not a mere
ticularly interesting character, as in its ontogenet- measure of "phylogenetic advancement" but
ically late form (occurring in bud or at anthesis) it reflects how a plant responds to the special
is found in Actinidiaceae (Fig. 4D, p. 17), Roridu- requirements of its environment. Even so, it is
laceae (Fig. lOSE, F, p. 340), Clethraceae (Fig. 22B, difficult to understand why Theaceae, Tern-
p. 70) and basal Ericaceae (see Fig. SSD, p. 166) stroemiaceae, Symplocaceae, Styracaceae, and
[but not in Cyrillaceae!], whereas its ontogeneti- Scytopetalaceae have scalariform vessel perfora-
cally early expression (Hermann and Palser 2000) tion, whereas the perforation is simple in families
characterises the higher Ericaceae. The ex- such as Sapotaceae and Ebenaceae.
10 K. Kubitzki
In Ericales the presence of bitegmic ovules Dahlgren, R. 1983. General aspects of angiosperm evolution
is usually correlated with nuclear endosperm, and macrosystematics. Nord. J. Bot. 3: 119-149.
Dahlgren, G. 1991. Steps toward a natural system of the
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"advanced" (Dahlgren 1991; Albach et al. 2001a) revised classification of the angiosperms with comments
cellular state. Unitegmic ovules and cellular on correlation between chemical and other characters.
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General References 13
combined analysis of plastid atpB gene sequences. Syst. Bioi. K.C., Farris, J.S. 2000. Angiosperm phylogeny inferred from
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14 S. Dressler and C. Bayer
Actinidiaceae
S. DRESSLER and C. BAYER
Actinidiaceae Gilg & Werderm. in Engler & Prantl, Nat. Vessels of the primary xylem are mostly solitary,
Pflanzenfam., ed. 2, 21: 36 (1925), nom. cons. often large, and often arranged in radial rows.
Saurauiaceae J. Agardh (1858).
They can have annular, helic~l, reticulate, or scal~
riform thickenings. PerforatiOn plates are scalan-
Trees shrubs or climbers, with simple or variously
form, more rarely simple. The peri cycle includes a
bran~hed trichomes. Leaves alternate, simple,
continuous ring of sclerenchyma. Young stems
usually serr(ul)ate or dentate, pinnatine~ved,yeti
have a solid pith, which may later disintegrate
olate; stipules absent or minute. Flowers maxillary
and/or become lamellate in some Actinidia
cymes or thyrso-paniculate in~orescenc~s, some-
species. The lamellae and a cylinder of outer pith
times solitary, pedicellate, actmomorphic, herm-
cells become sclerenchymatous with age.
aphroditic or unisexual; sepals (3- )~( -8),
The wood of Actinidia has dimorphic vessel ele-
imbricate-quincuncial in bud, usually persistent;
ments: few very large, moderately long vessel ele-
petals (3- )5( -9), longer than sepals, distinct or ±
ments with simple perforations, and numerous
fused at base, imbricate in bud; stamens
smaller, usually scattered or solitary, with oblique
{10-)20-240, filaments distinct, sometimes adnate
scalariform perforation plates. Saurauia has ex-
to petals and falling with these, a~thers ditheca!,
clusively solitary vessels with many-barred sc~
dorsifixed, versatile, extrorsely dehiscent by longi-
lariform perforation plates. Parenchyma . IS
tudinal slits or subapical pores; ovary superior,
diffuse apotracheal. Non-septate fibre tracheids
syncarpous, {3-)5- to many-locular, sometimes
have numerous bordered pits. Rays are uniseriate
incompletely septate, pubesce?-t or glabro_us,
and multiseriate, heterogeneous (Lechner 1914;
sometimes with apical depressiOn; placentatiOn
Metcalfe and Chalk 1950).
axile, placentae sometimes split; ovules numerous,
Wang et al. {1994) studied the root anatomy ~f
anatropous, unitegmic; stylodia distinct, as. many
five Actinidia species; cortex and endodermis
as locules, sometimes persistent, or style simple,
persist during secondary thickening.
stigma sometimes capitate; fruit usually a ~erry,
sometimes dehiscent; seeds numerous, exanllate,
REPRODUCTIVE STRUCTURES. Cauliflory is
enclosed by pulp, albuminous, embryo large,
reported for some species. Saurauia callithrix is
usually straight.
special by having leafless inflorescences at the base
A family of c. 360 spp. in three genera nat~v~ ~o of the trunk which spread in the soil and leaf litter,
tropical Asia and America, a few spp. of Actmzdza
elevating the flowers just above the forest floor
in temperate E Asia. (Gilg and Werdermann 1925). The flowers are
VEGETATIVE ANATOMY. Young organs are
often arranged in cymes, which are s';lbtended ?Y
foliage leaves or bracts. Some Saurauza ha~e axil-
covered with simple or multicellular, sometimes
lary thyrso-paniculate inflorescences that mclude
glandular hairs, which can be of diagnostic value
monochasia. Bracteose prophylls are frequently
(see Hunter 1966). Raphides contained in elon-
found.
gated idioblasts occur in most tissues. Crystal sand
The flowers are mostly pentamerous but a few
is reported for Clematoclethra (Lechner _1914).
Saurauia and Actinidia species have tetramerous
The leaves are dorsiventral and sometimes have
flowers, and other exceptions occur also. In pen-
an arm palisade mesophyll. Some species form a
tamerous flowers, quincuncial aestivation of
multilayered hypodermis. Stomata of the r~nun~u sepals is evident even in open flowers, since the
laceous type are restricted to the abaxial s~de
abaxial face is often pubescent in the two outer-
(Lechner 1914; Metcalfe and Chalk 1950). VenatiOn
most sepals, half pubescent in one other, and
is pinnate and of the camptodromous type. The
glabrous in the two inn~r~ost sepals. For
ultimate marginal veins are looped, but some
anatomical features see Dickison (1972) and
branches extend into the teeth (Yu and Chen 1991).
Schmid {1978a).
Actinidiaceae 15
Petals and stamens may be fused to various (Erdtman 1952; Dickison et al. 1982; Zhang 1987;
degrees and often fall together. The microsporan- Li et al. 1989; Kang et al. 1993). Similar pollen
gia of each theca merge. Pollen is released through occurs in Theaceae, Ochnaceae and Clethraceae
longitudinal slits or pores that become extrorse by (Erdtman 1952; Zhang 1987), but there are some
inversion of the anthers. differences to Dilleniaceae (Dickison et al. 1982).
Brown (1935) reported nectar-secreting tissue at Functionally female flowers of Actinidia deli-
the base of the petals of Saurauia subspinosa. ciosa shed nonviable pollen, which is usually enu-
According to his study, the androecium is basically cleate and shrivelled but otherwise similar to
diplostemonous. The outer whorl is formed by viable pollen (Schmid 1978a; White 1990).
single stamens in front of the sepals, whereas the
majority of stamens develop in centrifugal succes- KARYOLOGY. Chromosome counts are known for
sion by splitting of antepetalous primordia. For many Actinidia species (see Yan et al. 1997). The
Actinidia deliciosa, Brundell (1975) described two basic chromosome number is x = 29; diploids,
or three whorls of stamina! initials, depending tetraploids, hexaploids and octoploids occur.
on the cultivar. In a more detailed study of A. Intraspecific variation appears to be common
melanandra and A. deliciosa, van Heel (1987) (mainly diploid and tetraploid cytotypes, 4X and
found a single whorl of staminal primordia in the 6x in A. valvata, and 4x, 6x and 8x in A. arguta).
former, and centrifugal multiplication of such For Saurauia, counts of n = 30 (South American
primary primordia in the latter. The numerous species: Soejarto 1969, 1970) and n = 20 (Asian
carpel primordia arise in a single whorl. species: Mehra 1976) were reported.
EMBRYOLOGY. The anther wall consists of epider- REPRODUCTIVE BIOLOGY. Actinidia is usually
mis, fibrous endothecium, 2-3 ephemeral middle dioecious, Clematoclethra and Saurauia have
layers, and a secretory tapetum of multinucleate mostly bisexual flowers. American Saurauia,
cells the nuclei of which may fuse and become however, is described as functionally dioecious
polyploid. Raphides are present in the anther wall with dimorphic flowers: long-styled, functionally
and connective. Meiotic division of microspore female flowers with malformed, sterile pollen, and
mother cells is simultaneous. Pollen is shed at the short-styled ones with fertile pollen (Soejarto
two-celled stage. 1969). Brown (1935) reported protandry.
The ovules are anatropous, unitegmic and tenu- In Actinidia deliciosa, staminate flowers open
inucellar. A hypostase is present. The nucellus is about seven days before pistillate ones. Plants of
thin and ephemeral. A hypodermal archesporia! the pistillate cultivar 'Hayward' in New Zealand
cell forms the chalazal megaspore mother cell bloom 10-18 days, staminate clones usually flower
which develops into a linear tetrad. Embryo sac 3-5 days longer (Hopping 1990). Because of this
development is of the Polygonum type (Vija- limited overlap in flowering and the linear rela-
yaraghavan 1965; An et al. 1983). Inverted embryo tionship between seed number and fruit weight,
sac polarity is reported of Saurauia nepalensis fruit set is improved by artificial pollination.
(Rao 1953). Endosperm formation is cellular; Pollen tubes from the distinct stylar branches are
embryogeny corresponds to the Solanad type evenly distributed to the numerous carpels and
(Crete 1944b; Vijayaraghavan 1965). Polyembry- ovules by a compitum ("pollen tube distributor
ony due to the proliferation of suspensor cells was cup": Howpage et al. 1998). For pollen-pistil inter-
observed in Actinidia deliciosa (Crete 1944a). action, pollen tube growth and fertilisation of
Actinidia, see Hopping and Jerram (1979), Harvey
PoLLEN MoRPHOLOGY. Pollen of Actinidiaceae is et al. (1987) and Gonzalez et al. (1996).
remarkably uniform and rather unspecialised. Actinidia deliciosa is said to be bee- or wind-
Pollen is usually shed in monads; Saurauia elegans pollinated. Despite the abundance of literature
has tetrads. The grains are usually 3( 4)-colporate, on the floral biology of cultivated kiwifruit (e.g.
oblate-spheroidal to prolate, the longest axis Schmid 1978a; Harvey et al. 1987; Harvey and
13-26( -33) J..Lm. The colpi are long, crassimar- Fraser 1988; Hopping 1990; Howpage et al. 1998),
ginate and usually exhibit an equatorial bridge of there is only little information on pollination in
ektexine over the endoaperture. Exine is ( 1-) 1.5-2 the wild. Gilg and Werdermann (1925) suspect
J..Lm thick. Sexine is as thick as nexine. The com- insect pollination for Actinidia and Clemato-
plete tectum is psilate, (micro-)granulate or rugu- clethra. For Saurauia, observations on insect visits
late; columellae are reduced. Exine stratification (e.g. Hymenoptera) are reported (Soejarto 1969).
and sexine pattern are obscure in light microscopy The flowers are showy, often fragrant and at least
sometimes nectariferous (Brown 1935 for S. Actinidiaceae, Roridulaceae and Sarraceniaceae is

subspinosa). the presence of a hypostase. Other features con-
sidered by Anderberg et al. (2002), such as the
FRUIT AND SEED. Fruits are usually berries with presence of stylar branches and a fibrous endothe-
massive placentae, and the seeds embedded in a cium, are not found in all members of this clade.
mucilaginous pulp which is mostly greenish,
sticky, sweet and edible. Since this pulp originates DISTRIBUTION AND HABITATS. Actinidia is cen-
from the placenta, it does not correspond to an aril tred in hilly S and E China (between 25 and 30 °N),
(Schmid 1978a). Berries of Actinidia may include but some species occur in the cold-temperate and
more than 40 locules and 1500 seeds. In some arctic forests of Siberia, Korea and Japan, and the
Saurauia species, the pericarp is dehiscent and tropics. They inhabit the lower forest storey.
more or less dry (Soejarto 1969; Dickison 1972). Clematoclethra occupies similar habitats up to
The fruits of Clematoclethra have been described 3100m but is confined to China. In contrast,
as capsular (Gilg and Werdermann 1925), indehis- Saurauia species are trees and shrubs mainly of
cent (Ying et al. 1993) or drupaceous (Lechner humid montane forests of tropical Asia and
1914). Generally, endozoochory seems most likely. America, and occur in altitudes up to 3600 m.
In Saurauia, dispersal is by rain and animals such Some species are rheophytic (van Steenis 1981).
as birds (Soejarto 1969).
The seed coat is thin and finely reticulate. As in PALAEOBOTANY. Seeds resembling those of
other Ericales, the seeds of Actinidiaceae are extant Saurauia were recorded from the Maas-
exotestal, with thickened inner walls of the outer trichtian onwards in Europe (Knobloch and Mai
epidermis (Crete 1944b; Corner 1976; Huber 1991; 1986), and Actinidia-like seeds are known from
Takhtajan 1991). the Upper Eocene in Europe, sometimes in abun-
dance (Mai and Gregor 1982; Friis 1985; Mai 2001).
PHYTOCHEMISTRY. Iridoid compounds were Leaves similar to those of Saurauia were found in
found in Actinidia and Roridula (Jensen et al. the Middle Eocene of North America (Taylor
1975). Webby et al. (1994) studied leaf fl.avonoids 1990), and Actinidiophyllum was described from
of several Actinidia, which are based on common the Tertiary of Japan (Nathorst 1888). A Pliocene
fl.avonols including myricetin. Procyanidin and wood from the German Westerwald was described
prodelphinidin point to the presence of condensed as Actinidioxylon (Miiller-Stoll and Miidel-
tannins (Hegnauer 1964). Actinidin, a proteinase Angeliewa 1969). Flowers from the early Campan-
similar to papain, was detected in kiwifruit ian of Georgia, North America, were described as
(McDowall1970), which makes it a potential cause Parasaurauia, mainly differing from modern
of contact dermatitis. The same name was applied Saurauia in having only 10 stamens (Keller et al.
to a terpenoid pseudoalkaloid found in Actinidia 1996).
polygama (Hegnauer 1964). The mucilage of
Actinidia contains acidic polysaccharides ECONOMIC IMPORTANCE. Kiwifruit, produced by
(Redgwell1983). Actinidia deliciosa cultivars, are an economically
important crop, especially in New Zealand but also
AFFINITIES. Actinidiaceae have been variously in Italy, Spain, China and other countries. Since
placed; especially Theaceae, Dilleniaceae, Cornales this species was formerly treated mostly as a
and Ericales were considered as closest relatives variety of A. chinensis, most literature on culti-
(see, for example, Schmid 1978a). Ericalean affini- vated A. chinensis refers to what is now A. deliciosa
ties are now mostly accepted, which is supported (Liang and Ferguson 1986; Ferguson 1990). All
by agreement in embryological characters (Crete commercial plantations in New Zealand can be
1944b), floral features (Dickision 1972) and results traced back to a single introduction of seed from
of molecular analyses (Kron and Chase 1993; China in 1904 (Ferguson and Bollard 1990). There
Chase et al. 1993 and subsequent studies). Accor- are indications that the hexaploid A. deliciosa
ding to recent analyses, Actinidiaceae are sister to originated from diploid A. chinensis (Crowhurst
Roridula, both being sister to Sarraceniaceae et al. 1990; Atkinson et al. 1997). Breeding of A.
(Anderberg et al. 2002). In all analyses, nonethe- chinensis, up to 1997 collected only in the wild and
less, statistical support for this topology is not industrially processed in China, led to the recent
strong, and some studies have found different introduction of the Kiwi Gold (cv. 'Hort 16/\) from
topologies (e.g. Savolainen, Fay et al. 2000). A New Zealand. Relatively hardy species such as A.
potential synapomorphy of a clade comprising arguta and A. kolomikta are cultivated as orna-
Actinidiaceae 17
mentals in Europe and North America. In the

former Soviet Union, attempts were made to use
their aromatic fruits, which contain even more
vitamin C than do kiwifruit. Berries of some
Saurauia species are sold at local markets in South
America. Wood is occasionally used for construc-
tion, fire wood and charcoal, but is of no com-
mercial importance. Some uses in folk medicine
are reported (Soejarto 1980).
KEY TO THE GENERA
1. Trees or shrubs; stamens basally connate with petals; Asia

and America 3. Saurauia
- Woody climbers; stamens free; Asia 2
2. Flowers usually functionally unisexual; stamens numer-
ous; stylar branches distinct 1. Actinidia
- Flowers usually hermaphroditic; stamens 10(-30); style
simple 2. Clematoclethra
GENERA OF ACTINIDIACEAE
1. Actinidia Lindl. Fig. 3

Actinidia Lind!., Intr. Nat. Syst., ed. 2: 439 (1836); Li, J. Arnold
Arbor. 33: 1-61 (1952), rev.; Liang, Fl. Reip. Pop. Sin. 49(2):
196-268 (1984), reg. rev.; Wei, Higher plants of China 4:
657-672 (2000), reg. rev.
Woody climbers, dioecious or polygamous, ± Fig. 3. Actinidiaceae. A, F, G Actinidia strigosa. B-E A.

pubescent; sepals distinct or somewhat fused at polygama. A Flower. B Pistil. C Fruit. D, ESame, vertical and
base; petals white or yellow to reddish; stamens transverse section. F Seed. G Same, longitudinal section.
(Schneider 1912)
numerous, anthers longitudinally dehiscent; ovary
pubescent or glabrous, many-locular; stylar
branches distinct, persistent; fruit a globose to
oblong berry, sometimes pubescent; seeds numer-
ous, embedded in pulp. n = 29 or multiples. About
60 spp.; E Asia, mostly W to E China, north to
Sakhalin and Kuril Is., south to Taiwan, Himalayas,
NE India, Indochina, Malaysia.
2. Clematoclethra (Franch.) Maxim. Fig. 4

Clematoclethra (Franch.) Maxim., Trudy Imp. S.-Peterburgsk.
Bot. Sada 11: 36 (1890); Tang & Xiang, Acta Phytotax. Sin.
27: 81-95 ( 1989), rev.; Wei, Higher plants of China 4: 672- 674
(2000), rev.
Woody climbers, deciduous, ± pubescent; flowers

in up to 12-flowered cymes or solitary, hermaph-
roditic or unisexual; sepals united at base; petals
white or reddish; stamens c. 10-30, filaments
dilated at base; anthers longitudinally dehiscent;
ovary globose, 5-angled, (4)5-locular; style simple, Fig. 4. Actinidiaceae. Clematoclethra /asioclada. A Flowering
persistent, stigma small, swollen; ovules numer- twig. B Flower. C Androecium and gynoecium of young
ous; fruit berry-like, blackish. One species, C. scan- flower, longitudinal section. D Androecium a ndgynoecium at
anthesis, note inversion of anthers. E Young anthers. FMature
dens (Franch.) Maxim. (or 5), in montane forests anther. G Ovary in cross section. (Gilg and Werdermann
above 1000 m in W and C China. 1925)
3. Saurauia Willd. Friis, E.M. 1985. Angiosperm fruits and seeds from the Middle
Miocene of Jutland, Denmark. Bioi. Skr. 24: 1-165.
Saurauia Willd., Ges. Naturf. Freunde Berlin Neue Schriften 3: Gilg, E., Werdermann, E. 1925. Actinidiaceae. In: Engler &
407 (1801), nom. cons.; Hunter, Ann. Missouri Bot. Gard. 53: Prantl, Die natiirlichen Pflanzenfami!ien, ed. 2, vol. 21.
47-89 (1966), reg. rev.; Soejarto, Fieldiana Bot. N.S. 2: 1-141 Leipzig: W. Engelmann, pp. 36-47.
(1980), reg. rev.; Wei, Higher plants of China 4: 674-677 Gonzalez, M.V., Coque, M., Herrero, M. 1996. Pollen-pistil
(2000), reg. rev. interaction in kiwifruit (Actinidia deliciosa; Actinidiaceae).
Am. J. Bot. 83: 148-154.
Trees or shrubs, usually pubescent; flowers herm- Guedes, M., Schmid, R. 1978. The peltate (ascidiate) carpel
theory and carpel peltation in Actinidia chinensis (Actini-
aphroditic or sometimes functionally unisexual; diaceae). Flora 167: 525-543.
sepals fused at very base, petals white or pink; Harvey, C. F., Fraser, L.G. 1988. Floral biology of two species of
stamens IS-numerous, filament bases fused with Actinidia (Actinidiaceae). II. Early embryology. Bot. Gaz.
petals, pubescent, anthers dehiscing by pores; 149: 37-44.
ovary (3-)5( -8)-carpellate, usually glabrous, stylar Harvey, C.F., Fraser, L.G., Pavis, S.E., Considine, J.A.1987. Floral
biology of two species of Actinidia (Actinidiaceae). I. The
branches entirely or distally distinct, stigmas stigma, pollination, and fertilization. Bot. Gaz. 148: 426-432.
simple to capitate or discoid; fruit a berry, rarely a Heel, W.A. van 1987. Androecium development in Actinidia
leathery capsule; seeds numerous, minute, embed- chinensis and A. melanandra (Actinidiaceae). Bot. Jahrb.
ded in pulp. n = 30 (20 in Asian spp.). About 300 Syst.109: 17-23.
Hegnauer, R. 1964. See general references.
spp., Asia (Himalayas to E and SE Asia) to Fiji Hopping, M.E. 1976. Structure and development of fruit and
and tropical America (C Mexico to Chile but seeds in Chinese gooseberry (Actinidia chinensis Planch.).
absent from Antilles, Guianas, Brazil); 1 sp. in New Zeal. J. Bot. 14: 63-68.
Queensland. Hopping, M.E. 1990. Floral biology, pollination, and fruit set.
In: Warrington, I.J., Weston, G.C. (eds.) Kiwifruit: science
and management. Auckland: Ray Richards, pp. 71-96.
Hopping, M.E., Jerram, E.M. 1979. Pollination of kiwifruit
Selected Bibliography (Actinidia chinensis Planch.): stigma-style structure and
pollen tube growth. New Zeal. J. Bot. 17: 233-240.
An, H.-X., Cai, D.-R., Wang, J.-R., Qian, N.-F. 1983. Investiga- Howpage, D., Vithanage, V., Spooner-Hart, R.1998. Pollen tube
tions on early embryogenesis of Actinidia chinensis Planch. distribution in the kiwifruit (Actinidia deliciosa A. Chev.
var. chinensis. Acta Bot. Sin. 25: 99-104,2 pl. C.F. Liang) pistil in relation to its reproductive process. Ann.
Anderberg, A.A. et a!. 2002. See general references. Bot. II, 81: 697-703.
Atkinson, R.G., Cipriani, G., Whittaker, D.J., Gardner, R.C.1997. Huber, H. 1991. Angiospermen. Leitfaden durch die Ordnun-
The allopolyploid origin of kiwifruit, Actinidia deliciosa gen der Familien der Bedecktsamer. Stuttgart, New York:
(Actinidiaceae). Pl. Syst. Evol. 205: 111-124. G. Fischer.
Brown, E.G.S. 1935. The floral mechanism of Saurauia sub- Hunter, G.E. 1966. Revision of Mexican and Central American
spinosa Anth. Trans. Proc. Bot. Soc. Edinburgh 31: 485-497. Saurauia (Dilleniaceae). Ann. Missouri Bot. Gard. 53: 47-
Brundell, D.J. 1975. Flower development of the Chinese goose- 89.
berry (Actinidia chinensis Planch.) II. Development of the Jensen, S.R., Nielsen, B.J., Dahlgren, R. 1975. Iridoid com-
flower bud. New Zeal. J. Bot. 13: 485-496. pounds, their occurrence and systematic importance in the
Chase, M.W. eta!. (1993). See general references. angiosperms. Bot. Notiser 128: 148-180.
Corner, E.J.H. 1976. See general references. Johri, B.M. et a!. 1992. See general references.
Crete, P. 1944a. Polyembryonie chez !'Actinidia chinensis Kang, N., Wang, S., Huang, R., Wu, X. 1993. Studies on the
Planch. Bull. Soc. Bot. France 91: 89-92. pollen morphology of nine species of genus Actinidia. J.
Crete, P. 1944b. Recherches anatomiques sur Ia seminogenese Wuhan Bot. Res. 11: 111-116,5 pl.
de I' Actinidia chinensis Planch. Affinites des Actinidiacees. Keller, J.A., Herendeen, P.S., Crane, P.R. 1996. Fossil flowers
Bull. Soc. Bot. France 91: 153-160. and fruits of the Actinidiaceae from the Campanian (Late
Crowhurst, R.N., Lints, R., Atkinson, R.G., Gardner, R.C. 1990. Cretaceous) of Georgia. Am. J. Bot. 83: 528-541.
Restriction fragment length polymorphisms in the genus Knobloch, E., Mai, D.H. 1986. Monographie der Friichte und
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Dickison, W.C. 1972. Observations on the floral morphology of Geol. 47: 1-219.
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Elisha Mitchell Sci. Soc. 88: 43-54. Empetraceae, Epacridaceae and related taxa based upon
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20 E. Fischer
Balsaminaceae
E. FISCHER
Balsaminaceae A. Rich. in Bory, Diet. Class. Hist. Nat. 2: 173 forming large globose tubers (Cheek and Fischer
(1822), nom. cons. 1999). Suffruticose species are present in Africa
and India; they produce robust and woody stems
Annual or perennial herbs, sometimes with tubers and can reach up to 4 m in height. Leaf arrange-
or rhizomes, occasionally subshrubs; stems erect ment in most species is spiral; verticillate leaves
or procumbent, succulent, rarely woody below. with up to 10 leaves per whorl are found in several
Leaves spirally arranged, rarely decussate or species, whereas decussate leaves occur more
verticillate, simple, petiolate or sessile, pinnately rarely.
veined, margins crenate, dentate or serrate, teeth
or crenations apiculate, the lowermost often VEGETATIVE AND FLORAL ANATOMY. Calcium-
gland-tipped, petiole often with short capitate oxalate raphides occur in bundles within special
glands or fimbriae, rarely with extra-floral nec- cells in the cortex. They are also present in the
taries. Flowering shoots truncate, proliferating; anthers walls (raphid pollen). Idioblasts with pos-
inflorescences axillary racemes or pseudoumbels, sible mucilage content are found in stems and
often epedunculate and fascicled in leaf axils; leaves. The stems of most Impatiens species have
flowers zygomorphic, usually resupinate through a pericycle, which is devoid of sclerenchyma. The
180°, not resupinate in some Chinese Impatiens; rigidity of the stem is maintained by the turges-
sepals 3 or 5, free, the lower one (by resupination) cence of the parenchyma (Metcalfe and Chalk
larger, navicular to saccate, usually tapering or 1950). Only few species like I. sodenii, I. niamnia-
abruptly constricted into a nectariferous spur; mensis or the epiphytic I. paucidentata have
petals 5, dorsal petal free, flat or cucullate, often basally woody stems and show a special type of
crested dorsally, lower 4 petals free or united into primary thickening.
lateral pairs; stamens 5, connate into a ring sur-
rounding ovary and stigma, ripening and usually INFLORESCENCE STRUCTURE. The inflorescence
falling off in one piece before the maturity of consists of a frondose main stem, which prolifer-
the stigma; ovary superior, syncarpous, 5-locular ates vegetatively, and lateral racemes. Due to the
with axile placentation; ovules 5-numerous, anat- suppression of apical growth, as in I. palpebrata,
ropous, bitegmic or unitegmic, tenuinucellate; these racemes sometimes appear to be terminal.
style 1, very short or± absent; stigmas 1-5. Fruit The racemes may be elongate, as in Impatiens
a berry or a loculicidal fleshy explosive capsule; teitensis, or subumbellate as in I. stuhlmannii.
seeds exalbuminous, seed coat smooth, warted or Reduction of flowers may result in single-flowered
with simple hairs. lateral racemes with the appearance of single axil-
Two genera (one of which monotypic) and c. lary flowers (e.g. I. pseudoviola). By reduction of
1000 species, Europe, Africa, Asia, North and the peduncle, axillary flower clusters result (I.
Central America. niamniamensis, I. keilii, I. clavicalcar). In I. pauci-
dentata, the corresponding part is uniflorous. In I.
VEGETATIVE MORPHOLOGY. Most species of acaulis, the main stem is congested and reduced,
Impatiens are perennial herbs with thin rhizomes, forming a basal rosette with a pseudoterminal
adapted to constantly humid conditions in moist raceme.
forests. Relatively few species are annuals with
thin erect stems. Plants growing under temporar- FLORAL MORPHOLOGY. The flower is zygomor-
ily dry conditions have fusiform, fleshy subter- phic and usually resupinate through twisting of
ranean tubers. Large globose tubers are found in the pedicel. Few species (e.g. Hydro cera triflora,
the Madagascan I. tuberosa, which inhabits cal- Impatiens tinctoria, I. teitensis, I. quadrisepala)
careous rocks. Impatiens etindensis is an epiphyte have five sepals. Usually, each flower has only two
Balsaminaceae 21
reduced lateral sepals and one petal-like lower Govindaran and Subramanian 1986; Rao et al.
sepal, which is modified into a nectary-tipped 1986; Oginuma and Tobe 1991; Akiyama et al.
spur; both upper sepals are lacking. There is a 1992; Sugawara et al. 1995, 1997). Four basic
wide range of variation in form and size of the numbers, x = 7 8,
, 9, and 10, are more common than
spurs, from shallowly navicular to bucciniform or others. African species have mostly the basic
deeply saccate, short or long filiform, straight, numbers x = 7and 8. In southern India, x = 7, 8,
twisted or curved, obviously associated with dif- and 10 are predominant. In the western Himalaya
ferent pollinators. In a group of Madagascan taxa, x = 7 and 8 are common, whereas in the eastern
the spur is lacking. There are five petals, one of Himalaya and adjacent SE Asia basic numbers of
which, the upper dorsal one, is usually hood-like, x = 9and 10 are prevalent. In Japan, northern Asia,
whereas the others are united into two lateral Europe and North America, x = 10 predominates.
pairs, each with two unequal lobes. The assump- Jones and Smith (1966) and Akiyama et al. (1992)
tion of Rama Devi (1991) that the dorsal petal is considered x = 7 as the ancestral condition
the product of fusion of two sepals and one petal whereas Rao et al. (1986) proposed x = 8 as the
is questionable. The united lateral petals are also original base number.
very variable in shape and size, again in adapta-
tion to different pollinators, as they provide a suit- POLLINATION AND REPRODUCTIVE SYSTEMS. As
able landing platform and entrance guide to the far as floral diversity is concerned, Impatiens is the
spur and the nectar. The five stamens are united dicot counterpart of orchids. As in Orchidaceae,
by the upper part of the filaments and completely the evolutionary trend towards specialized polli-
cap the gynoecium. nation has implied abundant speciation and often
accounts for high rates of endemism in different
POLLEN MORPHOLOGY. The pollen is 4-colpate regions. Due to the strong expression of protandry,
and rectangular in the North American and most Impatiens are functionally monoecious.
Eurasian taxa (Fig. 5), 4-colpate rectangular, 4- During the male phase, whenthe flower just opens,
colpate square or 3-colpate triangular in the taxa the stamens are fused by their anther walls to
from tropical Africa and Asia (Huynh 1968; Lu form a "brush" which completely covers the pistil.
1991). The upper filaments are longer than the lower
ones, and the anthers project downwards and can
KARYOLOGY. The chromosome number of "brush" the pollinator's back or head and transmit
Hydrocera triflora was determined as 2n = 16 the pollen upon the next pollinator visiting the
(Govindaran and Subramanian 1986; Rao et al. flower. During this stage, the stigmas are non-
1986). Chromosomal variation in Impatiens is receptive and not exposed to come into contact
extensive; the over 170 species studied so far with the pollinator. At the end of the male phase,
range from n = 3 ton= 33 (Jones and Smith 1966; the stamens are shed as a single unit and the
Larsen 1981; Zinoveva-Stahevitch and Grant 1984; stigmas become receptive. Extra-floral nectaries,
usually in the form of stipitate glands, are often
present and may reward "safety guards" (usually
ants) protecting the flowers from damage by
non-pollinating animals.
The pollination biology and breeding systems of
the majority of species have not yet been studied,
and most available studies have been conducted
with the temperate species I. capensis and I.
pallida. Published reports and personal field
observations suggest the following animals as pos-
sible pollinators: honeybees (e.g. I. austrotanzan-
eY-
e
ica, I. gomphophylla), bumblebees, solitary bees,
butterflies (e.g. I. walleriana, I. hoehneliana ),
moths, hawkmoths (e.g. I. teitensis), flys (e.g. I.
elatostemoides), and sunbirds (e.g. I. niamniamen-
sis, I. keilii, I. paucidentata) (Arisumi 1974, 1980;
Fig. 5. Balsaminaceae. Impatiens jurpia, pollen grain. A Polar Beck et al. 1974; Rust 1977, 1979; Kato 1988;
view (x l600. B A mesh of the reticulum. C Equatorial view. Randall and Hilu 1990; Wilson 1995; Hurlbert et al.
(Huynh 1968) 1996).
22 E. Fischer
Whereas most Impatiens have mechanisms to

ensure outcrossing, various forms of self-
compatibility and cleistogamy have evolved. Grey-
Wilson (1980a) suspected self-pollination to be
limited to temperate species which, under
unfavourable conditions or the absence of
appropriate pollinators, occasionally switch to
cleistogamy. Meanwhile, self-compatibility and
facultative cleistogamy have been established for
species such as I. pallida, I. capensis, and I.
hypophylla. In these species, cleistogamy is envi-
ronmentally controlled by factors such as light
intensity, drought, flooding, and herbivore
damage. Even on the same individual, the upper
flowers have been observed to be chasmogamous,
whereas the flowers close to ground often are
cleistogamous (Schemske 1978, 1984; Antlfinger
1986; Gross et al. 1998; Paoletti and Holsinger
1999). This observation is also confirmed for the
Madagascan I. baroni (pers. obs.).An extreme case
is the Madagascan I. inaperta (formerly placed in
the monotypic genus Impatientella), which pro-
duces exclusively cleistogamous flowers.
EMBRYOLOGY AND SEEDS. Mature pollen is 2-

nucleate, with the generative nucleus remaining in
an arrested metaphase (Huynh 1970).
The ovules are anatropous and bitegmic,
unitegmic or intermediate and tenuinucellate;
they have a characteristic endothelium. Mature
ovules show an epistasis in the micropylar region
(Huynh 1970). The outer integument contains
raphid bundles. The growth processes involved in
the changeover from the bitegmic to the unitegmic
condition within Impatiens has been analysed by
Boesewinkel and Bouman (1991).
In most species, embryo sac development con-
forms to the Allium type or Polygonum type. The
straight embryo fills the seed almost completely.
Endosperm formation is ab initio cellular with ter- Fig. 6. Balsaminaceae. Seeds with smooth {A), warted (B) and
minal haustoria. The micropylar haustorium is long-haired (C) exotesta. A Impatiens stuhlmannii {x45).
B I. noli-tangere (xSO). C I. niamniamensis (x40). Photo E.
aggressive, and branches of it have been described Fischer
to reach and enter the raphe, funicle, or even the
placenta (Narayana 1965). In the mature seed, the
endosperm is restricted to one or only a few layers;
it is rich in lipids and poor in starch. The embryo and seeds with dub-shaped appendages (Fig. 6).
is large and in I. glandulifera is provided by well- The long-haired seeds bear long, helical, hair-like
developed lateral roots which may promote the structures that have spiral thickenings (Fig. 6C).As
rooting of the seedling (Boesewinkel and Bouman this seed type occurs in groups of Impatiens with
1991). The seed coat of Impatiens has a lignified facultative and obligate epiphytes (e.g. I. niamnia-
exotesta and more or less crushed middle layers. mensis aggregate), the hairs may play a role as
The testa cells have straight or undulating walls. adhesive structures. The seed coat of Hydrocera
Single cells are modified to papillae or hairs. consists of 5- 6 thickened layers but lacks a thick-
Grey-Wilson (1980a) distinguished five main seed ened exotesta (Venkateswarlu and Lakhshmi-
types: smooth, warted, short-haired, long-haired narayana 1957).
Balsaminaceae 23
FRUIT AND DISPERSAL. The mature fruit of Impa- walleriana, common as a weed throughout
tiens is fleshy and fusiform to cylindric in shape. tropical South America, may represent another
Dehiscence is caused by a tension in the convex example of such recent naturalization and rapid
fleshy walls of the ovary. This tension is released radiation.
by rupture of the walls along the septa, thus throw- The preferred habitat of Impatiens is the herb
ing the seeds for several meters by the elastic layer of montane rain forests. Epiphytes are found
force of the dehiscing fruit wall. At maturity, the in Africa (e.g. I. keilii, I. epiphytica, I. irangiensis,
slightest touch will cause the explosion of the I. paucidentata), Madagascar (e.g. I. purroi) and
fruit, hence the folk names "touch-me-not" or India (e.g. I. parasitica). Seasonally dry habitats
"busy lizzy". The fruits of Hydrocera are fleshy are usually avoided with the exception of some
pseudoberries with a hard endocarp that falls into specialists with particular morphological adapta-
five distinct units (partial endocarps, each with tions (e.g. tubers in I. tuberosa and I. cinnabarina,
one seed and two air sacs). These units are dis- Fig. 7).
persed by water and, due to their air sacs, are able
to float.
PHYTOCHEMISTRY. Impatiens spp. regularly con-

tain raphides of calcium oxalate and large amounts
of leucanthocyanins including prodelphinidin.
These compounds and raphides are rarely encoun-
tered in tender herbs (Hegnauer 1964, 1989).Apart
from the occurrence of naphtoquinones such as
lawsone, seed oils with acetate and parinarate
glycerides are highly characteristic.
DISTRIBUTION AND HABITATS. Hydrocera triflora

is a semi-aquatic herb native to tropical and sub-
tropical Indo-China and the Indian subcontinent.
Impatiens is essentially a montane genus centred
in tropical Africa (c. 120 spp.), Madagascar (c. 160
spp.), southern India and Sri Lanka (c. 150 spp.),
the eastern Himalaya (c. 120 spp.), Indo-China (c.
150 spp.) and adjacent SE Asia. Only a few species
of Impatiens are known from the temperate
montane regions of Europe, North Asia, and
Central and North America, and no species are
native to South America. A remarkable feature
is the high degree of endemism in the centres of
distribution. For example, over 90% of the Indian
species are endemic and most of them are limited
to the Western Ghats (Rao et al. 1986). All native
species of Madagascar are endemic, concentrated
in the eastern humid high-altitude regions. A
similar pattern is observed in SW China and
tropical Africa. The rapid radiation and possible
subsequent parallel evolution and hybridization
(Grey-Wilson 1980b; Merlin and Grant 1986) may
obscure the phylogenetic relationships among the
species. Recent rapid naturalizations of some
species offer further examples of the adaptability Fig. 7. Balsaminaceae. Hydrocera triflora. A H abit. B Flower
of the genus. For example, I. glandulifera, a bud. C F lower, front view. D Flower, lateralview. E Lower lateral
sepal. F Upper lateral sepal. G L ower sepal and spur, lateral
native of the Himalaya, is now widely naturalized
view. H Lower sepal, i nside. I D orsal p etal. J Upper l ateral petal.
in Europe and Japan. It probably escaped from K Lower l ate ral petal. L Androecium. M F r uit. N Fruit, cross
gardens and, within less than 100 years, success- section. 0 Single partial endocarp. Drawn by C .Grey-Wilson.
fully spread in Europe (Kurtto 1996). Impatiens (Grey-Wilson 1980c)
24 E. Fischer
KEY TO THE GENERA

1. Petals all free; lateral sepals 4, large and coloured like the
petals; fruit a pentagonal pseudoberry 1. Hydrocera
- Petals not all free, the lower 4 united laterally into pairs;
lateral sepals 2-4, small and rather inconspicuous, gener-
ally white or greenish; fruit a dehiscent capsule
2. Impatiens
1. Hydrocera Blume Fig. 7

Hydrocera Blume,Bijdr.: 241 (1825); Grey-Wilson, Kew Bull. 35:
213-219 (1980).
Semi-aquatic perennial herbs, rather succulent;

stems erect, with fleshy and fibrous roots. Leaves
alternate, petiolate, lamina linear-lanceolate to
elliptic, at base with pair of sessile glands. Flowers
pink with carmine to purplish, with 4lateral sepals
(2 pairs), almost as large as petals, lower sepal with
short, curved spur; petals 5, free, dorsal petal semi-
cucullate. Fruit a fleshy pseudoberry, with hard
endocarp which divides into 5 separate units
(partial endo carps, each with one seed and two air
sacs). One species, H. triflora (L.) Wight & Arn., Sri
Lanka, S India, S China to Thailand, Cambodia,
Laos, Malay Peninsula, Celebes and Java.
2. Impatiens L. Fig. 8
Impatiens L., Sp. Pl.: 937 (1753); Perrier de laBathie, Arch. Bot.
7, Mem. 1: 1-124 (1934); Perrier de la Bathie, Mem. Acad.
Fig. 8. Balsaminaceae. Impatiens cinnabarina. A Rootstock, Sci. Paris II, 67, 2: 1- 16 (1948); Shimizu, Acta Phytotax.
stem base and flowering shoot. B L ateral sepals. C Lower sepal Geobot. 24:43-51 (1969); Grey-Wilson, Impatiens of Africa
and spur. D Dorsal petal. E Lateral united petals. Drawn by C. (1980); Grey-Wilson, Kew Bull. 34: 661-668 (1980); 35:
Grey-Wilson. (Grey-Wilson 1980a) 203-211 (1980); 44: 61-66 (1989); 44: 67-106 (1989); 44:
115-122 (1989); 44: 711-716 (1989).
Trimorphopetalum Bak. (1887).
Impatientella H. Perr. (1927).
PHYLOGENY. Morphologically based classifica- Petalonema Peter (1928).
tions placed Balsaminaceae in Geraniales and sug-
gested close relationships to Tropaeolaceae and Terrestrial or epiphytic annual to perennial herbs;
Geraniaceae (e.g. Warburg and Reiche 1895; Cron- stems erect, ascending or decumbent. Leaves
quist 1981). The tenuinucellate ovule, endothe- alternate, verticillate or opposite, petiolate, lamina
lium, haustoria and exotestal seed coat of linear-lanceolate to orbicular. Flowers white,
Impatiens led Boesewinkel and Bouman (1991) to yellow, orange to purplish, red, greenish or dark
assume a link with sympetalous orders. Recent brown, lateral petals 2, rarely 4, much smaller than
molecular phylogenetic studies suggest that this petals, lower sepal with short or long, broad or fil-
family is a member of the newly circumscribed iform spur, or spur reduced and lacking (in subg.
order Ericales (Morton et al. 1997; APG 1998; Soltis Trimorphopetalum); petals 5, the 4 lateral united
et al. 2000). Within this broader alliance, plastid into pairs. Fruit an elastically dehiscent capsule,
(rbcL), nuclear (18SrDNA) and combined with few to numerous seeds. About 1000 spp. in
sequence data suggest a relationship between Bal- Europe, North and Central America, Northern
saminaceae and Marcgraviaceae, Pellicieraceae, Asia, Himalaya, India to China, New Guinea,
and Tetrameristaceae. It is not easy to understand tropical Africa and Madagascar.
how these woody families shared a common
ancestry with Balsaminaceae.
Balsaminaceae 25
Selected Bibliography Larsen, K. 1981. Chromosome numbers in Impatiens from

Thailand. Nord. J. Bot. 1: 43-44.
Lu, Y.-Q. 1991. Pollen morphology of Impatiens L. (Balsami-
Akiyama, S., Wakabayashi, M., Ohba, H. 1992. Chromosome naceae) and its taxonomic implications. Acta Phytotax. Sin.
evolution in Himalayan Impatiens (Balsaminaceae). Bot. J. 29: 352-357.
Linn. Soc. 109: 247-257. Merlin, C.M., Grant ,W.F. 1986. Hybridization studies in the
Antlfinger, A.E. 1986. Field germination and seedling gro~th genus Impatiens. Can. J. Bot. 64: 1069-1074.
of chasmogamous and cleistogamous progeny of Impatzens Metcalfe, C.R., Chalk, L. 1950. See general references.
capensis (Balsaminaceae). Am. J. Bot. 73: 1267-1273. Morton, C.M., Mori, S.A., Prance, G. T., Karol, K.G., Chase M. W.
APG (Angiosperm Phylogeny Group) 1998. See general 1997. Phylogenetic relationships of Lecythidaceae: a cladis-
references. tic analysis using rbcL sequence and morphological data.
Arisumi, T. 1974. Chromosome numbers and breeding behav- Am. J. Bot. 84: 530-540.
ior of hybrids among Celebes, Java and New Guinea species Narayana, L.L. 1965. Contributions to the embryology of Bal-
of Impatiens L. Hort. Sci. 9: 478-479. . saminaceae, part 2. J. Jap. Bot. 40: 104-116.
Arisumi, T. 1980. Chromosome numbers and comparative Oginuma, K., Tobe, H. 1991. Karyomorphology of two species
breeding behavior of certain Impatiens from Africa, India, of Impatiens from Kenya. Acta Phytotax. Geobot. 42: 67-71.
and New Guinea. J. Am. Soc. Hort. Sci. 105: 99-102. Paoletti, C., Holsinger, K.E. 1999. Spatial patterns of polygenic
Beck, A.R., Weigle, J.L., Kruger, E.W. 1974. Breeding behavior variation in Impatiens capensis, a species with an environ-
and chromosome numbers among New Guinea and Java mentally controlled mixed mating system. J. Evol. Bioi. 12:
Impatiens species, cultivated varieties, and their interspe- 689-696.
cific hybrids. Can. J. Bot. 52: 923-925. Rama Devi, D.1991. Floral anatomy of six species of Impatiens.
Boesewinkel, F.D., Bouman, F. 1991. The development of Feddes Repert. 102: 395-398.
bi- and unitegmic ovules and seeds in Impatiens (Balsami-
Randall, J.L., Hilu, K.W. 1990. Interference through improper
naceae). Bot. Jahrb. Syst.113: 87-104. pollen transfer in mixed stands of Impatiens capensis and I.
Cheek, M., Fischer, E. 1999. A tuberous and epiphytic new
pall ida (Balsaminaceae). Am. J. Bot. 77: 939-944.
species of Impatiens (Balsaminaceae) from Southwest Rao, R.V.S., Ayyangar, K.R., Sampathkumar, R. 1986. On the
Cameroon. Kew Bull. 54: 471-475.
karyological characteristics of some members of Balsami-
Cronquist, A. 1981. See general references. naceae. Cytologia 51: 251-260.
Govindarajan, T., Subramanian, D. 1986. Karyotaxonomy of Rust, R.W. 1977. Pollination in Impatiens capensis and Impa-
south Indian Balsaminaceae. Cytologia 51: 107-116. tiens pallida (Balsaminaceae). Bull. Torrey Bot. Club 104:
Grey-Wilson, C. 1980a. Impatiens of Africa. Rotterdam: A.A. 361-367.
Balkema. Rust, R. W. 1979. Pollination of Impatiens capensis: pollinators
Grey-Wilson, C. 1980b. Hybridization in African Impatiens. and nectar robbers. J. Kansas Entomol. Soc. 52(2): 297-308.
Studies in Balsaminaceae 2. Kew Bull. 34: 689-722.
Schemske, D. W. 1978. Evolution of reproductive characteristics
Grey-Wilson, C. 1980c. Hydrocera triflora, its floral morphol- in Impatiens (Balsaminaceae): the significance of cleis-
ogy and relationship with Impatiens. Kew Bull. 35: 213-
togamy and chasmogamy. Ecology 59: 596-613. .
219. Schemske, D.W. 1984. Population structure and local selectiOn
Gross, J., Husband, B.C., Stewart, S.C. 1998. Phenotypic selec- in Impatiens pallida (Balsaminaceae), a selfing annual. Evo-
tion in a natural population of Impatiens pallida Nutt. lution 38: 817-832
(Balsaminaceae ). J. Evol. Bioi. 11: 589-609.
Soltis, D.E. et a!. 2000. See general references.
Hegnauer, R. 1964, 1989. See general references. Sugawara, T., Akiyama, S., Murata, J., Yang, Y.-P. 1995. Karyol-
Hurlbert, A.H., Hosoi, S.A., Temeles, E.J., Ewald, P.W. 1996.
ogy of ten species of Impatiens (Balsaminaceae) from SW
Mobility of Impatiens capensis flowers: effect on pollen
Yunnan, China. Acta Phytotax. Geobot. 45: 119-125.
deposition and hummingbird foraging. Oecologia 105: 243-
Sugawara, T., Akiyama, S., Yang, Y.-P., Murata, J. 1997. Kary-
246.
ological characteristics of Impatiens (Balsaminaceae) in
Huynh, K.-L. 1968. Morphologie du pollen des Tropaeolacees Yunnan, China. Acta Phytotax. Geobot. 48: 7-14.
et des Balsaminacees I, II. Grana Palynol. 8: 88-184, 277- Venkateswarlu, J., Lakshminarayana, L. 1957. A contribution to
516.
the embryology of Hydrocera triflora W. et A. Phytomor-
Huynh, K.-L. 1970. Quelques caracteres cytologiques, phology 7: 194-203.
anatomiques et embryologiques distinctifs du genre Tropae-
Warburg, 0., Reiche, K. 1895. Balsaminaceae. In: Engler &
olum et du genre Impatiens, et position taxonomique de Ia Prantl, Nat. Pflanzenfam. Ill, 5: 383-392. Leipzig:
famille des Balsaminacees. Bull. Soc. Neuchatel. Sci. Nat. 93: Engelmann.
165-177.
Wilson, P. 1995. Selection for pollination success and the
Jones, K., Smith, J.B. 1966. The cytogeography of Impatiens L. mechanical fit of Impatiens flowers around bumblebee
(Balsaminaceae). Kew Bull. 20: 63-72.
bodies. Bioi. J. Linn. Soc. 55: 355-383.
Kato, M. 1988. Bumblebee visits to Impatiens spp. - pattern and
Zinoveva-Stahevitch, A.E., Grant W.F. 1984. Chromosome
efficiency. Oecologia 76: 364-370.
numbers in Impatiens (Balsaminaceae). Can. J. Bot. 62:
Kurtto, A. 1996. Impatiens glandulifera (Balsaminaceae) as an 2630-2635.
ornamental and escape in Finland, with notes on the other
Nordic countries. Symb. Bot. Upsal31: 221-228.
26 K. Kubitzki
Brunelliaceae
K. KUBITZKI
Brunelliaceae Engler (1897) in Engler & Prantl, Nat. Pflanzen- VEGETATIVE STRUCTURES. All Brunellia species
fam., Nachtr. und Register zu II-IV: 226 (1897), nom. are trees with a straight, cylindrical bole which
cons.
often reaches the canopy and can attain a height
of 40 m. Branching is copious and in dense stands
Evergreen trees, unarmed; twigs with angular
is confined to the upper third of the trunk. Young
internodes alternating with prominent nodes.
shoots have a ferrugineous, ochraceous or rufes-
Leaves simple or pinnate (when pinnate with
cent indumentum which may be lanate, appressed-
stipels on the rachis), opposite or ternate, mostly
pubescent, or tomentose and may fall off at an
dentate; stipules lateral, sometimes fragmented,
early stage or persist over a long time.
free; indumentum of unicellular hairs, ovary and
The leaves are simple or pinnate, the former con-
fruit usually bristly. Inflorescences axillary, thyrso-
dition probably being derived. In pin?a~e leaves,
paniculate, provided with small, usually caducous
the insertion of the leaflets on the rachis IS accom-
prophylls. Flowers hermaphrodite or through
panied by stipels. Leaf shape - pinnate vs. simple
abortion mostly unisexual and dioecious, 4, 5 or
- was the character on which Cuatrecasas {1970)
6( -8)-merous; sepals valvate, persistent in fruit;
based his two sections of Brunellia, but later
petals 0; nectary disk intrastaminal, adn~te to
{1985) he recognised that the unifoliate species
calyx, cupular, 8-1 0-lobed; stamens free, twice as
group was not monophyl~tic_. (His crit~rio~ for
many as sepals, in 2 whorls, rarely more; ~laments
distinguishing between umfohate and umfoholate
inserted in the notches of the nectary disk, those
leaves was the presence of stipels in the latter.)
of outer whorl alternate with, of inner opposite to
Venation of the leaves and leaflets is craspedodro-
sepals; anthers bithecate, introrse, dehiscing lo~gi
mous; the margins are serrate, crenate or biserrate
tudinally, the connective with a small protr~swn;
to even multi-indentate. The stipules are small,
disk intrastaminal, thick, fiat, concave, with as
subulate or lanceolate, falling off very early but
many indentations as stamens or staminodes,
leaving notable elliptic or circular scars. Often
tomentose or hispidulous; carpels free, basally
there seem to be more than two stipules which
immersed in disk, as many as sepals or fewer and
Cuatrecasas {1970) described as "paired" stipules
alternating with them; ovaries ovoid or e~lipsoi~, but this may rather be due to a splitting of the
hairy and mostly hispid, biovulate; stylodia _verti-
stipule primordia.
cal in flower, apically hooked or curled; stigmas
Nodes are tri(penta)-lacunar. The leaves have a
linear sutural-decurrent; ovules 2 per carpel, col-
hypodermis of one or two cell layers and conspic-
lateraL bitegmic, anatropous, epitropous, with
uous bundle sheath extensions which isolate the
micropyle directed upwards; obturator ~; fema~e
palisade parenchyma cells of the meshes ~f the leaf
flowers with sterile staminodia, male With rudi-
venation. Stomata seem to be anomocytic. In the
mentary pistil. Fruit polyfollicular; follicles
wood, growth rings are faintly visible or abse~t.
tomentose and mostly additionally hispid; stylo-
Vessel elements are solitary or arranged m
dia diverging horizontally; the hard e~docarp
radial multiples; both simple and scalarifo~m
detaching from softer exocarp at matunty and
perforation plates occur. Vessel members are qmte
expelling seeds which remain att~ched to a p~a
long {720-1480J.lm). Pitting between them and
centary stalk continuing the fumcle; seeds with
between vessels and rays varies from scalariform
hard, shiny testa and raised raphe; embryo lar~e,
through transitional to opposite. Libriform fibres
straight, embedded in carnose, mealy, white
are often septate and pitted on radial walls.
endosperm. n = 14. Rays are heterogeneous and up to 6 cells wide,
A monogeneric family comprising some 61
and usually have long uniseriate wings. Axial
species distributed from Me:Oco through ~es? parenchyma and crystals are absent (Eyde in
America and the Greater Antilles south to 18 S m
Cuatrecasas 1970).
Bolivia.
Brunelliaceae 27
Fig. 9. Brunelliaceae. A Brunellia standleyana, part of flores-

cence. B B. velutina, polyfollicle with exposed seeds. C B. lati-
folia, follicle structure with exocarp, endocarp, placental cord
and seeds. (Cuatrecasas 1970)
REPRODUCTIVE STRUCTURES. Brunellia has

thyrso-paniculate inflorescences (Fig. 10), differ-
ing mainly in the complexity of branching (Orozco
and Weberling 1999). The flowers have a cupuli-
form nectariferous disk interpreted as androgy-
nophore by Matthews and Endress (2002}. Notable
features of the flowers include the apocarpous
gynoecium partially adnate to the surrounding Fig. 10. Brunelliaceae. Brunellia integrifolia. A Habit.
disk, the partially open ventral carpel, the two col- B Flower. C Fruit. (Cuatrecasas 1970)
lateral ovules and the woody endocarp, and the
extended, sutural stigmas (also found in some
Rosaceae but not in Cunoniaceae). all formerly considered as possible relatives of
In most Brunellia, during fruit development the Brunelliaceae but which have small embryos. An
ventral side of the carpels grows faster than the aril, ascribed to Brunellia by Cuatrecasas (1970)
dorsal, pushing the stylodia into a horizontal posi- and others, has not been seen; the raphe is some-
tion and leading to the stellate arrangement what raised but relatively dry and hard.
attained in most species. To the reddish or ochra-
ceous indumentum of the fruits are added nearly PoLLEN MoRPHOLOGY. Pollen grains are tricol-
always hard, hispid, lignified trichomes. When the porate, prolate to oblate, 11 - 32.5 x 10.5 - 281J.m.
endocarp dehisces through transverse contraction The exine is incompletely to almost completely
of the endocarp, the seeds are expelled but remain tectate, and varies from coarsely reticulate to
attached to the fruit by a narrow ribbon of pla- rugulate and punctate (Orozco 2001; see also
cental and marginal exocarp tissue continuing the Marticorena in Cuatrecasas 1970).
funicle. This flexuose stalk holds the seeds upright
above the pericarp (Fig. 9), displaying the shining AFFINITIES. Engler's (1897) main reason for
seeds on the stellate polycarpic fruit. recognising Bunellia as an independent family was
Details of seed coat structure were given by the position of the ovules, which are epitropous
Lopez Naranjo and Huber (1971). Both testa and with a ventral raphe, whereas at his time Cunoni-
tegmen are initially 2-layered but eventually the aceae were considered to be apotropous through-
two layers of the tegmen fuse. The endosperm is out. (Cuatrecasas 1970: 45 observed, however, that
carnose, subhyaline; the embryo is nearly as long ovules in the basal Spiraeanthemoideae [Cunoni-
as the endosperm, a feature that distinguishes aceae] are also epitropous.) Perhaps more charac-
Brunelliaceae and Cunoniaceae from families such teristic may be the peculiar development of the
as Saxifragaceae, Escalloniaceae and Cornaceae, carpels, including their abaxial deformation, the
28 K. Kubitzki
sutural stigmas, the dissociation of exocarp and Selected Bibliography

endocarp, and the presentation of seeds on the
dehisced carpels. Bradford, J.C., Barnes, R.W. 2001. Phylogenetics and classifica-
The abovementioned similarities with Rosaceae tion of Cunoniaceae (Oxalidales) using chloroplast DNA
are probably parallelisms: the androecium in sequences and morphology. Syst. Bot. 26: 354-385.
Cuatrecasas, J. 1970. Brunelliaceae. Fl. Neotropica Monogr. 2.
Brunellia lacks the 10 + 5 + 5 pattern of Rosaceae, Darien: Hafner.
and the large embryo is oxalidalean rather than Cuatrecasas, J. 1985. Brunelliaceae. Fl. Neotropica Monogr. 2,
rosalean. Incidentally, also Engler (1897) referred Suppl. New York: The New York Botanical Garden.
to the diagrammatical similarity of Brunellia and Ehrendorfer, F., Morawetz, W., Dawe, J. 1984. The neotropical
Cephalotus, both now included in Oxalidales. This angiosperm families Brunelliaceae and Caryocaraceae: first
karyosystematical data and affinities. Pl. Syst. Evol. 145:
position is also confirmed by molecular analyses 183-191.
(e.g. Savolainen, Fay et al. 2000; Bradford and Engler, A. 1897. Brunelliaceae, pp. 182-184. In: Engler & Prantl,
Barnes 2001). Nat. Pflanzenfam., Nachtr. und Register zu II-IV. Leipzig: W.
Engelmann.
Engler,A.1930. Brunelliaceae, pp. 226-229. In: Engler & Prantl,
DISTRIBUTION AND HABITATS. Brunellia is exclu- Nat. Pflanzenfam. ed. 2, 18a. Leipzig: W. Engelmann.
sively American, with only six of the 61 known Lopez Naranjo, H., Huber, H. 1971. Anatomia comparada de las
species occurring north of Panama, of which a semillas de Brunellia y Weinmannia con respecto a su posi-
single, B. comocladiifolia, is native to the Greater cion sistematica. Pittiera 3: 19-28.
Antilles, where it has given rise to several well- Matthews, M.L., Endress, P.K. 2002. Comparative floral struc-
ture and systematics in Oxalidales (Oxalidaceae, Con-
defined subspecies. The South American species naraceae, Brunelliaceae, Cephalotaceae, Cunoniaceae,
are mostly Andean, with exception of four species Elaeocarpaceae, Tremandraceae). Bot. J. Linn. Soc. 140:
that extend to the Parfa Peninsula, Venezuela, and 321-381. With 104 figs.
the Roraima Massif in eastern Venezuela. The Orozco, C.I. 1997. Sobre Ia posicion sistematica de Brunellia
great majority of the species are narrow endemics, Ruiz & Pavon. Caldasia 19: 145-164.
Orozco, C.I. 2001. Pollen morphology of Brunellia (Brunelli-
and the majority of them (37) occurs in Colombia; aceae) and related taxa in the Cunoniaceae. Grana 40:
Ecuador has six species, Peru nine, and Bolivia 245-255.
five. The presence of Brunellia in the Greater Orozco, C. I., Weberling, F. 1999. A comparative study of inflo-
Antilles may indicate that the genus may have rescences in Brunellia Ruiz & Pav. (Brunelliaceae) and
related taxa. Beitr. Bioi. Pfl. 71: 261-279.
been represented north of Panama before the
Savolainen, V., Fay, M.F. et a!. 2000. See general references.
closing of the central American land bridge.
Most species grow in montane forests, either at
lower elevations such as from 600 to 2000 m, or
much higher, usually between 2800 and 3800 m. A
single species, B. hygrothermica, grows in the
superhumid sea-level region of Colombia.
A single genus:
Brunellia Ruiz & Pavon Fig. 10

Brune Ilia Ruiz & Pavon, Prodr. Fl. Peruv. Chi!.: 71, pl. 12 (1794);
Cuatrecasas, Fl. Neotropica 2 (1970), and Suppl. (1985).
Description as for family. A single genus of 61

described species.
Celastraceae 29
Celastraceae
M.P. SIMMONS
or angular, rarely lacerate or irregularly lobed;

Celastraceae R. Br., in Flinders, Voy. Terra Austral. 2: 554
(1814), nom. cons. stamens (2)3-5, rarely numerous, rarely 3long and
Hippocrateaceae Juss. (1811), nom. cons.
Stackhousiaceae R. Br. (1814), nom. cons.
2 short, rarely alternating with staminodes, sta-
Brexiaceae Loudon (1830). minodial or 0 in female flowers, alternate with
Siphonodontaceae (Croizat) Gagnep. & Tardieu (1951). petals when stamen number equals petal number,
Canotiaceae Airy Shaw (1965). anthers (1)2-celled, basifixed to dorsifixed, some-
Plagiopteraceae Airy Shaw ( 1965). times versatile, dehiscing longitudinally, obliquely,
Pottingeriaceae? (Engl.) Takht. (1987).
or transversely, introrse, latrorse, or extrorse,
rarely apical, connective sometimes apiculate,
M~noe~ious, andromonoecious, dioecious, gyn-
rarely pustular, with bilobed extension, or tipped
odwecwus, or polygamous, erect or scandent
by white gland, androgynophore occasionally +;
tr~es, shrubs, lianas with quickly-deciduous scales,
?vary sup:rio~ to half-inferior, often partially
without tendrils, or annual or perennial herbs with
Immersed m disk, present as pistillode or essen-
erect or prostrate stems, rarely suffrutices, rhi-
tially absent in male flowers, completely or incom-
zomatous shrubs, ericoid subshrubs, or epiphytic
plet:ly (1)2-5(10)-locular, rarely each locule
shrubs, gl~brous or glabrescent, rarely puberulent,
ho~IZonta~ly divided i.nto !-ovulate locelli, placen-
densely y!lose, or hirsute and stellate pubescent,
tatiOn axile, rarely mtruded parietal or basal,
often with elastic threads in soft tissues that are
ovules erect, axile, or pendulous, 1-12( -numer-
evident when broken, unarmed or with thorns
o~s) per !ocule; style terminal, simple, short to 0,
rarely ~ith stems terminating in sharp points:
stlg~a. Simple or lobed, rarely ovary with stig-
rarely With glandular stems, rarely with buttressed
matic hn~s on eac~ carpel on margin of an apical
trunks. Leaves simple, alternate, opposite, or sub-
hollow With style-hke central column arising from
opposite, rarely whorled, subverticillate, or irregu-
base of h~llow. Fruit a loculicidally and/or septici-
larly scattered, fasciculate on short shoots or
dally dehiscent capsule, rarely beaked, schizocarp
opposite on mature branches and alternat; on
of 2-5 i~dehis.cent m:ricarps, drupe, berry, or
juv~nile branches, pet.iolate or rarely sessile, rarely
samar~ With a smgle ~pi cal wmg, 3-5lateral wings,
gemculate, blade lammar, rarely needle- or scale-
or a smgle surroundmg wing, rarely an indehis-
like, venation pinnate, rarely acrodromous,
cent capsule or nut with lateral style, pericarp
secondary veins reticulate or rarely distinct
woody, bony, coriaceous, fleshy, or chartaceous,
crossbars, rarely with abaxial domatia in axils
rarely fibrous, capsules smooth, angular, deeply
or larger veins, margins entire, crenate, serrate,
lobed, transversely flattened and lobed to base, or
dentate, spinose, glandular-toothed, spinose-
rarely lobe.d ± halfway to base or entirely connate,
dentate, rarely notched; stipules small and
rarely echmate, laterally winged, with lateral or
ca?ucous, rarel~ 0. Inflorescences axillary or ter-
oblique horn-like outgrowths, or flattened along
mmal, rarely. epiphyllous or cauliflorous, thyrsoid,
each locule. Seeds !-numerous, smooth or
~ymose, fasciCulate, or flowers solitary, rarely pan-
occasionally furrowed, with unbranched or occa-
Iculate, umbellate-cymose, umbellate, racemose
sionally branched raphe, albuminous or exalbu-
or in s~ike~. Flowers actinomorphic, rarely± zygo~
minous, so~etimes winged, wing membranous,
morphic With 4 of 5 petals arched, bisexual or uni-
basal, someti.mes reduced to narrow stipe, apical,
sexual, rarely perigynous with short or cupular
or surroundmg seed, exarillate or aril basal to
hypanthium, perianth (3)4-5(6)-merous, sepals
completely enveloping seed, aril membranous
~nd petals. free, rarely petals medially connate; disk
fleshy, rarely with basal or apical filamentou~
mtrastammal, stamens on disk, or extrastaminal,
extensions, or mucilaginous.
annular, margins upturned, pulvinate, or cupular,
A subcosmopolitan family of 98 genera and
fleshy or membranous, sometimes indistinct or
about 1211 species that is most diverse in the
0, continuous, rarely discontinuous, entire, lobed,
tropics and subtropics, with fewer temperate
30 M.P. Simmons
species. One genus (Pottingeria) is tentatively erally petiolate, though they may be sessile (Apato-
included, another genus (Nicobariodendron) is phyllum, Empleuridium). Bhesa is unique in the
insufficiently known. family with geniculate petioles (at the laminar
end; Fig. 11A). Although leaves are generally
VEGETATIVE MORPHOLOGY. Most members of laminar, they may also be needle-like (Apatophyl-
Celastraceae are shrubs to small trees, although lum, Empleuridium), or even reduced to scales
members of some genera reach up to 50 m tall and (Canotia, Psammomoya). The scale-shaped leaves
have buttressed trunks (Bhesa, Kokoona, and of Canotia and Psammomoya do not appear
Lophopetalum spp.). Most Celastroideae are erect, similar and the two genera are not closely related
but some are scandent (Allocassine, Celastrus, (Simmons et al. 2001b), indicating that laminar
Euonymus spp., Maytenus spp., Monimopetalum, leaves have been reduced to scales independently
Ptelidium scandens, Tripterygium), whereas most in the two genera. Leaf dimorphism between juve-
Hippocrateoideae and Salacioideae are Hanas. nile and mature leaves occurs in Elaeodendron
Scandent taxa may climb with the aid of persist- spp., Pleurostylia pachyphloea, P. putamen, and
ent, sharp, downward-pointed bud scales (e.g., Brexia. Leaf dimorphism is particularly dramatic
Celastrus; Velenovsky 1910; Ding Hou 1955), in Elaeodendron orientale with entire, narrowly-
others use adventitious roots (Euonymus spp.; lanceolate juvenile leaves and serrulate, elliptic
Troll 1943). Members of Hippocrateoideae and mature leaves.
Salacioideae that grow as Hanas do so by thig-
motropism and quickly-deciduous scales on VEGETATIVE ANATOMY. Solereder (1908) and
dimorphic branches (Troll 1937; Loesener 1942b; Metcalfe and Chalk {1950) have provided
Halle 1962). Scandent Celastrus act as stranglers overviews of the vegetative anatomy of Celas-
(Troll1943). Atypical growth forms for the family traceae, and Stant {1951) has examined the
include: annual or perennial herbs with erect or anatomy of Stackhousioideae. Wood anatomy of
prostrate stems (Stackhousioideae), suffrutices Celastraceae has been examined by Record (1943),
(Gymnosporia spp.), rhizomatous shrubs (Paxis- Mennega (1972, 1994, 1997), Carlquist (1987;
tima), ericoid subshrubs (Empleuridium), and epi- Stackhousioideae) and Archer and van Wyk
phytic shrubs ( Quetzalia standleyi). (1993a). Growth rings are inconspicuous to dis-
Most genera are unarmed. However, stems ter- tinct. Growth ring borders are marked by the
minating in sharp points occur in Acanthotham- presence of unlignified ray cells in some
nus and Canotia, and thorns occur in Gloveria, Hippocrateoideae (Anthodon, Apodostigma,
Gymnosporia, Moya, and Putterlickia. In the same Cuervea, Pristimera, Reissantia, Simicratea,
four genera with thorns (and in two genera Simirestis). Vessels are solitary or in radial multi-
without thorns: Schaefferia spp., Wimmeria spp.), ples of 2-3(4). Perforation plates are generally
leaves are often fasciculate on short shoots, fre- simple, rarely scalariform (Bhesa, Brexia, Elaeo-
quently on the thorns themselves. Leaves (or fas- dendron, Perrottetia). Rays are often strictly
cicles of leaves) and inflorescences are borne on uni-biseriate, sometimes of two distinct sizes of
the thorns (except in Moya). These leaves on the 1-2 cells wide and 3-6 cells wide (Celastrus,Elaeo-
thorns are often opposite, even when the rest of dendron, Microtropis, Schaefferia, Siphonodon), or
the plant has alternate phyllotaxy. A similar uniseriate and 4-40 cells wide (Hippocrateoideae ).
pattern is found in Catha edulis and Lydenburgia Fibers are thin- to thick-walled, with numerous
cassinoides, which are closely related to Gym- bordered pits. In some taxa, parenchyma-like
nosporia and Putterlickia (Simmons et al. 2001b). bands of thin-walled, septate wood fibers occur
In these two species that lack thorns, leaves are (e.g., Elaeodendron). Included phloem, generally
opposite on mature (flowering) branches and as conspicuous concentric bands, occasionally as
alternate on juvenile branches. Based on the isolated strands, occurs in Salacioideae.
similar pattern and close relationship, the thorns In stems, the cork generally develops in the
of Gymnosporia and Putterlickia may be homolo- subepidermis. However, the cork develops in
gous to the mature, unarmed branches of Catha the epidermis of Euonymus and Microtropis. Cork
edulis and Lydenburgia cassinoides. wings occur in Euonymus alatus and E. phel-
Leaves are simple, and generally alternate, oppo- lomanus. Phloem and xylem are in closed bundles
site, or subopposite. Leaves may also be whorled separated by narrow rays. Laticifers containing
(Brexiella ilicifolia, Crossopetalum spp., Euonymus gutta-percha occur at the interface of the cortex
spp., Euonymopsis spp.), subverticillate (Salaciop- and phloem (Drennan et al. 1987), but perhaps not
sis), or scattered (Empleuridium). Leaves are genin Stackhousioideae (Stant 1951). Pith is homage-
Celastraceae 31
neous or heterogeneous, absent in Tripterygium stamens) and Salacioideae (except some Cheilo-
wilfordii. clinium spp. with 5 stamens). Exceptions are 2
Leaf epidermal characters have been examined stamens in Nicobariodendron and Salacia spp.,
by Pant and Kidwai (1966), den Hartog and Baas and numerous stamens in Plagiopteron. Loesener
(1978), and Archer (1990). Leaf anatomy has been ( 1942a, 1942b) used stamen number as the only
examined by Jansen and Baas (1973) and Muller character to differentiate Celastraceae s.str. (4-5
(1995). Stomatal types are very diverse (den stamens) from Hippocrateaceae (2-3 stamens).
Hartog and Baas 1978; Archer 1990). Stomata are However, there appear to have been multiple
often strictly abaxial, sometimes also on the changes in stamen number from 5 to 3 (inde-
midrib of the adaxial surface, rarely adaxial and pendently in Dicarpellum, and one or more origins
abaxial. In some genera, crystals occur in small in Hippocrateoideae and Salacioideae), and from
epidermal cells as druses (some Salacia spp., 3 to 5 (independently in Campylostemon and
Siphonodon, some Hippocrateoideae) or solitary, Cheiloclinium; Simmons et al. 2001b). In Siphon-
rhomboidal crystals (Elaeodendron, some odon, stamens alternate with staminodes. In Stack-
Lophopetalum spp., some Hippocrateoideae). housia and Tripterococcus, the stamens are of
Perrottetia alpestris and P. sandwicensis have unequal lengths with 3 long stamens and 2 short
mucilaginous epidermal cells (Solereder 1908). stamens (Fig. 17c).
Laticifers containing gutta-percha occur in acres- Anthers are generally 2-celled, but can be 1-
cent-shaped layer on the abaxial sides of vascular celled in some Euonymus spp. Anthers may
bundles (Drennan et al. 1987), and sometimes in dehisce longitudinally (most Celastroideae),
the mesophyll (some Wimmeria spp., many obliquely (e.g., Glyptopetalum, Plenckia), or trans-
Hippocrateoideae and Salacioideae). The petiole versely (Hypsophila, most Hippocrateoideae and
generally has one arc-shaped or circular vascular Salacioideae). The direction of anther dehiscence
bundle, rarely three (Bhesa, Brexia) or two to may be introrse (most Celastroideae), latrorse
several vascular bundles (Lophopetalum, and (e.g., Hedraianthera, Kokoona), or extrorse (most
some Hippocrateoideae and Salacioideae). Hippocrateoideae and Salacioideae), or rarely
apical (Plagiopteron [Fig. 19E], Salacia spp.). Con-
FLORAL MoRPHOLOGY. Flowers are actinomor- nectives are generally apiculate or not extended
phic, but ± zygomorphic with 4 of 5 arched (often variable in a single inflorescence), are pus-
petals in Apodostigma. The perianth is generally tular in some Kokoona spp., have bilobed exten-
4-5-merous, but rarely 3-merous (Elaeodendron sions in some Peritassa spp., or are tipped by a
spp., Plagiopteron) or 6-merous (rarely in white gland in Macgregoria racemigera. Androg-
Gymnosporia). ynophores are generally absent, but they are
The nectariferous disk may be intrastaminal present in some Hippocrateoideae (Fig. 18c) and
(most Celastroideae, Figs. 12-14), extrastaminal Salacioideae.
(Brassiantha, Dicarpellum, Hypsophila, Kokoona, Ovaries are 2-5-locular (3-locular in Hippocra-
Sarawakodendron, Salacioideae, most Hippocra- teoideae and Salacioideae except some Cheiloclin-
teoideae), or may have stamens on the disk (e.g., ium and Salacia spp.), but may be 1-locular
Glyptopetalum, Lophopetalum). In Celastroideae, (Empleuridium,Pottingeria) or 10-locular (Siphon-
the disk is generally annular or has the margins odon ). Placentation is axile, rarely intruded parietal
upturned, whereas in Hippocrateoideae and Sala- (Pottingeria) or basal (Empleuridium). Ovules
cioideae the disk is generally pulvinate or cupular. may be erect (most Celastroideae), axile (most
The disk is generally fleshy or membranous, but Hippocrateoideae and Salacioideae), or pendulous
sometimes indistinct (Arnicratea, Plagiopteron, (Euonymus spp., Glyptopetalum, Gyminda,
Schaefferia spp., Simirestis) or absent (Bequaertia, Maurocenia, Tetrasiphon). Styles are simple and
Campylostemon, Microtropis, Schaefferia spp., Tris- range from short to obsolete (e.g., Brassiantha),
temonanthus). The disk is generally continuous, and the stigmas simple or 2-5-lobed.
but is discontinuous in Apodostigma and Cheilo- Flowers of Plagiopteron, Pottingeria, and
clinium. The disk margin may be entire, lobed, Siphonodon are unique within the family: Pla-
or angular, rarely lacerate (Brexia) or irregularly giopteron (Fig. 19) with its numerous stamens that
lobed (Fig. 18, Helictonema). In Stackhousioideae, have apically-dehiscent anthers; Pottingeria with
a thin disk lines the hypanthium. its 1-locular, 3-carpellate, paracarpous ovaries
There are 4-5 stamens in Celastroideae (except with numerous ovules per carpel and septicidally
3 stamens in Dicarpellum) and 3 stamens in dehiscent capsules; Siphonodon with its 10-locular
Hippocrateoideae (except Campylostemon with 5 ovaries in which each locule is horizontally
32 M.P. Simmons
divided into !-ovulate !ocelli. In Siphonodon, a or polyads have triporate grains. Individual grains
stigmatic line is located at the upper inner margin are suboblate to subprolate, small to medium sized
of each carpel around a central, apical hollow (longest axis 11-35Jlm), with reticulate (rarely
within the ovary; a sterile, style-like central crotonoid) sculpturing (Lobreau-Callen 1977). An
column arises from the base of the hollow (Hooker endexine fold (also described as a conduplication
1857). Although Plagiopteron and Siphonodon are or replication) in the aperture is characteristic of
well-supported members of Celastraceae, Pottin- Celastraceae s.l., but is absent in Hippocratea and
geria has not been incorporated in a phylogenetic Lauridia (Lobreau-Callen 1977; Archer and van
analysis, and its inclusion within the family is Wyk 1992). An "annulus" (thickening around the
uncertain (Airy Shaw et al. 1973). internal margin of the pore) occurs in Kokoona,
Lophopetalum, and Hippocrateoideae (Halle 1960;
FLORAL ANATOMY. In his study of the vascular Lobreau-Callen 1977).
traces of Celastrus, Euonymus, and Paxistima
flowers, Berkeley (1953) concluded that Celas- KARYOLOGY. Relatively few chromosome num-
traceae have a floral tube, and the nectariferous bers have been published, mostly in Celastrus
disk is homologous to modified stamens. In Celas- and Euonymus. Base chromosome numbers in
trus, the vascular traces of the disk diverge from Celastraceae include x = 8, 9, 10, 12, 14, 15, 17,and
stamen and petal traces. In Euonymus and Paxis- 23. Polyploidy appears to be relatively common in
tima, no vascular traces lead to the disk. Celastraceae. For example, in Euonymus, gameto-
phytic chromosome numbers range from eight
EMBRYOLOGY. The embryology of Celastraceae (E. echinatus; Mehra 1976), 16 (e.g., E. radicans;
has been examined in several species by Maurit- Bowden 1940), and 24 (E. bullatus; Mehra 1976) to
zon (1936a, 1936b), Adatia and Gavde (1962), 32 (e.g., E. europaeus; Wulff 1937). Although base
Narang (1965), and Tobe and Raven (1993). chromosome numbers fluctuate in Celastroideae,
Anthers are tetrasporangiate, have a fibrous the base chromosome number has stabilized in
endothecium, crushed middle layers, and a Hippocrateoideae and Salacioideae at x = 14
glandular tapetum. Tapetal cells are two- or (except n = 30 in Semialarium mexicanum; Bawa
more-nucleate. Cytokinesis in meiosis is simulta- 1973), with n = 14 and 28.
neous. Microspore tetrads are tetrahedral or
decussate. Mature pollen is 2- or 3-celled PoLLINATION. Few taxa have been investigated to
(Brewbaker 1967). determine their specific pollinators. Nectariferous
Ovules are anatropous, bitegmic, crassinucellate disks, present in most genera, attract bees,
or tenuinucellate, and have the Polygonum-type of beetles, flies, wasps, and even ants (Knuth 1908;
embryo sac formation. Endosperm formation is Ding Hou 1962; Brizicky 1964a; Sebsebe 1985).
nuclear. Endosperm may be present or absent in Insects are attracted to the thin layer of nectar
mature seeds. In mature seeds, the embryos are secreted by the disks of flowers of Euonymus
straight and symmetrical. Polyembryony has been europaeus, generally resulting in cross-pollination
found to be common in several species of Euony- (Darwin 1877; Miiller 1883; Knuth 1908). In addi-
mus and in Celastrus paniculatus (Adatia and tion to the nectariferous disks, the carrion odor
Gavde 1962; Brizicky 1964b). produced by flowers of Gymnosporia buxifolia
attracts insects, especially bluebottle flies (Palmer
PoLLEN MoRPHOLOGY. The most thorough and Pitman 1972). Flowers of Stackhousioideae
survey of pollen in Celastraceae was completed by produce a sweet smell that attracts insects
Lobreau-Callen (1977). Within Celastroideae, (Mattfeld 1942).
pollen is isopolar and shed in monads, whereas
in Hippocrateoideae, Salacioideae, and REPRODUCTIVE SYSTEMS. Most Celastraceae
Lophopetalum, pollen may also be shed as tetra- have bisexual flowers. Among those with unisex-
hedral tetrads, or polyads of eight or 16 grains ual flowers, plants may be monoecious (Elaeoden-
(Ding Hou 1969; Lobreau-Callen 1977). The aper- dron spp., Gymnosporia spp.), andromonoecious
tures of grains shed in tetrads or polyads may be (Peripterygia), gynodioecious (some Euonymus,
irregularly positioned. Grains are generally tricol- Maurocenia [though perhaps dioecious; Archer
porate, but bicolporate and tetracolporate grains and van Wyk 1998a]), polygamous (Elaeodendron
have been reported (Ikuse 1956; Ding Hou 1969). spp., Gymnosporia, Maytenus spp., Orthosphenia,
Some genera in which grains are shed as tetrads Plenckia, Tricerma, Tripterygium), or dioecious
Celastraceae 33
(Celastrus, Dicarpellum, Elaeodendron spp., 1983). The thickened margin of the wing has also
Euonymus spp., Gyminda, Gymnosporia spp., been described as the raphe (Miers 1872; Smith
Maytenus spp., Menepetalum, Microtropis, Nico- 1940). However, the figures by Halle (1983) con-
bariodendron, Perrottetia, Quetzalia, Salaciopsis, clusively demonstrate that the raphe generally
Schaefferia, Tetrasiphon). traverses the wing near its center.
Within Celastraceae, the reproductive system of The fruits of Salacioideae have been described
Euonymus europaeus, a gynodioecious species, as baccate (e.g., Lindley 1853; Bentham and
has been most-intensively studied (Darwin 1877; Hooker 1862; Halle 1986), drupaceous (e.g., Miers
Webb 1979). Bisexual flowers are protandrous, 1872; Smith 1940; Ding Hou 1964), or drupaceous
making self-fertilization less likely (Muller 1883; or baccate (Loesener 1942b). I contend these fruits
Knuth 1908). Hermaphrodites are significantly are properly described as baccate because I have
more common than females, although the neither seen nor read of a stony endocarp in these
number of flowers per inflorescence is about fruits. The seeds are surrounded by mucilaginous
the same. Females produce significantly more pulp that is derived from the funiculus and is
fruits and seeds per inflorescence than hermaph- homologous to an aril (Miers 1872; Baillon 1880;
rodites. Furthermore, the females produce larger Simmons and Hedin 1999).
seeds. Planchon (1845) described the putative arils of
Hybrid individuals and putative hybrid species Celastrus scandens and Euonymus latifolius as
have been reported in Euonymus (Staszkiewicz arillodes - false arils. Planchon cited these false
1997) and Celastrus (Ding Hou 1955; Dreyer et al. arils as derived from the exostome of the outer
1987). However, artificial hybridization between integument rather than the funiculus. However,
Celastrus orbiculatus and C. scandens has Miers (1856) disputed Planchon's conclusion and,
been relatively unsuccessful (White and Bowden based on his investigation of Euonymus europaeus,
1947). he concluded that the aril is derived from the
funiculus and is therefore a true aril. Pfeiffer
FRUIT AND SEED. Fruit types in Celastroideae (1891) described the arils of Celastrus, Euonymus,
include: capsules with loculicidal dehiscence (e.g., and Gymnosporia cassinoides as derived from the
Maytenus), loculicidal and septicidal dehiscence exostome and the hilum. Corner (1976) described
(Canotia ), or septicidal dehiscence (Pottingeria ), the aril of Euonymus glandulosus as derived
schizocarps of indehiscent mericarps (Stack- entirely from the funiculus, and the aril of other
housioideae ), drupes (e.g., Elaeodendron ), berries species as derived from the exostome and the
(e.g., Cassine), nuts (Pleurostylia), or samaras with funiculus (Catha edulis, Celastrus paniculatus,
a single apical wing (Plenckia, Rzedowskia, Sarawakodendron filamentosum). Van der Pijl
Zinowiewia), 3-5 lateral wings (Platypterocarpus, (1972) concluded that Euonymus has an arillode,
Tripterygium, Wimmeria), or a single surrounding not an aril.
wing (Ptelidium). Alate seeds may have a basal wing with medial
The unusual, capsular fruits of Hippocra- attachment of the funiculus vasculature ( Canotia
teoideae are transversely flattened and trilobed, holacantha, Catha edulis), a wing surrounding the
with the lobes generally extending all the way to seed with medial attachment of the funiculus vas-
their base (Fig. 18H). Each of these three segments culature (Kokoona [appearing primarily as an
of the fruit has been referred to as separate cap- apical wing], Lophopetalum, Peripterygia), and
sules (Miers 1872; Smith 1940) or mericarps (Halle basal wings with basal attachment of the funicu-
1962, 1983; Gorts-van Rijn and Mennega 1994). lus vasculature (Hippocrateoideae). The wing of
However, these segments are not separate capsules Catha edulis has been described as an aril derived
because they all develop from a single ovary, and from the funicle and side of the exostome
they are not mericarps because they do not split (Loesener 1942a; Corner 1976). The winged seeds
into separate seed-containing parts upon dehis- of other Celastraceae may also be interpreted as
cence (Brizicky 1964a). The seeds of Hippocra- homologous to arils or arillodes (van der Pijl1972;
teoideae generally have membranous basal wings Simmons and Hedin 1999; but see Miers 1872, and
that may be reduced to narrow stipes (Halle 1983). Halle 1983). Seed wings have been interpreted as
The raphe generally traverses the wing near its being derived from arils in other families (e.g.,
center (e.g., Hippocratea) or rarely along the Forman 1965; Tobe and Raven 1984), and Corner
margin of the wing opposite the thickened margin (1954: 156) concluded that "all winged seeds are
(Helictonema velutinaum, Fig. 18K; Halle 1962, prima facie indications of arillate ancestry". In
34 M.P. Simmons
phylogenetic analyses of Celastraceae, genera with teoideae). The winged capsular fruits of some
winged seeds have been consistently resolved as Euonymus spp. are unlikely to serve for wind dis-
nested within clades of obviously-arillate taxa persal, but may serve to make the fruits more con-
(Simmons and Hedin 1999; Simmons et al. 2001a, spicuous to animals (Blakelock 1951). Birds and/ or
2001b). mammals disperse berries and drupes, as has been
Seed anatomy of Celastraceae has been reported for drupes of Elaeodendron australis
described by Corner (1976). The testa is 5-12 cells (Benson and McDougall 1995), E. croceum
thick, generally lignified, and unspecialized, often (Phillips 1927), E. glaucum, E. viburnifolium
with a palisade exotesta. The tegmen is 3-10 cells (Ridley 1930), Mystroxylon aethiopicum, and M.
thick, generally with ribbon-like fibers for the burkeanum, and berries of Cassine peragua and
outer layer, making the seeds exotegmic. However, Lauridia tetragona (Palmer and Pitman 1972).
the tegmen is crushed and absorbed in Drupes of Elaeodendron croceum have been
Salacioideae. In Microtropis, Perrottetia, and reported to be dispersed by elephants grazing on
Quetzalia, the pulpy sarcotesta may be mistaken the foliage (Ridley 1930). The berries of Sala-
for an aril. In the testa of Perrottetia, prismatic cioideae, with their seeds surrounded by mucilagi-
Malpighian cells project to different lengths nous arils, have been reported to be dispersed by
causing the furrowed appearance of the seeds. The monkeys (Garber 1986).
embryo has a short to long radicle, and flat, gen- Capsular fruits with seeds that have bright-
erally green cotyledons that are free to united. colored arils that contrast with the color of the
In Salacioideae, the cotyledons are generally capsule and seed occur in many members of the
described as massive (e.g., Miers 1872; Loesener family. These seeds are bird dispersed (reported
1942b; Halle 1962), and may be free or fused (Ding for Celastrus, Euonymus, and Maytenus; Pfeiffer
Hou 1964; de Vogel1980). However, Corner (1976: 1891; Ridley 1930; Martin et al. 1951; Ding Hou
94) described the embryo of Salacia sp. as "com- 1962; Brizicky 1964a; Palmer and Pitman 1972;
posed entirely of swollen hypocotyl ... without Dreyer et al. 1987; Benson and McDougall 1995),
trace of cotyledons". Corner presumably inter- and both small and large mammal dispersed
preted the fused cotyledons as the swollen (reported for Celastrus, Euonymus, and May tenus;
hypocotyl. Endosperm, when present (it is absent Martin et al. 1951; Dreyer et al. 1987; Rodrigues
in Goniodiscus, Kokoona, Lophopetalum, Polycar- et al. 1993; Kollmann et al. 1998). Additionally,
dia, Hippocrateoideae and Salacioideae), is oily, the seeds of Maytenus silvestris have been
generally thin-walled, and ranges from minimal to reported to be ant dispersed (Benson and
abundant. McDougall1995).
Seed germination of Celastraceae has been
studied by Lubbock (1892), Halle (1962), Rudolf PHYTOCHEMISTRY. The phytochemistry of Celas-
(1974), Wendel (1974), and de Vogel (1980). Seeds traceae has been reviewed by Hegnauer (in Ding
of Salacioideae have hypogeal germination. Seeds Hou 1962), Hegnauer (1964, 1966, 1989), Gibbs
of Hippocrateoideae have hypogeal (Cuervea (1974), and Bruning and Wagner (1978). Dulcitol,
macrophylla, Elachyptera sp., Loeseneriella accumulated in leaves and bark, is thought to
africana, Simirestis unguiculata) or epigeal occur in all members of the family. Dulcitol may
(Apodostigma pallens, Reissantia astericantha) be "a highly characteristic biochemical feature" of
germination. Other Celastraceae (Bhesa robusta, Celastraceae (Hegnauer in Ding Hou 1962: 229).
Celastrus scandens, C. paniculatus, Euonymus spp., Two alkaloids, (L(S)-(- )-alpha-aminopropiophe-
Lophopetalum javanicum, and Siphonodon celas- none (cathinone) and d-norpseudoephedrine
trineus) have epigeal germination. In all but (cathine), concentrated in young leaves and twigs
Lophopetalum and Salacia, the seedlings become of Catha edulis, act as stimulants of the central
free from their seedcoats. nervous system, similar to amphetamine (Kriko-
rian and Getahun 1973; Szendrei 1981; Zeiger et al.
DISPERSAL. Several genera are adapted for wind 1981). Maytansine and mayteine, isolated from
dispersal with samaras (Platypterocarpus, Gymnosporia, Maytenus, and Putterlickia, have
Plenckia, Ptelidium, Rzedowskia, Tripterygium, shown tumor-inhibitory activity and have been
Wimmeria, and Zinowiewia), winged mericarps investigated in clinical trials (summarized by
(Stackhousia spp., Tripterococcus [Fig. 17E]), or Sebsebe 1985, and Raintree Nutrition, Inc. 2000b).
winged seeds (Canotia holacantha, Catha edulis Many taxa have conspicuous, yellow pigments in
[Fig. 12G], Kokoona, Lophopetalum [Ridley 1930; their bark (of stems and/or roots) that are triter-
Sinha and Davidar 1992], Peripterygia, Hippocra- pene derivatives (celastrol, pristemerin or tin-
Celastraceae 35
genin) and may be ubiquitous within the family supported as monophyletic groups, although the
(Bruning and Wagner 1978; R. Archer, pers. comm. tribes Campylostemoneae, Helictonemateae,
2000). and Hippocrateeae have not been supported
Gutta-percha (E-1,4-polyisoprene) has been iso- (Simmons and Hedin 1999; Savolainen, Fay et al.
lated from many species in Celastraceae and is 2000; Simmons et al. 2001a, 2001b).
apparently universal in the family. Gutta-percha Stackhousia and Tripterococcus (Stackhousi-
is the trans-configuration of 1,4-polyisoprene, aceae) have been supported as nested within
whereas rubber is the cis-configuration. The pres- Celastraceae, using rbcL (Savolainen, Fay et al.
ence of gutta-percha is generally evident as elastic 2000), a simultaneous analysis of atpB, rbcL, and
threads when soft tissues are broken and pulled 18S nrDNA (Soltis et al. 2000), and 26S nrDNA
apart (but not, for example, in Allocassine; Archer and a simultaneous analysis of morphological and
and van Wyk 1998b). Gutta-percha is produced in molecular characters (Simmons et al. 2001b).
laticifers and occurs in the cytoplasm of most However, Stackhousiaceae have been resolved as
types of cells (Drennan et al. 1987). excluded from Celastraceae using the 5' flanking
region of rbcL (Savolainen et al. 1997) and mor-
SUBDIVISION AND RELATIONSHIPS WITHIN THE phology (Simmons and Hedin 1999). Note that
FAMILY. The most-recent comprehensive taxo- these two latter studies were poorly sampled for
nomic treatments of Celastraceae s.str. and Hip- taxa or characters, respectively, and that the best-
pocrateaceae were conducted by Loesener ( 1942a, supported hypothesis is that Stackhousiaceae
1942b) and Halle (1962, 1986, 1990), respectively. should be included within Celastraceae.
Loesener (1942a) recognized five subfamilies and The most-comprehensive phylogenetic analysis
five tribes of Celastraceae s.str. However, Loe- of Celastraceae sampled 71 species representing 53
sener's subfamilies and tribes have been found to genera that have been assigned to Celastraceae for
be heterogeneous in wood anatomy (Metcalfe and 26S nrDNA, phytochrome B, rbcL, atpB, and mor-
Chalk 1950), pollen structure (Lobreau-Callen phology (Simmons et al. 2001b). Although this
1977), and leaf anatomy (Den Hartog and Baas analysis has been useful to test earlier taxonomic
1978). Furthermore, Loesener's subfamilies and assertions regarding generic circumscriptions
tribes have been consistently resolved as unnatu- and intergeneric relationships, it is insufficiently
ral groups in phylogenetic analyses of the family, sampled to use as a basis to propose a new classi-
using morphological and molecular characters fication of Celastraceae to replace the classifica-
(Simmons and Hedin 1999; Simmons et al. 2001a, tions by Loesener (1942a) and Halle (1962, 1986,
2001b). 1990).
Loesener (1942b) and Halle (1962) recognized The four subfamilies in the classification used
Hippocrateaceae as a family, separate from Celas- here are the three monophyletic subfamilies Stack-
traceae. Loesener (1942b) did not formally subdi- housioideae, Hippocrateoideae, and Salacioideae
vide Hippocrateaceae. Halle (1962) described two that are independently derived from the para-
subfamilies (Hippocrateoideae, Salacioideae) and phyletic Celastroideae. Each of the three mono-
two tribes of the subfamily Hippocrateoideae: phyletic subfamilies are readily diagnosable using
Campylostemonae [sic] (properly Campyloste- morphological characters. This is an informal
moneae) and Hippocrateae [sic] (properly Hip- classification because of the obvious paraphyly of
pocrateeae). Halle (1986) added a third tribe, the subfamily Celastroideae.
Helictonemae [sic] (properly Helictonemateae).
Halle later recognized Hippocrateaceae as a tribe AFFINITIES. In broad-scale phylogenetic analyses
(e.g., Halle 1978) or as a subfamily (Halle 1986, that have been undersampled for Celastraceae,
1990) of Celastraceae. Lepuropetalon and/or Parnassia (Parnassiaceae)
The Hippocrateaceae have been consistently have been resolved as the sister group of Celas-
supported as nested within Celastraceae s.str., traceae using rbcL (Chase et al. 1993; Morgan and
using morphological and molecular characters Soltis 1993; Savolainen, Chase et al. 2000;
(Savolainen et al. 1997; Simmons and Hedin 1999; Savolainen, Fay et al. 2000), the 5' flanking region
Savolainen, Chase et al. 2000; Savolainen, Fay et al. of rbcL (Savolainen et al. 1997), and 18S nrDNA
2000; Soltis et al. 2000; Simmons et al. 2001a, (Soltis et al. 1997). In a comparatively well-
2001b). With the inclusion of Lophopetalum and sampled phylogenetic analysis of Celastraceae,
Plagiopteron, and the exclusion of Dicarpellum, using 26S nrDNA and a simultaneous analysis
the family Hippocrateaceae and the subfamilies of morphological and molecular characters,
Hippocrateoideae and Salacioideae have been Lepuropetalon and Parnassia were resolved, in a
36 M.P. Simmons
weakly-supported clade, as part of an early- none) and d-norpseudoephedrine (cathine), are

derived lineage within Celastraceae (Simmons concentrated in young leaves and twigs. Kat, cathi-
et al. 2001 b). However, because of their distinctive none, and cathine are now controlled drugs in the
morphology, and because the clade from Simmons U.S.A. (Federal Sentencing Guidelines Manual
et al. (2001b) was weakly-supported and based 1998).
only on characters from 26S nrDNA, Lep- Celastrus paniculatus, Kokoona zeylanica, Mys-
uropetalon and Parnassia are here treated as a troxylon aethiopicum, and species of Elaeoden-
separate family, Parnassiaceae. dron, Euonymus, Gymnosporia, Hippocratea,
Afrostyrax (Huaceae) has been resolved as the Maytenus, and Salacia have been used for making
sister group of the clade of Celastraceae and Par- traditional medicinal extracts (Blakelock 1951;
nassiaceae [but Parnassiaceae has not been Ding Hou 1962; Palmer and Pitman 1972; Burkill
sampled in a simultaneous analysis of atpB and 1985; Sebsebe 1985; Chant 1993; Raintree Nutri-
rbcL (Savolainen, Chase et al. 2000), and a simul- tion, Inc. 2000a, 2000b ). Euonymus atropurpureus,
taneous analysis of atpB, rbcL, and 18S nrDNA E. europaeus, E. latifolius, Elaeodendron
(Soltis et al. 2000)], but not in a comparatively buchananii and E. croceum are poisonous (Blake-
well-sampled analysis of rbcL (Savolainen, Fay lock 1951; Palmer and Pitman 1972; Burkill1985).
et al. 2000). In the latter study, Lepidobotrys and Pristimera celastroides and Tripterygium wilfordii
Ruptiliocarpon (Lepidobotryaceae), which had not have been used as insecticides (Smith 1940;
been previously sampled in broad-scale molecular Loesener 1942a). Oil has been extracted from the
analyses, were resolved as the sister group of the seeds of Euonymus europaeus, Goniodiscus
clade of Celastraceae and Parnassiaceae. elaeospermus, and Kokoona zeylanica (Kuhlmann
1933; Chant 1993). Human food sources have
DISTRIBUTION AND HABITATS. The family has a included the arils of Bhesa, members of Sala-
subcosmopolitan distribution (from tropical to cioideae (Smith 1940; Ding Hou 1962), the seeds of
temperate regions; absent in arctic regions) and Arnicratea grahamii, Hippocratea volubilis, and
is most diverse in the tropics and subtropics. Hylenaea comosa (Smith 1940; Loesener 1942b),
Although numerous species of Celastroideae are the drupes of Mystroxylon aethiopicum, and the
native to temperate regions, species of Hippocra- berries of Lauridia tetragona and Maurocenia
teoideae and Salacioideae are restricted to the frangula (Palmer and Pitman 1972). Various
tropics and subtropics. Major centers of diversity species of Catha, Celastrus, Elaeodendron, Euony-
for the family are Australia (for Stackhousioideae ), mus, Maytenus, Paxistima, Putterlickia, and
tropical America and Africa (for Hippocrateoideae Tripterygium are cultivated as ornamentals
and Salacioideae), South Africa (for Celas- (Huxley et al. 1992). Wood of Bhesa paniculata,
troideae), Madagascar, Macronesia, and Queens- Euonymus javanicus, Kokoona littoralis, K. reflexa,
land, Australia (for Celastroideae). Lophopetalum wightianum, L. multinervium,
Species of Celastraceae generally grow in wood- Maytenus spp., and Siphonodon celastrineus has
lands, from dry, gallery forests to swamps, from sea been used for timber (Record and Hess 1943; Ding
level to 3350 m elevation. Wooded habitats range Hou 1962; Palmer and Pitman 1972). Wood of
from thickets and bush to primary rain forests. Euonymus europaeus and E. hians has been used
Other members of the family grow in grasslands for turnery (Chant 1993). Several lianas, such as
(Gymnosporia spp., Salacia spp.), deserts (Acan- Loeseneriella apocynoides, are used for binding
thothamnus, Canotia, Mortonia), and coastal and basket weaving (Muhwezi 1999). Gutta-percha
dunes (Crossopetalum spp., Lauridia, Robsonoden- has been extracted from the bark of E. verrucosus
dron, and Salvadoropsis). At least one species, (Blakelock 1951). Bark of Kokoona and
Elaeodendron viburnifolium, grows in tidal man- Lophopetalum, which contain a thin oily layer, has
grove swamps (Ding Hou 1962) and another been used to start fires, even in wet conditions
species, Quetzalia standleyi, may grow as epiphytic (Ding Hou 1962). Bark extracts of Lophopetalum
shrubs (Lundell1939). javanicum and L. pallidum have been used in dart
and arrow poisons (Loesener 1942a; Ding Hou
EcoNOMIC IMPORTANCE. The leaves and twigs of 1962).
Catha edulis ("kat" or "khat") are chewed or drunk
as a tea for their stimulant properties (Krikorian CoNSERVATION. Although no species of Celas-
and Getahun 1973), similar to amphetamine traceae is listed in CITES (Convention on Inter-
(Zeiger et al. 1981). The two active components of national Trade in Endangered Species), several
kat, L(S)-(- )-alpha-aminopropiophenone (cathi- species are considered threatened. In Australia,
Celastraceae 37
Apatophyllum constablei is endangered and Apato- arils basal to partially enveloping seeds. Tropics and
phyllum olsenii, Denhamia parvifolia, Hexaspora subtropics 88
7. Plants with thorns or stems terminating in sharp points
pubescens, and Stackhousia annua are vulnerable 8
(Briggs and Leigh 1995). In China, Bhesa sinica is - Plants unarmed 13
known from only a single individual (Sheng-ye 8. Thorns 0; stems terminating in sharp points, glandular.
1992 ). In India, Euonymus angulatus, E. assamicus, SW North America 9
E. serratifolius, Salacia jenkinsii, and Salacia mal- - Thorns +; stems not terminating in sharp points, not
glandular. Old World, South America 10
abarica are endangered (Nayar and Sastry 1987). 9. Fruit drupaceous; leaves laminar, few
In South Africa, Empleuridium juniperinum, 1. Acanthothamnus
Gymnosporia bachmannii, Lydenburgia abbottii, - Fruit capsular, beaked; leaves reduced to scales
Maytenus abbottii, M. oleosa, and Pseudosalacia 8. Canotia
10. Leaves not borne on thorns; inflorescence fasciculate.
streyi are vulnerable (Hilton-Taylor 1996; Oldfield
South America 39. Moya
et al. 1998; R. Archer, pers. comm. 2000). Other - Leaves often borne on thorns; inflorescence generally
endangered and vulnerable species include Bhesa cymose. Old World 11
ceylanica, B. nitidissima, Gyminda orbicularis, 11. Ovules 2 per locule; flowers unisexual rarely bisexual. Old
Euonymus lanceifolia, E. morrisonensis, E. pallidi- World 24. Gymnosporia
- Ovules 3-12 per locule; flowers bisexual. South Africa, S
folius, E. paniculatus, E. serratifolius, E. thwaitesii, Mozambique 12
E. walkeri, Gymnosporia curtisii, Kokoona cori- 12. Nodes > 1 per thorn; ovules 3-6 per locule 20. Gloveria
acea, K. leucoclada, K. sabahana and K. sessilis, - Nodes 1 per thorn; ovules (4-)6(-12) per locule
Lophopetalum sessilifolium, Maytenus cymosa, M. 53. Putterlickia
harenensis, M. harrisii, M. jefeana, M. matudai, 13. Flowers 3-4-merous 14
- Flowers 5-merous 37
M. microcarpa, M. ponceana, M. stipitata, M. 14. Ovary 4-locular and ovules 1 or± 10 per locule 15
williamsii, Microtropis argentea, M. borneensis, - Ovary 1-5-locular and ovules 1-12 per locule 20
M. densiflora, M. fascicularis, M. grandifolia, M. 15. Ovules ± 10 per locule; fruit capsular, woody, oblong.
keningauensis, M. rigida, M. sabahensis, M. New Guinea 67. Xylonymus
sarawakensis, M. tenuis, Peritassa killipii, Perrotte- - Ovules 1 per locule; fruit drupaceous or capsular, fleshy
or coriaceous, spheroid, obovoid, or deeply lobed. Asia,
tia excelsa, Salacia miegei, Wimmeria acuminata, Macronesia, New World 16
and W. chiapensis (Oldfield et al. 1998). 16. Stamens on disk. Asia, Macronesia 17
- Disk intrastaminal. West Indies, tropical America 18
KEY TO THE GENERA 17. Ovules pendulous; capsule spheroid; seeds with branched
raphe. Asia, Macronesia 21. Glyptopetalum
1. Annuals or perennials with woody rootstocks; petals par- - Ovules erect; capsule deeply lobed; seeds with
tially connate or free; stamens 5, of unequal or equal unbranched raphe. China 37. Monimopetalum
length. Australasia 2 18. Ovules pendulous; leaves strictly opposite, entire. Jamaica
- Trees, shrubs, or Hanas; petals free; stamens 2-numerous, 62. Tetrasiphon
of equal length. Old and New World, Australia 4 - Ovules erect; leaves alternate, opposite, or whorled, entire
2. Petals free; stamens of equal length. Australia or toothed. West Indies, tropical America 19
69. Macgregoria 19. Fruit drupaceous, obovoid; seeds with branched raphe.
- Petals partially connate; stamens of unequal length. Aus- West Indies, tropical America 12. Crossopetalum
tralasia 3 - Fruit capsular, deeply lobed; seeds with unbranched
3. Style attached between cocci in fruit. Australasia raphe. Cuba, Hispaniola, Puerto Rico 63. Torralbasia
70. Stackhousia 20. Leaves alternate or irregularly scattered 21
- Style attached terminally in fruit. SW Australia - Leaves opposite 27
71. Tripterococcus 21. Leaves needle-like, irregularly scattered; ericoid sub-
4. Stamens numerous. China, Myanmar 83. Plagiopteron shrub. South Africa 16. Empleuridium
- Functional stamens 2-5. Old and New World, Australia - Leaves with laminar blade, alternate; trees or shrubs. Old
5 and New World, Australia 22
5. Functional stamens 2; disk intrastaminal. Andaman- 22. Leaves generally with small abaxial domatia in axils of
Nicobar Islands, India 97.Nicobariodendron larger veins; fruit baccate; seeds furrowed
- Functional stamens 3-5, or if 2, disk extrastaminal 6 44. Perrottetia
6. Disk intrastaminal or stamens on disk; flowers unisexual - Leaves without domatia; fruit samaroid, capsular or
or bisexual; fruit drupaceous, baccate, samaroid, or cap- drupaceous; seeds smooth 23
sular; seeds with or without surrounding wings, arillate 23. Disk indistinct or 0; flowers unisexual, rarely dioecious;
or exarillate. Tropics, subtropics, and temperate 7 fruit drupaceous. Americas, West Indies 60. Schaefferia
- Disk extrastaminal; flowers bisexual (unisexual in New - Disk fleshy; flowers bisexual or unisexual; fruit capsular,
Caledonia); fruit baccate with seeds surrounded by samaroid, or drupaceous. Old and New World 24
mucilaginous arils, transversely-flattened, trilobed cap- 24. Fruit samaroid; ovules 4-9 per locule. C America,
sules with seeds with membranous basal wings or narrow Mexico 66. Wimmeria
stipes, or capsules that are circular or trigonous in cross - Fruit capsular or drupaceous; ovules 1-2(-12) per locule.
section with seeds with membranous apical wings or Old and New World, Australia 25
38 M.P. Simmons
25. Fruit drupaceous 15. Elaeodendron 46. Fruit drupaceous. Old and New World, Australia 47
- Fruit capsular 26 - Fruit baccate. Africa, Madagascar 48
26. Capsule with lateral appendages or unornamented; 47. Drupes dry, trigonous; seeds 2-3. Madagascar
ovules erect and/or pendulous, 2( -12) per locule 25. Hartogiopsis
18. Euonymus - Drupes fleshy or coriaceous, not trigonous; seeds 1-2(3).
- Capsule unornamented; ovules erect, 1-2 per locule Old and New World, Australia 15. Elaeodendron
34. Maytenus 48. Ovules pendulous. S Africa 33. Maurocenia
27. Low, rhizomatous shrubs. North America 42. Paxistima - Ovules erect. Africa, Madagascar 49
- Trees or erect, rarely scandent shrubs. Old and New 49. Seeds with branched raphes, arillate. Madagascar
World, Australia 25 7. Brexiella
28. Ovules 1 per locule, pendant. West Indies, tropical - Seeds with unbranched raphes, exarillate. Africa 50
America 23. Gyminda 50. Flowers pedicellate; inflorescence cymose 9. Cassine
- Ovules 2-12 per locule, erect, rarely pendant. Old and - Flowers sessile; inflorescence thyrsoid or cymose 51
New World, Australia 29 51. Inflorescence thyrsoid; berries spheroid 30. Lauridia
29. Fruit samaroid. Madagascar, Mexico 30 - Inflorescence cymose; berries ellipsoid 2. Allocassine
- Fruit capsular, drupaceous, or nuts. Old and New World, 52. Capsule 1-locular, laterally split along 1 side; seed 1. SE
Australia 31 Asia, Macronesia, C America, Mexico 53
30. Samara with 1 surrounding wing. Madagascar - Capsule 3-5-locular, loculicidally dehiscent; seeds
51. Ptelidium 1-several. Old and New World, Australia 54
- Samara with 1 apical wing. Mexico 56. Rzedowskia 53. Disk fleshy. C America, Mexico 54. Quetzalia
31. Fruit capsular 32 - Disk 0 or thin. SE Asia, Macronesia 36. Microtropis
- Fruit indehiscent 34 54. Capsule woody; seeds with wing surrounding seed;
32. Capsules 3-5-locular, loculicidally dehiscent; seeds stamens on disk; inflorescence thyrsoid. Asia, Macrone-
1-several. Old and New World, Australia sia, Australia 31. Lophopetalum
18. Euonymus - Capsule bony to coriaceous; seeds without wing or wing
- Capsules 1-locular, laterally split along 1 side; seed 1. SE not surrounding seed; disk intrastaminal or stamens on
Asia, Macronesia, C America, Mexico 33 disk; inflorescence cymose or fasciculate. Old and New
33. Disk fleshy. C America, Mexico 54. Quetzalia World, Australia 55
- Disk 0 or thin. SE Asia, Macronesia 36. Microtropis 55. Flowers unisexual (dioecious); disk intrastaminal;
34. Fruit nuts with lateral styles. Old World, Australia capsule without wings or protuberances. New Caledonia
47. Pleurostylia 35. Menepetalum
- Fruit drupaceous or baccate with styles terminal or 0. Old - Flowers bisexual, rarely unisexual; monoecious or dioe-
and New World, Australia 35 cious; disk intrastaminal or stamens on disk; capsules
35. Fruit drupaceous. Old and New World, Australia with or without wings or protuberances. Old and New
15. Elaeodendron World, Australia 56
- Fruit baccate. South Africa 36 56. Leaves strictly opposite; ovary 3-5-locular; capsules with
36. Flowers sessile; inflorescence compact thyrsoid or or without wings or protuberances; seeds without wings;
irregularly cymose 30. Lauridia disk intrastaminal or stamens on disk. Old and New
- Flowers pedicellate; inflorescence cymose 9. Cassine World, Australia 18. Euonymus
37. Leaves opposite or whorled (and subopposite) on mature - Leaves opposite on mature stems, alternate on juvenile
stems 38 stems or strictly opposite; ovary 3(4)-locular; capsules
- Leaves alternate (and subverticillate in New Caledonia) without wings or protuberances; seeds with or without
on mature stems 58 wings; disk intrastaminal. Africa 57
38. Leaves reduced to scales. Australia 49. Psammomoya 57. Seeds without wings; leaves strictly opposite or opposite
- Leaves laminar. Old and New World, Australia 39 on mature branches and alternate on juvenile branches.
39. Ovary 5-locular. Madagascar 40 South Africa 32. Lydenburgia
- Ovary 2-5-locular. Old and New World, Australia 42 - Seeds with basal wings; leaves opposite on mature
40. Fruit capsular, coriaceous 18. Euonymus branches and alternate on juvenile branches. Africa,
- Fruit baccate,± fleshy 41 Arabia 10. Catha
41. Disk intrastaminal; seeds 1-2, exarillate 58. Leaves needle-like, sessile. Australia 3. Apatophyllum
58. Salvadoropsis - Leaves laminar, petiolate. Old and New World, Australia
- Stamens on disk; seeds 5-l 0, arillate 17. Euonymopsis 59
42. Fruit indehiscent 43 59. Leaves with distinct crossbar secondary venation, petiole
- Fruit dehiscent 52 distally geniculate. Asia, Macronesia 4. Bhesa
43. Fruit samaroid. Madagascar, Mexico, C America, South - Leaves with reticulate secondary venation, petiole not
America 44 geniculate. Old and New World, Australia 60
- Fruits baccate, drupaceous, or nuts. Old and New World, 60. Leaves generally with small abaxial domatia in axils of
Australia 45 larger veins; ovary 2-locular, seemingly 4-locular at base;
44. Samara with 1 apical wing, oblanceolate or obovate. seeds furrowed 44. Perrottetia
Mexico, C America, South America 68. Zinowiewia - Leaves without domatia; ovary locule number not appar-
- Samara with 1 surrounding wing, ovate or ovate- ently different at apex and base; seeds smooth 61
lanceolate. Madagascar 51. Ptelidium 61. Leaves conspicuously glandular-serrate and small (<10
45. Fruits as nuts with lateral styles. Old World, Australia mm long). Mexico 41. Orthosphenia
47. Pleurostylia - Leaves without glandular-toothed margins, or if glandu-
- Fruits baccate or drupaceous, with styles terminal or 0. lar-toothed, large (>15mm long). Old and New World,
Old and New World 46 Australia 62
Celastraceae 39
62. Leaf venation acrodromous; ovary !-locular, 3-carpellate, 73. Fruit drupaceous or samaroid; ovules 1-2 per locule 74
with intruded parietal placentation; capsule septicidally - Fruit capsular; ovules 1-12 per locule 79
dehiscent. Myanmar, Thailand 98. Pottingeria 74. Fruit samaroid 75
- Leaf venation pinnate; ovary 2-7-locular, with axile pla- - Fruit drupaceous 76
centation; fruit indehiscent or capsular, if capsular, loculi- 75. Samara with 1 apical wing; erect trees or shrubs. South
cidally dehiscent. Old and New World, Australia 63 America 46. Plenckia
63. Plants pubescent; inflorescence racemose; fruit linear- - Samara with 3 lateral wings; scandent shrubs. Asia
oblong; seed 1. Brazil 19. Fraunhofera 65. Tripterygium
- Plants glabrous or pubescent; inflorescence thyrsoid, 76. Disk intrastaminal; ovules 2 per locule; drupe fibrous;
cymose, fasciculate, or flowers solitary; fruit spheroid to leaves entire. Brazil 22. Goniodiscus
ovoid; seeds !-numerous. Old and New World, Australia - Stamens on disk; ovules 1 per locule; drupe fleshy or
64 coriaceous; leaves entire or toothed. Old and New World,
64. Inflorescence epiphyllous, rarely axillary; ovary (4)5- Australia 77
locular. Madagascar 48. Polycardia 77. Inflorescence fasciculate; ovary 2-locular; plant glabrous.
- Inflorescence axillary and/ or terminal; ovary 2-10- South Africa 55. Robsonodendron
locular. Old and New World, Australia 65 - Inflorescence cymose or cymose-umbellate; ovary 2-4-
65. Ovary with apical hollow and style-like central column locular; plant pubescent or glabrous. Old and New World,
arising from base of hollow; stamens alternating with Australia 78
small staminodes; drupe with numerous stones. Asia, 78. Inflorescence cymose-umbellate; ovary 3-4-locular; plant
Macronesia, Australia 61. Siphonodon pubescent or glabrous. Africa, Madagascar
- Ovary with terminal style; stamens not alternating with 40. Mystroxylon
staminodes (although disk may be lacerate); fruit dehis- - Inflorescence cymose; ovary 2-4-locular; plant glabrous.
cent or indehiscent, not drupe with numerous stones. Old Old and New World, Australia 15. Elaeodendron
and New World, Australia 66 79. Capsule flattened along each locule, without wings or pro-
66. Disk lacerate, alternating with stamens; fruit baccate, tuberances; seeds discoid with annular wing. New
woody, becoming pulpy; inflorescence umbellate- Caledonia 43. Peripterygia
cymose; ovary 5-7-locular. Africa, Madagascar - Capsule not flattened along each locule, with or without
6.Brexia wings or protuberances; seeds not winged. Old and New
- Disk entire or lobed; fruit dehiscent or indehiscent, char- World, Australia 80
taceous to fleshy; inflorescence thyrsoid, cymose, fascic- 80. Stamens on disk 81
ulate, or flowers solitary; ovary I-S-locular. Old and New - Disk intrastaminal 82
World, Australia 67 81. Ovary 3-locular; ovules erect, 2 per locule; capsule
67. Ovary 5-locular 68 without wings or protuberances; leaves entire. South
- Ovary 2-4-locular 72 Africa 50. Pseudosalacia
68. Inflorescence terminal, thyrsoid; fruits drupaceous, dry. - Ovary 3-4-locular; ovules pendulous to erect, 2-12 per
U.S.A., Mexico 38. Mortonia locule; capsule with or without lateral wings or protu-
- Inflorescence axillary or terminal, cymose, fasciculate, or berances; leaves toothed or entire. Old and New World,
condensed thyrsoid; fruits capsular or samaroid, charta- Australia 18. Euonymus
ceous to bony. Old and New World, Australia 69 82. Leaves alternate and subverticillate; inflorescence
69. Fruit samaroid with 4-Slateral wings; ovules 2 per locule. fasciculate; ovary 3-locular; ovules 2-6 per locule. New
Tanzania 45. Platypterocarpus Caledonia 57. Salaciopsis
- Fruit capsular with or without lateral wings; ovules 2-12 - Leaves strictly alternate; inflorescence thyrsoid, cymose,
per locule. Old and New World, Australia 70 fasciculate, or flowers solitary; ovary 2-4-locular; ovules
70. Inflorescence fasciculate or condensed thyrsoid; anthers 1-12 per locule. Old and New World, Australia 83
obliquely dehiscent, latrorse; disk intrastaminal; leaves 83. Capsule with lateral or oblique horn to wing-like out-
entire; ovules 4 per locule; aril at base and one side of growths; arils nearly enveloping seeds; inflorescence
seed. E Australia 26. Hedraianthera cymose; ovules 2 per locule. SE Africa
- Inflorescence cymose (fasciculate); anthers longitudinally 52. Pterocelastrus
or obliquely dehiscent, introrse or latrorse; stamens on - Capsule with or without lateral wings or protuberances;
disk or disk intrastaminal; leaves entire or toothed; ovules arils basal to enveloping seeds; inflorescence thyrsoid,
2-12 per locule; aril basal to enveloping seed. Old and cymose, fasciculate, or flowers solitary; ovules 1-12 per
New World, Australia 71 locule. Old and New World, Australia 84
71. Inflorescence terminal or axillary; disk intrastaminal; 84. Ovules 1 per locule; plant puberulent to glabrescent;
anthers longitudinally dehiscent, introrse; capsule inflorescence axillary; arils enveloping seeds. Americas
without wings or protuberances. Australia 64. Tricerma
13. Denhamia - Ovules 1-12 per locule; plant glabrous to pubescent;
- Inflorescence axillary; stamens on disk or disk intrasta- inflorescence terminal or axillary; arils basal to envelop-
minal; anthers obliquely or longitudinally dehiscent, ing seeds. Old and New World, Australia 85
latrorse or introrse; capsule with or without lateral wings 85. Scandent shrubs; inflorescence terminal or axillary,
or protuberances. Old and New World, Australia thyrsoid, cymose, or flowers solitary; flowers unisexual,
18. Euonymus rarely bisexual, dioecious; ovules 1-2 per locule; arils
72. Plant densely pilose; anthers obliquely dehiscent; ovary nearly enveloping seeds 11. Celastrus
3-locular; erect trees. Queensland, Australia - Erect or scandent trees or shrubs; inflorescence axillary,
27. Hexaspora rarely terminal, cymose, fasciculate, or flowers solitary;
- Plant glabrous or pubescent; anthers longitudinally or flowers bisexual, rarely unisexual; monoecious or dioe-
obliquely dehiscent; ovary 2-4-locular; scandent or erect cious; ovules 1-12 per locule; arils basal to enveloping
shrubs or trees. Old and New World, Australia 73 seeds 86
40 M.P. Simmons
86. Stamens on disk or disk intrastaminal; capsule with or 103. Petals thickened, fleshy 75. Bequaertia
without lateral wings or protuberances; ovary 3-4- - Petals not thickened or fleshy 90. Tristemonanthus
locular; ovules pendulous to erect 18. Euonymus 104. Ovules 2 per locule 105
- Disk intrastaminal; capsule without lateral wings or pro- - Ovules 3-22 per locule 109
tuberances; ovary 2-4-locular; ovules erect 87 105. Androgynophore +.Tropical Africa 88. Simicratea
87. Inflorescence cymose; ovules 2-10 per locule. Australia - Androgynophore 0. Old and New World, Australia 106
13. Denhamia 106. Disk of 3 discontinuous lobes around stamens; 4 of 5
- Inflorescence cymose, fasciculate, or flowers solitary; petals arched, ± zygomorphic. Africa, Madagascar
ovules 1-2 per locule. Old and New World, Australia 73. Apodostigma
34. Maytenus - Disk continuous; flowers actinomorphic. Old and New
88. Fruit baccate with seeds surrounded by mucilaginous World, Australia 107
arils 89 107. Nodes on which inflorescence borne without axillary
- Fruit transversely-flattened, trilobed capsules with seeds branches 85. Pristimera
with membranous basal wings or narrow stipes, or - Nodes on which inflorescence borne with axillary
capsules that are circular or trigonous in cross section branches 108
96 108. Disk pulvinate or short cupular; seeds with membranous
89. New World 90 wings. Old World 86. Reissantia
- Old World, Australia 93 - Disk deeply cupular; seeds with narrow stipes or mem-
90. Disk discontinuous 91. Cheiloclinium branous wings. Americas, Africa, Madagascar
- Disk continuous 91 78. Elachyptera
91. Anthers obliquely or longitudinally dehiscent 109. Androgynophore +.Africa 110
92. Peritassa - Androgynophore 0. Old and New World, Australia 112
- Anthers transversely dehiscent 92 110. Plant hirsute and stellate pubescent 79. Helictonema
92. Disk pulvinate, fleshy 93. Salacia - Plant glabrous or puberulent 111
- Disk flattened or cupular, membranous or ± fleshy 111. Plant glabrous 89. Simirestis
96. Tontelea - Plant puberulent or glabrescent 82. Loeseneriella
93. Anthers obliquely or longitudinally dehiscent 94 112. Petals regularly serrate; capsule lobes entirely connate. C
- Anthers transversely dehiscent 95 and South America 72. Anthodon
94. Branches opposite; ovules 2 per locule; berry fusiform. - Petals entire, erosulous, or fimbriate; capsule lobes not
Tropical Africa 95. Thyrosalacia connate or connate ± half length. Old and New World,
- Branches alternate; ovules 2-9 per locule; berry spheroid, Australia 113
oblong, or fusiform. Old World, Australia 93. Salacia 113. Capsule lobes connate ± half length; inflorescence
95. Inflorescence cauliflorous on long pendant branches; thyrsoid; ovules 6-8 per locule. Mexico, C and South
leaves subopposite or alternate; androgynophore 0; berry America 87. Semialarium
spheroid; ovules 2 per locule. Tropical Africa - Capsule lobes not connate; inflorescence cymose or
94. Salacighia thyrsoid; ovules 4-22 per locule. Old and New World,
- Inflorescence axillary or cauliflorous, thyrsoid, cymose, or Australia 114
fasciculate; leaves opposite or subopposite, rarely alter- 114. Capsule woody; seeds with narrow stipes or rarely mem-
nate, androgynophore + or 0; berry spheroid, oblong, or branous wings. C and South America 81. Hylenaea
fusiform; ovules 2-9 per locule. Old World, Australia - Capsule coriaceous or chartaceous; seeds with membra-
93. Salacia nous wings or rarely narrow stipes). Old and New World,
96. Capsule circular or trigonous in cross section; seeds with Australia 115
membranous apical wings or arils basal to partially 115. Disk lobed and indistinct; ovary papillose; ovules 4-7 per
enveloping seeds. Asia, Macronesia, Australia, New locule. India, SE Asia, Macronesia 74. Arnicratea
Caledonia 97 - Disk entire or lobed, distinct, rarely indistinct; ovary
- Capsule transversely-flattened, trilobed; seeds with mem- smooth, rarely papillose; ovules 3-22 per locule. Old and
branous basal wings or narrow stipes. Old and New World New World, Australia 116
tropics and subtropics, Australia 101 116. Flowers large (8-20mm diameter) 117
97. Stamens 3; ovary 2-locular. New Caledonia - Flowers small (<8mm diameter) 118
14. Dicarpellum 117. Disk membranous, cupular; seeds with narrow stipes or
- Stamens 5; ovary 3- or 5-locular. SE Asia, Macronesia, membranous wings. Americas, Africa 77. Cuervea
Australia 98 - Disk fleshy, annular-pulvinate; seeds with membranous
98. Ovary 5-locular. New Guinea 5. Brassiantha wings. Old and New World 84. Prionostemma
- Ovary 3-locular. Asia, Macronesia, Australia 99 118. Adaxial surface of petals puberulent; ovules 4-8 per
99. Capsule oblong, trigonous, woody; seeds with apical locule; seeds with membranous wings. Americas, Africa
wing. Asia, Macronesia 29. Kokoona 80. Hippocratea
- Capsule fusiform to narrow-elipsoid, coriaceous or bony, - Adaxial surface of petals glabrous; ovules 3-33 per locule;
seeds without wings. Australia, Borneo 100 seeds with narrow stipes or membranous wings. Old and
100. Inflorescence cymose or flowers solitary; arils without New World, Australia 119
basal filamentous extensions. Australia 28. Hypsophila 119. Petals ± valvate in bud, ovate to lanceolate. Old World,
- Inflorescence racemose; arils with basal filamentous Australia 82. Loeseneriella
extensions. Borneo 59. Sarawakodendron - Petals imbricate in bud, ovate, oblong, or suborbicular.
101. Disk 0; anthers introrse 102 Old and New World 120
- Disk +; anthers extrorse 104 120. Nodes on which inflorescence borne without axillary
102. Stamens 5 76. Campylostemon branches; ovules 3-10 per locule; seeds with membranous
- Stamens 3 103 wings. Old and New World 85. Pristimera
Celastraceae 41
- Nodes on which inflorescence borne with axillary Shrubs, glabrous. Leaves alternate, rarely sub-
branches; ovules 4 per locule; seeds with narrow stipes or opposite or subverticillate, sessile, needle-like,
membranous wings. Americas, Africa, Madagascar
78. Elachyptera
entire. Flowers axillary, solitary. Flowers bisexual,
5-merous; disk annular or margins upturned,
GENERA OF CELASTRACEAE
intrastaminal; anthers longitudinally dehiscent,
introrse; ovary 2(3)-locular; ovules erect, 2 per
I. SUBFAMILY CELASTROIDEAE Burnett (1835). locule. Capsule compressed, obovoid or pyriform,
loculicidally dehiscent, seeds 1(2), oblong or
Erect or scandent trees, shrubs, suffrutices, rhi- ovoid, albuminous, aril basal. Five spp., Australia,
zomatous shrubs, ericoid subshrubs, or epiphytic rocky hillsides and open forests.
shrubs. Flowers unisexual or bisexual, (3)4-5-
merous; petals free; disk intrastaminal or ex- 4. Bhesa Buch.-Ham. ex Arn. Fig. 11
trastaminal or 0, rarely stamens on disk;
Bhesa Buch.-Ham ex Arn., Edinburgh New Philos. J. 16: 315
androgynophore 0; stamens 3-5; ovary (1)2- (1834); Ding Hou, Blumea 4: 149-153 (1958), rev.
5(10)-locular. Capsule, drupe, berry, samara, or Kurrimia Wall. (1831).
nut; seeds albuminous or exalbuminous. Included Kurrimia Wall. ex Meisn. (1837).
phloem 0.
Trees with buttressed trunks, glabrous. Leaves
1. Acanthothamnus Brandegee alternate, with distinct crossbar secondary vena-
tion, entire, petiole geniculate. Inflorescences
Acanthothamnus Brandegee, Univ. Calif. Publ. Bot. 3: 383 axillary, paniculate or racemose. Flowers bisexual,
(1909). 5-merous; disk fleshy, cupular, subentire or
Scandivepres Loes. (1910).
Shrubs, glabrous; stems with dark glands, termi-

nating in sharp points. Leaves few, alternate, entire.
Inflorescences axillary, fasciculate, or flowers soli-
tary. Flowers bisexual, 5-merous; disk cupular,
lobed, intrastaminal; anthers longitudinally dehis-
cent, introrse; ovary 2-locular; ovules erect, 2 per
locule. Drupe ovoid,± fleshy; seed 1, albuminous.
Only one sp., A. aphyllus (Schltdl.) Standi., Mexico,
deserts and subdeserts.
2. Allocassine N. Robson
Allocassine N. Robson, Bol. Soc. Brot. 39: 30 (1965); Archer &
van Wyk, S. Afr. J. Bot. 64: 189-191 (1998), rev.
Scandent shrub, glabrous. Leaves opposite on

flowering branches and alternate on vegetative
branches, entire or ± glandular-crenate. Inflores-
cences axillary, cymose. Flowers bisexual, 5-
merous; disk fleshy, margins upturned, entire,
introrse; ovary 2-locular; ovules erect, 2 per locule.
Berry ellipsoid, fleshy; seeds 2-3( 4), narrowly
ellipsoid, albuminous. Only one sp., A. laurifolia
(Harv.) N. Robson, eastern S Africa and Zimbabwe,
Mozambique, dry forests.
Fig. I I. Celastraceae-Celastroideae. Bhesa archboldiana. A

3. Apatophyllum McGill. Fruiting branchlet. B Flower. C Same, vertical section, pistil
removed. D Pistil showing basal placentation. E Ovary, trans-
Apatophyllum McGill., Kew Bull. 25:401 (1971); Bean & Jessup, verse section. F Seed with aril. Drawn by R. van Crevel.
Austrobaileya 5: 691-697 (2000), rev. (Ding Hou 1962)
42 M.P. Simmons
5-lobed; anthers longitudinally dehiscent, introrse 7. Brexiella H. Perrier

or extrorse; ovary 2-locular, glabrous or apically
Brexiella H. Perrier, Bull. Soc. Bot. France 80: 204 (1933);
pubescent; ovules erect, 2 per locule. Capsule Perrier de Ia Bathie, Fl. Madag. 116: 1-76 (1946).
fusiform or deeply 2-lobed, coriaceous, loculici-
dally dehiscent; seeds 1-2, oblong, albuminous, Trees or shrubs, glabrous. Leaves opposite, rarely
aril basal or enveloping seed. Five spp., India, Sri whorled, entire, serrulate, or spinose-dentate.
Lanka, SE Asia, Macronesia, humid forests, Inflorescences axillary, fasciculate or cymose.
0-2150 m. The genus is properly Bhesa, not Kur- Flowers bisexual, 5-merous; disk fleshy, annular, 5-
rimia because the original generic description of angled or lobed, intrastaminal; anthers longitudi-
Kurrimia by Meisner (1837) is based on a taxo- nally dehiscent, introrse; ovary 2-3-locular; ovules
nomical synonym of Itea (Ding Hou 1958). 2 per locule. Berry spheroid, ± fleshy; seeds 3-4,
subspheroid, with branched raphe, albumino~s,
5. Brassiantha A.C. Sm. enveloped by aril. Two spp., Madagascar, humid
and littoral forests.
Brassiantha A.C. Sm., J. Arnold Arbor. 22: 389 (1941); Ding
Hou, Fl. Males. I, 6: 227-291 (1962).
8. Canotia Torr.
Trees, glabrous. Leaves alternate, entire. Inflores-
Canotia Torr., Pacific Railroad Reports 4: 68 (1857); Johnston,
cences axillary, cymose. Flowers bisexual, Brittonia 27: 119-122 (1975), rev.
5-merous; disk fleshy, pulvinate, lobed, extrasta-
minal; anthers transversally dehiscent, extrorse; Trees or shrubs, glabrous; stems with dark glands,
ovary 5-locular; ovules erect, 2-5 per locule. terminating in sharp points. Leaves alternate,
Capsule subspheroid, coriaceous, locul~cid~lly reduced to scales. Inflorescences axillary, cymose
dehiscent; seeds (1-)2-4(-5) per locule, ellipsOid- or thyrsoid. Flowers bisexual, 5-merous; disk
oblong, albuminous, partially enveloped by aril. fleshy, annular, intrastaminal; anthers longitudi-
Only one sp., B. pentamera A.C. Sm., New Guinea, nally dehiscent, introrse; ovary 5-locular; ovules
lowland forests, to lOOm (1800-1950m). axile, 2-6 per locule. Capsule oblong, beaked,
woody, septicidally and loculicidally dehiscent;
6. Brexia Noronha ex Thouars seeds 1(2) per locule, oblong, flattened, with or
without membranous basal wing, albuminous.
Brexia Noronha ex Thouars, Gen. Nov. Madag. 20 (1806); Two spp., Mexico, SE U.S.A., deserts.
Perrier de Ia Bihhie, Bull. Soc. Bot. France 89: 219-221
(1942); Verdcourt, Fl. Trop. E. Afr. 108A: 1-3 (1968). Canotia has been assigned to Rutaceae (Gray
1877), Koeberliniaceae (Barnhart 1910), Canoti-
Trees or shrubs, glabrous. Leaves alternate, entire aceae (Cronquist 1981), and Celastraceae
or spinose-dentate. Inflorescences axillary, umb~l (Hutchinson 1969) as an anomalous genus (Loe-
late-cymose. Flowers bisexual, 5-merous; disk sener 1942a), or as closely related to Acan-
fleshy, lacerate, alternating with stamens; anthers thothamnus (Johnston 1975). Phylogenetic
longitudinally dehiscent, introrse; ovary . 5-7- analyses support the sister-group relationship of
locular; ovules axile, numerous. Berry, ovo1d or Acanthothamnus and Canotia (Simmons and
oblong-fusiform, woody, becoming pulpy; seeds Hedin 1999; Simmons et al. 200la, 200lb).
numerous, irregularly compressed-ellipsoid, albu-
minous. Only one sp., B. madagascariensis 9. Cassine L.
Thouars ex Ker Gawl., Madagascar, Seychelles,
Cassine L., Sp. Pl. ed. 1: 268 (1753); Archer & van Wyk, S. Afr.
coastal east Africa, coastal bush and swamp
J. Bot. 63: 146-157 (1997), rev.
forests. Eight species of Brexia have been recog- Hartogiella Codd (1983).
nized by Perrier de la Bathie (1942), but these are
tentatively lumped into one species (Verdcourt Trees or shrubs, glabrous. Leaves opposite, rarely
1968). subopposite, entire or glandular-crenate. Inflores-
Brexia has been assigned to Escalloniaceae cences axillary, cymose. Flowers bisexual, rarely
(Hutchinson 1967), Brexiaceae (Verdcourt 1968), unisexual, 4-5-merous; disk fleshy, lobed or entire,
and Grossulariaceae (Cronquist 1981). However, intrastaminal; anthers longitudinally dehiscent,
phylogenetic analyses consistently support Brexia introrse, rarely extrorse; ovary 2-3-locular; ovules
as a member of Celastraceae (e.g., Savolainen et al. erect, 2 per locule. Berry spheroid, fleshy or coria-
1997; Simmons et al. 2001b). ceous; seeds 1-2( -6), spheroid or ellipsoid, albu-
Celastraceae 43
minous. Three spp., S Africa, forests, woodlands, Capsule narrowly oblong, ± bony, loculicidally
fynbos, and bush. dehiscent; seeds 1-3, with membranous, basal
Cassine is narrowly defined following Archer wing, albuminous. Only one sp., C. edulis (Vahl)
and van Wyk (1997). Cassine is distinct from Forssk. ex Endl., Africa, Arabia, evergreen
Elaeodendron, based on morphology and anatomy medium-altitude forest. Catha abbottii A.E. van
(Archer and van Wyk 1992, 1993a, 1993b, 1997), Wyk & M. Prins will be transferred to Lydenbur-
and a cladistic analysis (Simmons et al. 2001b). gia (van Wyk, in press).
Catha edulis ("kat" or "khat") is intensely
cultivated as a cash crop in Ethiopia (Getahun
10. Catha Forssk. ex Scop. Fig. 12
and Krikorian 1973). Two alkaloids, L(S)-(-)-
Catha Forssk. ex Scop., Intr. Hist. Nat. 228 (1777). alpha-aminopropiophenone (cathinone) and d-
norpseudoephedrine (cathine), concentrated in
Trees or shrubs, glabrous. Leaves opposite on the young leaves and twigs of kat act as stimulants
mature branches and alternate on juvenile of the central nervous system (Krikorian and
branches, glandular-crenulate-denticulate. Inflo- Getahun 1973; Szendrei 1981; Zeiger et al. 1981).
rescences axillary, cymose. Flowers bisexual, 5- The leaves and twigs of kat are chewed or drunk
merous; disk thin, margins upturned, 5-lobed, as a tea in northeastern Africa, the Arabian
intrastaminal; anthers longitudinally dehiscent, Peninsula, and Madagascar.
11. Celastrus L. Fig.13A-D
Celastrus L., Sp. Pl. ed. 1: 196 (1753); Ding Hou, Ann. Missouri
Bot. Gard. 42: 215-302 (1955), rev.
Scandent shrubs, glabrous, rarely pubescent.

Leaves alternate, subentire or serrate. Inflores-
cences axillary or terminal, thyrsoid, cymose, or
flowers solitary. Flowers rarely bisexual, unisexual,
rarely dioecious, 5-merous; disk membranous
or fleshy, annular to cupular, entire or 5-lobed,
introrse; ovary 3-locular; ovules erect, 1-2 per
locule. Capsule subspheroid, rarely oblong, coria-
ceous, loculicidally dehiscent; seeds 1-6, ellipsoid,
albuminous, nearly enveloped by aril. 2n = 46.
Thirty-one spp., Americas, Madagascar, India, SE
Asia, Macronesia, Queensland, Australia, humid
forests to dry thickets. Two species, C. orbiculatus
Thunb. and C. scandens L., are widely cultivated as
ornamentals for the winter color of their fruits and
arillate seeds (Ding Hou 1955).
12. Crossopetalum P. Browne

Crossopetalum P. Browne, Civ. Nat. Hist. Jamaica ed. 1: 145
(1756); Brizicky, J. Arnold Arbor. 45: 206-234 (1964), part.
rev.
Rhacoma L. (1759).
Myginda Jacq. (1 760).
Trees or shrubs, glabrous. Leaves alternate, oppo-

site, or whorled, entire, serrulate or spinose-
Fig. 12. Celastraceae-Celastroideae. Catha edulis. A Flowering
dentate. Inflorescences axillary, cymose. Flowers
branchlet. B Flower. C Same, vertical section. D Ovary, trans- bisexual, 4-merous; disk annular to cupular, 4-
verse section. E Infructescence. F Dehiscing fruit. G Seed. lobed, intrastaminal; anthers longitudinally dehis-
(Robson et al. 1994) cent, introrse; ovary 4-locular; ovules erect, 1 per
44 M.P. Simmons
Fig. 13. Celastraceae-Celastroideae. A-D Celastrus scandens. Flowers unisexual, rarely dioecious, (2-)5-merous;
A Fruiting branch. B Female flower. C Male flower. D Capsules. disk fleshy, cupular, entire, extrastaminal; stamens
E-G Euonymus americana. E Flowering branch. F Flower.
G Capsule. (Takhtajan 1981) 3; anthers obliquely or longitudinally dehiscent,
Capsule obovoid or ellipsoid, bony, loculicidally
dehiscent; seeds 1-2 per locule, ellipsoid, albu-
locule. Drupe obovoid, ± fleshy; seeds 1(2), minous, aril basal. Four spp., New Caledonia,
obovoid, with branched raphe, albuminous. About humid forests and infrequently in maquis, 75-
26 spp., West Indies, tropical America, coastal sand 1250m.
dunes and scrub, hummocks, and pinelands. Dicarpellum is recognized as distinct from
Salacia (Smith 1941; Simmons, in press).
13. Denhamia Meisn.
Denhamia Meisn., Pl. Vase. Gen.18 (1837); Jessup, Fl. Austr. 22:
15. Elaeodendron Jacq. Fig. 14
150-180 (1984). Elaeodendron Jacq., Icon. Pl. Rar. t. 48 (1782); Archer & van
Wyk, S. Afr. J. Bot. 64: 93-109 (1998), reg. rev.; Kostermans,
Trees or shrubs, glabrous. Leaves alternate, entire Gard. Bull. Sing. 39: 177-191 (1986), reg. rev.
or serrate. Inflorescences axillary or terminal, Crocoxylon Eckl. & Zeyh. (1834/5).
Telemachia Urb. (1916).
cymose. Flowers bisexual, 5-merous; disk fleshy,
annular, 5-lobed, intrastaminal; anthers longitudi-
Shrubs or trees, glabrous. Leaves opposite, sub-
nally dehiscent, introrse; ovary 2-5-locular; ovules
opposite, or alternate, entire, crenate, dentate, or
erect, 2-10 per locule. Capsule ovoid or spheroid,
spinose. Inflorescences axillary, cymose. Flowers
bony, loculicidally dehiscent; seeds 1-several,
bisexual or unisexual (dioecious, polygamous, or
ellipsoid or ovoid, albuminous, aril basal to
monoecious), 3-5-merous; disk fleshy, annular,
enveloping seed. Seven spp., eastern and northern
entire, 4-5-angled, or lobed, intrastaminal or
Australia, humid forests to dry thickets.
stamens on disk; anthers longitudinally dehiscent,
introrse or extrorse; ovary 2-4-locular; ovules
14. Dicarpellum (Loes.) A.C. Sm. erect, 2 per locule. Drupe spheroid or ellipsoid,
fleshy or coriaceous; seeds 1-2(-3), ellipsoid or
Dicarpellum (Loes.) A.C. Sm., Am. J. Bot. 28: 443 (1941);
Simmons, Fl. Nouv.-Caled. (in press). ovoid, flattened or triangular, albuminous. 2n =34.
Salada L. subgenus Dicarpellum Loes. (1907). About 40 spp. (R. Archer, pers. comm. 2000), West
Indies, Africa, Madagascar, India, Macronesia,
Trees or shrubs, glabrous. Leaves alternate, entire. Australia, forests, woodlands, scrub.
Inflorescences axillary, condensed thyrsoid.
Celastraceae 45
sions. Only one sp., E. juniperinum Sond. & Harv.,

S Africa, rocky sandstone slopes, above 600 m.
When first described, Empleuridium was
doubtfully assigned to Rutaceae (Sonder 1860).
However, based on its embryology, morphology,
and pollen, the genus has been transferred to
Celastraceae (Goldblatt et al. 1985).
17. Euonymopsis H. Perrier

Evonymopsis H. Perrier, Notul. Syst. (Paris) 10: 202 (1942).
Trees or shrubs, glabrous. Leaves opposite or

whorled, entire, serrulate, spinose-dentate, or
notched. Inflorescences axillary, rarely cauliflor-
ous, fasciculate or cymose. Flowers bisexual, 5-
merous; disk fleshy, annular, 5-lobed, stamens
inserted on disk; anthers longitudinally dehiscent,
introrse; ovary 5-locular; ovules axile, 4-12 per
locule. Berry ovoid, ellipsoid, or oblong, fleshy;
seeds 5-10, spheroid or ovoid, albuminous,
enveloped by aril. Four spp., Madagascar,
tropophilous forest and xerophytic scrub, to
400m.
18. Euonymus L. Fig.13E-G

Euonymus L., Sp. Pl.: 197 (1753); Ma, Thaiszia 11: 1-264 (2001),
rev.
Quadripterygium Tardieu (1948).
Fig. 14. Celastraceae-Celastroideae, Elaeodendron buch-
ananii. A Flowering branchlet. 8 Male flower. C Female flower.
D Fruiting branch. Drawn by Jill Lowett. (Robson et al. 1994) Trees or shrubs, rarekly scandent, glabrous, rarely
pubescent. Leaves opposite, rarely alternate or
whorled, entire, serrulate, or spinose-denticulate.
Inflorescences axillary, cymose, rarely fasciculate.
Many species of Elaeodendron have been Flowers bisexual, rarely unisexual and dioecious
described as Cassine. Although most species have or gynodioecious, 4-5-merous; disk fleshy,
been described as both Elaeodendron and Cassine, annular, 4-5-lobed, intrastaminal or stamens on
some Austral-Asian species have only been disk; anthers longitudinally or obliquely dehis-
described as Cassine, and new combinations are cent, introrse or latrorse; ovary 3- 5-locular; ovules
required for them (R. Archer, pers. comm. 2000). erect to pendulous, 2( -12) per locule. Capsule sub-
spheroid or obovoid, smooth or echinate, round,
16. Empleuridium Sond. & Harv. angular, deeply lobed, or laterally winged, coria-
ceous, loculicidally dehiscent; seeds 1-several,
Empleuridium Sond. & Harv. in Harvey, Thes. cap. 1: 4 9 (1859); ellipsoid, albuminous, aril basal to enveloping
Goldblatt et a!., Ann. Missouri Bot. Gard. 72: 167-183 (1985),
rev.
seed. 2n = 16, 32, 48, 64. About 129 species, Old
and New World, Australia, humid forests to mesic
Ericoid subshrub, glabrous. Leaves irregularly thickets.
scattered, sessile, needle-like, entire. Flowers axil- Several species (especially E. alatus [Thunb.]
lary, solitary. Flowers bisexual, 4-merous; disk Siebold, E. europaeus L., and E. fortunei [Turcz.]
annular, 4-angled or lobed, intrastaminal; anthers Hand.-Mazz.) are cultivated as ornamentals for
longitudinally dehiscent, introrse; ovary 1-locular; the winter color of their fruits and arillate seeds
ovules erect, 4. Capsule linear-oblong, laterally and/or autumn color, as groundcovers or hedges.
split along one side, seed 1, oblong-fusiform, albu- This large, variable genus seems unlikely to be a
minous, aril basal with apical, filamentous exten- monophyletic group, and some genera, such as
46 M.P. Simmons
Glyptopetalum, Monimopetalum, and Torralbasia, merous; disk fleshy, 5-angled, stamens on disk;
may be nested within it. anthers longitudinally dehiscent, introrse; ovary
3-4-locular; ovules erect, 1 per locule. Drupe
obovoid or ovoid, fibrous; seed 1, oblong, exalbu-
19. Fraunhofera Mart.
minous. Only one sp., G. elaeospermus Kuhlm.,
Fraunhofera Mart., Nov. Gen. Sp. Pl. 3: 85, t. 235 (1831). Brazil, Amazon forest.
Trees or shrubs, pubescent. Leaves alternate,

23. Gyminda Sarg.
serrate. Inflorescences axillary and terminal, race-
mose. Flowers bisexual, 5-merous; disk thin, Gyminda Sarg., Gard. & Forest 4: 4 (1891); Brizicky, J. Arnold
cupular, 5-lobed, intrastaminal; anthers longitudi- Arbor. 45: 206-234 (1964), part. rev.
nally dehiscent, introrse; ovary 2-locular; ovules
erect, 2 per locule. Drupe? (only immature fruit Trees or shrubs, glabrous. Leaves opposite, entire
known) coriaceous, linear-oblong; seeds 1, linear- or crenate. Inflorescences axillary, cymose. Flowers
oblong. Only one sp., F. multiflora Mart., Brazil, unisexual (dioecious), 4-merous; disk fleshy,
deciduous forests. annular, 4-angled or lobed, intrastaminal; anthers
longitudinally dehiscent, introrse; ovary 2(3)-
locular; ovules pendulous, 1 per locule. Drupe
20. Gloveria M. Jordaan obovoid or ellipsoid, ± fleshy; seeds 1-2(3),
Gloveria M. Jordaan, S. Afr. J. Bot. 64: 299 (1998), rev. oblong-ellipsoid, albuminous. Four spp., West
Indies, tropical America, hummocks, rocky thick-
Shrubs with thorns, glabrous. Leaves alternate and ets, and woodlands.
fasciculate on short shoots, entire. Inflorescences
axillary, cymose. Flowers bisexual, 5-merous; disk 24. Gymnosporia (Wight & Arn.) Hook. f.
fleshy, annular, undulate, intrastaminal; anthers
longitudinally dehiscent; ovary 3-locular; ovules Gymnosporia (Wight & Arn.) Hook f. in Bentham & Hooker,
erect, 3-6 per locule. Capsule obovoid-trigonous, Gen. Pl. 1: 365 (1862); Sebsebe, Symb. Bot. Upsal. 25: 1-98
(1985); Jordaan and van Wyk, S. Afr. J. Bot. 65: 177-181
chartaceous, loculicidally dehiscent; seeds ellip- (1999), part. rev.
soid, enveloped by aril. Only one sp., G. integrifo- Celastrus L. sect. Gymnosporia Wight & Arn. (1834).
lia (L.f.) M. Jordaan, S Africa, karroid broken veld.
Trees, shrubs, or suffrutices with thorns, glabrous
21. Glyptopetalum Thwaites or pubescent. Leaves alternate (opposite if on
thorns) and fasciculate on short shoots, entire or
Glyptopetalum Thwaites, Hooker's J. Bot. Kew Gard. Misc. 8: serrulate. Inflorescences axillary, cymose, fascicu-
267, t. 7B (1856); Ding Hou, Fl. Males. I, 6: 227-291 (1962), late, or flowers solitary. Flowers rarely bisexual,
reg. rev.
unisexual (dioecious, polygamous, or monoe-
cious), 5(6)-merous; disk fleshy, annular, 5-lobed,
Trees or shrubs, glabrous. Leaves opposite, rarely
subopposite, entire or crenulate. Inflorescences introrse; ovary (2)3(4)-locular; ovules erect, 2(3)
axillary, cymose. Flowers bisexual, 4-merous; disk
per locule. Capsule ovoid, spheroid, or obovoid,
fleshy, annular, 4-angled or lobed, stamens on disk;
chartaceous, coriaceous, or woody, loculicidally
anthers obliquely dehiscent, latrorse; ovary 4-
dehiscent; seeds 1-4, ellipsoid, albuminous, aril
locular; ovules pendulous, 1 per locule. Capsule
basal to enveloping seed. 2n = 20, 24, 36, 56. About
spheroid, coriaceous, loculicidally dehiscent; seeds
80 sp., Africa, Madagascar, India, Sri Lanka, SE
1-4, oblong, pendulous, with branched raphe, aril
Asia, Macronesia, Queensland, Australia, forests,
basal. About 20 spp., India, Sri Lanka, SE Asia,
woodlands, thickets, scrub, and grasslands.
Macronesia, forests and thickets, 0-1400 m.
Gymnosporia is recognized as distinct from
Maytenus s.s. (Jordaan and van Wyk 1999;
22. Goniodiscus Kuhlm. Simmons and Hedin 1999; Simmons et al. 200la,
2001b). The affinity of the three American species
Goniodiscus Kuhlm., Arch. Jard. Bot. Rio de Janeiro 6: 109
(1933).
(Lundell1985; Hammel1997) needs to be critically
examined (R. Archer, pers. comm.).
Trees, glabrous. Leaves alternate, entire. Inflores-
cences axillary, cymose. Flowers bisexual, 5-
Celastraceae 47
25. Hartogiopsis H. Perrier albuminous, aril basal. Two spp., Queensland,

Australia, mesic forests and thickets.
Hartogiopsis H. Perrier, Notul. Syst. (Paris) 10: 194 (1942), rev.
Tree, glabrous. Leaves opposite, serrulate. 29. Kokoona Thwaites

Inflorescences axillary, cymose. Flowers bisexual,
Kokoona Thwaites, Hooker's J. Bot. Kew Gard. Misc. 5: 379
5-merous; disk fleshy, pulvinate, lobed, (1853); Ding Hou, Fl. Males. I, 6: 227-291 (1962).
introrse; ovary 3-locular; ovules erect, 2 per Trees, glabrous. Leaves opposite, rarely suboppo-
locule. Drupe ellipsoid, trigonous, dry; seeds 2-3, site or alternate, entire or crenulate. Inflorescences
oblong, albuminous. Only one sp., H. trilobocarpa axillary, paniculate. Flowers bisexual, 5-merous;
(Baker) H. Perrier, Madagascar, humid forests, disk fleshy, cupular, corrugated, extrastaminal;
500-2000m. anthers longitudinally dehiscent, latrorse, gener-
ally with triangular, pustular connective; ovary 3-
26. Hedraianthera F. Muell. locular; ovules axile, 6-16 per locule. Capsule
oblong, trigonous, woody, loculicidally dehiscent;
Hedraianthera F. Muell., Fragm. 5: 58 (1865); Jessup, Fl. Austr.
22: 150-180 (1984).
seeds 6-10(-16?) per locule, flattened, with mem-
branous, apical wing, exalbuminous. Eight spp., Sri
Lanka, southern India, Myanmar, Malaysia,
Trees or shrubs, glabrous. Leaves alternate, entire.
Borneo, Philippines,\ to dry forests, 0-1500m.
Inflorescences axillary, condensed thyrsoid or fas-
ciculate. Flowers bisexual, 5-merous; disk± fleshy,
annular, 5-lobed, intrastaminal; anthers obliquely 30. Lauridia Eckl. & Zeyh.
dehiscent, latrorse; ovary 5-locular; ovules erect, 4
Lauridia Eckl. & Zeyh., Enum. Pl. Afr. Austral. 1: 124 (1834/5);
per locule. Capsule ovoid or spheroid, bo~y, lo~uli Archer & van Wyk, S. Afr. J. Bot. 63: 227-232 (1997), rev.
cidally dehiscent; seeds 1-4 per locule, elhpsmd or
ovoid, triangular, albuminous, aril at base and one Scandent shrubs or rarely trees, branchlets often
side. Only one sp., H. porphyropetala F. Muell., sharply deflexed, glabrous. Leaves opposite, entire
eastern Australia, humid forests. or glandular-crenate. Inflorescences axillary,
compact thyrsoid or irregularly cymose. Flowers
27. Hexaspora C.T. White bisexual, 4(5)-merous; disk fleshy, entire, intrasta-
minal; anthers longitudinally dehiscent, introrse;
Hexaspora C.T. White, Contrib. Arnold Arbor. 4: 58 (1933);
ovary 2-locular; ovules erect, 2 per locule. Berry
Jessup, Fl. Austr. 22: 150-180 (1984).
spheroid, fleshy; seeds 1-2, spheroid, albuminous.
Two spp., S Africa, bush, woodlands, and dune
Trees, densely pilose. Leaves alternate, serrulate.
scrub.
5-merous; disk fleshy, annular, 5-angled, intrasta-
minal; anthers obliquely dehiscent; ovary 3- 31. Lophopetalum Wight ex Arn.
locular; ovules erect, 2-4 per locule. Fruit
Lophopetalum Wight ex Arn., Ann. Mag. Nat. Hist. I, 3: 150
unknown. Only one sp., H. pubescens C.T. White, (1839); Ding Hou, Fl. Males. I, 6: 227-291 (1962).
Queensland, Australia, humid forest.
Trees, glabrous. Leaves opposite or subopposite,
28. Hypsophila F. Muell. entire. Inflorescences axillary, thyrsoid. Flowers
bisexual, 5-merous; disk fleshy, annular, 5-angled
Hypsophila F. Muell., Victorian Nat. 3: 168 (1887); Jessup, Fl.
Austr. 22: 150-180 (1984).
or lobed, stamens on disk; anthers longitudinally
dehiscent, introrse; ovary 3-locular; ovules axile,
4-18 per locule. Capsule oblong or fusiform, trigo-
Trees or shrubs, glabrous. Leaves alternate or
nous or flattened along each locule, woody, loculi-
opposite, entire. Inflorescences axillary, cymose or
flowers solitary. Flowers bisexual, 5-merous; disk cidally dehiscent; seeds many, oblong, flattened,
medially attached, membranous wing surround-
fleshy, pulvinate, lobed, extrastaminal; anthers
transversely dehiscent; ovary 3-locular; ovules ing seed, exalbuminous or scanty. 2n = 40. About
20 spp., India, SE Asia, Macronesia, Northern Ter-
erect, 6-10 per locule. Capsule fusiform, bony,
loculicidally dehiscent; seeds several, oblong, ritory, Australia, humid to dry forests, swamp
forests, 0-1500m.
48 M.P. Simmons
32. Lydenburgia N. Robson

Lydenburgia N. Robson, Bol. Soc. Brot. 39: 34 (1965).
Trees or shrubs, glabrous. Leaves strictly opposite

or opposite on mature branches and alternate on
juvenile branches, crenulate-denticulate or glan-
dular-crenate. Inflorescences axillary, cymose.
Flowers bisexual, 5-merous; disk fleshy, annular,
5-angled, intrastaminal; anthers longitudinally
dehiscent, introrse; ovary 3(4)-locular; ovules
erect, 2 per locule. Capsule oblong, loculicidally
dehiscent; seeds 1-3(4), trigonous, albuminous.
Two spp., S Africa, bush or woodland on rocky
hills.
Lydenburgia is recognized as distinct from
Catha following Robson (1965) and Robson et al.
(1994). Catha abbottii A.E. van Wyk & M. Prins will
be transferred to Lydenburgia (R. Archer and Y.
Steencamp, pers. comm. 2000; van Wyk, in press).
33. Maurocenia Mill.

Maurocenia Mill., Gard. Diet. ed. 4: 859 (1754); Archer & van
Wyk, Bothalia 28: 7-10 (1998), rev.
Shrub, glabrous. Leaves opposite, entire. Inflores-

cences axillary, fasciculate or cymose. Flowers
bisexual or unisexual, rarely gynodioecious (or Fig. 15. Celastraceae-Celastroideae. Maytenus magellanica.
dioecious?), 5-merous; disk fleshy, undulate, A Fruiting branch. B Part of male inflorescence, showing
intrastaminal; anthers longitudinally dehiscent, flower buds with prophylls. C Female flower. D Petal of female
flower. E Female flower, vertical section. F Male flower bud.
introrse; ovary 2(3)-locular; ovules pendulous, 2 G Male flower. H Dehiscing fruit. (Dimitri 1972)
per locule. Berry spheroid, fleshy; seeds 1-2 per
locule, spheroid, albuminous. Only one sp., M.
frangula Mill., S Africa, mountain kloofs and tropics of Old and New Worlds, Australia, humid
coastal bush. forests to dry thickets.
Maytenus is narrowly circumscribed here and
does not include Gymnosporia, Moya, or Tricerma.
34. Maytenus Molina Fig. 15 Maytenus s.s. is a heterogeneous genus that is,
Maytenus Molina, Sagg. Stor. Nat. Chili ed. 1: 177 (1782); perhaps, best divided into segregate genera
Carvalho-Okano, Ph.D. Thesis, Instituto de Biologia, Uni- (Jordaan and van Wyk 1999). Maytenus s.s. has
versidade de Campinas, Campinas, Brazil (1992), reg. rev. been resolved as polyphyletic in phylogenetic
analyses (Simmons et al. 2001a, 2001 b). Additional
Trees or shrubs, erect, rarely scandent, glabrous, work is needed to re-circumscribe this genus.
rarely pubescent. Leaves alternate, entire or
serrate. Inflorescences axillary, fasciculate,
cymose, rarely racemose, or flowers solitary. 35. Menepetalum Loes.
Flowers bisexual or unisexual, rarely polygamous Menepetalum Loes. in R. Schlechter, Bot. Jahrb. Syst. 39: 163
or dioecious, (4)5-merous; disk fleshy, annular, (1906); Miiller, Fl. Nouv.-Caled. 20: 3-74 (1996).
(4)5-angled or lobed, intrastaminal; anthers longi-
tudinally dehiscent, introrse; ovary 2-3-locular; Trees or shrubs, glabrous. Leaves opposite, crenate
ovules erect, 1-2 per locule_ Capsule spheroid or or serrate. Inflorescences axillary, rarely cauliflor-
obovoid, coriaceous, loculicidally dehiscent; seeds ous, cymose. Flowers unisexual, rarely bisexual,
1- 6, ellipsoid or ovoid, albuminous, aril basal to dioecious, 5-merous; disk fleshy, annular, 5-angled
enveloping seed. About 200 spp., tropics and sub- or lobed, intrastaminal; anthers longitudinally
Celastraceae 49
dehiscent, introrse; ovary 3-locular; ovules erect, 39. Moya Griseb.

2-6 per locule. Capsule subspheroid, bony, loculi-
Moya Griseb., Pl. Lorentz.: 13, 63 (1874); Lourteig & O'Donell,
cidally dehiscent; seeds 1-5 per locule, ovoid, Natura 1: 181-233 (1955), reg. rev.
albuminous, aril basal to enveloping seed. Four
spp., New Caledonia, humid forests and maquis, Shrubs with thorns, glabrescent or pubescent.
500-1600m. Leaves alternate and fasciculate on short shoots,
entire or denticulate. Inflorescences axillary, fasci-
36. Microtropis Wall. ex Meisn. culate or flowers solitary. Flowers unisexual (or
bisexual?), 5-merous; disk fleshy, annular, entire or
Microtropis Wall. ex Meisn., Numer. List 152, n. 4337-40 (1831); 5-lobed or angled, intrastaminal; anthers longitu-
Merrill & Freeman, Proc.Am.Acad.Arts 73:271-310 (1940),
rev. dinally dehiscent, introrse; ovary 2-locular; ovules
Otherodendron Makino (1909). erect, 2 per locule. Capsule ellipsoid or subspher-
oid, loculicidally dehiscent; seeds 1-2, ellipsoid,
Trees or shrubs, glabrous. Leaves opposite, entire. albuminous, nearly enveloped by aril. Three spp.,
Inflorescences axillary, fasciculate or cymose. Argentina, Bolivia, Paraguay. Moya has been
Flowers bisexual, rarely unisexual, dioecious, (4)5- reduced to Maytenus by Lourteig and O'Donell
merous; disk 0 or annular, (4)5-angled, intrasta- {1955), but has been recognized as distinct by
minal; anthers longitudinally dehiscent, introrse, Jordaan and van Wyk {1999).
rarely extrorse; ovary 2-3-locular; ovules erect, 2
per locule. Capsule ellipsoid or oblong, coriaceous, 40. Mystroxylon Eckl. & Zeyh.
!-locular, laterally split along one side; seed 1,
ovoid, furrowed, albuminous, sarcotestal. About Mystroxylon Eckl. & Zeyh., Enum. Pl. Afr. Austral.: 125
66 spp., SE Asia, Macronesia, forests, 0-2700m. (1834/5); Robson et al., Fl. Trop. E. Afr. 108: 1-78 (1994);
Archer et al., Fl. Pl. Afri. 55: 76-80 (1997), part. rev.
New World Microtropis are recognized as
Quetzalia following Lundell {1970), contra Lundell
Trees or shrubs, glabrous or pubescent. Leaves
{1939).
alternate, entire or crenulate or glandular-
denticulate. Inflorescences axillary, cymose-
37. Monimopetalum Rehder umbellate. Flowers bisexual, 5-merous; disk thin,
annular or margins upturned, 5-angled or lobed,
Monimopetalum Rehder, J. Arnold Arbor. 7: 233 (1926).
introrse; ovary 3-4-locular; ovules erect, 2 per
Scandent shrubs, glabrous (except inflorescence
locule. Drupe spheroid or ellipsoid, ± fleshy; seed
minutely pubescent). Leaves alternate, serrulate.
1, spherical or ovoid, albuminous. Only one spp.
(R. Archer, pers. comm. 2000), M. aethiopicum
4-merous; disk annular, 4-lobed, stamens on disk;
(Thunb.) Loes., Africa, Madagascar, forests, open
anthers?; ovary 4-locular; ovules erect, 1 per
woodlands, and scrub, 0-2550m.
locule. Capsule deeply 1-2( -4) lobed, loculicidally
dehiscent; seeds 1-2{ -4), oblong, albuminous,
aril basal. Only one sp., M. chinense Rehder, 41. Orthosphenia Standi.
China.
Orthosphenia Standl., Contrib. U.S. Natl. Herb. 23: 684 (1923);
Rzedowski, Ciencia (Mexico) 16: 139-142 (1956), rev.
38. Mortonia A. Gray
Shrubs, glabrous. Leaves alternate, glandular-
Mortonia A. Gray, Pl. Wright. 1: 34, t. 4 (1852); Prigge, Ph.D serrate. Inflorescences axillary, fasciculate, or
Thesis, Claremont Graduate School: 1-136 (1983), rev.
flowers solitary. Flowers bisexual or unisexual
(polygamous), 5-merous; disk small, intrastami-
Shrubs, glabrous. Leaves alternate, entire. Inflores-
nal; anthers longitudinally dehiscent, introrse;
cences terminal, thyrsoid. Flowers bisexual, 5-
ovary 2-locular; ovules erect, 2 per locule. Berry
merous; disk fleshy, margins upturned, 5-lobed,
ovoid-spheroid, dry; seeds 1-2, ellipsoid, albu-
minous. Only one sp., 0. mexican a Standi., Mexico,
introrse; ovary 5-locular; ovules erect, 2(4) per
deserts.
locule. Drupe oblong, dry; seed 1, oblong, albu-
minous. Five spp., southwestern U.S.A., Mexico,
deserts.
50 M.P. Simmons
42. Paxistima Raf. America, C America, SE Asia, Macronesia, Queens-

land, Australia, Hawaii, humid forests and thickets.
Paxistima Raf., Sylva Tellur.: 42 (1838); Navaro & Blackwell,
Sida 14: 231-249 (1990), rev. Cuatrescasas (1948) noted that the species differ
Pachistima Raf. (1818). by only minute characteristics, and Ding Hou
Pachystima 'Raf.' ex End!. (1841). {1962) suggested that there may be fewer species
than are recognized.
Low, rhizomatous shrubs, glabrous. Leaves oppo-
site, entire or serrulate. Inflorescences axillary,
cymose or flowers solitary. Flowers bisexual, 4- 45. Platypterocarpus Dunkley & Brenan
merous; disk fleshy, annular, 4-angled, intrastami- Platypterocarpus Dunkley & Brenan, Kew Bull. 1948: 47 (1948);
nal; anthers longitudinally dehiscent, introrse; Robson et al., Fl. Trop. E. Afr. 108: 1-78 (1994).
ovary 2-locular; ovules erect, 2 per locule. Capsule
oblong, chartaceous, loculicidally dehiscent; seeds Trees, glabrous. Leaves alternate, crenate-dentate.
1-2, oblong, albuminous, partially enveloped by Inflorescences axillary, cymose. Flowers bisexual,
lacerate aril. 2n = 32. Two spp., N America, forests, 5-merous; disk thin, annular, 5-lobed, intrastami-
open hillsides, and bluffs, 600-3350 m. This genus nal; anthers longitudinally dehiscent, introrse;
is properly spelled "Paxistima", not "Pachistima" ovary (4?)5-locular; ovules erect, 2 per locule.
or "Pachystima" (Wheeler 1943; Uttall1986). Both Samara with 4-5 lateral wings, chartaceous or
species are cultivated as groundcovers. subcoriaceous; seed 1, flattened. Only one sp., P.
tanganyikensis Dunkley & Brenan, Tanzania,
transition between wet and dry forest, 1500-
43. Peripterygia (Baill.) Loes. 1900m.
Peripterygia (Bail!.) Loes. in R. Schlechter, Bot. Jahrb. Syst. 39:
168 (1906); Muller, Fl. Nouv.-Caled. 20: 3-74 (1996).
Pterocelastrus Meisn. sect. Peripterygia Baill. (1874). 46. Plenckia Reissek
Plenckia Reissek in Martius, Fl. Bras. 11: 30 (1861).
Shrubs, glabrous. Leaves alternate, serrate. Inflo- Viposia Lundell (1939).
rescences axillary or terminal, cymose. Flowers
bisexual or unisexual, rarely andromonoecious, 5- Trees or shrubs, glabrous. Leaves alternate, entire.
merous; disk fleshy, annular, 5-angled or lobed, Inflorescences axillary, cymose. Flowers bisexual
intrastaminal; anthers longitudinally dehiscent, or unisexual (polygamous), 5-merous; disk fleshy,
introrse; ovary 3-locular; ovules erect, 1-2(3) per pulvinate, 5-angled, intrastaminal; anthers
locule. Capsule flattened along each locule, bony, obliquely dehiscent, latrorse; ovary 2-locular;
loculicidally dehiscent; seeds discoid with annular, ovules erect, 2 per locule. Samara with 1 apical
membranous wing, albuminous. Only one sp., P. wing, obovate or oblanceolate, chartaceous; seeds
marginata (Baill.) Loes., New Caledonia, maquis, 1-2, oblong, albuminous. Four spp., S America.
0-1050 m.
47. Pleurostylia Wight & Arn.
44. Perrottetia Kunth
Pleurostylia Wight & Arn., Prod. Fl. Ind. Orient.: 157 (1834).
Perrottetia Kunth in Humboldt, Bonpland and Kunth, Nov. Herya Cordem. (1895).
Gen. Sp. 7:73, t. 622 (1824); Ding Hou,Fl.Males. !,6: 227-291
(1962); Lundell, Phytologia 57: 231-238 (1985), reg. rev. Trees or shrubs, glabrous. Leaves opposite, entire.
Trees or shrubs, puberulent or glabrescent. Leaves {4)5-merous; disk fleshy, annular or margins
alternate, entire or serrulate, with generally small upturned, {4)5-lobed, intrastaminal; anthers
abaxial domatia in axils of larger veins. Inflores- longitudinally dehiscent, introrse; ovary 2-locular,
cences axillary, cymose or thyrsoid. Flowers bisex- rarely becoming 1-locular by abortion; ovules
ual or unisexual (dioecious), (4)5-merous; disk erect, 2-8 per locule. Nut obovoid or ellipsoid,
annular or margins upturned, entire or lobed, bony, with lateral style; seeds 1(-2), ovoid, albu-
intrastaminal; anthers longitudinally dehiscent, minous. About 8 spp., Africa, Madagascar, India,
introrse; ovary 2-locular (seemingly 4-locular at Sri Lanka, SE Asia, Macronesia, Queensland,
base); ovules erect, 2 per locule. Berry spheroid,± Australia, New Caledonia, forests and woodlands,
fleshy; seeds 2-4, subspheroid, furrowed, albu- 0-1600m.
minous, sarcotestal. About 17 spp., northern S
Celastraceae 51
48. Polycardia Juss.

Polycardia Juss., Gen. Pl.: 377 (1789); Perrier de Ia Bathie, Fl.
Madag. 116: 1-76 (1946), rev.
Trees or shrubs, glabrous or pubescent. Leaves alter-

nate, entire, denticulate, or spinose-dentate. Inflo-
rescences epiphyllous, rarely axillary, fasciculate.
Flowers bisexual, 5-merous; disk fleshy, annular,
entire or± 5-lobed, intrastaminal; anthers longitu-
dinally dehiscent, introrse; ovary 5( 4)-locular;
ovules axile, numerous. Capsule ovoid, coriaceous-
woody, loculicidally dehiscent; seeds several per
locule, oblong, albuminous or exalbuminous,
partially enveloped by lacerate aril. Four spp.,
Madagascar, littoral and dry forests, 0-1200 m.
49. Psammomoya Diels & Loes.

Psammomoya Diels & Loes., Bot. Jahrb. Syst. 35: 339 (1904);
Jessup, Fl. Austr. 22: 150-180 (1984), rev.
Shrubs, glabrous. Leaves opposite, reduced to

scales. Inflorescences axillary, fasciculate. Flowers
bisexual, 5-merous; disk thin or± fleshy, annular,
5-angled, intrastaminal; anthers longitudinally
dehiscent, introrse; ovary 2-3-locular; ovules
erect, 2 per locule. Capsule obovoid or ellipsoid, Fig. 16. Celastraceae-Celastroideae. Pterocelastrus echinatus.
bony, loculicidally dehiscent; seeds 1-2, ovoid or A Flowering branchlet. B Flower. C Same, vertical section. D
Ovary, transverse section. E Fruit. F Seed. G Seed with aril
spheroid, albuminous, aril basal. Two spp., western removed. (Robson 1966)
Australia, heath.
2-locular; ovules erect, 2(1 ?) per locule. Samara

50. Pseudosalacia Codd
with 1 surrounding wing, widely ovate or ovate-
Pseudosalacia Codd, Bothalia 10: 565 (1972). lanceolate, membranous; seeds 1-2, oblong, albu-
minous. Two spp., Madagascar, tropophilous
Trees, glabrous. Leaves alternate, entire. Inflores- forest, 250-400 m.
cences axillary, fasciculate or cymose. Flowers
bisexual, 5-merous; disk fleshy, annular, 5-angled,
stamens on disk; anthers apically dehiscent, 52. Pterocelastrus Meisn. Fig. 16
extrorse; ovary 3-locular; ovules erect, 2 per Pterocelastrus Meisn., Pl. Vase. Gen.: 68 (1837); Robson, Fl.
locule. Capsule spheroid, coriaceous, loculicidally Zambesiaca 2: 355-418 (1966).
dehiscent; seeds 2-5, trigonous, albuminous. Only
one sp., P. streyi Codd, S Africa, coastal, rocky river Trees or shrubs, glabrous. Leaves alternate, entire.
banks. Inflorescences axillary, cymose. Flowers bisexual,
5-merous; disk thin, margins upturned, 5-lobed,
51. Ptelidium Thouars introrse; ovary (2)3-locular; ovules erect, 2 per
Ptelidium Thouars, Hist. Veg. Isles Austr. Afr. l : 11,29 (1805); locule. Capsule ellipsoid with lateral or oblique
Perrier de Ia Bathie, Fl. Madag. 116: 1- 76 (1946), rev. horn to wing-like outgrowths, loculicidally dehis-
cent; seeds 1-3, ovoid, albuminous, nearly
Scandent or erect shrubs, glabrous. Leaves oppo- enveloped by aril. Four spp., SE Africa, evergreen
site, entire or crenulate-dentate Inflorescences forests and rocky slopes.
axillary, cymose. Flowers bisexual, 4-5-merous; Pterocelastrus needs to be conserved against
disk margins upturned, 4-5-lobed; anthers longi- Asterocarpus Eckl. & Zeyh. (Enum. Pl. Afr. Austral.
tudinally dehiscent, introrse or extrorse; ovary 1: 122 (1834/5); R. Archer, pers. comm. 2000).
52 M.P. Simmons
53. Putterlickia Endl. Flowers bisexual, 4-merous; disk annular, intrast-

aminal; anthers longitudinally dehiscent; ovary
Putterlickia End!., Gen. Pl. 1086 (1840); Jordaan & van Wyk, S.
Afr. J. Bot. 64: 322-329 (1998), rev. 2-locular; ovules erect, 2 per locule. Samara with 1
apical wing, elliptical, chartaceous; seed 1, fur-
Shrubs or lianas with thorns, glabrous. Leaves rowed, albuminous. Only one sp., R. tolantonguen-
alternate (opposite if on thorns) and fasciculate on sis Medrano, Mexico, montane scrub, 1400-
short shoots, entire or serrulate. Inflorescences 2000m.
axillary, cymose. Flowers bisexual, 5-merous; disk
fleshy, annular, 5-lobed, intrastaminal; anthers lon- 57. Salaciopsis Baker f.
gitudinally dehiscent, introrse; ovary 3-locular;
Salaciopsis Baker f., J. Linn. Soc. Bot. 45: 287 (1921); Miiller, Fl.
ovules {4-)6{-12) per locule. Capsule obovoid-
Nouv.-Caled. 20: 3-74 (1996).
trigonous, bony, loculicidally dehiscent; seeds Lecardia J. Poiss. ex Guillaumin (1927).
6-18, enveloped by aril. Four spp., S Africa, south-
ern Mozambique, forests to scrub. Trees or shrubs, glabrous. Leaves alternate and
subverticillate, entire. Inflorescences axillary,
54. Quetzalia Lundell rarely cauliflorous, fasciculate. Flowers unisexual
(dioecious), 5-merous; disk fleshy, annular, 5-
Quetzalia Lundell, Wrightia 4: 137 (1970); Lundell, Contrib. lobed, intrastaminal; anthers longitudinally
Univ. Michigan Herb. 3: 5-46 (1939), rev. (under
Microtropis).
dehiscent, introrse; ovary 3-locular; ovules erect,
2-6 per locule. Capsule ellipsoid or subspheroid,
Trees or shrubs, rarely lianas, glabrous. Leaves bony, loculicidally dehiscent; seeds 1-6, ovoid,
opposite, entire. Inflorescences axillary, cymose. albuminous, aril basal to enveloping seed. Six
Flowers unisexual (dioecious), (4)5-merous; disk spp., New Caledonia, humid and gallery forests,
fleshy, annular or margins upturned, (4)5-angled, 100-1400m.
introrse; ovary 2-locular; ovules erect, 2 per locule. 58. Salvadoropsis H. Perrier
Capsule ellipsoid or oblong, coriaceous, !-locular,
laterally split along one side; seed 1(2), ovoid, fur- Salvadoropsis H. Perrier, Bull. Soc. Bot. France 91: 96 (1944).
rowed, albuminous, sarcotestal. Eleven spp., C
America, Mexico, humid forests, to 3100 m. Quet- Trees or shrubs, glabrous. Leaves opposite, entire.
zalia is recognized as distinct from Microtropis Inflorescences axillary, cymose. Flowers bisexual,
following Lundell {1970), contra Lundell {1939). 5-merous; disk 5-angled, intrastaminal; anthers
longitudinally dehiscent, introrse; ovary 5-locular;
ovules axile, 12 per locule. Berry spheroid,± fleshy;
55. Robsonodendron R.H. Archer seeds 1-2 per locule, oblanceolate, albuminous.
Robsonodendron R.H. Archer in Archer & van Wyk, S. Afr. J.
Only one sp., S. arenicola H. Perrier, Madagascar,
Bot. 63: 116 (1997). coastal dunes.
Trees or shrubs, glabrous. Leaves alternate, entire 59. Sarawakodendron Ding Hou
or glandular-crenate. Inflorescences axillary, fasci-
culate. Flowers bisexual, 5-merous; disk fleshy, Sarawakodendron Ding Hou, Blumea 15: 141 (1967).
margins upturned, entire, intrastaminal; anthers
longitudinally dehiscent, introrse; ovary 2-locular; Trees, glabrous. Leaves alternate, entire. Inflores-
ovules erect, 2 per locule. Drupe spheroid, fleshy; cences axillary, racemose. Flowers bisexual, 5-
seeds 1-2, spheroid, albuminous. Two spp., S merous; disk fleshy, annular, 5-angled,
Africa, coastal forest and forest margins, coastal extrastaminal; anthers transversally dehiscent,
dunes. extrorse; ovary 3-locular; ovules horizontal, 8 per
locule. Capsule narrow-ellipsoid, coriaceous,
loculicidally dehiscent; seeds 6-8 per locule,
56. Rzedowskia Medrano narrow-lanceolate, descending; aril basal, cushion-
Rzedowskia Medrano, Bol. Soc. Bot. Mexico 41: 41 (1981). shaped, with basal, filamentous extensions. Only
one sp., S. filamentosum Ding Hou, Borneo, humid
Shrubs, glabrous. Leaves opposite, glandular- forests.
denticulate. Inflorescences axillary, cymose.
Celastraceae 53
60. Schaefferia Jacq. one sp., T. jamaicensis Urb., Jamaica, thickets on

limestone.
Schaefferia Jacq., Enum. Syst. Pl.: 10 ( 1760); Brizicky, J. Arnold
Arbor. 45: 206-234 (1964), part. rev.
63. Torralbasia Krug & Urb.
Trees or shrubs, glabrous, rarely pubescent. Leaves
Torralbasia Krug & Urb. in Segui, Flora Med. T6x. Cuba: 60
alternate or fasciculate on short shoots, entire, (1900); Liogier, Flora Hispaniola 1: 12-25 (1981).
rarely serrulate. Inflorescences axillary, fasciculate
or flowers solitary. Flowers unisexual (dioecious),
Trees . or shrubs, glabrous. Leaves alternate ,
4-merous; disk indistinct or 0; anthers longitudi-
opposite, or subopposite, entire or crenulate. In-
nally dehiscent, introrse; ovary 2-locular; ovules
florescences axillary, cymose. Flowers bisexual,
erect, 1(2) per locule. Drupe subspheroid, dry;
4-merous; disk cupular, 4-lobed, intrastaminal;
seeds 1-2, ellipsoid or ovoid, albuminous. About
anthers longitudinally dehiscent, introrse; ovary 4-
23 spp., Americas, West Indies, dry, rocky wood-
locular; ovules .erect, 1 per locule. Capsule deeply
lands, hummocks, thickets.
1-4-lobed, conaceous, longitudinally dehiscent;
seeds 1-4, ellipsoid, albuminous, aril basal. Only
61. Siphonodon Griff. one sp., T. cuneifolia (Wright) Krug & Urb., Cuba,
Hispaniola, Puerto Rico, dwarf forests, 800-1500 m.
Siphonodon Griff., Calcutta J. Nat. Hist. 4: 246, t.14 (1844); Ding
Hou, Fl. Males. I, 6: 227-291 (1962); Jessup, Fl. Austr. 22:
150-180 (1984). 64. Tricerma Liebm.
Trees, glabrous. Leaves alternate, entire or serru- Tricerma Liebm., Vidensk. Meddel. Dansk Naturhist. Foren.
Kjobenhavn: 97 (1853); Lundell, Wrightia 4: 153-172 (1971),
late. Inflorescences axillary, cymose. Flowers
rev.
bi~exual, 5-~erous; 5 stamens often alternating
With 5 stammodes, anthers obliquely dehiscent,
Trees or shrubs, puberulent or glabrescent. Leaves
latrorse; ovary with apical hollow with style-like
alternate, entire or serrulate. Inflorescences axil-
central column arising from base of hollow, 10-
lary, fasciculate, condensed thyrsoid, or flowers
locular with each locule horizontally divided into
solitary. Flowers bisexual or unisexual (polyga-
1-ovulate locelli; ovules in upper locelli erect, in
mous),. 5-merou~; disk fleshy, annular, 5-angled or
lower !ocelli pendulous, 1 per locule. Drupe sphe-
lobed: mtrastammal; anthers longitudinally dehis-
roid or ob.ovoid, hard to fleshy; seeds few to many,
cent, mtrorse; ovary 3(4)-locular; ovules erect, 1
fla~tened, m 1-s~eded, bony pyrenes. Seven spp., SE
per locule. Capsule obovoid or fusiform, coria-
Asia, Macronesm, Queensland, Australia, forests
ceous, lo~ulicidallf dehiscent; seeds 1-3, ellipsoid
and thickets, 0-1650m.
or obov01d, albummous, enveloped by aril. 2n = 80.
Siphonodon has been considered unusual
Seven spp., southern North America, South
within Celastraceae based on the structure of
America, dry thickets.
its gynoecium (Croizat 1947), wood anatomy
Tricerma is recognized as distinct from
(Metcalfe and Chalk 1950), and pollen morphol-
Maytenus (Lundell1971; Simmons et al. 2001b).
ogy (Erdtman 1952). However, Siphonodon
has been supported as a derived member of
Celastraceae (e.g., Savolainen, Fay et al. 2000; 65. Tripterygium Hook. f.
Simmons et al. 2001b).
Tripterygium Hook f. in Bentham & Hooker, Gen. Pl. 1: 368
(1862); Ma, Edinburgh J. Bot. 56: 35-46 (1999), rev.
62. Tetrasiphon Urb.
Scandent shrubs, glabrous or tomentose. Leaves
Tetrasiphon Urb., Symb. Antill. 5: 83 (1904); Fawcett & Rendle, alternate, serrate. Inflorescences axillary or termi-
Fl. Jamaica 5: 24-35 (1926).
nal, thyrsoid. Flowers bisexual or unisexual
(polygamous), 5-merous; disk fleshy, margins
Tre~s or shrubs, glabrous. Leaves opposite,
upturned, s.-lobed, intrastaminal; anthers longitu-
entire. Inflorescences axillary, cymose. Flowers
dmally dehiscent, introrse; ovary 3-locular; ovules
unisexual (dioecious), 4-merous; disk pulvinate, 4-
erect, 2 per locule. Samara with 3 lateral wings,
lobe~, intrastaminal; anthers longitudinally dehis-
chartaceous; seed 1, trigonous, albuminous. 2n =
cent, mtrorse; ovary 4-locular; ovules pendulous, 1
24. ~ne sp., China, Taiwan, Korea, Japan, thickets.
per locule. Drupe spheroid, ± fleshy; seeds 2-4,
Cultivated as ornamentals for their winged fruits.
oblong-ellipsoid, ± flattened, albuminous. Only
54 M.P. Simmons
66. Wimmeria Schltdl. & Cham. to scales. Inflorescences terminal, racemose, in

spikes, umbellate, or flowers solitary. Flowers
Wimmeria Schltdl. & Cham., Linnaea 6: 427 (1831); Lundell,
Contrib. Univ. Michigan Herb. 3: 5-46 (1939), rev. actinomorphic or± zygomorphic, bisexual, perig-
ynous with short or cupular hypanthium; 5-
Trees or shrubs, glabrous or pubescent. Leaves merous; petals free or medially connate; disk thin,
alternate, some fasciculate on short shoots, lining hypanthium; stamens 5, sometimes 3 long
serrulate. Inflorescences axillary, cymose. Flowers and 2 short; anthers longitudinally dehiscent,
bisexual, 4-merous; disk fleshy, annular or margins introrse; ovary 2-5-locular, with basal placenta-
upturned, 5-angled, intrastaminal; anthers longi- tion; ovules 1 per locule, basal, erect. Schizo carp of
tudinally dehiscent, introrse; ovary (2)3-locular; 2-5 indehiscent mericarps; seeds albuminous.
ovules axile, 4-9 per locule. Samara with (2)3(4)
lateral wings, chartaceous; seed 1(2), linear, 69. Macgregoria F. Muell.
albuminous. Twelve spp. (C. Clevinger, pers.
comm. 2000), C America, Mexico, humid to dry Macgregoria F. Muell., Fragm. 8: 160 (1874).
forests.
Erect to prostrate annual herbs, glabrous. Leaves
linear. Inflorescences racemose. Flowers sub-
67. Xylonymus Kalkman ex Ding Hou tended by one bract each; hypanthium shallow;
Xylonymus Kalkman ex Ding Hou, Fl. Males. I, 6: 243 (1962). petals free; stames equal in length; anthers longer
than filaments, tipped by white glands; ovary 5-
Shrub, glabrous. Leaves alternate, entire. Inflores- locular; style with membranous cup at base.
cences axillary, cymose. Flowers bisexual, 4- Schizocarp of 5 mericarps, mericarps densely
merous; disk fleshy, annular, 4-angled, papillose with hooked hairs. Only one sp., M.
intrastaminal; anthers obliquely dehiscent, racemigera F. Muell., central Australia, arid grass-
latrorse; ovary 4-locular; ovules axile, ± 10 per lands and woodlands.
locule. Capsule, oblong, woody, loculicidally dehis-
cent; seeds several per locule, ellipsoid, albu- 70. Stackhousia Sm.
minous, partially enveloped by aril. Only one sp.,
X. versteeghii Kalkman, New Guinea, humid Stackhousia Sm., Trans. Linn. Soc. Lond. 4:218 (1798); Barker,
Fl. Austr. 22: 186-199 (1984).
lowland forests.
Perennial or annual herbs, sometimes rhizoma-
68. Zinowiewia Turcz. tous. Leaves obovate to linear, sometimes reduced
to scales. Inflorescences racemose, in spikes,
Zinowiewia Turcz., Bull. Soc. Imp. Nat. Moscou 32: 275 (1859);
Lundell, Contrib. Univ. Michigan Herb. 3: 5-46 (1939), rev. umbellate or flowers solitary. Flowers bracteolate;
hypanthium cupular; petals medially connate;
Trees or shrubs, glabrous. Leaves opposite, entire. stamens 3 long and 2 short; anthers as long or
Inflorescences axillary, cymose. Flowers bisexual, shorter than filaments; ovary 3-5-locular.
5-merous; disk fleshy, annular, 5-angled or lobed, Schizocarp of (1-)3-5 mericarps; style inserted
intrastaminal; anthers longitudinally dehiscent, between mericarps; mericarps attached only at
introrse; ovary 2-locular; ovules erect, 2 celled. base, sometimes each with 3 wings. 2n = 18, 20, 30,
Samara with 1 apical wing, oblanceolate or 36. About 16 spp., Australasia, dunes, grasslands to
obovate, chartaceous; seeds 1(2), linear-oblong, forests.
albuminous. Seventeen spp., Mexico, C America, Barker (1984) suggested a number of sub-
northern S America, humid forests to mesic thick- generic groups but did not formalize them. In
ets, to 3150m. particular, Barker noted the group of species
related to S. viminea with clustered flowers
subtended by numerous bracts on the inflores-
II. SUBFAMILY STACKHOUSIOIDEAE cence, and the group of species related to S.
Burnett (1835). monogyna with separate tri-bracteate flowers on
the inflorescence.
Stackhousiaceae R. Br. (1814), nom. cons.
71. Tripterococcus Endl. Fig. 17
Annual or perennial herbs with erect or prostrate
stems. Leaves alternate, entire, sometimes reduced Tripterococcus Endl., Enum. Pl. Hueg.: 17 (1837).
Celastraceae 55
been published. Recognition of Tripterococcus

probably makes Stackhousia paraphyletic.
Ill. SUBFAMILY HIPPOCRATEOIDEAE

(Juss.) Lindl. (1836).
Hippocrateaceae Juss. (1811), nom. cons.
Lianas or scandent, rarely completely erect trees

or shrubs. Inflorescences axillary or terminal.
Flowers bisexual, (3-4)5-merous; petals free; disk
extrastaminal or 0, rarely stamens on disk;
androgynophore wide, narrow, or 0; stamens 3,
rarely 5 or numerous; anthers transversely dehis-
cent, introrse or extrorse (apical); ovary 3-locular;
ovules axile, 2-16(-22) per locule. Capsule trans-
versely flattened, lobed to base, rarely lobed ±
halfway to base or locules entirely connate, coria-
ceous, rarely woody or chartaceous, loculicidally
dehiscent; seeds 2-numerous, with membranous
basal wing or narrow stipe, exalbuminous.
Included phloem 0.
72. Anthodon Ruiz & Pav.

Anthodon Ruiz & Pav., Fl. Peruv. 1: 45, t. 74 (1798); Giirts-van
Rijn & Mennega, Fl. Guianas, 16: 3-81 (1994).
Lianas, glabrous. Leaves opposite or subopposite,

crenulate or serrulate. Inflorescences axillary,
cymose. Flowers bisexual, 5-merous; petals regu-
larly serrate; disk fleshy, cupular, entire, extrasta-
minal; stamens 3; anthers transversely dehiscent,
extrorse; ovary 3-locular; ovules axile, 8-14 per
locule. Capsule transversely flattened, locules
Fig.17. Celastraceae-Stackhausioideae. Tripterococcus bruno- entirely connate, coriaceous, loculicidally dehis-
nis. A Plant. B Flower with two of the three sub tending bracts cent; seeds 8-14 per locule, with membranous
visible. C Flower with corolla removed. D Fruit. E Mericarp. basal wing, exalbuminous. Two spp., C and S
F Gynoecial column after shedding mericarps. (Barker 1984) America, humid forests, 100-900 m.
73. Apodostigma R. Wilczek

Perennial herbs, glabrous. Leaves linear to ovate Apodostigma R. Wilczek, Bull. Jard. Bot. Etat 26: 402 (1956);
or sometimes reduced to scales. Inflorescences in Robson eta!., Fl. Trop. E. Afr. 108: 1-78 (1994).
one-sided spikes. Flowers subtended by three
bracts each; hypanthium cupular; petals medially Lianas, glabrous. Leaves opposite or subopposite,
connate; stamens 3 long and 2 short; anthers entire or denticulate. Inflorescences axillary,
shorter than filaments; ovary 3-locular. Schizocarp cymose. Flowers bisexual, 5-merous; 4 of 5 petals
of ( 1-) 3 mericarps; style terminal, persistent; arched, ± zygomorphic; disk of 3 discontinuous
mericarps each with 3 prominently-veined wings, lobes around stamens, extrastaminal; stamens 3;
glabrous. 2n = c. 30, c. 32. Two spp., SW Australia, anthers transversely dehiscent, extrorse; ovary 3-
heath, sclerophylous forests, woodlands. locular; ovules axile, 2 per locule. Capsule trans-
Barker (1984) cited two species in the genus, one versely flattened, lobed to base, coriaceous,
undescribed. Only one sp., T. brunonis Endl., has loculicidally dehiscent; seeds 2 per locule, with
56 M.P. Simmons
membranous basal wing, exalbuminous. Only one 77. Cuervea Triana ex Miers
sp., A. pallens (Oliv.) R. Wilczek, Africa, Madagas-
Cuervea Triana ex Miers, Trans. Linn. Soc. Lond. 28: 370 (1872);
car, humid forests to dry thickets. Smith, Brittonia 3: 341-555 (1940), reg. rev.; N. Halle, Fl.
Cameroun 32: 3-243 (1990).
74. Arnicratea N. Halle
Trees, shrubs, or Hanas, glabrous. Leaves opposite
Arnicratea N. Halle, Bull. Mus. Natl. Hist. Nat., B, Adansonia 6: or subopposite, entire, crenulate, or denticulate.
12 (1984). Inflorescences axillary, rarely terminal, cymose or
thyrsoid. Flowers bisexual, 5-merous; disk mem-
Lianas, glabrous or pubescent. Leaves opposite, branous, cupular, entire or lobed, extrastaminal;
rarely subopposite, entire, rarely serrulate. Inflo- stamens 3; anthers transversely dehiscent,
rescences axillary, thyrsoid. Flowers bisexual, 5- extrorse; ovary 3-locular; ovules axile, 4-9 per
merous; disk indistinct, lobed, extrastaminal; locule. Capsule transversely flattened, lobed to
anthers transversely dehiscent, extrorse; ovary 3- base, coriaceous, loculicidally dehiscent; seeds
locular; ovules axile, (4-)6(7) per locule. Capsule 2-numerous, with membranous basal wing or
transversely flattened, lobed to base, coriaceous, narrow stipe, exalbuminous. Five spp., C and S
loculicidally dehiscent; seeds numerous, with America, West Indies, Africa, humid to gallery
membranous basal wing, exalbuminous. Three forests.
spp., India, SE Asia, Macronesia, lowland forests, to
700m.
78. Elachyptera A.C. Sm.
75. Bequaertia R. Wilczek Elachyptera A.C. Sm., Brittonia 3: 383 (1940); N. Halle, Fl.
Cameroun 32: 3-243 (1990).
Bequaertia R. Wilczek, Bull. Jard. Bot. Etat 26: 399 (1956);
Robson et al., Fl. Trop. E. Afr. 108: 1-78 (1994). Lianas or scan dent trees or shrubs, glabrous (inflo-
rescences sometimes puberulent). Leaves opposite
Lianas, glabrous. Leaves opposite, crenulate- or subopposite, entire, crenulate, or serrate. Inflo-
dentate. Inflorescences axillary, cymose. Flowers rescences axillary or terminal, cymose or thyrsoid.
bisexual, 5-merous; petals thick and fleshy; disk 0; Flowers bisexual, 5-merous; disk± fleshy, cupular,
stamens 3; anthers transversely dehiscent, lobed, extrastaminal; stamens 3; anthers trans-
extrorse; ovary 3-locular; ovules axile, 6-8( -12) versely dehiscent, extrorse; ovary 3-locular; ovules
per locule. Capsule transversely flattened, lobed axile, 2 or 4 per locule. Capsule transversely flat-
to base, coriaceous, loculicidally dehiscent; seeds tened, lobed to base, coriaceous, loculicidally
numerous, with membranous basal wing, exalbu- dehiscent; seeds 2-4 per locule, with membranous
minous. Only one sp., B. mucronata (Exell) R. basal wing or narrow stipe, exalbuminous. Seven
Wilczek, tropical Africa, humid forests and river spp., C and S America, Africa, Madagascar, humid
banks, 900-1000m. forests to dry thickets, mangrove swamps.
76. Campylostemon Welw. 79. Helictonema Pierre Fig. 18

Campylostemon Welw. in Bentham & Hooker, Gen. Pl. 1: 998 Helictonema Pierre, Bull. Mens. Soc. Linn. Paris N.S. 9: 73
(1862); N. Halle, Fl. Cameroun 32: 3-243 (1990). (1898); Robson et al., Fl. Trop. E. Afr. 108: 1-78 (1994).
Lianas, glabrous. Leaves opposite, entire or Lianas, hirsute and stellate pubescent. Leaves
serrate. Inflorescences axillary, cymose, rarely opposite, entire. Inflorescences axillary, cymose.
thyrsoid. Flowers bisexual, 5-merous; disk 0; Flowers bisexual, 5-merous; disk fleshy, annular,
stamens 5; anthers transversely dehiscent, irregularly lobed, extrastaminal; androgynophore
introrse; ovary 3-locular; ovules axile, 4-16 per wide; stamens 3; anthers transversely dehiscent,
locule. Capsule transversely flattened, lobed to extrorse; ovary 3-locular; ovules axile, numerous.
base, coriaceous, loculicidally dehiscent; seeds Capsule transversely flattened, lobed to base, cori-
numerous, with membranous basal wing, exalbu- aceous, loculicidally dehiscent; seeds numerous,
minous. 2n = 56. Eight or more species, Africa, with membranous basal wing, exalbuminous. Only
forests and thickets. one sp., H. velutinum (Afzel.) N. Halle, tropical
Africa, forests.
Celastraceae 57
81. Hylenaea Miers

Hylenaea Miers, Trans. Linn. Soc. Lond. 28: 366 (1872); Gi:irts-
van Rijn & Mennega, Fl. Guianas, 16: 3-81 (1994).
Trees or lianas, glabrous. Leaves opposite or sub-

opposite, entire. Inflorescences axillary or termi-
nal, thyrsoid. Flowers bisexual, 5-merous; disk
membranous or ± fleshy, cupular, entire; stamens
3; anthers transversely dehiscent, extrorse; ovary
3-locular; ovules axile, 4-8 per locule. Capsule
transversely flattened, lobed to base, woody,
loculicidally dehiscent; seeds 4-8 per locule,
enlarged embryoniferous portion with narrow
stipe (or membranous wing), exalbuminous.
Three spp., C and S America, forests.
82. Loeseneriella A.C. Sm.

Loeseneriella A. C. Sm., Am. J. Bot. 28: 438 (1941); N. Halle, Fl.
Cameroun 32: 3-243 (1990).
Lianas or scan dent, rarely erect shrubs, puberulent

or glabrescent. Leaves opposite, entire or crenulate.
5-merous; disk fleshy, pulvinate or cupular,
extrastaminal; androgynophore 0 or wide;
stamens 3; anthers transversely dehiscent,
extrorse; ovary 3-locular; ovules 4-15( -22) per
Fig. 18. Celastraceae-Hippocrateoideae. Helictonema veluti- locule. Capsule transversely flattened, lobed to
num. A Cymule of inflorescence. B Stellate leaf hair. base, coriaceous, loculicidally dehiscent; seeds
C Flower with artificially expanded petals, note disk and numerous, with membranous basal wing, exalbu-
androgynophore. D Flower, vertical section. E Anther. FOvary, minous. 2n = 56. Sixteen spp., Africa, Madagascar,
transverse section, with stamen filaments. G Ovary locule
opened, showing the ovules. H Trilobed capsule. I Basal part India, SE Asia, Macronesia, Queensland, Australia,
of capsule lobe showing two placentas. J Fruit lobe, opened. humid forests to dry thickets.
K Winged seed, the cotyledons transversely sectioned.
L Embryo. Drawn by N. Halle. (Halle 1990)
83. Plagiopteron Griff. Fig. 19
J. Nat. Hist. 4: 244, t. 13 (1843);
Plagiopteron Griff., Calcutta
Tang, Wu & Li, Acta Bot. Yunnan. 12: 126-128 (1990), rev.
80. Hippocratea L.
Hippocratea L., Gen. Pl. ed. 1: 363 (1737); Sp. Pl. ed. 1: 1191 Scandent shrubs or lianas, stellate pubescent.
(1753); Smith, Brittonia 3: 341-555 (1940), reg. rev. Leaves opposite, entire. Inflorescence axillary,
thyrsoid. Flowers bisexual, (3)4(5) merous; disk
Lianas, puberulent or glabrescent. Leaves opposite, indistinct, stamens on disk; stamens numerous;
entire, crenate, or serrate. Inflorescences axillary, anthers transversely, apically dehiscent; ovary 3-
cymose or thyrsoid. Flowers bisexual, 5-merous; locular; ovules erect, 2 per locule. Capsule trans-
disk fleshy, pulvinate or cupular, entire, extrasta- versely flattened, lobed to base, coriaceous; seeds
minal; stamens 3; anthers transversely dehiscent, unknown. Only one sp., P. suaveolens Griff., China,
extrorse; ovary 3-locular; ovules axile, 4-8 per Myanmar, forests.
locule. Capsule transversely flattened, lobed to Plagiopteron has been assigned to Tiliaceae
base, chartaceous or coriaceous, loculicidally (Bentham and Hooker 1862), Flacourtiaceae
dehiscent; seeds numerous, with membranous (Warburg 1893), and Plagiopteraceae (Airy Shaw
basal wing, exalbuminous. 2n = 28. Three spp., 1965). However, Plagiopteron has been consis-
Americas, West Indies, tropical Africa, humid to tently supported as related to Hippocrateoideae in
gallery forests, 0-1800m. phylogenetic analyses (e.g., Simmons and Hedin
58 M.P. Simmons
1999; Soltis et al. 2000; Simmons et al. 2001b), and

the anatomy of Plagiopteron is consistent with
Celastraceae (Baas et al. 1979).
84. Prionostemma Miers

Prionostemma Miers, Trans. Linn. Soc. Lond. 28: 354 (1872);
N. Halle, Bull. Mus. Natl. Hist. Nat., B, Adansonia 3: 5-14
(1981), rev.
Lianas, glabrous or glabrescent. Leaves opposite,

entire, rarely dentate. Inflorescences axillary, rarely
terminal, thyrsoid. Flowers bisexual, 5-merous;
disk fleshy, annular-pulvinate, entire, extrastami-
nal; stamens 3; ovary 3-locular; ovules axile, 6-20
per locule. Capsule transversely flattened, lobed
to base, coriaceous, loculicidally dehiscent; seeds
numerous, with membranous basal wing, exalbu-
minous. Five spp., C and South America, tropical
Africa, India, forests and savannas.
85. Pristimera Miers

Pristimera Miers, Trans. Linn. Soc. Lond. 28: 360 (1872);
N. Halle, Bull. Mus. Natl. Hist. Nat., B, Adansonia 3: 5-14
(1981), rev. Fig. 19. Celastraceae-Hippocrateoideae. Plagiopteron suave-
a/ens. A Flowering branchlet. B Flower bud. C Flower with
most stamens removed. D Flower from underside, showing
Lianas or scandent shrubs, glabrous or puberulent. unequal sepals. E Anthers showing transverse dehiscence.
Leaves opposite or subopposite, entire or serrate. F Pistil, ovary vertically sectioned, showing basal placentation.
Inflorescences axillary, rarely terminal, cymose, G Ovary, transverse section. H Infructescence. I Fruit. Drawn
rarely thyrsoid. Flowers bisexual, 5-merous; disk by R. van Crevel. (Baas et al. 1979)
fleshy, indistinct, annular or cupular, entire or
lobed; stamens 3; anthers transversely dehiscent,
extrorse; ovary 3-locular; ovules axile, 2-10 per
locule. Capsule transversely flattened, lobed to per locule. Capsule transversely flattened, lobed to
base, chartaceous or ± coriaceous, loculicidally base, coriaceous, loculicidally dehiscent; seeds 2
dehiscent; seeds 2-10 per locule, with membra- per locule, with membranous basal wing, exalbu-
nous basal wing, exalbuminous. Twenty-four spp., minous. 2n = 28. Six spp., Africa, Madagascar,
Old and New World tropics, humid forests to dry India, SE Asia, Macronesia, humid forests to dry
thickets. This genus, which has been resolved as thickets.
polyphyletic in a 26 S gene tree and a simultane-
ous analysis (Simmons et al. 2001b), will need
87. Semialarium N. Halle
further attention.
Semialarium N. Halle, Bull. Mus. Natl. Hist. Nat., B, Adansonia
5: 22 (1983).
86. Reissantia N. Halle
Reissantia N. Halle, Bull. Mus. Nat!. Hist. Nat. 30: 466 (1958); Tree, shrub, or Iiana, puberulent or glabrescent.
Robson et al., Fl. Trop. E. Afr. 108: 1-78 (1994). Leaves opposite, rarely subopposite, crenulate or
serrulate. Inflorescences axillary, rarely terminal,
Lianas or scandent shrubs, glabrous or puberulent. thyrsoid. Flowers bisexual, 5-merous; fleshy,
Leaves opposite or subopposite, entire, crenulate, annular-pulvinate, entire, extrastaminal; stamens
or serrate. Inflorescences axillary, rarely terminal, 3; anthers transversely dehiscent, extrorse; ovary
cymose, rarely thyrsoid. Flowers bisexual, 5- 3-locular; ovules axile, 6- 8. Capsule transversely
merous; disk ± fleshy, pulvinate or cupular, lobed, flattened, lobed ± halfway to base, coriaceous,
extrastaminal; stamens 3; anthers transversely loculicidally dehiscent; seeds 6-8 per locule, with
dehiscent, extrorse; ovary 3-locular; ovules axile, 2 membranous basal wing, exalbuminous. 2n = 30.
Celastraceae 59
Two spp., Mexico, C and South America, forests, to IV. Subfamily Salacioideae N. Halle (1962)
1300m. [published without Latin diagnosis].
Lianas or scan dent, rarely completely erect trees or

88. Simicratea N. Halle
shrubs. Inflorescences axillary, rarely terminal or
Simicratea N. Halle, Bull. Mus. Nat!. Hist. Nat., B, Adansonia 1: cauliflorous. Flowers bisexual, 5-merous; petals
18 (1983); Robson eta!., Fl. Trop. E. Afr. 108: 1-78 (1994). free; disk extrastaminal; androgynophore wide,
narrow, or 0; stamens (2)3(5); anthers longitudi-
Lianas, glabrous. Leaves opposite, entire or crenu- nally, obliquely, or transversely dehiscent, extrorse,
late. Inflorescences axillary, cymose. Flowers rarely apical; ovary (2)3(5)-locular; ovules axile,
bisexual, 5-merous; disk fleshy, annular, 5-angled, 2-9 per locule. Berry spheroid, oblong, or
extrastaminal; androgynophore narrow; stamens fusiform, coriaceous, rarely chartaceous; seeds
3; anthers transversely dehiscent, extrorse; ovary 1-many, oblong, angled, exalbuminous, sur-
3-locular; ovules axile, 2 per locule. Capsule trans- rounded by mucilaginous arils. Included phloem
versely flattened, lobed to base, coriaceous, generally present.
loculicidally dehiscent; seeds 2 per locule, with
membranous basal wing, exalbuminous. Only one
sp., S. welwitschii (Oliv.) N. Halle, tropical Africa, 91. Cheiloclinium Miers
forest, scrubland. Cheiloclinium Miers, Trans. Linn. Soc. Lond. 28: 420 (1872);
Giirts-van Rijn & Mennega, Fl. Guianas, 16: 3-81 (1994).
89. Simirestis N. Halle

Lianas or scandent trees or shrubs, glabrous
Simirestis N. Halle, Bull. Mus. Nat!. Hist. Nat. 30:464 (1958); N. or glabrescent. Leaves opposite, entire, crenate, or
Halle, Bull. Mus. Nat!. Hist. Nat., B, Adansonia 6: 3-18 (1984), serrate. Inflorescences axillary, thyrsoid or
rev. cymose. Flowers bisexual, 5-merous; disk of 3-5
discontinuous lobes around stamens, extrastami-
Lianas, glabrous. Leaves opposite or subopposite, nal; stamens 3 or 5; anthers transversely dehiscent,
entire or dentate. Inflorescences axillary, cymose, extrorse; ovary 3 or 5-locular; ovules axile, 2 or 4
rarely terminal and paniculate. Flowers bisexual, per locule. Berry subspheroid, coriaceous; seeds
5-merous; disk indistinct, lobed, extrastaminal; 2-6, oblong, angled, exalbuminous, surrounded by
androgynophore wide; stamens 3; anthers trans- mucilaginous arils. Eleven spp., C and South
versely dehiscent, extrorse; ovary 3-locular; ovules America, humid forests, 0-1300m.
axile, (4-)6-16 per locule. Capsule transversely
flattened, lobed to base, coriaceous, loculicidally
dehiscent; seeds numerous, with membranous 92. Peritassa Miers
basal wing, exalbuminous. Eight spp., Africa, Peritassa Miers, Trans. Linn. Soc. Lond. 28: 402 (1872); Smith,
forests. Brittonia 3: 341-555 (1940), rev.
90. Tristemonanthus Loes. Lianas or scandent trees or shrubs, glabrous (some

inflorescences pilose). Leaves opposite, subop-
Tristemonanthus Loes., Wiss. Ergebn. Deutsch. Zentr.-Afr.- posite, or alternate, entire, crenate, or serrate.
Exped. 1910-11, 2: 77 (1922) sine descr.; Loes., Feddes
Repert. 49: 226 (1940); N. Halle, Fl. Cameroun 32: 3-243
Inflorescences axillary, thyrsoid or cymose.
(1990). Flowers bisexual, 5-merous; disk± membranous,
cupular, lobed, extrastaminal; stamens 3; anthers
Lianas, glabrous. Leaves opposite, entire or dentic- obliquely or longitudinally dehiscent, extrorse,
ulate. Inflorescences axillary, cymose. Flowers with or without bilobed extended connective;
bisexual, 5-merous; disk 0; stamens 3; anthers ovary 3-locular; ovules axile, 2 or 4 per locule.
transversely dehiscent, introrse; ovary 3-locular; Berry spheroid or oblong-ellipsoid, chartaceous
ovules axile, 4-8 per locule. Capsule transversely or coriaceous; seeds 2-6, oblong, angled, exalbu-
flattened, lobed to base, coriaceous, loculicidally minous, surrounded by mucilaginous arils.
dehiscent; seeds numerous, with membranous Thirteen spp., C and South America, humid
basal wing, exalbuminous. Two spp., tropical forests.
Africa, swamp forests.
60 M.P. Simmons
93. Salacia L.
Salacia L., Mant. Pl. 2: 159,293 (1771); Loesener, Nat. Pflanzen-
faro. ed. 2, 20b: 198-231 (1942), part. rev.
Salacicratea Loes. (1910).
Lianas or scandent, rarely completely erect trees or

shrubs, glabrous or glabrescent. Leaves opposite
or subopposite, rarely alternate, entire, crenate, or
serrate. Inflorescences axillary or cauliflorous,
thyrsoid, cymose, or fasciculate. Flowers bisexual,
5-merous; disk fleshy, annular, margins upturned,
or cupular, entire or lobed, extrastaminal; androg-
ynophore wide, narrow, or 0; stamens (2)3; anth~rs
longitudinally, obliquely, or transversely dehis-
cent, extrorse or apical; ovary (2)3-locular; ovules
axile, 2-9 per locule. Berry spheroid, oblong, or
fusiform, coriaceous; seeds 1-many, oblong,
angled, exalbuminous, surrounded by mucilagi-
nous arils. 2n = 28. About 200 spp., Old and New
World tropics, Australia, humid forests to dry
scrub and grasslands.
This large, heterogeneous genus is unlikely to
be monophyletic and has been resolved as para-
phyletic in phylogenetic analyses (Simmons et al.
2001a, 2001b).
94. Salacighia Loes. Fig. 20. Celastraceae-Salacioideae. Thyrsosalacia pararace-

Salacighia Loes., Wiss. Ergebn. Deutsch. Zentr.-Afr.-Exped. mosa. A Flowering branchlet. B Flower bud. C Flower. D Calyx,
1910- 11 2: 77 (1922) sine descr.; Loes., Feddes Repert. 49: with quincuncial aestivation. E Anther, dorsal view. F Anther,
228 (1940); N. Halle, Fl. Cameroun 32: 3- 243 (1990). ventral view. G Dehiscing anther. H Flower, vertical section.
I Young flower, same. Drawn by N. Halle. (Halle 1990)
Lianas, glabrous. Leaves subopposite or alternate,

entire. Inflorescences cauliflorous, fasciculate
on long pendant branches. Flowers bisexual, 5- exalbuminous, surrounded by mucilaginous arils.
merous; disk fleshy, pulvinate, 5-lobed, extrasta- Four spp., tropical Africa, humid forests or rocky
minal; stamens 3; anthers transversely dehiscent, slopes, to 1000 m.
extrorse; ovary 3-locular; ovules axile, 2 per locule.
Berry spheroid; seeds 6, subspheroid, exalbu-
minous, surrounded by mucilaginous arils. 2n = 96. Tontelea Aubl.
28. Two spp., tropical Africa, dry to humid forests, Tontelea Aubl., Hist. Pl. Guiane 1: 31, t. 10 (1775); Smith, Brit-
10-1000m. tonia 3: 341-555 (1940), rev.
95. Thyrosalacia Loes. Fig. 20 Lianas or scandent trees or shrubs, glabrous or

glabrescent. Leaves opposite or subopposite,
Thyrosalacia Loes., Feddes Repert. 49: 229 (1940); N. Halle, Fl. entire, crenate or serrate. Inflorescences axillary,
Cameroun 32: 3- 243 (1990).
thyrsoid, rarely cymose. Flowers bisexual, 5-
merous; disk membranous or ± fleshy, cupular,
Scandent shrubs or lianas, glabrous. Leaves oppo- entire or lobed, extrastaminal; stamens 3; anthers
site, entire. Inflorescences axillary or terminal, transversely dehiscent, extrorse; ovary 3-locular;
thyrsoid or racemose. Flowers bisexual, 5-merou~; ovules 2-4( -8) per locule. Berry subspheroid or
disk fleshy, annular, entire or 5-lobed, extrastami- ellipsoid, coriaceous or woody; seeds few, rarely
nal; stamens 3; anthers obliquely or longitudinally many, oblong, angled, exalbuminous, surrounded
dehiscent, extrorse; ovary 3-locular; ovules axile, by mucilaginous arils. Thirty-one spp., C and
2 per locule. Berry fusiform; seeds 1-3, ellipsoid, South America, humid forests, to 1800m.
Celastraceae 61
Genera Doubtfully Included in Celastraceae Airy Shaw, H.K. 1965. Diagnosis of new families, new names,
etc., for the seventh edition of Willis's 'Dictionary'. Kew Bull.
18: 249-273.
97. Nicobariodendron Vasudeva Rao & Chakrab. Airy Shaw, H.K., Cutler, D.F., Nilsson, S. 1973. Pottingeria, its
Nicobariodendron Vasudeva Rao & Chakrab., }. Econ. Taxon. taxonomic position, anatomy and palynology. Kew Bull. 28:
Bot. 7: 513 (1985). 97-104.
Archer, R.H. 1990. The taxonomic status of Cassine L. s.l.
(Celastraceae) in southern Africa. M.Sc. Thesis. Pretoria:
Trees, glabrous. Leaves opposite, entire. Inflores- University of Pretoria.
cences axillary, racemose. Flowers unisexual Archer, R.H., van Wyk,A.E.1992. Palynology and intergeneric
(dioecious), 4-5-merous; disk ± fleshy, annular, relationships in some southern African species of subfam-
intrastaminal; stamens 2; anthers longitudinally ily Cassinoideae (Celastraceae). Grana 31: 241-252.
Archer, R.H., van Wyk, A.E. 1993a. Bark structure and inter-
dehiscent; ovary and ovules unknown. Drupe generic relationships of some Southern African Cassi-
oblong-ellipsoid or narrowly obovoid; seed 1, noideae (Celastraceae). IAWA J. 14: 35-53.
basal. Only one sp., N. sleumeri Vasudeva Rao & Archer, R.H., van Wyk, A.E. 1993b. Wood structure and generic
Chakrab., Andaman-Nicobar Islands, India, status of some southern African Cassinoideae (Celas-
traceae). IAWA }. 14: 373-389.
forests, to lOOm. Archer, R.H., van Wyk, A.E. 1997. A taxonomic revision of
This incompletely known genus is morphologi- Cassine L. s.str. (Cassinoideae: Celastraceae). S. Afr. }. Bot. 63:
cally unusual relative to other Celastraceae, with 146-157.
its racemose inflorescences and two stamens Archer, R.H., van Wyk, A.E. 1998a. A taxonomic revision of
(although these characteristics are not unique Maurocenia (Celastraceae), a Western Cape monotypic
endemic. Bothalia 28: 7-10.
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Cephalotaceae 65
Cephalotaceae
J.G. CONRAN
trap, these leaves alternating with the non-trap

Cephalotaceae Dumort., Anal. Fam. Pl.: 59, 61 (1829), nom.
cons. leaves in the shoot apex rosette. All leaves bear
unicellular hairs and numerous sessile glands.
Perennial carnivorous evergreen herbs. Rootstock Dickson (1883) also noted various intermediate
a thick, knotty rhizome, roots fibrous. Stems 0. forms between normal and trapping leaves.
Leaves borne in rosettes at rhizome apices, alter- The trapping leaves are highly modified, with
nate, exstipular, petiolate, fine-hairy, alternately the lamina replaced by a lidded pitcher (Dickson
spathulate-obovate or pitcher-shaped and insect- 1883). The pitcher is apical on the petiole and
trapping. Inflorescence a thyrse with scorpoid ovoid with two prominent anterior ribs and an
cymes. Flowers hermaphroditic, small, calyx well ovate, sub-apical opening covered by a protective
developed, basally connate, rotate, purplish-white; lid with conspicuous, areolate achlorophyllous
limb 6-lobed; lobes imbricate in bud; corolla 0; patterns. The outer pitcher surface is glandular,
stamens 12 in two whorls of six; petals 0; filaments with fluid (considered to be nectar) secreted over
free, divergent; anthers dorsifixed, versatile, with its surface (Hamilton 1904). The rim of the pitcher
a dorsal-connective appendage, 2-thecate, 4- is strongly ribbed with a series of large, inward-
sporangiate, introrse, opening by lateral slits; disk pointing teeth, and the upper third to half of the
intrastaminal, trichomatous-papillose; gynoecium inner pitcher surface is covered with small glands,
of 6 free carpels alternating with the inner stamens whereas the lower pitcher has two, generally red-
and sepals; ovaries superior, !-locular with 1-2 coloured patches of larger glands (Lloyd 1942).
erect anatropous ovules; stylodia terminal, Inside the rim, past the large teeth, there is a
straight, ventrally papillate near the apex. Fruit further region, the cornice, consisting of down-
1-2-seeded, hairy indehiscent leathery follicles; wardly-pointed, tooth-like epidermal cells and a
seeds small, ovoid, brown with a membranous smooth transition zone above the glands (Parkes
testa; endosperm copious, granulose; embryo and Hallam 1984).
minute, linear. The pitcher is not a simple epiascidium, and
A monotypic family with the single species ontogenetic studies have revealed that it is a
Cephalotus follicularis Labill. confined to far SW specifically modified rhachis of a bijugal pinnate
Western Australia. The plant is grown widely as an leaf (Froebe and Baur 1988).
ornamental curiosity.
VEGETATIVE ANATOMY. Raphides are absent.
VEGETATIVE MORPHOLOGY. Roots are fibrous Vessels occur in the roots and rhizome and are
and consist of a short-lived taproot and numerous heavily pitted with scalariform perforations
adventitious roots arising from the rhizome. The (Macfarlane 1911). Sieve-tube plastids are of
rhizome is spreading, much-branched and the dicotyledonous Ss-type (Behnke 1988, 1991).
sheathed with numerous scale leaves and bears Laticifers are absent.
apical rosettes of photosynthetic and trapping The roots are triarch with a 2-3-layered cortex
leaves. The flowering stems are erect, scapose, and thin-walled endodermis. A cork cambium is
bracteate; the lower bracts are leaf-like, the scape formed, resulting in 3-4 clear cork cell layers, but
bracts linear and deciduous, and the upper bracts the primary cortical tissues are not shed, instead
and those subtending the flowers are small, clus- turning brown with age. There is also a starch
tered and incurved. storage region of uncertain origin within the cork
Leaves are of three types: scale leaves on the rhi- of old roots (Macfarlane 1911).
zomes, aerial rosette leaves with non-sheathing, The rhizome epidermis is initially hairy, these
petiolate and spathulate-obovate photosynthetic hairs becoming tubercles with age. There is a
blades, and highly modified pitchers with the broad, starchy primary cortex and a partially
lamina modified into a pitcher-shaped, hooded incomplete ring of vascular tissue with a broad
66 J.G. Conran
phloem and xylem and a parenchymatous pith. EMBRYOLOGY. The ovules are anatropous,
Secondary development results in a 3-5-cell cork bitegmic and crassinucellate. The embryology of
layer, starchy secondary cortex and starch deposi- the family is the subject of ongoing research
tion in the pith, followed by eventual accumula- (Conran and Macfarlane, in prep.).
tion of tannins in the parenchymatous tissues
(Macfarlane 1911). POLLEN MORPHOLOGY. Pollen grains are spher-
Paracytic stomata occur on both leaf surfaces, as oidal, tricolpate (angulaperturate) with a reticu-
do so-called large and small glands and both late sexine and slightly thinner nexine and c. 18 f.tm
simple and complex hairs. The mesophyll consists diameter (Erdtman 1952). The tectum is perforate
of an adaxial and abaxial palisade layer, with a with dense pitting and the apertures are complex
loose, central spongy mesophyll. There is a larger (Hideux and Ferguson 1976).
midrib vein, two smaller lateral veins and a series
of smaller, sub-marginal veinlets (Macfarlane KARYOLOGY. Kondo (1969), Kress (1970) and
1911). Peng and Goldblatt (1983) all determined the
The pitchers are covered externally with numer- chromosome number in Cephalotaceae to be
ous stomata and large glands, the latter consisting 2n = 20.
of cells with dense cytoplasma arranged like a
tetracytic stoma, with a central inner cell pair Pollination. Cephalotaceae flowers are pollinated
surrounded by four symmetrically arranged cells, by small insects.
and another pair of internally associated cells
(Dickson 1878; Solereder 1908). In section, the Fruit and Seed Morphology. The fruit is a densely
pitchers have several layers of chlorenchyma hairy follicle, the hairs derived from the carpel
subtended by achlorophyllous mesophyll. The papillae which elongate after fertilisation. The
translucent areas are parenchymatous and non- small, single seed is enclosed inside the follicle and
vascularised (Parkes and Hallam 1994). The is thin-walled and ovoid. The copious endosperm
pitcher interiors have numerous non-functional cells are granular (Macfarlane 1911).
stomata, referred to by Goebel (1891) as water Germination is epigeal and phanerocotylar. The
stomata. Small glands in the upper pitcher consist seedlings have a well-developed but ephemeral
of 6-12 concentrically arranged cells, whereas the taproot and the hypocotyl is glabrous. The coty-
lateral glandular patches in the lower pitcher have ledons are ovoid and glabrous, the first leaves
numerous large, vascularised, multicellular glands are alternate, resemble the later ones and are
(Macfarlane 1911). These large glands are flask- glandular-hairy. The seedlings are unusual, with
shaped, with a ventral region of irregularly polyg- an expanded non-vascularised outgrowth of the
onal cells and an apical region of palisade-like cells hypocotyl growing into the follicle, filling the
(Solereder 1908), and they are bounded internally loculus and possibly acting as a non-starchy food
by transversely thickened, endodermal cells reserve for the seedling. In addition, the hairy fol-
(Parkes and Hallam 1984). licle coat may assist in keeping the seedling
hydrated (Conran and Denton 1996).
INFLORESCENCE STRUCTURE. The inflorescence
is a thyrse. The flowers are borne in short, con- Ecology. The Cephalotaceae are considered to be
densed, lateral scorpoid cymes. carnivorous, the modified pitcher leaves acting
to trap the prey (Lloyd 1942). The trapping leaves
FLORAL MORPHOLOGY. The sepals have three to have ultraviolet absorption patterns which are
seven anastomosing vascular traces (Chrtek et al. thought to attract prey in several families of car-
1989). The anthers are introrse and cruciform, nivorous plants (Joel et al. 1985). Dakin (1919)
dorsifixed medially, dehiscing laterally by slits. found no direct evidence of digestive enzymes,
There is a thin connective and prominent dorsal suggesting indirect digestion through putrefac-
connective of large cells. The endothecium is 1- tion, although Joel (in Juniper et al. 1989) reported
layered, discontinuous over the connective or the protease activity in the intercellular plugs of the
inner thecal surface. There is a septum between small glands within the pitchers. Brown (1866)
the pollen sacs (Endress and Stumpf 1991). Vogel reported the presence of live insect larvae and
(1998) reported that the floral nectary in Cephalo- coccoid unicellular green algae living inside the
tus forms a contiguous perigynous disk derived pitchers.
from a series of pillars, each with an apical stom- Cephalotus grows in the peat -soil swamps
atal pore. formed over granite substrata, often in seepage
Cephalotaceae 67
1998; Savolainen, Fay et al. 2000) place it instead in

the Oxalidales near the Cunoniaceae. Its floral
diagram is identical with that of Brunelliaceae.
DISTRIBUTION AND HABITATS. The family grows

only in the extreme SW of Western Australia
where the plant is widespread and locally abun-
dant. The plants are generally found growing in or
near Sphagnum mounds in well-drained areas of
swamps relatively close to the coast, in an area
extending over a distance of some 400 km.
ECONOMIC IMPORTANCE. Cephalotus is popular

with carnivorous plant enthusiasts and widely
cultivated. Although listed previously on CITES
Appendix II, the plant was recently removed from
CITES, given its relative abundance in protected
areas of SW Western Australia and its ease of veg-
etative propagation.
Only one genus:
Cephalotus Labill. Fig. 21

Cephalotus Labill., Nov. Hoi!. Pl. Sp. 2: 6, t. 145 {1806).
Description as for family.
Selected Bibliography
Albert, V.A., Williams, S.E., Chase, M.W. 1992. Carnivorous

plants: phylogeny and structural evolution. Science. 257:
Fig. 21. Cephalotaceae. Cephalotus follicularis. A Flowering 1491-1495.
plant. B Flower. C Flower, vertical section. (Baillon 1872) APG (Angiosperm Phylogeny Group) 1998. See general
references.
Bate Smith, E.C. 1962. See general references.
Baillon, H. 1872. Histoire des plantes, vol. 3. Paris: Hachette.
areas, along creeks and under grass tussocks Behnke, H.-D. 1988. Sieve-element plastids and systematic
relationships of Rhizophoraceae,Anisophyllaceae and allied
(Luffitz 1966). DeBuhr (1976) reported that the groups. Ann. Missouri Bot. Gard. 75: 1387-1409.
plants re-sprout from the rhizome following Behnke, H.D. 1991. Distribution and evolution of forms and
fire, but germination is not dependent on fire types of sieve-element plastids in the dicotyledons. Aliso 13:
(Piliciauskas 1989). Plants propagate readily from 167-182.
rhizome pieces, as well as from single leaves or Brown, R. 1866. General remarks on the botany of Terra
Australis. Misc. Bot. Works. 1: 76-78.
pitchers grown in Sphagnum. Chase et al. 1993. See general references.
Chrtek, J., Slavfkova, Z., StudniCka, M. 1989. Beitrag zur Leit-
PHYTOCHEMISTRY. Tannin cells are present. biindelanordnung in den Kronblattern von ausgewahlten
Myricetin, ellagic acid, quercetin and gallic acid Arten der fleischfressenden Pflanzen. Preslia 61: I 07-124.
are present (Bate Smith 1962) but iridoids are Conran, J.G., Denton, M.D. 1996. Germination in the Western
Australian Pitcher Plant Cephalotus follicularis and its
absent (Jensen et al. 1975). unusual early seedling development. W.A. Nat. 21: 37-42.
Corner, E.J.H. 1976. See general references.
AFFINITIES. Cephalotus has traditionally been Dakin, W.J. 1919. The West Australian pitcher plant (Cephalo-
associated with the Saxifragaceae but with some tus follicularis), and its physiology. J. Roy. Soc. W.A. 4: 37- 53.
DeBuhr, L.E. 1976. Field notes on Cephalotus follicularis in
features linking it to the Crassulaceae (Macfarlane
Western Australia. Cam. Pl. Newslett. 5: 8- 9.
1911). More recent molecular studies of its affini- Dickson, A. 1878. The structure of the pitcher of Cephalotus
ties (e.g. Albert et al. 1992; Chase et al. 1993; APG follicularis. J. Bot. 16: 1-5.
68 J.G. Conran
Dickson, A. 1883. On the morphology of the pitcher of Kondo, K. 1969. Chromosome numbers of carnivorous plants.
Cephalotus fol/icu/aris. Trans. Proc. Edinburgh Bot. Soc. 14: Bull. Torrey Bot. Club 96: 322-328.
172-181. Kress, A. 1970. Zytotaxonomie Untersuchungen an einigen
Diels, L. 1928. Cephalotaceae. In: Engler, A., Prantl, K. Insektenfiingern (Droseraceae, Byblidaceae, Cephalotaceae,
Die natiirlichen Pflanzenfamilen, ed. 2, 18a. Leipzig: W. Roridulaceae, Sarraceniaceae). Ber. Deutsch. Bot. Ges. 83:
Engelmann, pp. 71-74. 55-62.
Endress, P.K., Stumpf, S. 1991. The diversity of stamen struc- Lloyd, F.M. 1942. The carnivorous plants, 2nd edn. Waltham,
tures in 'Lower' Rosidae (Rosales, Fabales, Proteales, Sapin- Mass.: Chronica Botanica Co.
dales). Bot. J. Linn. Soc. 107: 217-293. With 294 figures. Luffitz, F. 1966. The West Australian pitcher plant ( Cephalotus
Erdtman, G. 1952. See general references. follicularis Labill.). Austral. Pl. 12: 34-35.
Froebe, H.A., Baur, N. 1988. Die Morphogenese der Kannen- Macfarlane, J.M. 1911. Cephalotaceae. In: Engler, A. (ed.) Das
blatter von Cephaltus follicularis Labill. Akad. Wiss. Lit. Pflanzenreich IV, 116. Leipzig: W. Engelmann, pp. 1-15.
Mainz, Abh. Math.-Naturwiss. Kl. Jg. 1988,3, 19 pp. Nicholls, K.W., Bohm, B.A., Ornduff, R. 1985. Flavonoids and
Goebel, K. 1891. Pflanzenbiologische Schilderungen, part 2. affinities of the Cephalotaceae. Biochem. Syst. Ecol. 13:
Marburg: Elwert. 261-264.
Hamilton, A.G. 1904. Notes on the West Australian pitcher Parkes, D.M., Hallam, N.D. 1984. Adaptation for carnivory in
plant ( Cephalotus follicu/aris Labill.). Pro c. Linn. Soc. N.S. W. the West Australian pitcher plant ( Cepha/otus fol/icularis).
29: 36-53. Austral. J. Bot. 32: 595-604.
Hideux, M., Ferguson, I.K. 1976. The stereostructure of the Peng, C.-I., Goldblatt, P. 1983. Confirmation of the chromo-
exine and its evolutionary significance in Saxifragaceae some number in Cephalotaceae and Roridulaceae. Ann.
sensu lato. In: Ferguson, I.K., Muller, I. (eds.) The evolu- Missouri Bot. Gard. 70: 197-198.
tionary significance of the exine. Linnean Society Sympo- Piliciauskas, E. 1989. Cephalotus fol/icularis and how to grow
sium Series no. 1. London: Academic Press, pp. 327-377. them from seed. Vic. C. P. Soc. Newslett. 6: 12-15.
Jay, M., Lebreton, P. 1972. Chemotaxonomic research on vas- Savolainen, V., Fay, M.F. et al. 2000. See general references.
cular plants. XXVI. The flavinoids of the Sarraceniaceae, Schulze, W., Schulze, E.D., Pate, J.S., Gillison, A.N. 1997. The
Nepenthaceae, Droseraceae and Cephalotaceae; a critical nitrogen supply from soils and insects during growth of the
study of the Sarraceniales. Nat. Can. (Quebec) 99: 607-613. pitcher plants Nepenthes mirabilis, Cephalotus follicularis
Jensen, S.R., Nielsen, B.J., Dahlgren, R. 1975. Iridoid com- and Darlingtonia californica. Oecologia. 112: 464-471.
pounds, their occurrence and systematic importance in the Solereder, H. 1908. Systematic anatomy of the Dicotyledons,
angiosperms. Bot. Notiser 128: 148-180. vol. 1. Introduction, Polypetalae and Gamopetalae. Oxford:
Joel, D.M., Juniper, B.E., Dafni,A.1985. UV patterns in the traps Clarendon Press.
of carnivorous plants. New Phytol. 101: 585-594. Vogel, S. 1998. Remarkable nectaries: structure, ecology,
Juniper, B.E., Robins, R.J., Joel, D.M. 1989. Carnivorous plants. organophyletic perspectives. II. Nectarioles. Flora (Jena)
London: Academic Press. 193: 1-29.
Clethraceae 69
Clethraceae
J.V. ScHNEIDER and C. BAYER
Clethraceae Klotzsch, Linnaea 24: 12 (1851), nom. cons. deciduous. The Chinese-Indochinese C. faberi is
usually evergreen but appears to be deciduous in
Trees or shrubs, evergreen or deciduous; in- the southeastern part of its area. Leaves often
dumentum of simple, fasciculate and/or stellate appear apically crowded on vegetative shoots. In
trichomes. Leaves alternate, simple, entire or Purdiaea, the branches often exhibit a zigzag
(glandular-)serrate to dentate, petiolate or sessile, pattern due to sympodial growth (Thomas 1960).
exstipulate. Inflorescences terminal or axillary Leaf venation in Clethra corresponds to the
racemes, simple or compound in panicle-, fascicle-, eucamptodromous, brochidodromous (Yu and
or umbel-like aggregates; bracts present, often Chen 1990), or semi-craspedodromous patterns,
caducous, prophylls absent (sometimes rudimen- i.e. the secondaries are connected by a series of
tary?); pedicels articulated, sometimes very short. arching veins, sometimes forming prominent
Flowers actinomorphic or slightly zygomorphic, arches, or they branch near the margin with one
hermaphroditic or rarely functionally female branch running into the margin, the other one
with reduced anthers; sepals 5(6), quincuncial- towards the superadjacent secondary. Purdiaea
imbricate, distinct or fused up to 3/4, equal in size shows an acrodromous to palinactinodromous
or the outer conspicuously larger than the inner; pattern (Rodriguez and Berazain 1991). Domatia
petals 5(6), distinct or rarely fused up to 1/3, formed by hair tufts are found in several Japanese
imbricate; stamens 10(12), in 2 whorls, free or and Chinese Clethra species.
adnate to petals at their very base; anthers ventri- Most Clethraceae are pubescent, particularly
fixed, bilobed-sagittate to caudate, versatile, on buds, leaves, inflorescences and flowers.
tetrasporangiate, swinging at anthesis from the Single and fascicled trichomes are found. The
extrorse bud position into an introrse, inverted latter are longer, more robust, free or basally
position, dehiscing by eventually apical pore-like connate, in some tropical American species
slits; disk hypogynous or absent; ovary superior, forming a kind of a stipe. True stellate trichomes
sometimes with nectary at base, 3-5-locular; pla- occur as well.
centation axile on upper portion of ovary; ovules
numerous or 1 per locule, anatropous to pendent- VEGETATIVE ANATOMY. The nodes are unilacunar
orthotropous, unitegmic, tenuinucellate; style (Thomas 1960). Sections of the bifacial leaves
entire or apically branched into 3( 4) stylar show about two layers of palisade parenchyma.
branches. Fruit indehiscent, 1-5-seeded, 3-5- Stomata are paracytic, anomocytic or anisocytic,
ribbed or indistinctly lobed, or a 3-loculicidal, confined to the abaxial surface and, in Clethra,
many-seeded capsule, usually pubescent, some- slightly raised above the surface. Secretory cells
times ± enclosed by the persistent calyx. Seeds (in are found in the petiole and in the phloem of the
Clethra) flattened or subtrigonal, ellipsoid to midrib. Crystals, particularly clustered ones, are
ovoid, irregularly angled to winged, seed coat abundant (Metcalfe and Chalk 1950; Thomas 1960;
absent (Purdiaea) or with foveolate-reticulate Vales et al. 1988).
impressions or inconspicuously prominent cells; The wood exhibits a comparatively primitive
endosperm cellular, copious, fleshy; embryo short, structure. Vessel elements are predominantly soli-
straight, cylindrical. tary, angular, narrow (diameter mostly 25-50 )lm),
Two genera with about 95 species, mostly from and in Clethra twice as long as in Purdiaea (Record
tropical montane America and SE Asia (to India), 1932), sometimes with fine, spiral thickening in
some in temperate North America and Asia, one the tips. Perforation plates are simple or scalari-
species on Madeira. form with many bars (Thomas 1960; Giebel and
Dickison 1975). As in most Ericales, vestured pits
VEGETATIVE MORPHOLOGY. Tropical Clethraceae are absent (Jansen et al. 2001). Parenchyma is
are generally evergreen, temperate species are paratracheal or apotracheal and diffuse. Rays are
70 J. V. Schneider and C. Bayer
uni- to multiserial, up to 6 cells wide (Record 1932;

Vales et al. 1988). Fibers and moderately long to
long fiber-tracheids are present.
Kubota et al. (2001) reported the formation of
arbuscular mycorrhiza in Clethra.
INFLORESCENCES. In mature evergreen species of

Clethra, each branchlet usually produces an inflo-
rescence, whereas in deciduous species only the
vigorous shoots develop inflorescences. Inflores-
cences can be terminal and axillary, even on the
same individual. The basic type appears to be an
open raceme, which can be variously aggregated to c
form panicle-, fascicle-, or umbel-like inflores-
cences. Some Purdiaea have very short pedicels,
which may occasionally lead to dense spiciform
inflorescences. Each flower is sub tended by a bract,
which is sometimes broad and showy or even
almost foliose.
FLOWER STRUCTURE. The flowers are basically

pentamerous and perfect, but some Clethra exhibit
functionally unisexual flowers and gynoedioecy
caused by reduced anthers and pollen production.
In Clethra, the petals arise in the same quincuncial
succession as the sepals (Nishino 1983). In some
species the petals are basally coherent or connate D
up to the lower third, although easily separable
along a marked line (Sleumer 1967). A surface Fig. 22. Clethraceae. Purdiaea nutans. A Young flower, part of
analysis revealed the petals to be irregularly perianth removed. B Flower at anthesis with inverted anthers.
C Stamens, from bud (left) and open flower (right). DApical
striate, the adaxial side being more distinctly dif- part of inverted anther, showing pores and filament attach-
ferentiated (Christensen and Jensen 1998). ment. (Gustafsson 1992)
In Purdiaea, the outer staminal whorl is
alternipetalous, the androecium being diploste-
monous throughout the entire development
(Thomas 1960). According to Leins (1964), both EMBRYOLOGY. The data refer to Clethra. The
staminal whorls of Clethra originate from a single anthers are tetrasporangiate, with a fibrous
circle. The alternipetalous stamens are initiated endothecium, two ephemeral layers and a secre-
before the alternisepalous ones but become tory tapetum. Simultaneous divisions of the
pushed towards the centre by the petal margins. microspore mother cells follow meiosis; the
Thus, the androecium is diplostemonous in origin tetrads are tetrahedral. Ovules are unitegmic
and only secondarily appears obdiplostemonous. with a single layer of integumentary cells. The
In both genera, the anthers are extrorse in bud but megagametophyte develops according to the
become inverted at anthesis. As a consequence, the Polygonum type (Kavaljian 1952). Endosperm is
original base with the pore-like clefts becomes the cellular throughout its development; chalaza!
top, resulting in an introrse position (Clethra: and micropylar haustoria are weakly developed
Kavaljian 1952; Purdiaea: Stahl1992, Fig. 22). The (Johri et al. 1992). Embryogeny corresponds to
two pollen sacs of each theca merge into a single the Asterad type (Anderberg 1993).
cavity before the pollen is shed, which often occurs
before anthesis. POLLEN MORPHOLOGY. The pollen monads are
Brown (1938) reported nectar secretion from tricolporate and oblate to spheroidal. The exine is
the lower part of the ovary and speculated that the psilate, tectate with obscure stratification. The
secretory region of Clethra can be compared to the sexine is thinner than the nexine, the tectum is
disk of Ericaceae. psilate or rugulate, and the infratectum granular
(Erdtman 1952; Alvarado and Ludlow-Wiechers
Clethraceae 71
1982; Barros and Barth 1994; Zhang and considered mainly based on the (analogous)
Anderberg 2002). agreement in anther inversion at anthesis. On the
other hand, the trimerous ovary led to specula-
KARYOLOGY. The few data available suggest a tions about relationships with Rhododendroideae
basic chromosome number of n = 8 for Clethra. (see Sleumer 1967).
A polyploid series with diploid (Clethra arb ore a, Based on the pentacyclic, hermaphroditic,
C. lanata), tetraploid (C. alnifolia), and decaploid choripetalous, hypogynous flowers and the well-
(C. barbinervis) species is discerned (Hagerup developed endothecium, Clethra was considered
1928; Kyhos 1965; Tanaka and Oginuma 1980). to represent a basal clade within Ericales (Sleumer
1967). In a cladistic analysis based on morphology
FRUIT AND SEED. The fruit is indehiscent or a and anatomy (Anderberg 1993), Clethra was found
three-valved loculicidal capsule that may be more sister to Ericaceae s.l. and distinguished by its 3-
or less enclosed by the persistent calyx. Only few carpellate gynoecium and Asterad-type embryo-
of the numerous ± anatropous ovules per locule geny. Sequence analyses of various genes yielded
develop into seeds in Clethra, whereas in Purdiaea different topologies, with Clethra being either
there is a single, pendent, orthotropous ovule per sister to Styracaceae or Theaceae (Olmstead et al.
locule; most of these solitary ovules grow to a 1993; Savolainen, Chase et al. 2000; Savolainen, Fay
mature seed. The seeds are exotestal-theoid. et al. 2000) or sister to Cyrillaceae/Ericaceae (Kron
Mature seeds of Clethra are subovoid or irregu- 1996; Savolainen, Fay et al. 2000; Soltis et al. 2000).
larly angular to subtrigonous and show a foveo- This last placement is supported by a molecular
late-reticulate seed coat or are flat and more or less study that includes Purdiaea (Anderberg and
winged. In Purdiaea, due to vacuolization during Zhang 2002).
ripening, the seed coat is lacking in mature seeds.
The cotyledons are short and embedded in the DISTRIBUTION AND HABITATS. Clethra has a rare,
copious cellular (Purdiaea) or in the oily and pro- disjunct distribution. One centre is tropical SE
teinaceous endosperm (Clethra). The small, fove- Asia, with some species extending into temperate
olate or winged seeds may be wind-dispersed. In China, Japan and India. Another centre is tropical
Purdiaea the naked seeds are most likely dispersed Central and South America, with a southernmost
by wind with the entire fruit, the sepals function- limit at about 29° in SE Brazil. Two species are
ing as wings (Berazafn and Rodriguez 1992). known from SE North America, and one from
Madeira. Such a distribution pattern is known for
PHYTOCHEMISTRY. Hegnauer (1964, 1989) few other plant genera, such as Persea. Sleumer
reported high contents of galli- and ellagitannins (1967) postulated an earlier development from a
as well as proanthocyanidins in twigs, leaves and tropical mountain flora, although it remained
inflorescences of Clethra. Triterpenes and waxes unclear whether this was situated in Asia or in
were found in the cortex, and urolic acid in the America. The Macaronesian species seems to be
fruits of C. barbinervis. most closely related to the tropical American ones,
indicating an old transatlantic connection.
SuBDIVISION. Until recently, Clethraceae were Purdiaea extends from Belize to Peru with the
recognized as a monogeneric family. The recent centre in Cuba.
inclusion of Purdiaea, formerly assigned to Cyril- Clethra prefers acid soils. Some species grow in
laceae, is well supported by morphological and swamps; there are also records from volcanic soils
molecular data (Anderberg and Zhang 2002). and limestone. The tropical species are predomi-
A subdivision of Clethra was based on the length nantly found in humid montane habitats, reaching
of stamens and style, distribution and deciduous- their upper altitudinal limit at about 3800 m in
ness (De Candolle 1839). Sleumer (1967) found China as well as in the Andes. The few species
these criteria not useful and retained only sections reported from tropical lowlands may all be in sec-
Clethra and Cuellaria, based on seed shape and ondary vegetation. Conversely, the temperate
structure of the seed coat. The first comprises the species are generally found in lowlands or lower
Asian and temperate American, the second the montane vegetation. Clethra species are light-
tropical American and Macaronesian species. demanding and thus well-adapted invaders of
cleared areas and secondary forests.
AFFINITIES. A placement in Ericales is gen-
erally accepted. Relationships with Ericaceae- PALAEOBOTANY. Fossil records assigned to
Arbutoideae, especially the tribe Arbuteae, were Clethra and the extinct genus Disoclethra would
72 J. V. Schneider and C. Bayer
point to a widespread occurrence of the family in outer whorl alternipetalous; disk hypogynous;
Europe, with the first records dating back to the ovary (3-)5-locular; ovules 1 per locule, pendent,
Middle Eocene (Friis 1985; Knobloch and Mai orthotropous; style unbranched. Fruit indehiscent,
1986; Mai 2001). In the Japanese flora, records of I-S-seeded, 3-5-ribbed or indistinctly lobed; seed
Clethra are from the Late Miocene and Pliocene coat absent, endosperm copious, embryo straight,
(Ozaki 1991). Other fossils from the Tertiary were cylindrical. About 12 spp., Belize to Peru, most
reported for East Asia and North America, but diverse in Cuba.
some of them are doubtful as to their identity
(Sleumer 1967).
ECONOMIC IMPORTANCE. Economic importance
is very minor. Several temperate Clethra species Alvarado, J.L, Ludlow-Wiechers, B. 1982. Cata!ogo palinologico
are cultivated as ornamental plants; especially C. para Ia flora de Veracruz, no.10. Familia Clethraceae. Biotica
alnifolia is popular for its fragrant flowers. In 8: 619-629.
Anderberg, A.A. 1993. Cladistic interrelationships and major
China, the leaves of Clethra barbinervis are eaten. clades of Ericales. Pl. Syst. Evol. 184: 207-231.
The timber of some species is used for furniture Anderberg, A.A., Zhang, X. 2002. Phylogenetic relationships of
and constructions, mostly in Central and South Cyrillaceae and Clethraceae (Ericales) with special empha-
America (Sleumer 1967). sis on the genus Purdiaea Planch. Org. Divers. Evol. 2:
127-136.
Barros, M.A., Barth, O.M. 1994. Catalogo sistematico do polen
KEY TO THE GENERA das plantas arboreas do Brasil meridional, vol. 28. Burser-
aceae e Clethraceae. Revista Brasil. Bioi. 54: 317-322.
Sepals± equal in size; ovary 3(4)-locular; disk lacking; petals
Berazain, R., Rodriguez, S. 1992. Novedades taxonomicas en el
white, rarely pinkish to cream; fruit capsular 1. Clethra
genero Purdiaea Planchon (Cyrillaceae) en Cuba. Revista
- Sepals distinctly unequal, the outer much larger than the
Jard. Bot. Nac. Univ. Habana 13: 21-25.
inner; ovary (3- )5-locular, disk hypogynous; petals pink to
Brown, W.H. 1938. The bearing of nectaries on the phylogeny
violet; fruit indehiscent 2. Purdiaea
of flowering plants. Proc. Am. Philos. Soc. 79: 549-595.
Candolle, A.P. de 1839. Prodromus systematis naturalis, Vol. 7
(2). Paris: Treuttel & Wurtz,, pp. 588-590.
1. Clethra L. Christensen, K.I., Hansen, H. V. 1998. SEM -studies of epidermal
Clethra L., Sp. Pl. 396 (1753): Sleumer, Bot. Jahrb. Syst. 87: patterns of petals in the angiosperms. Opera Bot. 135: 1-91.
36-175 (1967), rev. Drude, 0. 1889. Clethraceae. In: Engler & Prantl, Die
natiirlichen Pflanzenfamilien IV, 1. Leipzig: Engelmann,
pp. 1-2.
Trees and shrubs with entire to serrate leaves. Erdtman, G. 1952. See general references.
Racemes single or in panicle-, fascicle-, or umbel- Friis, E.M. 1985. Angiosperm fruits and seeds from the Middle
like aggregates. Flowers actinomorphic, 5( 6)- Miocene of Jutland, Denmark. Bioi. Skr. Dansk Vid. Selsk.
merous, hermaphroditic, rarely functionally 24(3): 1-165.
Giebel, K.P., Dickison, W.C. 1975. Wood anatomy of
androdioecious; outer stamens alternisepalous; Clethraceae. J. Elisha Mitchell Sci. Soc. 91: 17-26.
disk absent; ovary 3-locular; ovules many per Gustafsson, C. 1992. Cyrillaceae. In: Harling, G., Andersson, L.
locule; style entire to branched. Fruit a 3-loculici- (eds.) Flora of Ecuador no. 45, pp. 29-34.
dal, many-seeded capsule; seeds irregularly angled Hagerup, 0. 1928. Morphological and cytological studies of
Bicornes. Dansk Bot. Ark. 6(1): 1-27.
to winged, seed coat with foveolate-reticulate
Hegnauer, R. 1964, 1989. See general references.
impressions or inconspicuously prominent cells; Jansen, S., Baas, P., Smets, E. 2001. Vestured pits: their occur-
endosperm fleshy, embryo short, straight, cylin- rence and systematic importance in eudicots. Taxon 50:
drical. Eighty-three spp., tropical America and SE 135-167.
Asia (to India), temperate North America and Johri, B.M., Ambegaokar, K.B., Srivastava, P.S. 1992. See general
references.
Asia, one sp. on Madeira. Kavaljian, L.G. 1952. The floral morphology of Clethra alnifo-
lia with some notes on C. acuminata and C. arborea. Bot.
Gaz. 113: 392-413.
2. Purdiaea Planch. Fig. 22 Knobloch, E., Mai, H.D. 1986. Monographie der Friichte und
Purdiaea Planch., Lond. J. Bot. 5: 250 (1846); Thomas, Contrib. Samen in der Kreide von Mitteleuropa. Rozpr. ustred. list.
Gray Herb. 186: 1-114 (1960), rev.; Berazain & Rodriguez, Geol. 47: 1-219,56 pl.
Rev. Jard. Bot. Nac. 13: 21-25 (1992), Cuban spp. Kron, K.A. 1996. Phylogenetic relationships of Empetraceae,
Epacridaceae, Ericaceae, Monotropaceae, and Pyrolaceae:
evidence from nuclear ribosomal 18S sequence data. Ann.
Small trees or shrubs with sessile entire leaves. Bot. Lond. 77: 293-303.
Racemes simple. Flowers slightly zygomorphic, Kron, K.A., Chase, M.W. 1993. Systematics of Ericaceae, Empe-
hermaphroditic, 5-merous, the outer sepals con- traceae, Epacridaceae, and related taxa based upon rbcL
spicuously larger than the inner; stamens 10, the sequence data. Ann. Missouri Bot. Gard. 80: 735-741.
Clethraceae 73
Kubota, M., McGonigle, T.P., Hyakumachi, M. 2001. Clethra Rodriguez, S., Berazain, R. 1991. Caracterizaci6n de Ia ner-
barbinervis, a member of the order Ericales, forms arbus- vadura foliar en el genero Purdiaea Planch. (Cyrillaceae).
cular mycorrhizae. Can. J. Bot. 79: 300-306. Revista Jard. Bot. Nac. Univ. Habana 12: 69-73.
Kyhos, D.W. 1965. Documented chromosome numbers of Savolainen, V., Chase, M.W. eta!. 2000. See general references.
plants. Madrofio 18: 122-126. Savolainen, V., Fay, M.F. et a!. 2000. See general references.
Lechner, S. 1914. Anatomische Untersuchungen tiber die Gat- Sleumer, H. 1967. Monographia Clethracearum. Bot. Jahrb.
tungen Actinidia, Saurauia, Clethra und Clematoclethra Syst. 87: 36-175.
mit besonderer Beriicksichtigung ihrer Stellung im System. Soltis, D.E. et a!. 2000. See general references.
Beih. Bot. Zentralbl. 32: 431-467. Stahl, B. 1992. Cyrillaceae. In: Harling, G., Andersson, L. (eds.)
Leins, P. 1964. Entwicklungsgeschichtliche Studien an Ericales- Flora of Ecuador, vol. 45, pp. 29-34.
Bliiten. Bot. Jahrb. Syst. 83: 57-88. Tanaka, R., Oginuma, K. 1980. Karyomorphological studies on
Mai, D.H. 2001. Die mittelmiozanen und obermiozanen Clethra barbinervis and two allied species. J. Jap. Bot. 55:
Floren aus der Meuroer und Raunoer Folge in der Lausitz, 65-72.
Teil2: Dicotyledonen. Palaeontographica, Abt. B., 257(1-6): Thomas, J.L. 1960. A monographic study of the Cyrillaceae.
35-176. Contrib. Gray Herb. 186: 1-114.
Metcalfe, C.R., Chalk, L. 1950. See general references. Vales, M.A., Moncada, M., Machado, S. 1988. Anatomia com-
Nishino, E. 1983. Corolla tube formation in the Primulaceae parada de Clethraceae en Cuba. Revista Jard. Bot. Nac. Univ.
and Ericales. Bot. Mag. Tokyo 96: 319-342. Habana 9: 69-73.
Olmstead, R.G., Bremer, B., Scott, K.M., Palmer, J.D. 1993. A par- Yu, C.H., Chen, Z.L. 1990. Leaf architecture of the woody
simony analysis of the Asteridae sensu Jato based on rbcL dicotyledons from tropical and subtropical China. Oxford:
sequences. Ann. Missouri Bot. Gard. 80: 700-722. Pergamon Press.
Ozaki, K. 1991. Late Miocene and Pliocene floras in Central Zhang, X.-P., Anderberg, A.A. 2002. Pollen morphology in the
Honshu. Bull. Kanagawa Pref. Mus. Nat. Sci. Spec. Issue, ericoid clade of the order Ericales, with special emphasis on
244 pp. Cyrillaceae. Grana 41: 201-215.
Record, S.J. 1932. Woods of the Ericales, with particular refer-
ence to Schizocardia. Trop. Woods 32: 11-14.
74 R.H.M.J. Lemmens et a!.
Connaraceae
R.H.M.J. LEMMENS, F.J. BRETELER and C.C.H. JoNGKIND
Connaraceae R. Brown in Tuckey, Narr. Exped. Congo: 431 the Asiatic mainland). The family is largely
(1818).
restricted to lowland rain forest, but some species
occur in mountains and others in thickets in
Small trees, shrubs, or Hanas, evergreen or some- savannas.
times deciduous. Leaves alternate, imparipinnate,
trifoliolate or unifoliolate, exstipulate, exstipellate; VEGETATIVE MORPHOLOGY. Connaraceae are all
petiole pulvinate at base; petiolules entirely pulv- woody plants, mostly Hanas climbing by means of
inate; leaflets entire, opposite or not. Inflorescen- the winding ends of young branches, sometimes
ces axillary, sometimes clustered near the tip of additionally provided with leaves transformed
the branches, occasionally cauliflorous, usually into woody hooks. Several species are present as
paniculate, sometimes less branched and re- small shrubs in the undergrowth of the forest
sembling a raceme, or condensed. Flowers actin- and may hardly show any growth for many years
omorphic, 5(4)-merous, bisexual (unisexual), but, when the forest canopy opens, they rapidly
heterodistylous or heterotristylous (homosty- produce long shoots and become Hanas. However,
lous); pedicels usually jointed, prophylls 0; sepals all species of the genera Burttia, Ellipanthus,
imbricate or valvate, free or united at base (in Hemandradenia,]ollydora and Vismianthus do not
Rourea solanderi almost entirely fused in bud), show any lianescent tendencies and are shrubs or
caducous to persistent and sometimes accrescent small trees, rarely medium-sized trees up to 30m
in fruit; petals imbricate, free to connivent near the tall (Ellipanthus). A few Cnestis, Connarus and
base; stamens in 2 whorls, free or united at base, Rourea always have a shrub- or tree-like habit.
the outer antesepalous, longer than the inner ante- The main stem of lianescent species is usually
petalous ones, the latter sometimes staminodial, (sub)cylindrical in cross section, but some species
anthers dorsifixed, dehiscing lengthwise, introrse; (e.g. Rourea minor, Agelaea pentagyna) have a
gynoecium 1- or 5-carpellate, superior; carpels strongly ridged stem showing a number of con-
free, each carpel with a terminal stylodium and centric rings. These latter species have included
capitate stigma; ovules 2 per carpel, collateral, phloem. The bark is usually smooth, more rarely
nearly basal to nearly apical, anatropous to fissured, and in a few species strongly suberized
hemitropous, micropyle always directed upwards. ( Connarus suberosus, Rourea coccinea and R.
Fruit consisting of 1-5 follicles, dry or more or less orientalis). Lenticels are usually present on the
fleshy, usually dehiscing by a ventral suture, some- branchlets. Some species produce a red sticky
times along the dorsal side as well, rarely circum- ( Connarus, Agelaea gabonensis) or colourless,
scissile at the base (some Rourea), or indehiscent slimy (Rourea myriantha, R. solanderi) exudate.
(Hemandradenia, ]ollydora). Seed 1 per follicle, The leaves are pinnate, trifoliolate or unifolio-
sometimes 2 Uollydora); testa partly to entirely late, with the petiole pulvinate at base (Fig. 24B)
fleshy (sarcotesta), yellow to red; endosperm and the petiolules entirely so; this implies that uni-
present or not; hilum lateral to basal; cotyledons foliolate species have a pulvinus at the base and at
thin and flat to planoconvex; radicle situated at the the apex of the petiole. In Connarus, Heman-
margin of the cotyledons and then apical, ventral dradenia and Vismianthus, glands can be found on
or dorsal, or central and covered by the cotyle- the surface of the leaves. The tertiary venation
dons. x = 14, 16. is open except in Manotes where it consists of
A family comprising 12 genera and 110-200 a closed pattern of very fine, parallel veinlets
species (depending on the species concept used), (Fig. 25C).
almost exclusively found in the tropics. Only a VEGETATIVE ANATOMY. The indumentum con-
few species surpass the Tropic of Capricorn (e.g. sists of various hair types: multicellular (Cnestis,
Cnestis polyphylla in southern Africa) and the Connarus, ]ollydora, Manotes and Rourea) or uni-
Tropic of Cancer (e.g. some Connarus species on cellular glandular hairs (Manotes), unicellular,
Connaraceae 75
!-armed non-glandular hairs (all genera except are usually imbricate and may have glandular
]ollydora), unicellular 2-branched hairs (Burttia, hairs or glands on their surface. Manotes flowers
Connarus, Hemandradenia, some species of have a distinct androgynophore (Fig. 25E). The
Rourea, Vismianthus) and chained, 2-armed uni- inner, antepetalous whorl of stamens is shorter or
cellular hairs (Connarus). For more details, see subequal to the outer antesepalous whorl, and is
Jongkind (in Breteler 1989). Stomata are paracytic staminodial in Ellipanthus, Hemandradenia and in
in Cnestis and in some species of Rourea, aniso- some Connarus spp.
cytic including helicocytic in Agelaea, Cnestidium A wide diversity in forms of heterostyly is
and some species of Rourea, anomo-cyclocytic in known to occur in Connaraceae (Figs. 23-25). Het-
Burttia, Connarus, Ellipanthus, Hemandradenia, erotristyly is found in most Agelaea, ]ollydora and
Manotes, Vismianthus and some species of Rourea, Manotes spp., heterodistyly (either with 10 or 5
and bicyclic in ]ollydora. A papillose lower epider- fertile stamens) in many genera and species, and
mis is present in many species of Cnestis, Pseudo- homostyly in Cnestis ferruginea. The flowers of
connarus and Rourea. Mucous cells are sometimes several species show a form of heterostyly transi-
present in the upper epidermis. Sometimes a tional between tristyly and distyly. Ellipanthus
pattern of more or less parallel lines is present beccarii and possibly some American Connarus
in the lower epidermis (in Rourea calophylla, R. spp. are dioecious; male plants have rudimentary
camptoneura and Manotes macrantha). The sec- pistils, female plants sterile stamens (lacking
ondary xylem is anatomically rather uniform and pollen).
moderately highly specialized. The wood is
diffuse-porous, the perforations are simple and EMBRYOLOGY. Ovules are hemi-anatropous,
slightly oblique to transverse, the ground tissue bitegmic and crassinucellate, and have a pro-
usually consists of libriform fibres together with nounced preraphe; the micropyle is formed by
fibre-tracheids and/or vasicentric tracheids or the outer integument, and a funicle is absent. A
only libriform fibres, parenchyma is mostly absent primary parietal cell divides at least once. The
or scanty, rays are usually exclusively uniseriate or embryo sac develops according to the Polygonum
biseriate, crystals are often present (usually in type, endosperm development is nuclear and the
fibres), latex tubes generally absent (but present tissue becomes cellular only after the formation of
in Connarus), vertical intercellular canals often numerous free nuclei (Mauritzson 1939; Corner
present in libriform tissue. For more details, 1976).
see Dickison (1972b) and den Outer and van
Veenendaal (in Breteler 1989). POLLEN MORPHOLOGY. The pollen is rather
uniform, with the exception of that of ]ollydora.
INFLORESCENCE STRUCTURE. The inflorescences Pollen in the 5-carpellate tribes Cnestideae and
are basically axillary panicles. They often arise Manoteae is spheroidal or subspheroidal and
from the axils of reduced leaves near the apex of small (20-30 11m), with a thin wall and a finely
young branches, giving the impression of a com- reticulate ornamentation. In Connareae it is sub-
pound terminal inflorescence, particularly in oblate to subprolate and larger (30-40 11m), with a
Agelaea, Connarus, Manotes, Pseudoconnarus and thicker wall and more coarsely reticulate.]ollydora
some Cnestis and Rourea species. Such flowering (Jollydoreae) shows a completely different pollen
branches always end in a vegetative bud which type; its pollen is large (40-50 11m), oblate and
sometimes continues the branch after flowering tetracolpate, with a thick wall and a coarse reticu-
but usually shrivels. The panicles are often lum, a most unusual type in Dicotyledons, other-
reduced to raceme-like or strongly condensed wise only known from Impatiens. For details, see
inflorescences (e.g. in Hemandradenia mannii and Dickison (1979) and van den Berg (in Breteler
Rourea obliquifoliolata). Cauliflory with large 1989).
nodose proliferations on the stem occurs in folly-
dora, some Cnestis spp., Rourea calophylloides and KARYOLOGY. There is karyological information
Manotes macrantha. on African taxa only. The chromosome numbers
known indicate that there are two groups, one
FLOWER STRUCTURE. The flowers have 5 sepals, 5 comprising taxa with 2n = 28 and the other taxa
petals, 10 stamens in 2 whorls and either 1 or 5 with 2n = 32. The first group comprises species
carpels; they usually have a jointed pedicel. The from the tribes Cnestideae and Manoteae, the
sepals are distinctly valvate (Manotes) to distinctly second group species from the tribe Connareae.
imbricate (most species), or in between. The petals The somatic chromosomes of the Connareae have
76 R.H.M.J. Lemmens eta!.
lengths varying in the range 0.5-1 1-1m, whereas less ventrally or dorsally (relative to the follicle)
those in Cnestideae and Manoteae are 1-2!-lm and, in some African Connarus (C. congolanus, C.
long. It appears that the karyotype in Connareae staudtii), the radicle is found in the centre of the
differs from the other tribes investigated in both seed and quite hidden between the peltate
number and length of the chromosomes. For more cotyledons.
details, see Arends (in Breteler 1989).
DISPERSAL. The strikingly coloured, dehiscing
POLLINATION. Observations on pollination are follicles showing the often two-coloured seed-coat
very scarce. Many species are sweet scented and seem to advertise the seeds to birds such as fruit
probably pollinated by insects. Bees have been pigeons and hornbills, which are probably the
observed visiting the flowers. main dispersers. Most Connaraceae are Hanas of
the canopy or savannah shrubs, and birds will
FRUIT AND SEED. In 5-carpellate species, usually readily detect the fruits. The fruits of Heman-
some carpels fail to produce seeds; often only a dradenia and ]ollydora, which are both under-
single follicle is formed. The follicles are mostly storey treelets in the rain forest, are indehiscent;
dehiscent, but indehiscent in Hemandradenia and they may escape the attention of birds and are
]ollydora, and usually orange to red. They are more likely to be dispersed by small mammals.
glabrous to hairy outside, sometimes with long,
stinging hairs (some Cnestis spp.), and usually PHYTOCHEMISTRY. There are indications that
glabrous inside except in Cnestis and several many species of the family contain chemically
Connarus spp. interesting substances. The seeds and sometimes
Usually only one of the two ovules per carpel also other parts of the plants (particularly roots)
develops into a seed, but in ]ollydora often both are reported as poisonous. They are often used as
ovules mature. The seeds are often two-coloured dog poison, and they seem to be poisonous for
with a black, glossy testa strongly contrasting with other animals like sheep, goats and rats as well.
the yellow to orange sarcotesta, as well as with the All parts of Cnestis polyphylla are reported to con-
follicle. In most species the larger part of the seed tain a neurotoxical compound which has been
is squeezed out of the follicle that retains a firm named glabrin (Jeannoda et al. 1985); this com-
grip on the part that remains inside (Fig. 24D), but pound is methionine sulphoximine (S-(3-amino-
in Manotes and Vismianthus the seed is almost 3-carboxypropyl)-S-methyl sulphoximine), and
completely released from the follicle, remaining also has ichthyotoxic properties. Methionine
attached by means of a threadlike appendix of the sulphoximine is probably also the toxic compound
sarcotesta, which remains wedged in the narrowly in several other species (e.g. of Agelaea and
funnel-shaped, bottom part of the follicle. In these Rourea). However, in most species the toxic com-
latter two genera, endocarp and exocarp separate pounds have not yet been identified.
from each other at maturity. Roots of some Connarus spp. were demonstrated
In the seed-coat the testa is unlignified but the to contain rapanon, embelin and bergenin, and
outer epidermis is developed as a thickened pal- roots of Rourea minor ~-sitosterol, its glucoside,
isade. It becomes sarcotestal in the raphe-chalaza hentriacontan and mesoinosit.
region or more or less all over the seed (Heman- Tannins are reported as common, and both
dradenia, folly dora, Manotes macrantha and some hydrolysable and condensed tannins may be
Rourea spp.); the sarcotesta is yellow to red, pulpy, present. Gallic acid has been isolated from fruits
and in Connarus often has a crenulate border or and leaves of South American Connarus spp. In
short limb. The tegmen usually is crushed or leaf extracts from the Asiatic Connarus semide-
disintegrates early, except for an outer layer of candrus, myricetin, quercetin, kaempferol, del-
lignified fibres. phinidin and cyanidin have been identified.
The structure of the seeds concerning
endosperm and cotyledons is very variable. The SUBDIVISION AND RELATIONSHIPS WITHIN THE
cotyledons can be flat and thin and embedded FAMILY. Planchon (1850) divided the family into
in abundant endosperm (Manotes, some Cnestis two tribes, the Connareae and the Cnestideae,
spp.) to planoconvex without endosperm based on differences in the aestivation of the
(Agelaea, Connarus, ]ollydora, most Rourea), but sepals and the presence or absence of endosperm.
intermediate conditions are common. The radicle Gilg (1891) maintained these tribes and, in 1897,
is not always located near the micropyle. In some after the description of ]ollydora, he divided the
Connarus and Rourea spp. it is situated more or Connaraceae in two subfamilies, Connaroideae
Connaraceae 77
and Jollydoroideae. Schellenberg (1910) proposed Asia. This must be an old disjunction, as their large
a new subdivision of the family, distinguishing the seeds are not likely to be dispersed over long dis-
subfamilies Connaroideae (including ]ollydora) tances and have hardly been dispersed by man.
and Cnestoideae; he divided the former subfamily Connaraceae are largely restricted to the low-
into two tribes, Connareae and Roureeae, the latter land rain forest. More rarely they are found in
of which was again divided into two sub tribes. In mountain vegetation; some species grow in thick-
1938 Schellenberg reverted to Gilg's subdivision ets or in remnants of forests in savannas.
into two subfamilies, and the number of genera
reached 24. Subsequently the number of genera ECONOMIC IMPORTANCE. Representatives of Con-
was reduced by Leenhouts (1958), and even naraceae are little used by man. The seeds and
further by Breteler (1989) to 12. Lemmens (in roots of several species are toxic and some-
Breteler 1989) proposed a tribal classification times used to poison dogs and rats (particularly
which is followed in this treatment. Cnestis and Rourea spp.). Diluted in small doses,
they are occasionally used as medicine, mainly
AFFINITIES. Relationships of Connaraceae with against stomach-ache and dysentery (Agelaea
other families have been ambiguous for a long macrophylla, Cnestis ferruginea and several
time. Most often a close relationship with Legu- Rourea species), and as anthelmintic (Connarus
minosae and Rosaceae has been postulated, but africanus). Ellipanthus species that reach the
some authors considered Oxalidaceae, Anacar- size of medium-sized trees are sometimes cut
diaceae, Meliaceae and Dilleniaceae as related in South-East Asia for their timber. The fatty oil
families. The pinnate leaves and apocarpous, present in seeds of some Connarus spp. has been
pod-like fruits are reminiscent of Leguminosae, used in India in soap production.
and the exstipulate leaves and the presence of a
sarcotesta would emphasize affinities with KEY TO THE GENERA
Sapindaceae. However, any preference expressed
1. Flowers with 5 carpels 2
as to the position of Connaraceae in any of the - Flowers with 1 carpel 8
orders Rosales, Sapindales and Dilleniales left 2. Follicles (nearly) glabrous 3
them with part of their characters in disagreement - Follicles covered with a continuous indumentum of (some-
with that position. times glandular) hairs 4
Phylogenies based on plastid gene sequences 3. Seeds with abundant endosperm; leaves always 3-foliolate;
America 9. Pseudoconnarus
place Connaraceae as sister of Oxalidaceae in an - Seeds with little or no endosperm; leaves pinnate (but
order Oxalidales (=Cunoniales) (APG 1998). The branches with unifoliolate or 3-foliolate leaves may occur);
two families share the presence of heterostyly, circum tropical 12. Rourea
the benzoquinone rapanone, exotegmic seeds and 4. Flowers with a distinct androgynophore; Africa
7.Manotes
some other morphological characters that have - Flowers without a distinct androgynophore 5
been listed by Nandi et al. (1998). 5. Follicles hairy inside; Africa, Asia 8. Cnestis
- Follicles glabrous inside 6
DISTRIBUTION AND HABITATS. The Connaraceae 6. Sepals in fruit strikingly accrescent; circumtropical
are a pantropical family of which only a few 12.Rourea
- Sepals in fruit inconspicuous or caducous 7
species are found beyond the tropics. The largest 7. Leaves 3- or 5-foliolate; Africa, Asia 10. Agelaea
number of genera (10), representing all four tribes, - Leaves pinnate with variable number of leaflets within one
is found in Africa, followed by Asia (6) and then plant; America 11. Cnestidium
America (4), each with representatives of two 8. Leaves unifoliolate 9
tribes. The main centre of distribution is Central - Leaves with more than one leaflet (except sometimes
uppermost leaves on flowering branches) 12
Africa, particularly Cameroon and Gabon. Two 9. Follicles glabrous; Africa, Asia 5. Vismianthus
genera, Cnestidium and Pseudoconnarus, are - Follicles velutinous 10
endemic to tropical America, where Connarus and 10. Petals glabrous; follicles dehiscent; seeds with abundant
Rourea (which are also well represented in tropi- endosperm, cotyledons thin and flat; Africa 4. Burttia
- Petals hairy; follicles dehiscent or indehiscent; seeds with
cal Asia) are very rich in species. Some species
or without endosperm, cotyledons flat to planoconvex 11
have a very large area of distribution; for example, 11. Follicles dehiscent; testa for about 1/4 fleshy; Africa, Asia
Rourea minor is found in Africa, Madagascar, 2. Ellipanthus
islands of the Indian Ocean and the Pacific, and in - Follicles indehiscent; testa almost entirely fleshy; Africa
tropical Asia. In genera like Agelaea, Connarus, 3. Hemandradenia
12. Understorey treelet; petals and filaments glabrous; follicles
Ellipanthus, Rourea and Vismianthus, groups of indehiscent, 1-2-seeded; Africa 6. ]ollydora
closely related species are common to Africa and
78 R.H.M.J. Lemmens et al.
Small trees, shrubs or lianas. Leaves trifoliolate or

imparipinnate (upper leaves unifoliolate ); leaflets
often glandular punctate. Inflorescence panicu-
late, often pseudoterminal. Flowers bisexual (uni-
sexual), 5-merous, heterodistylous; sepals, petals,
filaments and style usually glandular, carpel 1.
Follicle often stipitate, often glabrescent outside,
glabrous or pilose and often glandular inside; seed
solitary, attached to the ventral side of the follicle,
below the hilum with sarcotesta, endosperm 0,
J
cotyledons thick, planoconvex, radicle apical,
dorsal or almost in the centre of the seed between
the cotyledons. About 75 spp., circumtropical.
.
~
.
:
2. Ellipanthus Hook.£.
F "---· Ellipanthus Hook.f. in Benth. & Hook.f., Gen. Pl. 1, 1: 434
(1862); Leenhouts, Flora Males. I, 6: 520 (1958); Lemmens in
Breteler (ed.) The Connaraceae: 268 (1989); Lemmens, Bull.
Mus. Natl. Hist. Nat. Adansonia 14: 99 (1992).
Shrubs or small trees. Leaves unifoliolate. Inflores-

cence a small panicle. Flowers bisexual or unisex-
ual, 4-5-merous, heterodistylous, petals pilose
outside and usually also inside, filaments and
style pilose, carpel I. Follicle more or less stipitate,
densely tomentose outside, glabrous inside, dehis-
cent; seed 1(2) per follicle, attached to the ventral
side of the follicle, basal part with sarcotesta,
endosperm thin, cotyledons thick, radicle apical.
About six spp., East Africa, Madagascar and trop-
ical Asia.
Fig. 23. Connaraceae-Connareae. Connarus longistipitatus.
A Flowering branchlet. B Leaflet. C Detail leaf underside. D 3. Hemandradenia Stapf
Flower. E Flower, sepals and petals partly removed. F Short and
long stamen. G Follicle. H Detail inner side of follicle. I Seed. Hemandradenia Stapf, Bull. Misc. Inform. (Kew Bull.): 288
J Seed, vertical section. Drawn by W. Wessel-Brand (Orig.) (1908); Eimunjeze in Breteler (ed.) The Connaraceae: 275
(1989).
- Treelet, shrub or Iiana; petals and/or filaments with many

(sometimes glandular) hairs; follicles dehiscent, 1-seeded;
circumtropical 1. Connarus cence a glomerule or panicle. Flowers bisexual,
{4-}5(-7)-merous, heterodistylous, petals pilose
TRIBES AND GENE RA OF CONN ARACE AE outside and usually also inside, filaments and style
pilose, carpel 1. Follicle not stipitate, densely
I. TRIBE CONNAREAE Planchon (1850). tomentose outside, glabrous inside, indehiscent.
Seed solitary, attached to the ventral side of the
Lianas, shrubs or small trees; leaves unifoliolate to follicle, sarcotesta completely covering the seed,
pinnate; carpel 1; follicle dehiscent or (Heman- endosperm thin to copious, cotyledons thin or
dradenia) indehiscent with 1 ventrally attached thick, radicle apical. Two spp., West and Central
seed; pollen tricolporate. Africa.
1. Connarus L. Fig. 23 4. Burttia Baker f. & Exell Fig. 24

Connarus L., Sp. Pl.: 675 (1753); Leenhouts, Flora Males. I, 6: Burttia Baker f. & Exell, J. Bot. 69: 249 (1931); Breteler &
525 (1958); Forero, Flora Neotropica 36:36 (1983); Lemmens Brouwer in Breteler (ed.) The Connaraceae: 169 (1989).
in Breteler (ed.) The Connaraceae: 239 (1989).
Connaraceae 79

cence 1-3-flowered. Flowers bisexual, (4-)5-
merous, heterodistylous, petals, filaments and
style glabrous, carpel 1. Follicle not stipitate,
densely tomentose outside, glabrous inside, dehis-
cent. Seed solitary, attached near the top of the
ventral side of the follicle, sarcotesta covering
whole length of the seed on one side, endosperm
copious, cotyledons thin and narrow, radicle
apical. One sp., B. prunoides Baker f. & Exell, East
Africa.
5. Vismianthus Mildbr.
Vismianthus Mildbr., Notizbl. Bot. Gart. Mus. Berlin 12 (115):
706 (1935); Breteler & Brouwer in Breteler (ed.) The Con-
naraceae: 369 (1989).
Schellenbergia Parkinson (1936).
Shrubs or small trees. Leaves unifoliolate, glandu-

lar punctate. Inflorescence racemose. Flowers
bisexual, 5-merous, heterodistylous, petals
glabrous but glandular, filaments glabrous, style
hairy, carpel 1. Follicle more or less stipitate,
(sub)glabrous outside, glabrous inside, dehis-
cent, inner and outer pericarp separating. Seed
solitary, attached to the ventral side of the follicle,
sarcotesta fimbriate or undulate with a long
appendage and covering the base of the seed,
endosperm very thin, cotyledons thin and folded
or thick and plano-convex, radicle apical. Two
spp., one in Tanzania, the other in Myanmar.
II. TRIBE JoLLYDOREAE (Gilg) Lemmens (1989). B

F(;)
Fig. 24. Connaraceae-Connareae. Burttia prunoides. A Flow-
Treelets, usually unbranched; leaves large, pinnate; ering branchlet. B Fruiting branchlet. C Flower, sepals and
carpel 1; follicle indehiscent, seeds often 2, ven- petals partly removed. D Dehiscing fruit with seed. E Seed
trally attached; pollen tetracolpate. with sarcotesta. F Transverse section of seed. G Section of seed
lengthwise, sarcotesta removed. Drawn by W. Wessel-Brand
(Orig.)
6. ]ollydora Pierre
]ollydora Pierre, unprinted drawing by Delpy (1895); Breteler
& van Ziel in Breteler (ed.) The Connaraceae: 284 (1989). III. TRIBE MANOTEAE Lemmens (1989).
Usually unbranched treelets. Leaves impari- Lianas; leaves pinnate, leaflets striately veined;
pinnate. Inflorescence racemose, in cauliflorous androgynophore +; carpels 5; follicles dehiscent,
clusters or in axils of leaves. Flowers bisexual, 5- seed 1, ventrally attached; pollen tricolporate.
merous, heterotristylous, petals and filaments
glabrous, style sometimes with a few hairs, carpel 7. Manotes Planchon Fig. 25
1. Follicle stipitate or not, glabrous or glabres-
cent outside, glabrous inside, indehiscent; seeds 1 Manotes Planchon, Linnaea 23: 438 (1850); Jongkind in
or 2, attached to the ventral side of the follicle, Breteler (ed.) The Connaraceae: 294 (1989).
sarcotesta almost completely covering the seed,
endosperm 0, cotyledons thick and almost horny, Lianas or scandent shrubs. Leaves imparipinnate,
radicle apical. Three spp., Central Africa. leaflets with a dense pattern of very fine parallel
80 R.H.M.J. Lemmens et al.
IV. TRIBE CNESTIDEAE Planchon (1850).
Shrubs or lianas; leaves pinnate or trifoliolate;

carpels 5; follicles dehiscent follicles with 1 seed
basally attached; pollen tricolporate.
8. Cnestis A.L. Juss.

Cnestis A.L. Juss., Gen. Pl.: 374 (1789); Leenhouts, Flora
Malesiana I, 6: 497 (1958); Lemmens in Breteler (ed.) The
Connaraceae: 174 (1989).
Lianas, climbing shrubs (small trees). Leaves

imparipinnate. Inflorescence a panicle, raceme or
fascicle, axillary or cauliflorous. Flowers bisexual,
5-merous, heterodistylous or sometimes more or
less homostylous, petals glabrous or with some
hairs outside, filaments glabrous, styles often hairy
near base, carpels 5. Follicle not or indistinctly
stipitate, densely tomentose, sometimes also with
long, rigid hairs outside, velutinous, sometimes
with rigid hairs inside, dehiscent; seed solitary,
attached to the base of the follicle, basal part with
sarcotesta, endosperm copious, cotyledons thin,
radicle apical. Thirteen spp., 12 in tropical Africa
and one in tropical Asia.
9. Pseudoconnarus Radlk.
Pseudoconnarus Radlk., Sitzb. Math.-Phys. Cl. Akad. Miinch.
16: 356 (1886); Forero, Flora Neotropica 36: 130 (1983).
Fig. 25. Connaraceae-Manoteae. Manotes macrantha. A Flow- Lianas. Leaves trifoliolate. Inflorescence pan-
ering branchlet. B Leaflet. C Detail of leaflet. D Flower. E
Flower, sepals and petals partly removed. F Petal. G Branchlet
iculate. Flowers bisexual, 5-merous, more or less
with fruits. H Follicles. Drawn by Wessel-Brand (Orig.) distinctly heterodistylous, petals glabrous but
glandular, filaments glabrous, styles glabrous or
hairy, carpels 5. Follicle not stipitate, glabrous
outside and inside, dehiscent; seed solitary,
veinlets. Inflorescence racemose or paniculate. attached to the base of the follicle, endosperm
Flowers bisexual, 5-merous, heterotristylous or copious, cotyledons thin, radicle apical. Five spp.,
heterodistylous, with distinct androgynophore; tropical South America.
petals, filaments and style hairy, often also with
glandular hairs; carpels 5. Follicle more or less
stipitate, minutely velutinous to long-hairy 10. Agelaea Planchon
outside, glabrous inside, dehiscent, inner and outer Agelaea Planchon, Linnaea 23: 437 (1850); Leenhouts, Flora
pericarp splitting apart at maturity; seed solitary, Males. I, 6: 500 (1958); Jongkind in Breteler (ed.) The Con-
attached to the ventral side of the follicle, sar- naraceae: 136 (1989).
cotesta (almost) completely covering the seed and
with a thread-like appendix attaching the seed to Lianas. Leaves trifoliolate, sometimes 5-foliolate
the base of the follicle, endosperm copious, cotyle- and imparipinnate to almost palmately com-
dons thin, radicle apical. Four or five spp., humid pound. Inflorescence paniculate. Flowers bisexual,
tropical Africa. 5-merous, heterotristylous or heterodistylous,
petals and filaments glabrous, styles often hairy,
carpels 5. Follicle not or indistinctly stipitate,
velutinous outside, glabrous inside, dehiscent;
seed solitary, attached to the base of the follicle,
Connaraceae 81
basal part with sarcotesta, endosperm 0, cotyle- Selected bibliography

dons thick, planoconvex, radicle apical. Eight spp.,
tropical Africa and tropical Asia. APG (Angiosperm Phylogeny Group) 1998. See general
references.
Breteler, F.J. (ed.) 1989. The Connaraceae. A taxonomic study
11. Cnestidium Planchon with emphasis on Africa. Agric. Univ. Wageningen Papers
89-6, 403 pp.
Cnestidium Planchon, Linnaea 23: 439 (1850); Forero, Flora Corner, E.J.H. 1976. See general references.
Neotropica 36: 29 (1983). Dickison, W.C.1971.Anatomical studies in the Connaraceae 1:
carpels. J. Elisha Mitchell Sci. Soc. 87: 77-86.
Lianas. Leaves imparipinnate. Inflorescence panic- Dickison, W.C. 1972. Anatomical studies in the Connaraceae 2:
wood anatomy. J. Elisha Mitchell Sci. Soc. 88: 120-136.
ulate. Flowers bisexual, 5-merous, ±distinctly het- Dickison, W.C. 1973a. Anatomical studies in the Connaraceae
erodistylous, petals and filaments glabrous, styles 3: leaf anatomy. J. Elisha Mitchell Sci. Soc. 89: 121-138.
often hairy, carpels 5. Follicle not stipitate, pubes- Dickison, W.C. 1973b. Anatomical studies in the Connaraceae
cent outside, glabrous inside, dehiscent; seed soli- 4: the bark and young stem. J. Elisha Mitchell Sci. Soc. 89:
tary, attached to the base of the follicle, basal part 166-171.
Dickison, W.C. 1979. A survey of pollen morphology of the
with sarcotesta, endosperm scarce, cotyledons Connaraceae. Pollen Spores 21: 31-79.
thick, radicle apical. Two spp., Central and South Forero, E. 1976. Connaraceae. Flora Neotropica Monogr 36.
America. New York: New York Botanical Garden.
Gilg, E. 1891. Connaraceae. In: Engler & Prantl, Nat. Pfianzen-
fam. III, 3: 61-70.
12. Rourea Aublet Gilg, E. 1897. Connaraceae. In: Engler & Prantl, Nat. Pfianzen-
fam., Nachtrage 1: 189-190.
Rourea Aublet, Pl. Guiane 1: 467 (1775); Leenhouts, Flora Hegnauer, R. 1964, 1989. See general references.
Males. I, 6: 510 (1958); Forero, Flora Neotropica 36: 138 Jeannoda, V.L.R., Valisolalao, J., Creppy, E.E., Dirheimer, G.
(1983); Jongkind in Breteler (ed.) The Connaraceae: 310 1985. Identification of the toxic principle of Cnestis glabra
(1989). as methionine sulphoximine. Phytochemistry 24: 854-855.
Bernardinia Planchon (1850); Forero, Flora Neotropica 36: 25 Jongkind, C.C.H. 1991. A new section and a new species in
(1983). Agelaea Sol. ex Planch. (Connaraceae). Bull. Jard. Bot. Nat.
Roureopsis Planchon (1850); Leenhouts, Flora Males. I, 6: 505 Belgique 61: 71-75.
(1958). Leenhouts, P.W. 1958. Connaraceae. In: van Steenis, C.G.G.J.
(ed.) Flora Malesiana I, 5: 495-591.
Lianas, shrubs or small trees. Leaves trifoliolate or Lemmens, R.H.M.J. 1992. A reconsideration of Ellipanthus
(Connaraceae) in Madagascar and continental Africa, and a
imparipinnate, occasionally unifoliolate. Inflores-
comparison with the species in Asia. Bull. Mus. Nat!. Hist.
cence racemose or paniculate, sometimes nearly Nat. B Adansonia 14: 99-108.
globose. Flowers bisexual, 5-merous, heterodisty- Mauritzson, J. 1939. Contributions to the embryology of the
lous, petals glabrous or with some hairs apically, orders Rosales and Myrtales. Acta Univ. Lund 35: 1-121.
filaments glabrous, styles often hairy near base, Nandi, 0.1. et a!. 1998. See general references.
Planchon, J.E. 1850. Prodromus monographiae ordinis
carpels 5. Follicle not or indistinctly stipitate, Connaracearum. Linnaea 23:411-442.
glabrous or velutinous outside, glabrous inside, Schellenberg, G. 1910. Beitrage zur vergleichenden Anatomie
dehiscent; seed 1(2) per follicle, attached to the und zur Systematik der Connaraceen. Mitt. Bot. Mus. Univ.
base of the follicle, with a sarcotesta around the Zurich so, 158 pp.
basal part to almost completely covering the seed, Schellenberg, G. 1938. Connaraceae. In: Engler, A. (ed.) Das
Pfianzenreich IV: 127.
endosperm 0, cotyledons thick and plano-convex, Takhtajan, A. 1969. Flowering plants, origin and dispersal.
radicle apical to ventral. About 40-70 spp., Edinburgh: Oliver and Boyd.
circumtropical.
82 K. Kubitzki
Cornaceae
K. KUBITZKI
Cornaceae (Dumort.) Dumort., Anal. Fam. Pl.: 33, 34 (1829), the xylem of the stem and thence the roots. It is
nom. cons. likely that the swollen bases of aquatic tupelos also
Alangiaceae DC., Prodr. 3: 203 (1828).
Nyssaceae Juss. ex Dumort. (1829). provide intercellular space.
Mastixiaceae Calest. (1905). The complex branching pattern in Cornus was
Davidiaceae (Harms) H.L. Li (1955). analysed in a series of papers by Hatta (see Hatta
et al. 1999).
Trees or shrubs (rhizomatous halfshrubs). Hairs The leaves are alternate or opposite, mostly
simple, unicellular. Leaves alternate or opposite, constantly so for whole genera, but among the
simple, entire (dentate), exstipulate. Inflorescences otherwise opposite-leaved Cornus two species
many-flowered thyrso-panicles, heads, or few- have alternate leaves, and in Mastixia phyllotaxis
flowered cymes; flowers hermaphrodite to uni- changes from alternate to (sub)opposite, often in
sexual, actinomorphic, epigynous, mostly 4- or the same individual.
5-merous; calyx adnate to ovary with 4-5(-10)
lobes or teeth or obsolete; petals free (basally VEGETATIVE ANATOMY. Sertorius (1893) and
connate), imbricate or valvate, reduced or lacking Adams (1949) gave detailed descriptions of the
in pistillate flowers; stamens as many as and alter- anatomy of the family, and Noshiro and Baas
nating with petals, or in 2 isomerous whorls (in 1 (1998) studied the wood anatomy. The following
whorl of up to 40 stamens), mostly attached to or features are of particular interest. Stomates are of
around the edge of an epigynous nectary disk the ranunculid type. Non-glandular and glandular
(lacking in Davidia); filaments free; anthers hi- hairs are always 1-celled. Two-armed 1-celled hairs
thecate, dehiscing longitudinally; pistil 1-2( -9)- are found in Mastixia, Diplopanax, Cornus and in
locular; style with lobed or capitate stigma, or a slightly different form (with asymmetric arms)
with 2-3 style branches; ovary inferior, with 1 in one Alangium; in Cornus they are calcified. Ca-
pendulous ovule per locule; ovules anatropous oxalate is deposited in druses. The wood is of a
(hemitropous), unitegmic and crassinucellate primitive type, and markedly heterogeneous rays
(Nyssa and red-fruited Cornus tenuinucellate). and relatively long vessel members with scalari-
Fruits drupes, often with a germination valve, form perforations are general. Davidia has the
1(2-6)-seeded (syncarps); seeds small to medium- largest number of bars in its perforation plates,
sized, exotestal, with straight embryo and copious followed by Mastixia, Nyssa, and Cornus; in most
endosperm. Alangium perforations are simple. In cross sec-
A family of 7 genera and ca. 110 species, in tion, vessels appear solitary or in radial groups.
warm-temperate and humid tropical regions pre- In Mastixia and Diplopanax, ducts with a secre-
dominantly of the northern hemisphere, most tory epithelium containing resin are found in the
genera in East and Southeast Asia. periphery of the pith, along the vascular bundles
of the leaves, and in the flowers and fruits. Judging
VEGETATIVE MORPHOLOGY. Cornus canadensis from fruit remains, the earliest mastixioids such as
and C. suecica are the only herbaceous (or more Beckettia and Mastixiopsis lacked secretory ducts
precisely halfshrubby) representatives of the (Eyde 1988).
family, all other being woody. These range from Alangium has articulated laticifers which
shrubs to trees, sometimes vast, but rarely but- develop from parenchyma cells, the transverse
tressed; some Alangium are scandent. Tupelos walls of which disintegrate or are ruptured. They
(Nyssa sylvatica subsp. biflora and N. aquatica), are usually arranged in an arc or ring around the
growing in swamps, develop protruding under- phloem of the vascular bundles. They can occur in
water lenticels and roots proliferating from older any part of the Alangium flower except the ovule
roots or from the stem bases (Hook et al. 1970). In and style, but terminate at the base of the flower
N. aquatica intercellular spaces are present in the or extend only a short distance into the pedicel.
cambium. Through these spaces, oxygen can reach In the leaves, they accompany the major vascular
Cornaceae 83
bundles. They are also plentiful in fruits of sect. PoLLEN MoRPHOLOGY. Pollen of Mastixia,
Alangium but are hard to find there in flowers, Cornus and Curtisia has been described by Fergu-
whereas in sects. Marlea and Rhytididandra they son (1977} and Ferguson and Hideux (1978} as
are conspicuous at all stages of flower and fruit 3-colporate, oblate (Mastixia) or spheroidal (to
development. In sect. Conostigma they seem to prolate, some Corn us), with a (rounded-)triangu-
appear as the fruits develop (Eyde 1968a). lar amb, short columellae, and usually a perforate
tectum (complete in Curtisia).Although habitually
INFLORESCENCES. Their structure has been looking quite different, these pollen grains agree
studied in detail by Jahnke (1986}. Murrell (1993}, in having a characteristic, complex H-shaped
who used inflorescence characters in a phyloge- endoaperture with lamellated thinnings of the
netic analysis, made some additions. In most endexine (the lamellations are lacking in Mas-
Cornaceae the inflorescences are terminal, which tixia). An H-shaped endoaperture may also be
sometimes leads to the formation of sympodia. present in Camptotheca and Nyssa, although the
The basic structure is a thyrsoid. Especially in analysis of their pollen morphology by Sohma
taxa with alternate leaves, the inflorescences (1963, 1967} and Eramijan (1971) is not fully con-
appear (thyrso- )paniculate. Variations of this basic clusive in this respect, and hardly so in Alangium
scheme include the formation of condensed capit- (Reitsma 1970). Pollen of Recent Alangium species
ula, sometimes in combination with showy involu- is mostly 3-colporate and its ornamentation
cra; the suppression of bracts; metatopies (both ranges from reticulate to rugulate, striate and ver-
concaulescence and recaulescence occur); sup- rucate or gemmate (Reitsma 1970; for fossil pollen
pression to loss of terminal flowers on inflo- see Eyde et al. 1969; Eyde 1972; Morley 1982}. A
rescences and partial inflorescences; and the fossil Alangium pollen named Lanagiopollis regu-
formation of open racemes. laris Morley exhibits great similarity to pollen of
Recent Mastixia.
FLORAL MORPHOLOGY. Cornaceae have a pecu-
liar gynoecial bundle supply, which has been EMBRYOLOGY. The pollen of Davidia, Nyssa,
analysed by Eyde ( 1968b ). Central bundles, usually Alangium, Cornus is 2-celled when shed (Eyde
present in syncarpous gynoecia of other plant 1988}.
groups and corresponding to the ventral bundles The ovules are anatropous or rarely (Cornus)
of the individual carpels, are lacking. In Davidia hemitropous, unitegmic, and crassinucellate or
the ovules are supplied by ramifying bundles that rarely (Nyssa and red-fruited Cornus) tenuinucel-
extend longitudinally through the septa from the late (Tandon and Herr 1971; Eyde 1988}. An
base to the apex of the locules. In Nyssa, Alangium endothelium is reported from Davidia. In the past,
and Cornus, the pendent ovules are supplied by much stress has been laid on the orientation of the
ovular bundles that pass in an arc above each ovules. The micropyle is always directed upward
septum. These bundles connect the ovules with and, as far as I could ascertain, in all genera except
paired or fused longitudinal bundles that raise Alangium and Cornus the raphe is ventral. In
through the ovary wall opposite the septa. They Alangium and also in Cornus, at least initially, the
can be interpreted as ventral carpel bundles. raphe is dorsal. A developing Cornus ovule turns
Recent Mastixia is usually pseudomonomerous from the first apotropous position so that the
but, according to Eyde (1968b ), fossil fruits related raphe is to one side of the locule and the micropyle
to it reveal a vascular pattern similar to that of to the other (Eyde 1988}.
Corn us. The development of the embryo sac usually
follows the Polygonum type, although in Cornus
KARYOLOGY. In Cornus and Alangium, the game- the Fritillaria type and some aberrant modes have
tophytic chromosome number n = 11 is wide- been observed. Endosperm development is Cellu-
spread; some Corn us have n = 10 or 9; C. canadensis lar in Davidia and Cornus, Nuclear in Nyssa and
is tetraploid (2n = 44}; for one Alangium, n = 11, 9 Alangium (Johri et al. 1992}.
and 8 have been reported (Goldblatt 1978; Eyde
1988).Nyssa and Camptotheca haven= 22,Davidia FRUIT AND SEED. Germination valves are a char-
21. In Mastixia, M. arborea has n = 13 and M. tri- acteristic trait of all cornaceous fruit stones except
chotoma has n = 11. This may point to an ancient for Alangium (Fig. 26; Hill 1933; Eyde 1988). In
base number x = 11, which was retained in the Mastixia, Diplopanax, and Davidia the germina-
cornoids and doubled in the nyssoids, and has tion valves extend full length of the locules (Fig.
undergone some aneuploid change. 26K-N), whereas in Cornus, Nyssa, Camptotheca,
84 K. Kubitzki
excepted) is 2-locular (Fig. 26G) but C. quadriloc-

ularis Chandler from the Eocene was 4-locular;
Palaeocene Nyssa fruits were 3-4-locular, Recent
are 1(rarely 2)-chambered (Fig. 26A-E; Eyde and
Barghoorn 1963), except for the recently discov-
ered Central American N. talamancana, which is a
2-3-chambered leftover. Fossil mastixioids had
fruits with up to four chambers, each provided
with a sulcus surrounded by a germination valve,
and a protrusion of the fruit stone into the locule.
In Recent mastixioids, Davidia excepting, the
plurilocular condition has vanished altogether.
Recent Mastixia and Diplopanax have a deeply
indented stone, with a C- or U-shaped seed
chamber, which is not found in Nyssa or Cornus.
Fruit wall histology is very similar in Nyssa,Mas-
tixia, Curtisia, Cornus and Alangium (Reidt 1997);
a striking feature is the "giant" cells of the meso-
carp (amounting to 800 J.lm long) of Mastixia,
Alangium and some Cornus.
The seeds have copious endosperm, which in
Alangium is oily and contains hemicellulose.
The embryo is small (Mastixia) to large and, in
Alangium, has foliaceous cotyledons and an elon-
gated hypocotyl.
POLLINATION. Cornus seems to have a general-

ized pollination system; beetles, bees, butterflies,
and flies having been observed to be attracted by
their flowers (Eyde 1988).
SEED DISPERSAL. Both blue and red-fruited

Cornus attract many kinds of bird, especially
thrushes (Eyde 1988). Nyssa aquatica and the dry-
site ecotypes of N. sylvatica have juicy fruits and
are dispersed by many kinds of bird, including
white-throated sparrow, catbird, brown thrasher,
Fig. 26. Cornaceae. A-F Nyssa sylvatica. A Fruit. B Fruit stone,
thrushes, and scarlet tanager. Also many tetrapods
showing the valve in face view and side view (C), and in D the play an important role in fruit dispersal; bears,
valve pushed away by the emerging radicle. E Fruit stone in raccoons, foxes, and opossums are known to eat
transverse section, the valve is shown by hatching. F Germi- the fruits. Rodents gnaw the stones, and tooth-
nating seed, the valve pushed aside. G-1 Cornus officina/is. G marks on fossils show that they did so in the past
Bilocular fruit stone showing the two valves commencing to
split away. H Side view of fruit stone with a valve removed (Eyde 1997). The swamp dwellers, N. sylvatica
exposing the seed. I Detached valve. J-N Davidia involucrata. subsp. biflora, N. aquatica, and N. ogeche, seem to
J Fruit stone in transverse section, showing 7locules, only 2 in be efficiently water-dispersed, N. aquatica having
this case fertile; the valves shaded. K Fruit stone with one been observed floating for a period of 85 days. Less
germinating seed. L Later stage in germination, showing three is known about dispersal of the Asian species, in
seedlings and the cavities in which the seeds lie. M, N Detached
valves seen from outside and inside. (Hilll933) which birds certainly play an important role (Eyde
1997).
It is obvious that Cretaceous mastixioid and
and fossil Amersinia they are confined to the apical Corn us fruits were considerably smaller than their
portion of the locule. Fruit stone traits of the extant Tertiary relatives (Eyde 1988). This supports the
genera are well differentiated, but lessen when they notion that the shift from abiotic to mammal-
are traced back in time. Cretaceous and Palaeogene mediated dispersal in the Palaeogene may have
Cornaceae had fruits with more chambers than been causal for an increase of the fruits in fleshi-
presently; Recent Cornus (the 3-locular C. oblonga ness and mechanical and chemical protection, the
Cornaceae 85
latter including resin, latex, iridoids and the alka- as the sister-group of Hydrangeaceae + Loasaceae
loids based on them. within a re-modelled order Cornales. In the cir-
cumscription adopted in this volume, the family
PHYTOCHEMISTRY. Among the phenolic com- Cornaceae comprises two well-supported clades, a
pounds, ellagic acid, gallic acid, and the tannins Cornus-Alangium clade, and a nyssoid-mastixioid
based upon them are reported for Davidia, Nyssa, clade with Nyssa, Davidia, Camptotheca, Mastixia
Camptotheca, Cornus and Mastixia but not for and Diplopanax. In some molecular analyses,
Alangium. Iridoids are strongly diversified, the latter grouping dissociates into the pairs
comprising normal iridoids and secoiridoids of Nyssa/Camptotheca and Mastixia/Diplopanax,
route I sensu Jensen (1991), and are known from with Davidia in a vacillating position. In the desire
Davidia, Mastixia, Nyssa, Alangium, and Cornus, for basing families only on clades that have broad
cornin being the most widespread compound. statistic support, Xiang et al. (2002) suggest three
The alkaloids of Camptotheca (camptothecin) and digeneric families, Cornaceae, Nyssaceae, and
Alangium (alangiside and tubulosin) are based on Mastixiaceae, and the monogeneric Davidiaceae.
secoiridoids (Hegnauer 1964, 1989; Bate-Smith et Accordingly, they put into question the phyloge-
al. 1975). In the species of the blue-fruited line of netic value of the peculiar gynoecial vasculature,
Cornus including the two alternate-leaved species, the germination valves in the fruit stones, and the
iridoids are lacking and obviously are replaced H-shaped endoapertures.
by salidroside and its oxidised congener, whereas Here I prefer a broader circumscription of
red-fruited carnelian cherries and showy-bracted the family that is based on the combination of
dogwoods contain iridoids and secoiridoids epigyny, fruit stones with one seed per chamber
(Jensen et al. 1975a). dehiscing with valves, and the chromosome base
number x = 11. Transseptal bundles, which are an
PHYLOGENY. Relationships and circumscription attribute of many inferior ovaries (Eyde 1988),
of this group have always been problematic; may be less decisive, and H -shaped endoapertures,
progress in this field is largely due to the efforts of also found in Curtisia but perhaps not in nyssoids,
Eyde (1968b, 1988) and the results of molecular may be a cornalean plesiomorphy. That
studies. On the basis of morphological evidence, Diplopanax would go with Mastixia in the molec-
three groups of genera have been recognised: the ular analysis could be expected from the outset,
nyssoids (Davidia, Nyssa, and Camptotheca); whereas the close association of Cornus with
the mastixioids (Mastixia and Diplopanax); and Alangium expressed in all sequence analyses is the
the cornels (Cornus); Alangium, which possesses clarification of a contentious issue of long stand-
many traits in common with Cornus, has been ing. Now it becomes clear why the flowers, fruits
accepted as cornaceous since the time of Hooker and ovules of Alangium are much like those of
(1867) and Harms (1897), but not so by Eyde (1988). Cornus, in spite of the substantial differences
With increasing knowledge of their floral, pol- between the two genera: Hooker (1867) and
len and general morphology, genera formerly Harms (1897) had the right instinct in including
included in Cornaceae, such as Helwingia, the former in their Cornaceae.
Kaliphora, Melanophylla, Corokia, Garrya, Aucuba It is remarkable that this circumscription of
and Griselinia, have been excluded from them the Cornaceae coincides exactly with Takhtajan's
upon morphological arguments, as exposed by ( 1997) morphologically based concept of his order
Eyde (1988: 309 seq.). Since most of these genera Cornales, except for his inclusion of Curtisia.
contain iridoids (Jensen et al. 1975b), but none
ellagic acid or the tannins based upon it (Bate- FOSSILS AND DISTRIBUTIONAL HISTORY. The
Smith et al. 1975), and since otherwise they are oldest cornaceous fossils are mastixioid fruit
similar to Cornaceae in floral and embryological stones from the Upper Cretaceous and Palaeogene
traits, they had been retained in Cornales or their of Europe, where the "Mastixioidean Flora" seems
surroundings (Dahlgren 1980, 1989; Takhtajan to have originated. Most of these fruit stones are
1987). Only with the application of molecular tech- traversed by resin ducts and each locule opens
niques (e.g. Xiang et al. 1993, 1997, 1998) has it with a separate valve. Various form genera have
been possible to determine their true affinity, and been based on these fruit remains; some of the
now they are placed in two orders, Garryales and earliest were Eomastixia Chandl. and Beckettia
Apiales, forming part of two different, major Reid & Chandl., both of which were 2-4-locular
clades (Euasterids I and II; see APG 1998). but still lacked secretory ducts. Later forms
Several detailed molecular studies, including the included the !-locular Mastixiopsis Kirchh.,
most recent by Xiang et al. (2002), show Cornaceae Retinomastixia Kirchh., Mastixicarpum Chandl.
86 K. Kubitzki
and Tectocarya Kirchh. and, although Mai (1993) In Cornus, the blue-fruited line is known since
suggested they could easily be included into a the Upper Cretaceous, whereas the red-fruited line
broadly construed genus Mastixia, they may appeared only in the Tertiary. Early Corn us species
exhibit different affinities to extant cornaceous had more than 2-locular fruits, such as C.
genera. Thus, the fruit stones of extinct Mastix- quadrilocularis from the Eocene. The big-bracted
iopsis had the diagnostic infold of mastixioids but dogwoods with separate fruits, which today are
on the surface had vertical ribs and intervening purely American, were present in Europe dur-
bundles of modern Nyssa sinensis and N. sylvatica. ing the mid-Oligocene. Their compound-fruited
Mastixicarpum is considered as congeneric with descendents (Fig. 29), surviving in East Asia, are
the Recent Diplopanax. known from Pliocene beds from western Europe
In Europe the record of the Mastixioidean Flora and from the Pleistocene of Japan (Eyde 1985).
extends from the Upper Cretaceous into the Alangium has a fossil record based on fruits and
Pliocene; in North America these fossils are found pollen (Eyde et al. 1969; Reitsma 1970; Morley
in the Palaeocene and Eocene (Mai 1993). From 1982). In the Upper Cretaceous restricted to North
Asia, where the two extant genera live today, no America, by the Eocene Alangium had extended
fossils are known to me. until East Asia along the northern shore of the Sea
Among the Nyssoids, a fossil species of the genus of Tethys. From there it seems to have dispersed to
Davidia, which is still alive in eastern Asia, is Africa/Madagascar. During the Neogene, its main
known from the Palaeocene of mid-latitude North diversification took place in India and Southeast
America and eastern Russia (Manchester 2002); Asia; by the Pliocene, Alangium had retreated to
the fruits of the fossil species (D. antiqua) were its present range.
definitely smaller than those of the extant species.
The extinct genus Amersinia Manchester, Crane KEY TO THE GENERA
& Golovneva (Manchester et al. 1999) from the
1. Leaf margin serrate or dentate; perianth 0 or indistinct;
Palaeocene of North America and East Asia had ovary 6-10-locular 3. Davidia
fruit characters reminiscent of extant Camp- - Leaf margin entire (dentate in capitate-flowered Camp-
totheca, whereas associated leaves described as totheca); ovary 1-3-locular 2
Beringiaphyllum Manchester, Crane and Golovnea 2. Two-armed 1-cellular hairs + on leaves and flowers; petals
resemble those of extant Davidia. The erstwhile ± valvate 3
- Two-armed 1-cellular hairs 0 (but see Alangium
and present geographic restriction of Davidia grisolleoides); petals± imbricate 5
and Amersinia to East Asia and North America 3. Two-armed hairs calcified; petals not inflexed at apex;
may indicate that these plants dispersed across resin ducts 0; style with capitate or truncate stigma; fruit
Beringia in the late Cretaceous or early Tertiary. 2(3)-locular 6. Cornus
- Two-armed hairs not calcified; petals inflexed at apex; resin
Nyssa has an extensive fossil record which can
ducts + in vegetative parts, flowers and fruits; style with
be attributed to the preference of its species for punctiform or rarely bifid or lobed stigma; fruit 1-locular
swampy and lakeside habitats, where chances for 4
fossilisation were high. Although today disjunct 4. Inflorescence thyrso-paniculate; epicarp soft; fruit stone
between eastern Asia, eastern North America and sulcate 4. Mastixia
- Inflorescence± spicate; epicarp hard; fruit stone not sulcate
Central America, during the Tertiary the genus 5. Diplopanax
was widespread in the Northern Hemisphere. 5. Petals pronouncedly strap-shaped; stamens 4-40, in a single
The oldest fossils of Nyssa appeared in the lower whorl; articulated laticifers present in various tissues
Eocene of Europe, but the genus disappeared from 7.Alangium
this continent during the Pliocene. The first - Petals not strap-shaped; stamens 8-15, in 2 isomerous
whorls; laticifers lacking 6
records in North America are from the middle 6. Pistillate and staminate inflorescences in globular heads of
Eocene, whereas in Asia Nyssa appeared only in ca. 50 flowers, these compound in racemes; fruits dry, flat-
the Miocene (see also Wen and Stuessy 1993). tened, grouped in heads 2. Camptotheca
Three types of Nyssa endocarp sculpture can be - Pistillate flowers solitary, staminate in racemes or heads of
distinguished in the fossil material from the up to 20 flowers; fruits drupaceous with fleshy epicarp
I. Nyssa
Eocene of Europe and North America, and indicate
that considerable diversification of the genus had
GENERA OF CORNACEAE
been attained at that time. These types include
ridged fruits with sunken bundles, ridged fruits
1. Nyssa L. Fig. 26A-F
with raised bundles (today represented by N.
aquatica), and smooth fruits (today exemplified Nyssa L., Sp. Pl.: 1058 (1753); Burckhalter, Sida 15: 323-342
by N. javanica) (Eyde 1997). (1992) (rev. N Am. spp.).
Cornaceae 87
Polygamodioecious deciduous trees. Leaves alter·

nate. Inflorescences axillary, pedunculate; stami-
nate flowers in short racemes or heads, pistillate
flowers single or in few-flowered racemes; stami-
nate flowers: calyx lobes minute; petals 5-10,
imbricate; stamens 8-15, usually in 2
whorls; disk pulvinate; gynoecium abortive;
hermaphroditic or pistillate flowers: floral tube
adnate to ovary; stamens fewer than in staminate
flowers, or abortive, or 0; disk pulvinate or conical;
ovary 1( -3)-locular; style simple or with 2-3 styl~r
branches; stigma reflexed or revolute. Drupe ovo1d Fig. 27. Cornaceae. Davidia involucrata. A Inflorescence sub-
tended by showy bracts. B Same, without bracts. C Same, with
or ellipsoid, 1(-3)-chambered, blue-black, or more most male flowers removed, the single female flower to the
rarely purple or red; endocarp with longitudinal right. D Female flower, vertical section. E Ovary, transverse
ridges or wings, opening in apical part by a trian- section. F Fruit. GSame, transverse section. (Schneider 1912)
gular abaxial valve; embryo straight; endosperm
copious. n = 22. About 8 species, 3 in SE North
America, 1 in Costa Rica, and 4 in East and South-
Drupe 3-5-seeded, ellipsoid, mesocarp fleshy,
east Asia; preferably on flooded ground.
endo carp bony, longitudinally ca. 12-ridged and 6-
valved; seeds with fleshy endosperm and straight
2. Camptotheca Dcne. embryo subequal to endosperm. n = 21. A single
sp., D. involucrata Baill., C China, in montane
Camptotheca Dcne., Bull. Soc. Bot. France 20: 157 (1873).
mixed mesophytic forest.
Monoecious deciduous tree. Leaves alternate,
usually entire but occasionally dentate. Inflores- 4. Mastixia Bl. Fig. 28
cences of globular heads grouped in a raceme, the
Mastixia Bl., Bijdr.: 654 (1826); Matthew, Blumea 23: 51-93
heads composed of up to 50 flowers; flowers in (1976), rev.
dyads, 5-merous, in terminal head hermaphro-
ditic, in lateral heads staminate; petals overlapping Resinous evergreen trees with 2-armed hairs.
little if at all; stamens 5 + 5, inserted on disk; ovary Leaves alternate or (sub)opposite. Inflorescences
(0 in staminate flowers) 1-locular; style immersed thyrso-paniculate, many-flowered; flowers her-
in disk, halfway split into 2 style branches. Fruits maphroditic, 4-5( -7)-merous; calyx toothed or
grouped in heads, flattened-spatulate, crowned by lobed; petals usually valvate in bud (contorted),
persistent disk and style, epicarp fibrous, endo carp thick, inflexed at apex and bidentate or fimbriate;
leathery, 1-seeded; embryo elongated, endosperm stamens 1 or 2 whorls inserted at base of nectary
fleshy. n = 22. A single sp., C. acuminata Dcne., C, disk; ovary !-locular; ovule 1, pendulous; style
S, and SE China, in lowland warm-temperate short, thick, with punctiform or rarely 2-lobed
mixed mesophytic forest. stigma. Drupes with thin or thick epicarp, blue to
dark purple at maturity; endocarp woody, sulcate
3. Davidia Baillon Fig. 27 with an intrusive, incomplete, wedge-shaped
(sub gen. Mastixia) or swollen (sub gen. Manglesia)
Davidia Baillon, Adansonia 10: 114 (1871). septum on the germination valve; seeds with small
embryo in copious endosperm. n = 11, 13. About
Deciduous, andromonoecious tree. Leaves alter- 19 spp. or more (Matthew 1976); extending from
nate, serrate. Inflorescences on short lateral India through warm-temperate Southeast Asia
shoots, in solitary globose heads subtended by and Malesia to New Guinea and the Solomon Is.
2(3) large, creamy-white subopposite spreading The status of subgen. Manglesia and its relation-
bracts; staminate flowers apetalous, numerous, ship to Diplopanax requires further study.
densely packed on globose axis, 1-7-staminate;
hermaphroditic or pistillate flowers inserted at top
of head, with calyx adnate to ovary; stamens, if 5. Diplopanax Hand.-Mazz.
present, 15-26 arranged around the base of style; Diplopanax Hand.-Mazz., Sinensia 3: 197 (1933); Ying et a!.,
style short and conical, with 6-10 spreading style The endemic genera of seed plants of China: 142, fig. 36.1
branches; ovary 6-1 0-locular; nectary disk 0. (1993); Averyanov & Nguyen, Novon 12: 433-436 (2002).
88 K. Kubitzki
Fig. 28. Cornaceae. Mastixia pentandra. A Flowering twig. B

Part of inflorescence. C Flower, vertical section. D Stamens. E
Fruit. Drawn by R. van Creve!. (Matthew 1976)
Resinous deciduous tree with 2-armed hairs. Fig. 29. Cornaceae. Cornus japonica, flowering branch and
Leaves subverticillate. Inflorescence spicate(- coenocarp. (Big-bracted dogwoods with coenocarps in the
Tertiary extended to Europe, but today are restricted to E
paniculate); flowers sessile, hermaphroditic, 5-
Asia.) Drawn by T. Yamada. (Nakai 1927)
merous; sepals short; petals valvate in bud,
inflexed at apex; stamens 5 + 5; ovary 1-locular,
1-ovulate; style conical. Drupe (oblong-)ovoid;
exocarp hard; endocarp woody, with an intrusive,
swollen, incomplete septum on the germination terminal or axillary and thyrso-paniculate, or
valve; seed 1, compressed. Two species in moun- in heads surrounded by petal-like involucra!
tain forests of S China and Vietnam. bracts; flowers hermaphroditic, rarely dioecious,
tetramerous; calyx distinct or obsolete; petals
valvate; stamens alternipetalous; disk fleshy;
6. Cornus L. Figs. 26G-I, 29 anthers dorsifixed; style with capitate or truncate
Cornus L., Sp. Pl.: 117 (1753). stigma; ovary (1)2(3)-locular. Drupe areolate at
Chamaepericlymenum Hill (1756). apex; pericarp fleshy, blue, white, or red; endocarp
Swida Opiz (1838). bony or crustaceous, 2(1)-locular, opening with
Benthamidia Spach (1839).
valves. Seeds oblong, compressed, with elongate
Afrocrania (Harms) Hutch (1942).
Dendrobenthamia Hutch. (1942). embryo and fleshy endosperm. n = 9-11. About 60
Bothrocaryum (Koehne) Pojarkova (1950). spp., mainly of temperate regions of the northern
Yinquania Zhu (1984). hemisphere, 1 or 2 spp. in South America, 1 in
Africa.
Evergreen or deciduous trees or shrubs, or rarely The infrageneric classification is complex and
rhizomatous herbs, with 2-armed, calcified tri- controversial; no formal subdivision is given here.
chomes. Leaves usually opposite (alternate in C. However, the molecular data available support the
alternifolia and C. controversa). Inflorescences distinction of five groups in two major branches.
Cornaceae 89
The first major branch comprises the blue- or

white-fruited dogwoods with 1) alternate-leaved
and 2) opposite-leaved Cornus. The second major
branch represents the red-fruited dogwoods with
3) the cornelian cherries, 4) the dwarf dogwoods,
and 5) the big-bracted dogwoods.
7. Alangium Lamk. Fig. 30

Alangium Lamk., Encycl. Meth., Bot.1: 174 (1783), nom. cons.;
Bloembergen, Bull. Jard. Bot. Buitenzorg III, 16: 139- 235
(1939), monogr.
Evergreen (A. platanifolium summergreen) trees,

shrubs, or lianas, sometimes thorny, with stellate
and glandular, rarely 2-armed (A. grisolleoides)
hairs with unequal arms; articulated laticifers in
various tissues of the shoot. Leaves alternate, undi-
vided or lobed. Inflorescences axillary cymes;
flowers with articulated pedicels, hermaphroditic
or rarely (A. grisolleoides) dioeciously distributed;
calyx with 4-10 lobes or teeth, or truncate; petals
4-10, strap-shaped, valvate or rarely contorted,
sometimes connate at the base; stamens 4-40 in a
single cycle around a conspicuous nectary disk; fil-
aments barbate or villose at inner side; anthers
linear, basifixed or dorsifixed and versatile and
widely sagittate; style elongate, filiform, with
clavate, 2-4-lobed or capitate stigma or rarely style
short and with 2 elongate stylodia; ovary 1(2)-
locular. Drupe crowned with calyx and disk, with
crustaceous or woody endocarp, 1(2)-locular, each
locule with 1 seed or one locule empty; seeds with
large, straight embryo; endosperm copious, oily, Fig. 30. Cornaceae. Alangium chinense. A Flowering branch-
also containing hemicellulose. n = 8, 11. About 21 let. B Flower. C Corolla and stamens. D Gynoecium with disk.
spp., Africa, Madagascar, tropical and warm-tem- E Fruit. F Transverse section through fruit. G F
r uit, longitudi-
perate S and SE Asia from India to Japan, and nal section. (Cannon 1978)
extending further through Malesia to Queensland
and Fiji. Four sections distinguished by Bloem-
bergen (1939) and Eyde (1968a); sect. Conostigma Bloembergen, S. 1939. A r evision of the genus Alangium. Bull.
Jard. Bot. Buitenzorg III, 6: 139-235.
probably basal, position of A. grisolleoides Cannon, J.F.M. 1978. Alangiaceae. In: Launert, E. (ed.) Flora
Capuron from Madagascar still problematic (Eyde Zambesiaca vol. 4. London: Flora Zambesiaca Managing
1988). Flowering in the dry season, fruiting at the Committee, pp. 633- 635.
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Dahlgren, G . 1989. The last D ahlgrenogram. System of classi-
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Jensen, S.R. 1991. Plant iridoids, their biosynthesis and distri- logenetic relationships in Corn us based on chloroplast DNA
bution in angiosperms. In: Harborne, J.B., Tomas-Barberan, restriction sites: implications for biogeography and charac-
F.A. (eds.) Ecological chemistry and biochemistry of plant ter evolution. Syst.Bot. 21: 515-534.
terpenoids. Oxford: Clarendon Press, pp. 133-158. Xiang, Q.- Y., Soltis, D.E., Soltis, P.S. 1998. Phylogenetic rela-
Jensen, S.R., Kjaer, A., Nielsen, B.J. 1975a. The genus Cornus: tionships of Cornaceae and close relatives inferred from
non-flavonoid glucosides as taxonomic markers. Biochem. matK and rbcL sequences. Am. J. Bot. 85: 285-297.
Syst. Ecol. 3: 75-78. Xiang, Q.-Y., Moody, M.L., Soltis, D.E., Fan, C.-Z., Soltis, P.S.
Jensen, S.R., Nielsen, B.J., Dahlgren, R. 1975b. Iridoid com- 2002. Relationships within Cornales and circumscription
pounds, their occurrence and systematic importance in the of Cornaceae - matK and rbcL sequence data and effects
angiosperms. Bot. Not. 128: 148-180. of outgroups and long branches. Molec. Phylog. Evol. 24:
Johri, B.M. et a!. 1992. See general references. 35-57.
Cunoniaceae 91
Cunoniaceae
J.C. BRADFORD, H.C. FoRTUNE HoPKINS and R.W. BARNES
Cunoniaceae R.Br. in Flinders, Voy. Terra Austral. 2: 548 ( 1814), syncarpous or less often the carpels ± distinct;
nom. cons. stylodia free, often diverging, with terminal, rarely
Baueraceae Lind!. (1830).
Eucryphiaceae End!. (1841), nom. cons.
decurrent stigmas; ovules (1)2-many in each
Davidsoniaceae. (1952). locule, often in two rows, anatropous or hemiana-
tropous; apotropous or epitropous, bitegmic;
Trees, shrubs, occasionally hemiepiphytic or placentation axile to pendulous. Fruit dehiscent,
strangling (some Weinmannia); hairs generally follicular or capsular; seeds usually exposed from
simple, sometimes trichomes stellate and peltate, open carpel sutures, the carpels separating distally
long, stiff and irritating in Davidsonia; bark or (Cunonia) from the base, or fruit indehiscent,
usually grey to light brown; fissures longitudinal; drupe-like, or dry with wing-like expanded sepals
stems commonly with lenticels. Leaves evergreen, or carpel-walls, or carpels hairy, or carpels swollen
winter deciduous in Eucryphia glutinosa; and bladder-like (Platylophus). Seeds small,
opposite-decussate in pairs or sometimes 3 or 4 winged or not, glabrous or hairy ( Weinmannia and
leaves per node (spiral in Davidsonia); imparipin- Ackama); endosperm starchy, 0 in Davidsonia.
nate, trifoliolate, palmately compound, or unifoli- A family of 27 genera and c. 300 species, occur-
olate, pinnately veined, stipellate or not, firm, ring in South and Meso America, Australia,
often coriaceous; leaf margins toothed, often Malesia, Madagascar, and islands of the
glandular-serrate, sometimes entire; venation Caribbean, South Pacific, and western Indian
craspedodromous, semicraspedodromous, occa- Ocean, only two species in southern Africa and
sionally brochidodromous or reticufodromous no species in mainland Asia north of the Malay
(some Eucryphia); small tuft or pocket domatia Peninsula.
along midvein sometimes present; stipules often
conspicuous, often interpetiolar. Inflorescences VEGETATIVE MORPHOLOGY. All known species
terminal, axillary, sometimes cauliflorous, panicu- are potentially long-lived plants that flower repeat-
late, thrysoid or cymose and with straight pedun- edly over many years. Nearly all species require
cle, or capitate, rarely flowers solitary and axillary; moderate to high light intensity, and as a conse-
bracts usually stipulate; flowers often with pro- quence most are canopy species, but life form
phylls; pedicels present or sometimes 0. Flowers varies depending on the habitat. In lowland or pre-
actinomorphic, occasionally protandrous, com- montane rainforest, Cunoniaceae may be trees
monly hermaphroditic, plants dioecious or polyg- over 30m tall and 1 m in diameter. In montane
amodioecious in Pancheria, Vesselowskya, and forest, subalpine zones, or on poor soils, plants
some Weinmannia, andromonoecious in some are commonly small trees to rounded shrubs.
Schizomeria and some Cunonia; sepals (3)4 or Extremely windswept, cold or rocky conditions
5( -10}, valvate or imbricate, separate or basally produce ground-creeping shrubs. In neotropical
connate; petals as many as sepals and alternate cloud forests, some Weinmannia grow as hemiepi-
with them, sometimes 0, more numerous than the phytes and may even develop into large stranglers
sepals in Bauera; androecium usually diplostemo- (J. Bradford, pers. obs.; A. Gentry, pers. comm.).
nous, sometimes uniseriate or multiseriate, rarely Many Cunoniaceae coppice readily, especially
polyandric; filaments slender, exceeding petals; Anodopetalum biglandulosum, an endemic of
anthers dithecal, opening longitudinally, some- Tasmania that forms dense tangles of stems called
what apically in Bauera; disc free, annular or com- 'horizontal scrub' by sending vertical stems from
posed of segments, less often adnate to the ovary bent and fallen layers of once vertical stems
or 0; ovary usually superior or partially to mostly (Barker and Brown 1994}. Branch nodes in contact
inferior, usually 2- or 3-5-carpellate, 4-14- with the ground can root and sprout shoots,
carpellate in Eucryphia, effectively 1-carpellate in enabling an individual genet to disperse and dom-
Hooglandia with second carpel possibly vestigial, inate an area. The growth form of Schizomeria
92 J.C. Bradford et al.
ovata, from eastern Australia, differs depending on flakes, platelets, or coarse rods or rodlets. Very
its habitat (Stanley and Ross 1983). In tall Euca- prominent cuticular, papilla-like protrusions
lyptus woodlands, which generally have a light, occur on the abaxial surface cells of Eucryphia
open understorey, it grows as a spreading shrub, (Hill 1991; Barnes and Jordan 2000). Epidermal
whereas in shorter-statured rainforests, which cells often have periclinal divisions in addition to
generally have little understorey light penetration, being enlarged and mucilaginous. Abaxial epider-
it grows as a canopy tree. On ultramafic substrates mal cells vary from smooth to papillose (e.g. Acro-
in New Caledonia, several species have tall, spindly phyllum). An adaxial and sometimes abaxial
growth forms with branching rare above the base. hypodermis composed of thick-walled cells is
In general, members of the family are easy to present in some taxa. The mesophyll is tanninifer-
recognize based on vegetative characters such as ous and contains large solitary crystals and druses.
dentate, decussately arranged leaves (sometimes Young stem anatomy is characterized by a con-
whorled, or alternate in Davidsonia) and promi- tinuous ring of xylem. The secondary phloem
nent stipules. Stipules may arise as a pair of includes groups of fibres. The cortex is sometimes
primordia that remain separate throughout provided with sclereids and often a ring of
development, as pairs of primordia that fuse perivascular fibres. Nodal anatomy is trilacunar,
during growth, or as a single primordium 3-trace, multilacunar, multi-trace, or unilacunar,
(Rutishauser and Dickison 1989; Dickison and one-trace (Bauera). Species with whorled leaves
Rutishauser 1990). Mature stipular form depends typically possess 'split-lateral' traces, including the
upon the number of stipular primordia, their loca- very uncommon condition of multilacunar, multi-
tion, and degree of fusion during development. trace nodes with split-lateral traces in Pancheria
Most often stipules are interpetiolar or called confusa, P. hirsuta and P. robusta.
lateral if in unfused pairs (Figs. 35, 37) but axillary Wood anatomy exhibits a wide range of varia-
stipules occur in South Pacific Geissois species. tion. Vessel distribution is exclusively or predom-
Leaves are imparipinnate, unifoli( ol)ate, inantly solitary, or commonly a combination of
trifoli( ol)ate, or palmately compound. Stipels are solitaries, radial pore multiples, and clusters.
sometimes present. Tuft or pocket domatia may Vessel element perforation plate type is either (1)
occur on the abaxial leaf surface near the midvein. exclusively scalariform (less than ten to over 30
Hairs are unbranched, variable in length, density bars), (2) predominantly simple but with variable
and distribution on the plant, and may be straight frequencies of few-barred scalariform plates, or
or curled to various degrees. Stellate hairs are (3) exclusively simple (Pseudoweinmannia). Inter-
present on stems and sepals in Gillbeea. Spiraeop- vessel pitting ranges between scalariform and
sis has stellate hairs and orbicular, gland-like transitional to opposite and alternate. Spiral thick-
trichomes on leaves, stems and sepals, and Ackama enings are present in the vessels of some
has orbicular, gland-like trichomes on leaves. Eucryphia species. Imperforate tracheary elements
Davidsonia has long, glandular, urticating hairs. are tracheids or libriform fibres. Rays are hetero-
Extra pairs of axillary buds occur in some taxa. cellular or heterocellular and homocellular,
with both uniseriates and multiseriates. Axial
VEGETATIVE ANATOMY. (Refer to Ingle and parenchyma is variable, being absent, diffuse and
Dadswell 1956; Eyde 1970; Dickison 1975b, 1978, diffuse-in-aggregates, or distributed in continuous
1980a, 1980b; Rao and Dickison 1985a, 1985b; or discontinuous bands (1-5 cells wide), some-
Rutishauser and Dickison 1989; Dickison and times terminal. Prismatic crystals are common.
Rutishauser 1990). Leaf venation is craspedodro-
mous, semicraspedodromous, or less commonly INFLORESCENCE STRUCTURE. Inflorescence varia-
brochidodromous and reticulodromous (some tion occurs in ( 1) the number and position of
Eucryphia). Areolation ranges from incompletely flower-bearing axes within the stem system, (2)
closed to completely closed and well-developed. the timing of floral maturation within a flower-
Major veins are often ensheathed with fibres. Ulti- bearing axis, and (3) the pattern and orders of
mate veinlets often have specialized thick-walled, branching within the flower-bearing axis.
solitary or clustered, terminal elements. Elon- Flower-bearing axes (e.g. racemes, panicles) may
gated, tubular cells are a characteristic feature of be borne in leaf axils, terminal on shoots, or both;
the vein endings of Codia. Mature stomata can cauliflory is occasional (Pacific Geissois, Davidso-
be anomocytic, brachyparacytic, encyclocytic or nia, some Acsmithia). Within a species or higher-
anisocytic, and are confined to the abaxial lamina level clade, the position and number of
surface. Extracuticular wax occurs in the form of flower-bearing axes may be highly predictable or
Cunoniaceae 93
very plastic. For example, nearly all c. 75 species Tribes Geissoieae and Cunonieae have flowers
in Weinmannia sect. Weinmannia have racemes borne along unbranched axes, mainly racemes.
restricted to axils of distal leaves. In contrast, Flowers may arise either solitarily or in fascicles,
Davidsonia species may bear panicles from near and may be pedicellate or not. These racemose
the base of the trunk to the crown of the tree. structures are borne directly from leaf axils or,
Bradford and Barnes (2001) found a significant more often, in groups from a lateral or central
variation in the timing of floral maturation within peduncle (i.e. Inflorescence Module, sensu Brad-
a flower-bearing axis among tribes and genera of ford 1998). There are several different patterns of
Cunoniaceae. Floral maturation occurs either in raceme organization along Inflorescence Modules.
terminal flowers first (i.e. basipetal), or more or For example, in some groups the Inflorescence
less at the same time to slightly acropetally (i.e. Module only bears racemes laterally and produces
synchronous/acropetal). Tribe Spiraeanthemeae a vegetative bud in the terminal position, whereas
appears to have the derived form of synchronous in other groups the Inflorescence Module bears a
timing, whereas most outgroups and a basal grade terminal and lateral racemes.
of ingroup taxa, including tribe Schizomerieae, Three genera, Pancheria (Fig. 36), Codia and
Davidsonia, Hooglandia, Aistopetalum and Gill- Callicoma, have flowers arranged in capitula.
beea, have basipetalous flower sequence. A possi- These are borne at the ends of slender stalks
ble clade of four tribes, Geissoieae, Caldcluvieae, (peduncles), either directly from leaf axils, or
Codieae and Cunonieae, and the genus Acrophyl- peduncles are borne in pairs or series from an axil-
lum have the synapomorphy of synchronous lary shoot (i.e. within an Inflorescence Module).
flower sequence. Codia and Callicoma are closely related to Pullea,
In branched flower-bearing axes (e.g. paniculi- which has paniculate flower-bearing axes and
form or thrysoid to cymiform axes), phyllotaxis often clustered flowers (Bradford and Barnes
is usually basally decussate and distally spiral but, 2001), whereas Pancheria is more closely related to
depending upon the taxa, this transition may Cunonia and Weinmannia with racemose flower-
occur near the stem, not until the end of the axis, bearing axes (Bradford 2002).
or not at all. The flower-bearing axis often consists The inflorescence in Acrophyllum is unusual,
of a dominant, primary shoot that bears second- with the main axis alternating between produc-
ary lateral axes that are longer and more branched ing a sequence of vegetative nodes with whorled
basally, and shorter and less branched distally. leaves and a sequence of several reproductive
This typically leads to a conical outline, which may nodes with axillary inflorescence structures (Fig.
be called paniculiform, and typifies tribe Spi- 34). At each reproductive node is a whorl of small,
raeanthemeae and the genera Davidsonia, once-branched panicles bearing about a dozen
Hooglandia, Aistopetalum, Gillbeea, Ackama and flowers (Hoogland 1960).
Spiraeopsis. In some genera, no dominant inflo- Solitary flowers are found in Eucryphia,
rescence axis or only a weakly dominant one Anodopetalum and Bauera. In the former two
exists. For example, in tribe Schizomerieae and the genera, the flower is provided with a pedicel,
genera Caldcluvia and Opocunonia in tribe Cald- bracts, and a peduncle suggesting that solitary
cluvieae, there is no elongate main axis and, there- flowers are derived from reduced cymes. In
fore, the inflorescence has a rounded shape. The fact, three-flowered cymes rarely occur in
degree of branching varies from as little as one Anodopetalum (Barnes and Rozefelds 2000).
order in taxa with small inflorescences, such as in Bauera is the only genus in which pedicels arise
Acrophyllum, to very large and profuse branching directly from leaf axils.
systems with up to five or six orders, as in
Aistopetalum, Ceratopetalum, Schizomeria and FLORAL MoRPHOLOGY AND ANATOMY. Floral
Spiraeopsis. Within a tree, the branching order or morphology has been best described in the
profusion of inflorescence units may also differ various treatments of genera (see references in
among flower-bearing axes in relation to their taxonomic part), in the context of floral biology by
position along the stem (Bradford 1998). For Hopkins (see discussion re Pollination below) and
example, Ackama has more prolific panicles at ter- in a comparative phylogenetic context by Bradford
minal nodes than at subterminal ones (i.e. aero- and Barnes (2001). Flowers typically have two
tonic total inflorescence) whereas in some species perianth whorls, twice the number of stamens
of Weinmannia sect. Inspersae, inflorescence axes as petals (usually diplostemonous, sometimes
are more branched at subterminal nodes (i.e. obdiplostemonous; see Endress and Stumpf 1991),
basitonic total inflorescence). and a nectary surrounding a bicarpellate, superior
ovary with terminal stylodia. Variations include capitata and compare it to previous studies in
the presence of only one perianth whorl, stamen Cunoniaceae and Saxifragaceae in general (e.g.
number equal to petal number, numerous Davis 1966; Johri et al. 1992). Cunoniaceae anthers
stamens, absence of nectary, a few to several are tetrasporangiate with a glandular tapetum
carpels, a semi-inferior to inferior ovary, and and tanniniferous endothecium. Pollen is 2-celled.
decurrent stigmas. Perianth parts are distinct or Ovules are anatropous, crassinucellate, bitegmic,
only weakly fused. Petal or, when absent, sepal and the micropyle is zigzag in shape. The embryo
colour is usually white, yellow or pink, less often sac contains abundant starch at maturity. Seeds
red to purple. Filaments usually exceed the peri- have nuclear endosperm, lack haustoria, and have
anth and are slender, cylindrical or strap-like. a coat formed by both integuments. Embryos
Anthers are dithecal, with or without an apically are straight and usually embedded in abundant
protruding connective, and open longitudinally or, endosperm, but endosperm is lacking in Davidso-
rarely, somewhat apically (Bauera). The floral disc nia (Doweld 1998).
is most often distinct from the ovary but may be
adnate to it. When distinct, it may be entire or POLLEN MORPHOLOGY. Pollen grains are small
composed of inter- and intrastaminal segments. {10-13Jlm), generally tricolporate (Cunonia,
The carpels are usually fused into a single ovary Opocunonia, Spiraeanthemum, Weinmannia) or
but always have free stylodia (only at the tip in dicolp( or )ate (Ceratopetalum, Eucryphia, Geissois,
Gillbeea adenopetala), locule number is equal to Pullea, Schizomeria), but syncolpate in Gillbeea.
carpel number, except in the bicarpellate, uniloc- Grains vary in shape from sub oblate to subprolate,
ular Hooglandia; in some groups the carpels are generally with a circular equatorial outline, but
more or less distinct. sometimes dicolp(or)ate grains are bilaterally
Aspects of floral anatomy, including details flattened (Ceratopetalum, Schizomeria ). Only
of floral vascularization, were presented by Eyde Cunonia capensis has oblate, angulaperturate
(1970), Bensel and Falser {1975), Dickison {1975a, grains which differ markedly from the normal
1975c, 1978), and Matthews et al. {2001). Tannin suboblate grains with a circular outline found in
and druses are common throughout all floral the New Caledonian species of Cunonia.
parts, with mucilage-producing cells localized in Ectoapertures are colpate (except Gillbeea) or
some tissues. Sepals usually have three-trace vasu- indistinct, as in Pullea. Endoapertures vary
larization with fusion of lateral traces of adjacent from indistinct or simply constructed (Pullea,
sepals. Petals (when present) and stamens are vas- Ceratopetalum, Schizomeria) to simple-
cularized by one trace, except for Eucryphia which complex/complex ( Weinmannia, Aistopetalum,
has prominent stamen fascicle (trunk) bundles. Opocunonia, Spiaeopsis, Spiraeanthemum).
Gynoecia are predominantly vascularized by three Tectum sculpture also varies from uniformly per-
major bundles, a dorsal and a pair of ventral vas- forate (homogeneous) or finely reticulate (Cald-
cular bundles. The ventral bundles of adjacent cluvia, Spiraeopsis, Opocunonia, Pullea, some
(usually opposing) carpels may be fused in the Weinmannia) to partial and coarsely reticulate
ovary and separate into the stylodia of adjacent (heterogeneous) in the centre of the mesocolpia
carpels. Pullea is the only genus studied in which (Gillbeea, Ceratopetalum, some Schizomeria)
ventral carpel vasculature terminates in the ovules (Hideux and Ferguson 1976). In some genera the
and does not enter the stylodia. Ventral carpel sculpture in the centre of the mesocolpia tends
sutures are open at, or slightly above, the level towards rugulate (Schizomeria, some Eucryphia).
of placentation in most genera, or are closed
throughout (Aistopetalum). Carpels of the essen- KARYOLOGY. Published chromosome counts are
tially apocarpous genera Acsmithia and Spiraean- summarised in Fortune Hopkins and Hoogland
themum (Fig. 31) are vascularized by a dorsal {2002) and include species from 9 genera. Most
bundle and either three or four bundles constitut- taxa have 2n = 32 (Ackama, Bauera, Cer-
ing the ovular and wing vasculation in the ventral atopetalum, Cunonia, Geissois and Lamanonia).
position. Ovules are (1)2-many in each locule New Zealand species of Weinmannia have 2n = 30,
along the axile to pendulous placenta. Ovules are as does Eucryphia Iucida (Labill.) Baill. Pancheria
anatropous or hemianatropous, and the micropyle sebertii Guillaumin has 2n = 24. More chromo-
is either apotropous (derived) or epitropous (ple- some counts of species in tribe Cunonieae (Ves-
siomorphic) (Bradford and Barnes 2001). selowskya, Weinmannia, Pancheria and Cunonia)
may be warranted since they appear variable and
EMBRYOLOGY. Prakash and McAlister {1977) give relationships in this tribe are not well understood
a detailed account of the embryology of Bauera (Bradford 2002).
Cunoniaceae 95
FRUIT AND SEED. Dickison (1984) provides a tribution and Ecology). Indehiscent Cunoniaceae
review and description of most Cunoniaceae fruits fruits typically have only one or two seeds per
and seeds. Corrections are made to previous carpel. In Hooglandia, one seed develops in a
descriptions of Anodopetalum fruits by Barnes unilocular ovary, and the fruit is elongate and flat-
and Rozefelds (2000). Bradford and Barnes (2001} tened rather than orbicular as in other genera.
discuss evolution of fruit morphology. For varia- Davidsonia and Hooglandia seeds are the largest
tion in carpel number, see discussion of flowers. in the family, being about 2 em long.
Usually, all carpels develop into pericarps of Dickison (1984} describes seed morphology,
mature fruits, although locule and seed abortion seed coat development and mature seed anatomy
may occur in some indehiscent fruits. of most Cunoniaceae, and Doweld (1998}
Seventeen genera have capsular or multi- describes Davidsonia seeds. Wing-like outgrowths
carpellate follicular fruits. Free carpels are proba- are formed primarily by two epidermal layers and
bly the plesiomorphic condition in Cunoniaceae, have variable sizes and shapes. Wings form oppo-
being present in related families (Brunelliaceae site the hilum, or both chalazally and micropylarly
and Cephalotaceae) and in the most basal tribe with possible lateral extension. Seed hairs, found
Spiraeanthemeae (see Bradford and Barnes 2001, in Weinmannia and Ackama, are unicellular, thin-
their Fig. 5). Most other Cunoniaceae have a syn- walled and often wavy. Seed surfaces may be warty,
carpous gynoecium. Reversal to pseudo-follicular striate, smooth, papillate or a combination of these
fruits has occurred in Pancheria (Cunonieae), but features. At early stages of development, the
the carpels remain basally fused and their ventral epidermis of the outer integument forms a tan-
margins are inrolled. Capsular fruits are similar niniferous exotesta in mature seeds. In most taxa,
to follicular ones, in that they open through their the outer epidermis of the inner integument
septa (usually dehiscing basipetally) to expose gradually becomes sclerotized to produce
adaxial (ventral) surfaces of separate carpels that exotegmic mature seeds. However, seed coats in
release seeds from open sutures. Cunonia has a Pullea and Codia are composed of only three cell
unique circumbasal, often acropetal dehiscence, layers and lack a mechanical tissue. A sclerotic
but is also septicidal. Each carpel may contain one layer is also absent in Platylophus, Anodopetalum
to numerous small (c. 0.5-2.0mm long) seeds. and Gillbeea. Most taxa have a visible hilar scar,
Dispersal is frequently facilitated by appendages of an elongate, raised raphe, and a raphal bundle
wings or hairs, but a few genera have ovoid seeds that is usually unbranched or, in drupaceous
lacking prominent appendages. fruits, well-branched near the chalaza. The large
Indehiscent fruits occur in ten genera, including seeds of Davidsonia have an expanded pachycha-
members of tribes Schizomerieae, Geissoieae, laza layer.
Codieae, as well as Davidsonia, Hooglandia,
Aistopetalum and Gillbeea (Bradford and Barnes FRUIT AND SEED DISPERSAL. Capsular or follicu-
2001; Sweeney and Bradford, in prep.). Several lar fruits generally remain attached to the parent,
genera have dry, indehiscent fruits, but each is and seeds disperse individually. Plants are rarely
different from another or, if similar, this is due found with open fruits in which seeds remain.
to convergence. For example, Ceratopetalum (Open fruits with seeds on herbarium specimens
(Schizomerieae) and Pullea (Codieae) have an were likely collected when closed, and only opened
enlarged calyx that persists as wings. In Codia during drying). In 1995, Bradford and Hopkins
(Codieae) and Pseudoweinmannia (Geissoieae) (pers. obs.) found a tree of Weinmannia richii A.
the exocarp is covered with hairs, but these are Gray in Fiji that had open capsules with seeds. The
wavy and stiff in Pseudoweinmannia and wooly seeds were found blowing out of the fruit and
in Codia. In Gillbeea the carpel walls form wings being carried high by a light breeze of hot, dry air.
as lateral phalanges. Platylophus (Schizomerieae) Seeds of Weinmannia and Ackama are covered
fruits lack appendages and develop into swollen with light hairs, whereas in most genera they have
bladders (utricles). Drupaceous fruits, with a wing-like appendages and a few are simply ovoid.
range of exocarp toughness and mesocarp fleshi- In contrast to the dispersal potential of Weinman-
ness, are found in Davidsonia, Schizomeria, nia seeds,Acrophyllum, which is a narrow endemic
Aistopetalum and Hooglandia. Doweld (1998) on specialized substrates, has unappendaged seeds
reports that Davidsonia fruits are dry and split that may inhibit dispersal.
into two carpels at maturity, but field observations Most indehiscent fruits fall from the parent,
indicate otherwise. Ripe, syncarpous Davidsonia but some smaller, fleshy fruits may persist while
fruits fall from the parent when still fleshy (not ripe. Wind or water may disperse the fruits of
dry), and are eaten whole by vertebrates (see Dis- Pseudoweinmannia and Codia, which are covered
96 J.C. Bradford et a!.
by hairs. Fruits with wing-like appendages, includ- The genera Acsmithia and Spiraeanthemum
ing laterally expanded carpel walls in Gillbeea (tribe Spiraeanthemeae) from New Guinea and the
and enlarged calyces on fruits in Pullea and Cer- South Pacific form a small sister clade to the rest
atopetalum, may also be wind dispersed. Dry blad- of the family. Within this larger clade, Bauera and
ders in Platylophus may be water dispersed, and a group of genera with mostly thyrsoid to cymi-
consumption by wild pigs is reported (Palmer and form inflorescences and drupaceous or indehis-
Pitman 1961). Drupaceous fruits in Aistopetalum, cent dry fruits (including tribe Schizomerieae and
Davidsonia, Hooglandia and Schizomeria are Davidsonia, Aistopetalum and Hooglandia) form a
probably vertebrate dispersed. In SE Australia, basal grade. The largest, i.e. the 'Core Cunon', clade
Schizomeria fruits are eaten by pigeons (Water- includes four tribes representing genera such as
house 2001), in New Guinea by fruit bats, arboreal Caldcluvia, Callicoma, Geissois and Weinmannia,
marsupials and birds, including flightless cas- with paniculiform, racemose, or capitate inflores-
sowaries (Pratt 1983). Davidsonia fruits are eaten cences and usually capsular fruits. About two-
by cassowaries (Cooper and Cooper 1994). Most thirds of Cunoniaceae species richness is in a clade
species probably require moderate to high light of the three largest genera, Weinmannia, Cunonia,
levels for successful establishment and are long- and Pancheria (tribe Cunonieae).
lived in the canopy (e.g. Lusk 1999). Most recognized genera have clear apomorphies
(Bradford and Barnes 2001). Weinmannia was
PHYTOCHEMISTRY. Large amounts of aluminium weakly supported as monophyletic, based on the
and mucilage are typically found in the leaves, and presence of hairs on seeds (Bradford 1998). In
the family is characterized by a richly developed addition, most species studied have a multicellu-
phenyl propanoid metabolism. In the leaves and lar hair base unique in the family (Barnes, unpubl.
bark, both condensed and hydrolyzable tannins data). However, molecular data is thus far incon-
are prominent, the latter sometimes based on clusive as to the monophyly of Weinmannia with
mono- and di-0-methylellagic acid. Among respect to Cunonia (Bradford 1999, 2002; Bradford
flavonol glycosides, those based on kaempferol and Barnes 200 1), although the five sections of
and quercetin are widespread, whereas myricetin Weinmannia are clearly monophyletic. Although
glycosides are found in Ceratopetalum, Bauera, no known morphological character unites the
Davidsonia and some Weinmannia, often in com- species of Geissois, cladistic analyses of DNA
bination with prodelphinidin (Hegnauer 1964, sequences suggest that Geissois is monophyletic.
1989; Bate-Smith 1977). Wollenweber et al. (2000) However, greater sampling of species and loci is
found numerous flavonoids in the leaf and bud warranted (Bradford and Barnes 2001).
excretes of Eucryphia, some species of which also
share the possession of azaleatin with other FAMILY CIRCUMSCRIPTION AND AFFINITIES. At
Cunoniaceae. one time or another, Bauera, Eucryphia, Davidso-
nia and Brunellia have either been retained as dis-
SUBDIVISION AND RELATIONSHIPS WITHIN THE tinct families, or included within Cunoniaceae
FAMILY. Cladistic analyses of Cunoniaceae by s.l. Bauera is sometimes placed in Saxifragaceae,
Hufford and Dickison (1992) and Orozco (1997) where Cunoniaceae in general once resided. Until
used only morphology and could not resolve or recently, little has been known about how this
provide strong support for many clades. With group of families relates to other angiosperms.
outgroups from other members of Oxalidales, Large-scale cladistic analyses of DNA sequences
improved morphological data, and especially DNA have now placed Cunoniaceae s.l. in the order
sequences, strong support for some intrafamilial Oxalidales, along with Elaeocarpaceae (including
relationships was provided by Bradford and Tremandraceae), Cephalotaceae, Connaraceae and
Barnes (2001). Based on their phylogenetic analy- Oxalidaceae (APG 1998). On the basis of cladistic
ses, six tribes were circumscribed. Six genera could analyses of plastid DNA sequences and morphol-
not be placed in any tribe, but were placed infor- ogy (Bradford and Barnes 2001), Cunoniaceae are
mally into either a 'Basal Grade' (Davidsonia and here circumscribed to include Bauera, Eucryphia
Bauera) or a 'Core Cunon' clade (Eucryphia, Gill- and Davidsonia within the family, but Brunellia
beea and Acrophyllum). A slightly expanded analy- outside of Cunoniaceae and possibly near
sis to include Hooglandia showed that it is not a Cephalotus. Aphanopetalum, traditionally placed
member of any tribe but falls within the 'Basal in Cunoniaceae, belongs in Saxifragales (Dickison
Grade' and appears closely related to Aistopetalum et al. 1994; Bradford and Barnes 2001; Fishbein
(Sweeney and Bradford 2003). et al. 2001). Relationships within Oxalidales need
Cunoniaceae 97
further study, but Tremandraceae are nested tribes, and the fossil record. Caldcluvia paniculata
within Elaeocarpaceae; Brunelliaceae, Cephal- from South America is the sister taxon to
otaceae and Elaeocarpaceae are probably more Opocunonia, Ackama and Spiraeopsis from
closely related to Cunoniaceae than to Oxalidaceae Austro-Malesia and New Zealand (tribe Caldclu-
and Connaraceae. vieae); Lamanonia from South America is the
Only by examining a combination of morpho- sister group to Geissois and Pseudoweinmannia
logical characters can one distinguish Cunoni- from Australia and the South Pacific (tribe Geis-
aceae from the related families Brunelliaceae and soieae); and Platylophus from South Africa is the
Elaeocarpaceae. A single pair of interpetiolar stip- sister taxon to Anodopetalum and Ceratopetalum
ules is synapomorphic for a Cunoniaceae clade, from Australia and New Guinea (tribe
and is unique among Oxalidales families. However, Schizomerieae). Cunonia has one species endemic
while Brunelliaceae and Elaeocarpaceae have to South Africa and c. 25 species endemic to New
lateral stipules, which may be one way to differen- Caledonia. Two species in section Weinmannia
tiate among these families, lateral stipules are alsofrom the Mascarene Islands (Indian Ocean) are
derived within Cunoniaceae (Geissoieae, Bauera, most closely related to about 70 species from the
Caldcluvia and Gillbeea). Character combinations Americas (Bradford 1998, 2002; Bradford and
typical for these three families are: Barnes 2001).
Although the majority of species inhabit tropi-
- Cunoniaceae: leaves usually opposite, simple or cal montane rainforests, there are many lowland
compound, toothed; stipules often interpetio- rainforest species in Madagascar, eastern Malesia,
lar; filaments exceeding the petals; floral disc western Melanesia and Australia, and some in
inter- or intra-staminal. cool-temperate rainforests in Chile, Tasmania and
- Brunelliaceae: leaves usually opposite, simple or New Zealand. A few species occur in moderately
compound, toothed; stipules lateral; brown dry habitats, such as Lamanonia and Weinmannia
hairs on stems and leaves; dioecious; floral disc in South American cerrado vegetation, or Codia
intra-staminal; large decurrent stigmas; shiny, and Pancheria in seasonally dry maquis in New
arillate seeds exposed from follicles. Caledonia. In dry Australian heathlands, Bauera
- Elaeocarpaceae: leaves usually alternate, simple, may be found in riparian habitats.
toothed; stipules lateral; petals often fringed; Species occur on a diversity of soils, from
anthers open by terminal pores or slits; floral lowland clays to upland humus. Many species are
disc extra-stamina!. opportunists on poorly developed, shallow soils
derived from substrates such as sandstone, quartz
DISTRIBUTION AND HABITATS. Cunoniaceae are or lava, suggesting a good primary colonizing
predominantly found in tropical (especially ability. Acrophyllum australe is narrowly endemic
montane) and wet temperate regions of the South- on soils derived from exposed layers of shale along
ern Hemisphere, with most generic diversity in moist sandstone cliffs in Australia's Blue Moun-
Australia and Tasmania (16 genera), New Guinea tains (Cooper 1986). Cunoniaceae often suc-
(9 genera), and New Caledonia (7 genera). Some cessfully occupy ultramafic (e.g. serpentine)
groups are widespread in Malesia, Melanesia and substrates. Some species occur both on and off
Polynesia (especially Weinmannia). Two genera ultramafics, but many are substrate endemics,
are in South Africa (Cunonia and Platylophus). which is especially common in New Caledonia in
About 40 species of Weinmannia are located in the genera Cunonia, Pancheria, Codia, Acsmithia
Madagascar, the Comores and the Mascarenes. In and Geissois. Weinmannia clemensiae Steenis
the Americas only 4 genera occur, but the region and W. devogelii H.C. Hopkins are serpentine
is species rich because of the diversity of Wein- endemics from Mt. Kinabalu, Borneo and Lake
mannia. The American distribution is from Matano, Sulawesi respectively (Hopkins 1998a).
Mexico and the West Indies to Central America Some plants living on ultramafics tend to hyper-
(Weinmannia), and through tropical South accumulate heavy metals in their leaves (Jaffre
America (Weinmannia and Lamanonia) to tem- 1980).
perate South America (Caldcluvia, Eucryphia, Because many Cunoniaceae species tend to be
Lamanonia and Weinmannia). locally common and their pollen has a short
The Cunoniaceae lineage was probably wide- disperal radius (McGlone 1982), palaeoecologists
spread and diversified prior to the breakup of have used pollen records to track the movement of
Gondwana based on phylogenetic reconstruc- Cunoniaceae species, especially Weinmannia, and
tions, continental disjunctions of some genera and thereby infer the historic locations of vegetation
associations (Pocknall 1980; McGlone 1983; including wasps, flies, Lepidoptera and beetles,
Helmens and Kuhry 1986; Hooghiemstra 1989; although bees of various sorts are likely to be the
Villagran 1990; Heusser 1993). This indicates, for main pollinators in most cases (see Economic
example, that many species characteristic of upper Importance) and have been reported from genera
elevation Andean vegetation occurred in the lower with a variety of inflorescence types, including
Amazon basin during glacial periods (Bush et al. Codia, Cunonia, Eucryphia, Pancheria, Platylophus
1990). and Weinmannia.
Some flower-visiting vertebrates are probably
FLORAL AND POLLINATION BIOLOGY. Cunoni- part of a suite of generalist potential pollinators;
aceae flowers are generally small to medium sized for example, various birds have been reported
and, when not individually showy, they are fre- from the flowers Opocunonia, Weinmannia, Platy-
quently massed into highly visible inflorescences. lophus and Cunonia capensis, and geckos have
Exceptions to this include Eucryphia, with large, been seen licking the white, candle-like racemes of
white, solitary flowers, up to 6 em in diameter in C. balansae Brongn. & Gris in New Caledonia
E. glutinosa, and Anodopetalum which has in- (Hopkins, Bradford and Donovan, pers. obs. 2000).
conspicuous, solitary, green flowers (Barnes and A few Cunon species appear to be specialised for
Rozefelds 2000). bird-pollination, such as the Pacific species of
Floral colour is generally white or a pale pastel Geissois whose red, generally cauliflorous racemes
shade, including cream-yellow, greenish or pink, attract numerous Meliphagidae, Nectariniidae and
but magenta (some Bauera), purplish (some Spi- Zosteropidae, as well as some insects, geckos
raeopsis and Cunonia) or bright scarlet (Pacific (Bavary 1869; S. Zona, pers. comm.) and perhaps
Geissois, some Cunonia) also occur. A range of pteropodid bats (MacKee, specimen labels). In
floral structures contributes to the colour, includ- New Caledonia, Cunonia macrophylla Brongn.
ing the sepals, especially when the petals are small, & Gris has bright green, scentless flowers in
fugaceous or absent. Filaments are usually white, one-sided, bottle-brush racemes that attract
sometimes with contrasting yellow or pink Meliphagidae. Anemophily is possible in the dioe-
anthers, and the disc and gynoecium sometimes cious Vesselowskya from eastern Australia, which
contrast sharply with the pale tepals and stamens. has catkin-like racemes, decurrent stigmas and
The numerous, often exserted stamens in which grows gregariously along streams and rivers
Eucryphia, Bauera, Lamanonia and Geissois (Bradford, pers. obs.; Rozefelds et al. 2001).
may also enhance visibility. Included stamens As inferred from floral morphology, most species
occur in Davidsonia. have bisexual, homogamous flowers. However,
Where present, the floral disc appears nectarif- dioecy occurs in Spiraeanthemum, Hooglandia,
erous but the abundance and composition of the Pancheria, Vesselowskya and some groups of
nectar have not been assessed. Large nectar Weinmannia, where it may have evolved three
droplets are produced in red-flowered Geissois, but times (Bradford 1998), being present in sections
in some genera the volumes per flower may be Weinmannia, Fasciculatae and Leiospermum. In
very small indeed. For instance, honey bees at the latter genus at least, dioecy is frequently in com-
capitula of Pancheria gatopensis Guillaumin plete ('leaky'), with geographical variation in
(Plateau de Tango) inserted their tongues into sexual expression occurring in W. fraxinea, and
flowers, suggesting foraging for nectar, but none temporal switches, usually from male to bisexual
was visible to the human eye and it was scarcely or vice versa, are seen in several Pacific species
detectable to the human tongue (Donovan, (Hopkins 1998a, 1998b; Hopkins and Florence
Bradford and Hopkins, pers. obs. 2000). The 1998). Protandry is marked in Opocunonia and
small, rounded, shiny structures at each tip of Spiraeopsis, and perhaps occurs in Ackama (all
the crescent-shaped petals in Gillbeea are also tribe Caldcluvieae), and andromonoecy has been
apparently secretory (Endress 1994). Where infor- reported in some species of both Schizomeria and
mation is available, flowers are generally sweetly Cunonia.
scented and sometimes cloyingly so (e.g. Codia),
with notable exceptions in Pacific Geissois and PALAEOBOTANY. The fossil record of Cunoni-
several species of Cunonia. aceae has been reviewed by Barnes and colleagues
Cunoniaceae flowers are generally cup-, dish- or (1999, 2001). Confirmed macrofossils of Cunoni-
bowl-shaped and most species are probably ento- aceae represent 11 genera from a broad spectrum
mophilous, although data are few and largely anec- of morphological diversity in the family. Most of
dotal. A wide range of visitors has been noted, these records are leaf compressions or impressions
Cunoniaceae 99
or preserved cuticles and, with the exception of a Elaeocarpaceae and Cunoniaceae leaf macrofos-
single, fossil infructescence from New Zealand sils may be difficult to distinguish, especially when
(?Weinmannia sp. indet.), all accepted leaf and specimens lack preserved cuticle, or are incom-
reproductive macrofossils of Cunoniaceae are plete. This is the case with many historical records.
from Australian Cainozoic deposits. The oldest It is also difficult to distinguish between Elaeo-
of these records are from the Late Palaeocene carpaceae and Cunoniaceae pollen, which are
(Eucryphia ), Eocene (Acsmithia/Spiraeanthemum, both very small and not easily identifiable to the
Ceratopetalum, Codia, Weinmannia) and Oligo- generic level.
cene (Acsmithia, Callicoma, Ceratopetalum, Given the above caveats, the oldest pollen
Schizomeria, Vesselowskya), suggesting that the records of Cunoniaceae date from the Late
family had diversified by the mid to late Palaeocene, Eocene and Oligocene. These ancient
Cretaceous or Early Palaeocene. deposits are evidence of regional extinction of
A Late Cretaceous fossil flower from Sweden some taxa, such as Cunoniaceae-dominated
has been assigned to a fossil genus within forests in the Late Palaeocene of South Australia
Cunoniaceae, Platydiscus Schonenberger & Friis (Sluiter 1991), Weinmannia from Australia (Hill
(Schonenberger et al. 2001), although strong and Macphail 1983), and Gillbeea (Concolpites
similarities to Anisophylleaceae (Matthews et al. leptos fossil pollen type) from South America
2001) make this placement less certain. (Romero and Castro 1986) and southern Australia
Fossil wood identifications include Weinman- (Stover and Partridge 1973).
nioxylon eucryphioides from Eocene sediments at
King George Island (Poole et al. 2001), which has EcoNOMIC IMPORTANCE. References to human
strong affinities to Eucryphia, and two Weinman- use of Cunoniaceae species come mainly from
nioxylon species from Late Cretaceous sediments local or regional economic botany texts or floras,
in Antarctica (Poole et al. 2000; Poole and Cantrill and industry sources (e.g. timber, apiculture). The
2001). The fossil Cunonioxylon parenchymatosum wood of Cunoniaceae species is used in general
from the Eocene of Lower Saxony in Germany was construction as poles, wall panelling, flooring,
directly compared to wood of extant Cunonia plywood, furniture, cabinetry, joinery, tools, boat
(Gottwald 1992). There is significant taxonomic keels, canoes, packing boxes, musical instruments
confusion generated by Cunoniaceae fossil wood and, historically, to make wagons and coaches
identifications, as a single fossil genus may contain ( Ceratopetalum apetalum, Cunonia capensis ). The
species with affinities to numerous extant genera. most important commercial timber regions for
On this basis, Cunoniaceae fossil wood species the family are Malesia, Australia, and the western
should not be considered to represent extant Pacific, principally of the genera Ackama, Calli-
genera. However, Antarctic specimens suggest coma, Ceratopetalum, Eucryphia, Geissois,
that the family has been a common element of Pseudoweinmannia, Schizomeria and Weinman-
southern temperate rainforest for over 100 million nia. In the Americas, Weinmannia species are
years. mostly of local construction and cabinetry use.
Leaf and reproductive macrofossils indicate that The Chilean Eucryphia cordifolia is valued for the
the extant geographic range of some genera has high quality of its wood, which has diverse uses.
been reduced or is different to that in the past. Mascarene island and South African species are
Acsmithia, Callicoma, Schizomeria, Vesselowskya used mainly for furniture making. Where the
and Weinmanniaphyllum (form genus encom- human population relies on wood for fuel, espe-
passing leaf macrofossils indistinguishable from cially in the Andes and Madagascar, Weinmannia
Weinmannia and Cunonia) were present in trees are often chopped down or whole forests are
Tasmania during the Early Oligocene but are no burned to make charcoal. Several species have
longer locally extant (Carpenter and Buchanan been used in local tanning industries (e.g.
1993; Barnes 1999; Barnes and Hill1999b; Barnes Weinmannia tinctoria).
et al. 2001). Ceratopetalum, Codia and Spiraean- Bee keepers use Weinmannia forests for com-
themum!Acsmithia are known from Eocene- mercial production of honey in Madagascar and
Oligocene deposits in southern and south-western New Zealand. Eucryphia-dominated forests are
Australia (Carpenter and Pole 1995; Barnes 1999; honey resources in Tasmania and Chile. Platylopus
Barnes and Hill 1999a, 1999b; Barnes et al. 2001), trifoliatus honey is popular in South Africa. David-
which are regions presently too dry for extant sonia is a highly regarded 'bush fruit' in Australia
species to occur, based on the climate envelope of where it is used for jams and wine, and Schizome-
their modern habitat. ria fruits may also be used in jams and pies.
Horticultural use is greatest in Australia, including disturbed habitats. Outside of New Caledonia, the
Ceratopetalum gummiferum (the New South Wales most imperilled, cunon-rich Pacific flora is Fiji,
Christmas Bush). Because a variety of ethnomed- which has five genera and 13 species with 12
icinal sources indicate that Cunoniaceae extracts endemic (Smith 1985). The spectacular Geissois
may treat infections, a research program in New species are most endangered because they live at
Caledonia has begun on the pharmacology of relatively low elevations where there is the great-
Cunoniaceae (Fogliani et al. 2002). est loss of habitat from agriculture, grazing,
and the invasion of exotics.
CoNSERVATION. Little has been done generally to Madagascar is home to about 40 endemic
identify species of Cunoniaceae at risk of extinc- species of Weinmannia that are members of two
tion, although 28 taxa are listed as threatened in endemic and diverse sections. Species occur pri-
the most recent global assessment (Walter and marily in the northern and eastern regions of the
Gillett 1998). Unfortunately, many species occur in island, from sea level (which is unusual in Wein-
areas with extremely high deforestation rates, such mannia) to near the summit of the highest peaks.
as portions of the Andes and Madagascar. These Burning and clearing forest to grow hill rice is the
are also the areas where systematic understanding major cause of habitat loss for Weinmannia, espe-
is poorest. The greatest threats are the conversion cially in the lowlands and central plateau. Com-
of forested land to subsistence agricultural land, prehensive systematic information is lacking and
habitat degradation from logging, grazing, fire, several species are undescribed (Bradford and
mining and invasion of exotic species, and Miller 2001). Very little habitat remains for two
expanding suburban development. In the follow- endemic Mascarene island species of Weinmannia
ing, some regions are selected where high diver- (Cadet 1984), and this is being maintained by
sity and/or endemism of Cunoniaceae coincide manual weeding of non-native, invasive plants
with elevated rates of environmental interference. (W. Strahm, pers comm.).
The tropical rainforest in the Atherton Tableland In tropical America, Weinmannia is highly
of NE Queensland harbours 13 endemic species in diversified in the Andes where natural vegetation
eight genera (Hyland and Whiffin 1993) and there- is being destroyed rapidly in some regions, espe-
fore it is especially critical for Cunoniaceae con- cially Ecuador and Colombia. Presently, montane
servation. Small areas of upland habitat in forest is being eliminated by fires being set to clear
north-eastern and eastern Australia support two farmland below, and to improve grazing and create
narrow endemic Eucryphia and Ceratopetalum more grasslands above. Lamanonia occurs in
species (Forster and Hyland 1997; Rozefelds and moist, evergreen, semideciduous forests and
Barnes 2002). Bauera sessiliflora and Acrophyllum savannah-like vegetation of southern Brazil,
australe are lowland forest/heath endemics, the northern Argentina and northern Paraguay, in
latter especially being at risk of extinction from habitats that are some of the most endangered
frequent fires and encroaching human distur- in South America due to expanding soybean cul-
bance. Davidsonia johnsonii and D. jerseyana tivation and cattle grazing. Eucryphia glutinosa
are restricted to a small region of north-eastern occurs in riparian habitats in the Mediterranean
New South Wales and south-eastern Queensland climate region of Chile and is threatened by
heavily impacted by small-scale agriculture, and grazing and water management (Munoz Pizarro
the former species may not reproduce sexually 1973).
(Harden and Williams 2000).
New Caledonia, with seven genera and over 90 KEY TO THE GENERA
endemic species occurring in a relatively small
1. Leaves alternate 5. Davidsonia
area, is a major conservation priority for Cunoni- - Leaves decussate or whorled 2
aceae. Many species are restricted to a single or 2. Flowers solitary in leaf axils 3
a few populations, often on top of mountains. - Flowers borne on an inflorescence axis 5
Species occurring at the highest elevations are 3. Flowers subtended by a pedicel only 6. Bauera
somewhat protected from the anthropogenic fires - Flowers subtended by a pedicel, pair of bracteoles and a
peduncle 4
that frequently overrun vegetation at lower eleva- 4. Polyandric; carpels usually >3 11. Eucryphia
tions (Lowry 1998), but may be more susceptible - Diplostemonous; bicarpellate 9. Anodopetalum
to vanishing climatic envelopes due to global 5. Inflorescences condensed spherical heads (capitula) 6
warming (Still et al. 1999). Currently, the most - Inflorescences not capitate, e.g. racemose, paniculiform or
cymiform 9
endangered species have narrow ranges at low to
6. Flowers unisexual; fruits capsular; leaves (2)3-4 per
mid elevations and do not establish well in fire- node 25. Pancheria
Cunoniaceae 101
- Flowers bisexual; fruits indehiscent or capsular; leaves 2 - Leaves pinnately compound; petals 0 27
per node, rarely more 7 27. Flowers hermaphroditic; gynoecium of 4-6 locules
7. Fruits capsular; leaves deeply serrate 23. Callicoma 3. Aistopetalum
Fruits indehiscent; leaves entire or weakly serrate 8 - Flowers unisexual (plants dioecious); gynoecium
8. Leaves entire, rarely toothed; ovary inferior; calyx small unilocular 4. Hooglandia
22. Codia 28. Ovary half inferior; sepals enlarged and often colourful in
- Leaves toothed; ovary half inferior; calyx prominent fruit; petiole swollen and articulated at junction of blade
21. Pullea 8. Ceratopetalum
9. Flowers maturing synchronously to acropetally in an inflo- - Ovary superior; sepals inconspicuous in fruit; petiole not
rescence 10 swollen or articulated at junction of blade
- Flowers maturing at different times in basipetalous 10. Platylophus
sequence 23
10. Polyandric (more than twice as many stamens as perianth GENERA OF CUNONIACEAE
parts); leaves palmately compound or trifoliate; inflores-
cences racemes 11 Tribes and genera are mainly organized according
- Diplostemonous, rarely haplostemonous, leaves imparip- to the branching sequence of cladograms in Brad-
innate, unifoliolate or rarely palmately compound; inflo- ford and Barnes (2001 with Figs. 4 and 5), who also
rescences racemes, capitula, panicles or cymes 13 give descriptions of tribes, and with some modifi-
11. Leaves 3-foliolate; fruit indehiscent, dry, densely covered in
wavy hairs 15. Pseudoweinmannia
cation based on Sweeney and Bradford {2003),
- Leaves 3- or more-foliolate; fruit capsular, glabrous or notably the inclusion of Hooglandia and the new
hairy 12 placement of Aistopetalum. Several genera with
12. Stipules lateral and unfused; flowers never red uncertain relationships are not placed in a tribe,
14. Lamanonia but informally included in either a 'Basal Grade'
- Stipules axillary or lateral and fused; flowers red or not
16. Geissois
or 'Core Cunon Clade'.
13. Inflorescence axes unbranched (racemose) 14
- Inflorescence axes branched (e.g. cymes and panicles) Characters applying to all genera unless
16 stated otherwise
14. Leaves palmately compound or trifoliolate; flowers 3-
merous; stigma decurrent 24. Vesselowskya Leaves decussate; non-stipellate; margin toothed;
- Leaves simple or imparipinnate; flowers 4-5-merous; hairs simple; stipules interpetiolar; one pair of
stigma terminal 15 stipules borne per node. Flowers bisexual, pedi-
15. Seeds with hairs; fruits dehisce basipetally; nectary disc
distinct from ovary 27. Weinmannia cellate; perianth 4-5-merous, of sepals and petals,
- Seeds without hairs; fruits dehisce circumbasally; nectary calyx valvate; androecium diplostemonous; ovary
disc adnate to ovary 26. Cunonia bicarpellate, syncarpous, superior, stylodia free,
16. Carpels fused; petals + or 0 17 stigmas terminal.
- Carpels unfused or only basally so; petals 0 22
17. Petals 0; ovary half-inferior; fruit indehiscent; leaves uni-
foliolate 21. Pullea
- Petals +; ovary superior; fruit capsular; leaves unifoliolate I. TRIBE SPIRAEANTHEMEAE Engl. {1928).
or imparipinnate 18
18. Leaves 3 per node, unifoliolate 13. Acrophyllum 1. Acsmithia Hoogl.
- Leaves 2 per node, unifoliolate or imparipinnate 19
19. Stipules lateral; leaves unifoliolate; flowers not protan- Acsmithia Hoogl., Blumea 25: 492-501 (1979), rev.; Hoogland,
drous 17. Caldcluvia Bull. Mus. Nat!. Hist. Nat., B Adansonia 9: 393-397 (1987).
- Stipules interpetiolar; leaves imparipinnate, rarely unifoli- Spiraeanthemum A.Gray (1854) p.p.
olate; flowers often protandrous 20
20. Stellate hairs + on leaves 20. Spiraeopsis
- Stellate hairs 0 on leaves 21
Trees and shrubs; nodes of stems knobby. Leaves
21. Inflorescence conical in outline (paniculate); seeds with whorled, unifoliolate, margin entire or toothed,
hairs 19. Ackama pocket domatia along the midrib often present.
- Inflorescence rounded in outline (cymiform); seeds Inflorescence axillary, axillary and terminal, or
without hairs 18. Opocunonia sometimes cauliflorous, paniculate; floral matura-
22. Leaves 2 per node; flowers unisexual
2. Spiraeanthemum tion synchronous; perianth of one whorl; nectary
- Leaves 3-4 per node; flowers bisexual 1. Acsmithia of small segments; carpels 2-5, free almost to base.
23 Inflorescence a 3-flowered cyme 9.Anodopetalum Fruits follicular; seeds winged. Fourteen species:
- Inflorescence of numerous flowers 24 4 from E. Malesia, 1 from Australia, 8 from New
24. Stellate hairs + on floral parts; fruit dry with laterally Caledonia, and 1 from Fiji.
expanded carpel walls 12. Gillbeea
- Stellate hairs 0 on floral parts; fruit fleshy or dry, not as
above 25
2. Spiraeanthemum A. Gray Fig. 31
25. Fruit drupaceous 26
- Fruit dry and indehiscent 28 Spiraeanthemum A. Gray, Proc. Am. Acad. Arts 3: 128 (1854);
26. Leaves simple; petals+ 7. Schizomeria Hoogland, Blumea 25: 501-505 (1979), rev.
Fig. 32. Cunoniaceae. Aistopetalum viticoides. A Flowering

branch. B Flower. C Same, longitudinally sectioned. D Fruit.
Drawn by R. van Creve!. (Hoogland 1960)
Fig. 31. Cunoniaceae. Spiraeanthemum katakata. A Flowering perianth of one 4-6-merous whorl; floral disc
branch. B Male flower, vertical section. C The same, transverse large, adnate to the ovary; gynoecium of 4-6
section of disk. D Female flower. E Same, pistil removed. F
Same, vertical section. G Fruit. H Seed. Drawn by R. van Creve!. carpels. Fruit drupaceous, one seed per carpel. Two
(Hoogland 1960) species (A. multiflorum Schltr. and A. viticoides
Schltr.), endemic to New Guinea.
Trees and shrubs. Leaves unifoliolate, often with 4. Hooglandia McPherson & Lowry
pocket domatia along the midvein. Inflorescence Hooglandia McPherson & Lowry, Ann. Missouri Bot. Gard. (in
axillary; paniculate; floral maturation synchro- prep.); Sweeney and Bradford, Ann. Missouri Bot. Gard. (in
nous. Flowers unisexual, dioecious; perianth of prep.).
one whorl, nectary of small segments; gynoecium
of (2- )4- 5(6) free carpels. Fruit follicular; seeds Trees. Leaves imparipinnate; stipules interpetiolar
winged, 2 per carpel. Six species from the or occasionally divided partially or completely to
Bismarck Archipelago, Bougainville, Solomons, the base. Inflorescence axillary; paniculate; floral
Vanuatu, Fiji, and Samoa. maturation subsynchronous with terminal flowers
opening first. Flowers unisexual, dioecious; peri-
anth of one whorl, 4-merous, imbricate; nectary
II. UNPLACED 'BASAL GRADE' GENERA adnate to the ovary; gynoecium bilaterally sym-
metrical, with a single locule and a single bent
3. Aistopetalum Schltr. Fig. 32 style, but with a small bump opposite the style sug-
Aistopetalum Schltr., Bot. Jahrb. Syst. 52: 1 42-144 (1914); Perry, gesting the fusion of two carpels, two ovules. Fruit
J. Arnold Arb. 30: 158- 159 ( 1949); Hoogland, Austral. J. Bot. drupaceous, flattened and elongate, with a thin
8: 333- 337 (1960), rev. fleshy exocarp, hard endocarp, and a single large
seed. One species, Hooglandia ignambiensis
Trees. Leaves imparipinnate. Inflorescence a xil- McPherson and Lowry, from upper elevation
lary; paniculiform, large, with decussate branch- montane forest in the Panie range of north-eastern
ing throughout; floral maturation centrifugal; New Caledonia.
Cunoniaceae 103
5. Davidsonia F. Muell. floral maturation basipetalous. Flowers bisexual

or male and plants hermaphrodite or
Davidsonia F. Muell., Fragm. 4: 4 (1867); Bange, Blumea 7:
293-296 (1952); Harden & Williams, Telopea 8: 413-428 andromonecious; perianth (4)5( 6)-merous; petals
(2000), rev. tridentate; floral nectary annular, often with large
lobes. Fruit drupaceous. Ten species distributed in
Trees. Leaves alternate, imparipinnate with pro- the Moluccas, New Guinea, the Solomons, and
minent teeth and a jagged rachis; hairs rigid, eastern Australia.
caducous, mildly urticating. Inflorescence axillary,
terminally, and adventitiously along branches 8. Ceratopetalum Sm.
and the trunk, an elongate panicle; inflorescence
very large in D. pruriens F.Muell. but smaller in Ceratopetalum Sm., Specim. Bot. New Holl. 1: 9, t. 3 (1793);
other taxa and sometimes nearly racemose in D. Hoogland, Austral. J. Bot. 8: 318-328 (1960), rev.; Hoogland,
Brunonia 4:213-215 (1981); Stanley & Ross, Flora of South-
jerseyana (F.M. Bailey) Harden & Williams and D. eastern Queensland 1: 225 (1983); Harden, Flora of New
johnsonii Harden & Williams; floral maturation South Wales 1: 519 (1990); Rozefelds & Barnes, Int. J. Plant
basipetalous. Flowers with a single perianth whorl, Sci.163: 651-673 (2002).
the lobes fused half their length; nectary of small
segments. Fruit drupaceous, with fleshy, edible Trees and shrubs, resinous. Leaves unifoliolate
pulp containing 4 seeds. Three species from or trifoliolate. Inflorescence terminal or axillary;
Eastern Australia. Used as an ornamental and fruit thrysoid to cymiform, often retaining decussate
tree to make jams and wine, with the common branching, or somewhat helical distally; floral
name 'Davidson's Plum'. maturation basipetalous; petals 0 or small and
deeply incised (C. gummiferum Sm.); nectary
large, adnate to the ovary; ovary half inferior.
6. Bauera Banks ex Andrews
Fruit dry, indehiscent, with enlarged, often reddish
Bauera Banks ex Andrews, Bot. Repos. 3: t. 198 (1801); Bailey, calyx lobes. About 9 species occurring in Eastern
The Queensland Flora, 2: 543 (1900); Stanley & Ross, Flora Australia and New Guinea, the 'Christmas Bush'
of South-eastern Queensland 1: 219 (1983); Everett in (C. gummiferum Sm.) used ornamentally, and
Harden, Flora of New South Wales 1: 521 (1990).
'Coachwood' (C. apetalum D.Don) a valued timber
species.
Scrambling shrub with narrow stems. Leaves uni-
foliolate (appearing 3-foliolate, but lateral 'leaflets'
considered to be modified stipules, see Dickison 9. Anodopetalum A. Cunn. ex Endl.
and Rutishauser 1990), sessile or nearly so; stip-
Anodopetalum A. Cunn. ex End!., Gen. Pl. 11: 818 (1839);
ules leaf-like, borne beside each leaf. Flowers soli- Curtis, The Student's Flora of Tasmania 1: 176 (1956);
tary in leaf axils; perianth parts variable in Barnes and Rozefelds,Austral. Syst. Bot.13: 267-282 (2000).
number, corolla of one or sometimes two whorls;
stamens numerous; floral nectary 0; ovary of 2- Trees and shrubs. Leaves unifoliolate or trifolio-
3 carpels. Fruit capsular; seeds elliptical, late. Inflorescence axillary; reduced to a single
appendages 0. Four species in eastern and south- flower, (a 3-flowered cyme); floral maturation
ern Australia and Tasmania. basipetalous; calyx large, petals slender and
apically notched; anthers with prominent horn;
nectary annular. Fruit capsular. One species, A.
III. TRIBE SCHIZOMERIEAE J.C. Bradford & biglandulosum (A. Cunn ex. Hook.) Hook. f.,
R.W. Barnes (2001). endemic to Tasmania. The plant's unusual growth
form can produce dense, overlaying thickets of
7. Schizomeria D.Don branches (Barker and Brown 1994).
Schizomeria D. Don, Edinburgh New Philos. J. 9: 94- 95 (1830);
Perry, J. Arnold Arb. 30: 151-158 (1949); Whitmore, Guide 10. Platylophus D. Don
to the Forest of the British Solomon Islands (1966); Stanley
& Ross, Flora of South-eastern Queensland 1: 225 (1983); Platylophus D. Don, Edinburgh New Philos. J. 9: 92 (1830);
Harden, Flora of New South Wales 1: 520 (1990); Fortune Coates Palgrave, Trees of Southern Africa (1983).
Hopkins & Hoogland, Fl. Males. I, 16: 53-165 (2002).
Cremnobates Rid!. (1916).
Trees. Leaves 3-foliolate. Inflorescence axillary;
Trees and shrubs. Leaves unifoliolate. Inflores- thrysoid; floral maturation basipetalous; petals
cence terminal or less often axillary; thrysoid; small and tridentate; nectary annular and lobed.
104 J.C. Bradford et a!.
Fruits indehiscent, swelling into a dry bladder; unifoliolate or impanpmnate; margin entire or
seeds few small, wingless. One species, P. trifolia- toothed. Inflorescence an axillary peduncle with a
tus (L.) D.Don, in South Africa. single, pedicellate flower. Flowers large, enclosed
by imbricate, apically coherent sepals that are shed
at maturity; petals 4, large and white; stamens
IV. UNPLACED 'CORE CUNON' CLADE GENERA numerous, fasciculate, at the margins of a broad
receptacle; ovary of 4-14 fused carpels. Fruit
11. Eucryphia Cav. Fig. 33 capsular; seeds winged, numerous. Seven species
from E Australia (3 spp.), Tasmania (2 spp.),
Eucryphia Cav., Icon. 4: 48, t. 372 (1797}; J. Bausch, Bull. Misc.
Inform. 8: 317-349 (1938}; Curtis, The Student's Flora of and southern S America (2 spp.). Cultivated as an
Tasmania 1: 179 (1956}; Harden, Flora of New South Wales ornamental.
1: 523 (1990}; P.I. Forst. & B. Hyland, Austrobaileya 4:
589-596 (1997).
12. Gillbeea F. Muell.
Trees and shrubs; resinous. Leaves evergreen and Gillbeea F. Muell., Fragm. 5: 17 (1865}; Hoogland, Austral. J.
peltate with white undersurface, or deciduous and Bot. 8: 328-333 (1960), rev; Rozefelds & Pellow, Nordic J. Bot.
lacking peltations in E. glutinosa (P. & E.) Baill.; 20: 435-441 (2000}.
Trees. Leaves imparipinnate; stipules in lateral

pairs; stellate hairs restricted to inflorescence axes
and flowers. Inflorescence axillary and terminal;
paniculiform; floral maturation basipetalous;
petals bifid with apical glands; nectary annular,
large; gynoecium of three fused carpels. Fruit
indehiscent, dry, with carpel walls expanding to
form lateral wings. Three species, two from north-
eastern Australia (G. adenopetala F. Muell. and
G. whypallana Rozefelds & Pellow) and one from
New Guinea (G. papuana Schltr.).
13. Acrophyllum Benth. Fig. 34

Acrophyllum Benth. in Maund, Botanist 2: t. 95 (1838};
Hoogland, Aust. J. Bot. 8: 318-341 (1960}; Hoogland,
Brunonia 4: 216 (1981).
Calycomis D. Don (1830).
Long-stemmed shrub. Leaves in whorls of 3,

unifoliolate, margin deeply serrate, indumentum
on undersurface white, peltate. Inflorescence at
several successive nodes along the main stem, pro-
ducing small, axillary panicles, the first node
sometimes branching to form a lateral inflores-
cence, and the main axis resuming vegetative
growth; floral maturation synchronous; perianth
5-merous; nectary small, annular at base of ovary,
or 0. Fruit capsular; seeds tiny, ellipsoid,
unwinged, with a granular surface. One species, A.
australe (A. Cunn.) Hoogland, restricted to a small
area of moist sandstone and shale cliffs between
520 and 660 m above sea level in the lower Blue
Mountains of New South Wales, Australia (Cooper
1986).
Fig. 33. Cunoniaceae. Eucryphia cordifolia. A Branch with
fruits and flower buds. B Flower. C Pistil. D Fragment of
dehisced fruit. E Seed. Drawn by E. Gasipi. (Dimitri 1972}
Cunoniaceae 105
Fig. 34. Cunoniaceae. Acrophyllum australe. A Flowering Ic .

branch. B Fruiting branch. C Flower. D Flower, vertically
sectioned outside the ovary. E Ovary, vertical section. F Fruit. Fig. 35. Cunoniaceae. Lamanonia speciosa. A Flowering
G Seeds. Drawn by R. van Crevel. (Hoogland 1960) branch. BFlower, vertical section. C Stamens. D Pistil. E Ovary,
transverse section. F Ovary, vertical section. Drawn by P.J.F.
Turpin. (St.-Hilaire 1828)
V. TRIBE GEISSOIEAE Endl. ex Meisn. (1838).
14. Lamanonia Vell. Fig. 35 Trees. Leaves 3-foliolate; stipels minute, caducous.
Lamanonia Veil., Fl. Flum. 228 (1825); Zickel & Leitao, Revista Inflorescence axillary, often proximal to current
Brasil. Bot. 16: 73-91 (1993), rev. leaves; racemose, racemes solitary, sometimes
branching into 2-3 axes, or in pairs from axillary
Small trees. Leaves palmately compound with 3-5 short-shoots; floral maturation synchronous; peri-
leaflets, stipellate, sometimes with weakly-formed anth of one, usually 6-merous whorl; androecium
tuft domatia along midribs; stipules in lateral of numerous (c. 20) stamens, strap-like basally and
pairs. Inflorescence axillary; racemose; floral mat- imbricate; nectary annular and adnate to ovary;
uration synchronous; perianth of a single, 5-7- gynoecium of 2(3) fused carpels, with false septa
merous whorl; androecium polystemonous, c. to appear four-locular. Fruits indehiscent, ovary
20-60 stamens; disc annular, adnate to ovary. covered with long, dense, wavy hairs, easily dis-
Fruits capsular; seeds winged, numerous. Five persed by wind; seeds small, unwinged. Two
species in seasonally dry forest and woodland in species, P. apetala F.M.Bailey and P. lachnocarpa
southern Brazil, Argentina and Paraguay. (F. Muell.) Engl. in E Australia.
15. Pseudoweinmannia Engl. 16. Geissois Labill.

Pseudoweinmannia Engl., Nat. Ptlanzenfam. ed. 2, 18a: 249 Geissois Labill., Sert. Austro-Caledon.: 50, t. 50 (1825); Bailey,
(1928); Bailey, The Queensland Flora 2: 542 (1900); Stanley The Queensland Flora 2: 541 (1900); Guillaumin, Bull. Soc.
& Ross, Flora of South-eastern Queensland 1: 226 (1983); Bot. France 87: 242- 245 {1940); Stanley & Ross, Flora of
Harden, Flora of New South Wales 1: 518 (1990). South-eastern Queensland 1: 226 (1983); Harden, Flora of
New South Wales 1:518 (1990); Schimaski & Rozefelds,Aust. variable in many characters that Schlechter (Bot.
Syst. Bot. 15: 221-236 (2002), reg. rev. Jahrb. Syst. 52, 1914) considered it to be several
species and even a separate genus.
Trees or shrubs. Leaves palmately compound, 3-9
leaflets; stipellate; tuft domatia occasional along
midrib; stipules axillary, rarely connate into a cup- 19. Ackama A. Cunn.
like phytotelm, or lateral, foliaceous, and often Ackama A. Cunn., Ann. Nat. Hist. 2: 358 (1839); Bailey, The
fused across the leaf axil and node. Inflorescence Queensland Flora 2:540 (1900); Allan, Flora of New Zealand
axillary and coeval to current leaves, or cauli- v. 1 (1961); Salmon, The Native Trees of New Zealand 188
florous, or medial and distal to current leaves; (1986); Stanley & Ross, Flora of South-eastern Queensland
1: 226 (1983); Harden, Flora of New South Wales 1: 519
racemose, racemes solitary or grouped into a (1990).
compound structure; floral maturation synchro-
nous; perianth a single whorl, large, 4-merous and Tree. Leaves, imparipinnate, stipellate or not,
red in Pacific species, and 5-6-merous and cream- tuft domatia along midvein, orbicular trichomes
coloured in Australian ones; stamens several to present. Inflorescence axillary, often in a series;
numerous (c. 8-32), curled in bud in Pacific paniculate; floral maturation synchronous.
species, erect in bud in Australian species; nectary Flowers sometimes protandrous, sessile or nearly
annular, adnate to base of ovary. Fruit capsular; so; nectary segmented; ovary 2- or 3-4-carpellate.
seeds winged, numerous. About 18 species distrib- Fruit capsular, with hairy seeds that lack wings.
uted in eastern Australia, Fiji, New Caledonia, Four species, 2 in New Zealand and 2 in Australia.
Vanuatu, and Santa Cruz (Solomon Islands).
20. Spiraeopsis Miq.

VI. TRIBE CALDCLUVIEAE J.C. Bradford & Spiraeopsis Miq., Fl. Ind. Bat. 1, 1: 719 (1856); Hoogland,
R.W. Barnes (2001). Blumea 25: 481-490 (1979); Fortune Hopkins & Hoogland,
Fl. Males. I, 16: 53-165 (2002).
17. Caldcluvia D.Don
Caldcluvia D. Don, Edinburgh New Philos. J. 9: 92 (1830);
Trees. Leaves imparipinnate; indumentum stellate
R. Rodriguez eta!., Flora Arborea de Chile (1983). and of small, orbicular-glandular trichomes.
Inflorescence axillary; paniculate, often large;
Trees. Leaves unifoliolate; small, tuft domatia floral maturation synchronous; disc segmented;
along the midvein; stipules in lateral pairs. Inflo- ovary of 2-5 fused carpels. Fruit capsular; seeds
rescence axillary; cymiform; floral maturation winged, small. Six species, E Malesia and Solomon
synchronous; perianth 5-merous; floral nectary Islands.
segmented. Fruit capsular; seeds small with
minute wings and a papillate surface. One species,
C. paniculata (Cav.) D. Don, temperate rain forests VII. TRIBE CODIEAE G. Don {1834).
of Chile and Argentina.
21. Pullea Schltr.
18. Opocunonia Schltr. Pullea Schltr., Bot. Jahrb. Syst. 52: 164-166 (1914); Bailey, The
Queensland Flora 2: 536 (1900); Perry, J. Arnold Arb. 30:
Opocunonia Schltr, Bot. Jahrb. Syst. 52: 154-155 (1914); Perry, 163-165 (1949); Hoogland, Blumea 25: 490-492 (1979), rev.
J. Arnold Arb. 30: 143-145 (1949); Hoogland, Blumea 25: Stutzeria F. Muell. (1865).
486-487 (1979); Fortune Hopkins & Hoogland, Fl. Males. I,
16: 53-165 (2002). Trees. Leaves decussate or in whorls of 3; unifolio-
late. Inflorescence axillary, terminal, or cauliflor-
Trees. Leaves imparipinnate; tuft domatia often ous; paniculate, often in a series from a common
present along midrib. Inflorescence axillary or ter- peduncle, flowers well dispersed or grouped in
minal; cymiform; floral maturation synchronous. glomerules; floral maturation synchronous.
Flowers protandrous; perianth (5)6-merous; Flowers sessile or with a short pedicel; perianth a
nectary annular. Fruit capsular; seeds winged, single, 5-merous whorl, imbricate; filaments
numerous (c. 40). According to Hoogland {1979), strap-like; nectary of 5 segments; gynoecium half-
there is one species, 0. nymanii (K. Schum.) inferior. Fruit dry, indehiscent, with enlarged, per-
Schltr., endemic to New Guinea but common and sistent sepals. There are at least three species: P.
widespread there. The species is apparently so stutzeri (F. Muell.) Gibbs in NE Australia; P. mollis
Cunoniaceae 107
Schltr. in New Guinea, and the highly variable P. growing gregariously in cool-temperate rain-
glabra Schltr. from eastern Malesia and Fiji. forests along rocky streams.
22. Codia J.R. Forst. & G. Forst. 25. Pancheria Brongn. & Gris Fig. 36
Codia J.R. Forst. & G. Forst., Char. Gen. Pl.: 59, t. 30 (1776); Pancheria Brongn. & Gris, Bull. Soc. Bot. France 9: 74-76
Brongn. & Gris, Bull. Soc. Bot. France 9: 76-77 (1862); (1862); Brongn. & Gris, Ann. Sci. Nat. Bot. V, 1: 367-368,
Brongn. & Gris,Ann. Sci. Nat. Bot. V, 1:377-378 (1864); Guil- 374-377 (1864); Guillaumin, Bull. Soc. Bot. France 87:
laumin, Bull. Soc. Bot. France 87: 254-256 (1940). 249-254 (1940).
Shrubs and trees. Leaves decussate in pairs Trees and shrubs. Leaves whorled or decussate (P.
(whorled in C. albifrons Vieill. ex Guillaumin), uni- confusa Guillaumin), unifoliolate or imparipin-
foliolate, margin usually entire. Inflorescence axil- nate. Inflorescence axillary; spherical heads (i.e.
lary, spherical heads (i.e. capitula), on slender capitula), capitula on slender peduncles from leaf
peduncles, or peduncles borne from short-shoots; axils, or peduncles borne from axillary short-
floral maturation synchronous; petals slender or 0;
ovary inferior. Fruits indehiscent with an outer
covering of lanate hairs; seeds with a thin, undif-
ferentiated seed coat. About 12 species endemic to
New Caledonia.
23. Callicoma Andrews

Callicoma Andrews, Bot. Repos. 9: t. 566 (1809); Kennedy,Aust.
J. Bot. 29: 721-731 (1981).
Trees. Leaves unifoliolate, deeply toothed; glossy

green upper surface, lower surface with minute,
white trichomes. Inflorescence axillary; spherical
heads (i.e. capitula), on slender peduncles from
leaf axils, or peduncles borne from axillary
short-shoots, often in a series; floral matura-
tion synchronous; perianth of one 4-6-merous
whorl, valvate or imbricate; stamens 11-15;
nectary 0; ovary of 2-3 carpels. Fruit capsular;
seeds unwinged with short papillae. One species,
C. serratifolia Andrews, common in Eastern
Australia.
VIII. TRIBE CUNONIEAE (R.Br.) Schrank &

Mart. (1829).
24. Vesselowskya Pamp.

Vesselowskya Pamp., Ann. Bot. (Rome) 2: 93 (1905); Rozefelds
et al., Aust. Syst. Bot. 14: 175- 192 (2001), rev.
Small trees. Leaves palmately compound, 3-5-

foliolate. Inflorescence axillary; racemose, 3-5
racemes borne on a peduncle; floral maturation
synchronous. Flowers unisexual, dioecious; peri-
anth 3-merous, calyx valvate, petals present in
male flowers, 0 in female flowers; nectary seg- Fig. 36. Cunoniaceae. Pancheria elegans. A Flowering branch.
mented; stigmas decurrent. Fruit capsular; seeds B Fruiting branch. C Male flower. D Same, opened to show pis-
tillode. E Female flower. F Dehiscing fruit. G Seed. Drawn by
winged. Two species from eastern Australia, Riocreux. (Brongniart and Gris 1868)
shoots; floral maturation synchronous. Flowers

unisexual; perianth 3- or 4-merous, calyx imbri-
cate; nectary annular or segmented; gynoecium of
two, mostly free carpels. Fruits capsular/pseudo-
follicular; seeds winged, small. About 30 species,
endemic to New Caledonia.
26. Cunonia L.
Cunonia L., Syst. Nat., ed. 10, 2: 1013, 1025, 1368 (1759);
Guillaumin, Bull. Soc. Bot. France 87:246-248 (1940); Coates
Palgrave, Trees of Southern Africa (1983); Hoogland eta!.,
Adansonia III, 19: 7-20 (1997).
Trees and shrubs. Leaves unifoliolate or imparip-

innate. Inflorescence axillary or terminal; race-
mose; racemes borne in pairs or threes from a
peduncle, occasionally arising directly from leaf
axils; floral maturation synchronous. Flowers
bisexual, or male and bisexual on a raceme; peri-
anth 5-merous, calyx imbricate; diplostemonous
(haplostemonous); floral nectary adnate to ovary.
Fruits capsular, opening circumbasally and septi-
cidally, stylodia often remaining fused so that
dehiscence is acropetal; seeds small, winged. A
genus of 25 species, with 24 in New Caledonia and Fig. 37. Cunoniaceae. Weinmannia heterophylla. A Flowering
one in South Africa. The South African species, C. branch; note stipules on lateral branch to the right. B Flower.
capensis L., is morphologically similar to two dis- C Petal. D Pistil with petals and stamens removed. E Young
tinctive species in New Caledonia, C. macrophylla pistil with disk. F Ovary, vertical section. G Ovary, transverse
Brong. & Gris and C. schinziana Daniker (Bradford section. Drawn by P.J.F. Turpin. (Kunth in Humboldt et al.1823,
tab. 522)
1998).
27. Weinmannia L. Fig. 37 in compound units of various forms; floral matu-

ration synchronous. Flowers bisexual or unisexual,
Weinmannia L., Syst. Nat. ed. 10: 1005 (1759), nom. cons.;
Bernardi, Candollea 17: 123-189 (1961), reg. rev.; Bernardi,
dioecious or polygamodioecious; solitary or fasci-
Candollea 18: 285- 334 (1963), reg. rev.; Bernardi, Bot. Jahrb. culate, pedicellate or sessile; calyx imbricate, petals
Syst. 83: 126-184 (1964), reg. rev.; Bernardi, Flore de Mada- persistent or caducous; nectary annular or seg-
gascar et des Comores (1965), reg. rev.; Hopkins et a!., Adan- mented. Fruits capsular; seeds with hairs, lacking
sonia Ill, 20: 5- 130 (1998), reg. rev.; J.C. Bradford, Ann. wings. This large genus has five sections: sect.
Missouri Bot. Gard. 85: 565-593 (1998); G. Harling, Flora of
Ecuador. Cunoniaceae (1999), reg. rev.; Bradford & Miller,
Weinmannia (Americas and Mascarene Islands;
Adansonia III, 23: 219-236 (2001), reg. rev. includes sect. Simplicifoliae ), sect. Fasciculatae
Arnoldia Blume (1826) = section Fasciculatae Bernardi ex (Malesia and western South Pacific), sect. Spicatae
Hoogland & H.C. Hopkins. Bernardi ex J.C. Bradford (Madagascar and
Leiospermum D.Don (1830) = section Leiospermum. Comores ), sect. Inspersae Bernardi ex J.C. Bradford
Ornithrophus Bojer ex Engl. (1870) =section Spicatae Bernardi
ex J.C.Bradford
(Madagascar), and sect. Leiospermum (South
Pterophylla D.Don (1830) = section Fasciculatae Bernardi ex Pacific). About 150-160 species with the following
Hoogland & H.C. Hopkins. distribution: c. 75 Americas, c. 40 W Indian Ocean
Windmannia P.Browne (1756) =section Weinmannia (Madagascar, Comores and Mascarenes), 40
Malesia and South Pacific.
Trees and shrubs, sometimes hemiepiphytic and
strangling. Leaves usually decussate in pairs but
whorled in W. commersonii Bernardi, unifoliolate Excluded Genera
or imparipinnate, rachis winged or not, margins
usually toothed (entire). Inflorescence axillary or Adenilema Bl. is now commonly included in Neillia D. Don
terminal, racemose, borne solitary in leaf axils or (Rosaceae).
Cunoniaceae 109
Aphanopetalum End!. will probably be recognized as its own Bradford, J.C., Miller, J.S. 2001. New taxa and nomenclatural
family in Saxifragales (Fishbein et a!. 200 1.; Bradford and notes on the flora of the Marojejy massif, Madagascar. V.
Barnes 2001). Cunoniaceae: Weinmannia. Adansonia III, 23: 219-236.
Gummellia Ruiz & Pav. is now part of Simaroubaceae Brongniart, A., Gris, A. 1868. Description de quelques plantes
(Hoogland in Gunn et a!., U.S.D.A. Tech. Bull. no. 1796 remarquables de Ia Nouvelle-Caledonie. Nouv. Arch. Mus.
[1992]). Hist. Nat. Paris 4: 1-48, pl. 1-15.
Kaernbachia Schltr. is synonymous with Turpinia Vent. Bush, M.B., Colinvaux, P.A., Wiemann, M.C., Piperno, D.R.,
(Staphyleaceae; Linden in Van Stennis, Fl. Males. I, 6: 49 Liu, K. 1990. Late Pleistocene temperature depression and
[1960]). vegetation change in Ecuadorian Amazonia. Quat. Res. 34:
Paracryphia Baker f. was originally included in Eucryphi- 330-345.
aceae, but most subsequent workers have placed it else- Cadet, T. 1984. Plantes rares ou remarquables des
where. Dickison and Baas ( 1977) suggested affinities with Mascareignes. Paris: Agence de Cooperation Culturelle et
Sphenostemonaceae, Theaceae and Actinidiaceae. It is now Technique.
considered a monotypic family. Molecular data place it in Carpenter, R.J., Buchanan, A.M. 1993. Oligocene leaves, fruits
the Euasterid II clade and in a subclade with Quintiniaceae and flowers of the Cunoniaceae from Cethana, Tasmania.
and Sphenostemonaceae (Savolainen, Fay et a!. 2000; Aust. Syst. Bot. 6: 91-109.
Lundberg 2001). Carpenter, R.J., Pole, M. 1995. Eocene plant fossils from the
Lefroy and Cowan paleodrainages, Western Australia. Aust.
Syst. Bot. 6: 91-109.
Cooper, M.G. 1986. A pilot survey of six rare plants in New
Selected Bibliography South Wales. Sydney, NSW: Parks and Wildlife Service.
Cooper, W., Cooper, W.T. 1994. Fruits of the Rain Forest.
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112 K. Kubitzki
Curtisiaceae
K. KUBITZKI
Curtisiaceae (Harms) Takht., Sist. Magnoliof.: 214 (1987). found in Cornus and Mastixia and possibly also
other genera of Cornaceae.
Evergreen tree. Leaves opposite, coarsely dentate,
rusty velvety beneath; hairs unicellular. Inflores- KARYOLOGY. Curtisia and Mastixia arborea share
cences terminal, tricussately thyrso-paniculate, the gametophytic chromosome number n = 13
many-flowered, prophylls present; flowers minute, (Goldblatt 1978).
4-merous, hermaphroditic, or rarely plants dioe-
cious; calyx tube turbinate with open, triangular FRUITS. Curtisia fruits are red at maturity and are
lobes; petals (sub)valvate, triangular-ovate; eagerly sought after by birds, wild pigs, monkeys
stamens 4, alternipetalous; filaments subulate; and baboons. Fruit opening seems to be with
anthers cordate at base, introrse, longitudinally valves, just as in Cornaceae, as Eyde (1988: 296 [in
dehiscing; disk epigynous, broad, 4-angled, legend to Curtisia fruit]) writes: "If boiled and
densely barbate; ovary inferior, 4-locular; ovules 1 dried repeatedly, this stone would open like a
per locule, with ventral raphe and micropyle Corn us stone".
directed upwards; style short, subconical, with 4-
lobed stigma. Fruits drupaceous, small, subglo- PHYTOCHEMISTRY. The red wood of Curtisia
bose, 4-locular, 4-seeded, areolate at apex, crowned quickly darkens in fresh air and the bark contains
by the calyx, eventually dehiscing; seeds elongate- 3-21.5% tannin (Watt and Breyer-Brandwijk
oblong, subcylindric, with elongate embryo in 1962). Proanthocyanins, ellagitannins and two
copious endosperm. n = 13. unidentified iridoids have been reported from
A single sp., C. dentata (N.L. Burm.) C.A. Smith, leaf tissue by Bate-Smith et al. (1975).
southern and eastern S Africa, in forest and
bushland. AFFINITIES. Most plant systematists have
included Curtisia in Cornaceae, which made sense
FLORAL STRUCTURE. Most Cornaceae, in which in view of its pollen morphology and the valvate
Curtisia has always been included, have a peculiar fruit stone, apart from the general agreement with
gynoecial bundle supply that was studied by Eyde Cornaceae in floral and vegetative structures. Eyde
(1968). In Davidia, Cornus, Alangium and Nyssa, (1988) excluded Curtisia because its gynoecium
ovular bundles pass through the septa or opposite lacks the apomorphic transseptal bundles of the
to them and enter the placentae arching above cornaceous genera. In contrast, I would not object
each septum. The syncarpous, 4-carpellate inferior against including the plesiomorphic condition in
ovary of Curtisia differs from them in having an the family, and would like to follow taxonomists
axile bundle supply to the ovules, just as would be such as Hooker (1867) and Harms (1897) in the
expected for a syncarpous ovary, and which is placement of Curtisia in Cornaceae, had not the
probably plesiomorphic in relation to the corna- molecular analyses of Xiang et al. (2002) resolved
ceous condition. Grubbia and Curtisia in a well-supported associa-
tion and sharing a deletion, the latter making
PoLLEN MoRPHOLOGY. Pollen is sphaeroidal and an artifactial long branch attraction of the two
relatively small (12-20 x 12-17 J.lm, i.e. half the size less probable (see also under Grubbiaceae, this
of most Cornus), tricolporate, provided with colpi volume).
costae, a complete tectum and a lamellated ektex-
ine in the areas of the endoapertures; the tectum DISTRIBUTION AND HABITATS. Curtisia extends
has impressions that appear as perforations but from the Cape Peninsula eastward to Natal and
do not connect with the intercolumellar space from there to the north through the Transvaal into
(Ferguson and Hideux 1978). Similar pollen is the mountains of eastern Zimbabwe. In closed
Curtisiaceae 113
One genus:
Curtisia Aiton Fig. 38

Curtisia Aiton, Hort. Kew. 1: 162 (1789), nom. cons.
Description as for family.
Bate-Smith, E.C., Ferguson, I.K., Hutson, K., Jensen, S.R.,

Nielsen, B.J., Swain, T. 1975. Phytochemical interrelation-
ships in the Cornaceae. Biochem. Syst. Ecol. 3: 79-89.
Cannon, J.F.M. 1978. Cornaceae. In: Launert, E. (ed.) Flora
Zambesiaca vol. 4. London: Flora Zambesiaca Managing
Committee, pp. 635-638.
Eyde, R.H. 1968. The peculiar gynoecial vasculature of Cor-
naceae and its systematic significance. Phytomorphology
17: 172-182.
Eyde, R.H.1988. Comprehending Cornus: puzzles and progress
in the systematics of the dogwoods. Bot. Rev. 54: 233-351.
Ferguson, I.K. 1977. Cornaceae. World pollen and spore flora
6. Stockholm: Almqvist and Wiksell, 34 pp.
Ferguson, I.K., Hideux, M.J. 1978. Some aspects of the pollen
morphology and its taxonomic significance in Cornaceae
s.l. In: 4th Int. Palynol. Conf., Lucknow, 1976-77, 1: 240- 249.
Goldblatt, P. 1978. A contribution to cytology in Cornales. Ann.
Missouri Bot. Gard. 65: 650-655.
Harms, H.1897. Cornaceae. In: Engler, A., Prantl, K. Die natiir-
lichen Pflanzenfamilien III, 8. Leipzig: W. Engelmann, pp.
250-270.
Fig. 38. Curtisiaceae. Curtisia dentata. A Flowering/fruiting Hooker, J.D., 1867. Cornaceae. In: Bentham, G., Hooker, J.D.
twig. B Hairs. C Group of flowers. D Fruit. E Transverse section Genera Plantarum, vol. 1 ( 3). London: Reeve & Co., pp.
of fruit. (Cannon 1978) 947-952.
Marais, W. 1985. Curtisia. Kew Mag. 2: 368.
Watt, J.M., Breyer-Brandwijk, M.G. 1962. The medicinal and
poisonous plants of southern and eastern Africa. Edinburg:
vegetation it is a tree up to 20m high and in open E. & S. Livingstone.
vegetation it has a more shrubby habit; it grows up Xiang, Q.-Y. et a!. 2002. See general references.
to an altitude of 1800 m.
114 K. Kubitzki
Cyrillaceae
K. KuBITZKI
Cyrillaceae Endl., Ench. Bot.: 578 (1841), nom. cons. unknown homology but, according to Thomas
(1960), may be vestigial stipules.
Deciduous or evergreen and glabrous small trees The root system is shallow. Along the horizon-
or shrubs. Leaves alternate, simple, entire, short- tally running roots numerous adventitious roots
petioled or sessile, exstipulate. Inflorescences ter- arise, by means of which Cyrilla and Cliftonia are
minal or axillary racemes; pedicels with paired able to build up clones of vast extension. Thomas
prophylls. Flowers perfect, regular, hypogynous; (1961) excavated a group of Cyrilla bushes cover-
sepals 5( -7), imbricate, sometimes quincuncially ing an area 15 feet wide and 30 feet long, com-
arranged, persistent, the outer larger than the prising over 50 bushes. These plants were
inner, often accrescent in fruit; petals as many as interconnected with each other by their root
sepals, distinct, imbricate; androecium diploste- system and remained so for several years.
monous or (Cyrilla) haplostemonous; anthers Sieve element plastids have protein filaments
dorsifixed, tetrasporangiate, not inverted at and crystals but no starch (Pcf type; Behnke 1982,
anthesis, dehiscing by longitudinal slits; disk 1991), and these are probably autapomorphic.
intrastaminal, surrounding the base of the
ovary; gynoecium syncarpous; ovary (2-)5- EMBRYOLOGY. In the anthers the epidermis,
locular with axile placentas; style very short or endothecium and middle layer contain tannin; the
nearly suppressed with lobed stigma or with 2-4 tapetum is of the secretory type. Ovules are
very short stylar branches with apical stigmas; unitegmic and tenuinucellate; both normal Poly-
ovules 1-3 per locule (pendulous from near the gonum type and aposporous embryo sacs develop
top of the locule or from a pendulous, stalk-like in the same ovule (Cliftonia), the latter remaining
placenta), anatropous, unitegmic, tenuinucellar. functionless. Haustoria develop at both ends of the
Fruit indehiscent, 1-seeded, dry and drupaceous embryo sac. Degeneration of the egg and zygote is
or samaroid; seed coat completely reduced; frequent. During embryo development, the integu-
embryo slender, straight, with small, slightly mentary cells become vacuolised and degenerate;
expanded cotyledons, embedded in a copious, they have vanished in the mature seed. The rela-
fleshy endosperm. x = 10. tively large embryo is straight and nearly as long
A family of two probably monotypic genera, as the seed. The pericarp is tanniniferous and con-
extending from the coastal plains of SE North tains druses (Copeland 1953; Vijayaraghavan 1969;
America to Central and N South America, includ- Vijayaraghavan and Dhar 1978).
ing the West Indies.
POLLEN MORPHOLOGY. Pollen is tri-(to hexa-
VEGETATIVE STRUCTURES. Young stems are pro- )colporate, spheroidal to suboblate with wide
vided with ridges decurrent from the leaf bases; colpi, the tectum psilate and microperforate, the
these vanish at the end of the second year. Perid- infratectum granular, the equatorial diameter
erm develops early in the inner layers of the 18-291Jm (Zhang and Anderberg 2002).
cortex. The secondary xylem is composed of vessel
elements with scalariform perforation plates with REPRODUCTIVE BIOLOGY. The vegetative repro-
numerous bars, tracheids and fiber-tracheids, duction by root sprouting of Cyrilla and Cliftonia
uniseriate and multiseriate rays, and diffuse axial is often observed in cut-over or otherwise dis-
parenchyma; nodes are unilacunar throughout turbed areas, but disturbance or injury is not a
(Thomas 1961). necessary prerequisite for the formation of clones
Leaf venation is pinnate. Stomata are anomo- by root sprouting. Within these clones, the abun-
cytic. Solitary crystals are found in the leaves and dant fruits annually produced rarely contain
druses are abundant. Ligulate, glandular struc- seeds, and parthenocarpoy has been observed to
tures on each side of the buds and of bracts are of be more frequent towards the centre of large
Cyrillaceae 115
clones. This may indicate a pronounced self-steril- the Maestrichtian onwards have been referred to
ity; Thomas (1961) concluded that self-fertiliza- Clethraceae and/or Cyrillaceae, but the distinction
tion may provide sufficient stimulus to initiate between the two remains problematic (Muller
fruit development, whereas seed development 1981). The same may be true of several fossil fruit
seems to require pollination from a different remains from the European Upper Cretaceous and
individual. Tertiary that have been related to Cyrillaceae (see
In North America Uphof (1942) observed that Knobloch and Mai 1986), the taxonomic position
Cyrilla and Cliftonia are protogynous. The flowers of which is contentious (Friis 1985).
are showy and produce scent and nectar; they are
visited by several classes of insects. KEY TO THE GENERA
1. Stamens 10; petals not glandular; fruit distinctly 3-5-
FRUIT AND SEED. The complete lack of a seed coat winged 1. Cliftonia
has been noted by several students of the family - Stamens 5; petals glandular on adaxial surface; fruit
and there is not the slightest rudiment of a testa (sub )globose, longitudinally bisulcate or trisulcate
(Gilg 1892; Thomas 1960). 2. Cyrilla
PHYTOCHEMISTRY. Bate-Smith (1962) found 1. Cliftonia Banks ex Gaertn.f.

myricetin, ellagic acid and procyanidin; i.e. both
condensed and hydrolysable tannnins are likely to Cliftonia Banks ex Gaertn.f., Fructus 3: 246 (1805); Thomas,
Contrib. Gray Herb. 186: 74-76 (1960), rev.
be present.
FAMILY CIRCUMSCRIPTION. The South and Evergreen. Pedicels with two caducous prophylls;
Central American genus Purdiaea, usually stamens 5 + 5; filaments laterally expanded and
included in Cyrillaceae, has on the basis of mor- petaloid in lower half; ovary 3-5-angled and
phological data and DNA sequence analyses been -locular, each locule containing a single, pendu-
transferred to Clethraceae, with which it agrees lous ovule; stigma massive, subsessile, 2-5-lobed.
in slight sympetaly and versatile, ventrifixed, Fruit 2-5-winged, up to 5-seeded but usually
inverted and poricidal anthers (Anderberg and devoid of seeds. A single sp., C. monophylla (Lam.)
Zhang 2002). Britton ex Sarg., on the coastal plains of the SE
U.S.A.
AFFINITIES. Formerly associated with Aquifoli-
aceae or Celastraceae, DNA sequence analyses 2. Cyrilla Garden ex L.
place Cyrillaceae in the broadly circumscribed
Ericales sensu APG (1998), in which they are the Cyrilla Garden ex L., Mant. Pl. 1: 5 (1767); Thomas, Contrib.
Gray Herb. 186: 76-104 (1960), rev.
sister group of Ericaceae, whereas Ericaceae +
Cyrillaceae are sister to Clethraceae (Anderberg
Evergreen or deciduous. Pedicels with two persist-
et al. 2002; Anderberg and Zhang 2002).
ent prophylls; petals thickened and glandular on
upper surface medially and below the middle;
DISTRIBUTION AND HABITATS. Cyrillaceae prefer
stamens 5, antepetalous; filaments subulate; ovary
the wet and acidic habitats on the coastal plains of
2-4-locular; ovules pendulous, 1-3 per locule;
the SE USA; Cyrilla further extends into Mexico,
style short, persistent, with 2-4 short stylar
Central America, the Caribbean, the Roraima
branches. Fruit longitudinally 2-3-sulcate with no
sandstone formation in Venezuela and Guiana,
more than 1 seed in each locule, often devoid of
Colombia, and N Brazil, ranging there from the
seeds. 2n = 40. A single sp., C. racemiflora L., N
lowland to the mountain summits, and growing
South America, Central America, Caribbean, and
in wet heath formations.
SE U.S.A. Up to eleven spp. have been dis-
tinguished, but Thomas (l.c.), who studied their
PALAEOBOTANICAL RECORD. In the Brandon
variation pattern, was unable to detect real gaps or
Lignite of Vermont (USA) of Upper Oligocenic
discontinuities separating them.
age, the most abundant wood and second most
abundant pollen was found to be that of Cyrilla
(Barghoorn and Spackman 1949). Cyrilla-like
wood is also known from Tertiary brown coal beds
in western Europe (van der Burgh 1973), accom-
Anderberg, A.A. et al. 2002. See general references.
panied by ample pollen. Other pollen records from Anderberg, A.A., Zhang Xiaoping 2002. Phylogenetic relation-
116 K. Kubitzki
ships of Cyrillaceae and Clethraceae (Ericales) with special Muller, J. 1981. See general references.
emphasis on the genus Purdiaea Planch. Org. Div. Evol. 2: Thomas, J.L. 1960. A monographic study of the Cyrillaceae.
127-137. Contrib. Gray Herb. 186: 1-114.
APG (Angiosperm Phylogeny Group) 1998. See general Thomas, J.L. 1961. The genera of the Cyrillaceae and
references. Clethraceae of the southeastern United States. J. Arnold
Barghoorn, E.S., Spackman, W. 1949. A preliminary study of Arbor. 42: 96-106.
the flora of the Brandon Lignite. Am. J. Sci. 247: 33-39. Uphof, J.C.Th. 1942. Cyrillaceae. In: Die natiirlichen Pfianzen-
Bate-Smith, E.C. 1962. See general references. familien, 2nd edn., 20b. Leipzig: W. Engelmann, pp. 1-12.
Behnke, H.-D. 1982. Sieve element plastids of Cyrillaceae, Ery- van der Burgh, J. 1973. Holzer der niederrheinischen
throxylaceae and Rhizophoraceae: description and signifi- Braunkohlenformation. 2. Holzer der Braunkohlengruben
cance of subtype PV plastids. Plant Syst. Evol. 141: 31-39. "Maria Theresia" zu Herzogenrath, "Zukunft West" zu
Behnke, H.-D. 1991. See general references. Eschweiler und "Viktor" (Ziilpich Mitte) zu Ziilpich. Nebst
Copeland, H.F. 1953. Observations on the Cyrillaceae, particu- einer systematisch-anatomischen Bearbeitung der Gattung
larly on the reproductive structures of the North American Pinus. Rev. Palaeobot. Palynol. 15: 73-275.
species. Phytomorphology 3:405-411. Vijayaraghavan, M.R. 1969. Studies in the family Cyrillaceae.
Friis, E.M. 1985. Angiosperm fruits and seeds from the Middle 1. Development of male and female gametophyte in Clifto-
Miocene of Jutland (Denmark). Bioi. Skr. Vid. Selsk. nia monophylla (Lam.) Britton ex Sarg. Bull. Torrey Bot.
K0benhavn 24 (3): 1-66. Club 96: 484-489.
Gilg, E. 1892. Cyrillaceae. In: Engler & Prantl, Nat. Pfianzenfam. Vijayaraghavan, M.R., Dhar, U. 1978. Embryology of Cyrilla
III, 5. Leipzig: W. Engelmann, pp. 179-182. and Cliftonia (Cyrillaceae). Bot. Not. 131: 127-138.
Knobloch, E., Mai, D.H. 1986. Monographie der Friichte und Zhang, X.-P., Anderberg, A. 2002. Pollen morphology in the
Samen in der Kreide von Mitteleuropa. Rozpr. Ustt. list. ericoid clade of the order Ericales, with special reference on
geol., Praha 47: 1-220. Cyrillaceae. Grana 41: 201-215.
Diapensiaceae 117
Diapensiaceae
P.J. ScoTT
Diapensiaceae (Link) Lind!., Nat. Syst. Bot., ed. 2: 233 (1836), ate regions. The domes have the surface covered
nom. cons. with living leaves and the interior is packed with
dead leaves which decompose to duff (Day and
Low-growing, generally mat-forming, evergree.n Scott 1981).
herbs or shrublets with ecto- and endotroph1c
mycorrhizae. Leaves alternate, simple, linear- VEGETATIVE ANATOMY. The leaf venation pattern
lanceolate, exstipulate, usually crowded on stems, of Diapensia and Pyxidanthera is unique among
or leaves round-cordate, petiolate, and fewer. dicotyledons. It is inverted-eucamptodromous,
Flowers solitary or in compact racemes, perfect, having the secondary veins curve basally and
actinomorphic, hypogynous, slightly protogynous become attenuated in that direction without
or homogamous; prophylls usually present; peri- rejoining the primary vein or other secondaries.
anth biseriate, pentamerous, imbricate; sepals free Galax and Shortia have actinodromous or camp-
or connate into a 5-lobed tube, persistent; petals 5, todromous patterns, and Berneuxia has a brochi-
white, purple, or rarely yellow, caducous, connate dodromous pattern (Murphy and Hardin 1976).
into a sympetalous corolla or (Galax) almost free; Leaf epidermal cells have undulating lateral walls
stamens 5, attached to corolla tube and alternate and, particularly on the upper surface, a very thick
with its lobes, distinct or connate basally in a ring, outer wall traversed by numerous pits. On the
in all but Diapensia and Pyxidanthera alternating lower leaf surface, Berneuxia thibetica has papil-
with 5 staminodia; anthers tetrasporangiate or lose epidermal cells and Diapensia lapponica has
(Galax) disporangiate, dehiscing longitudinally or a swollen, thick cuticle over the middle of each
(Galax, Pyxidanthera) transversely; nectary disk epidermal cell. Mesophyll is either bifacial or iso-
absent; gynoecium 3-carpellate; ovary syncar- diametric in Shortia and Galax. Chlorophyll is
pous, 3-locular; ovules few to many on more or present in the epidermis of Diapensia and Galax.
less intrusive placentas; ovules anatropous to Stomata are found only on the lower surface in
campylotropous, unitegmic, tenuinucellate; style Berneuxia, some Shortia, and Diapensia lappon-
simple, hollow; stigma 3-lobed (Diapensia with 3 ica, but on both surfaces in other species of
style branches). Fruit a loculicidal capsule; seeds Diapensia, Galax, Pyxidanthera, and Shortia.
thin, with exotestal seed coat, a central straight or Subsidiary cells are absent. The number of layers
slightly curved embryo and abundant fleshy of palisade cells varies from 3 in Berneuxia, 2-3 in
endosperm. Diapensia lapponica, and 1 in Shortia uniflora. The
A family of 5 genera and 13 species distributed spongy mesophyll in Diapensia lapponica has
in North America and eastern Asia, and one some thick-walled cells (Solereder 1908; Metcalfe
cultivar. and Chalk 1950).
Calcium oxalate is excreted along the leaf
VEGETATIVE MORPHOLOGY. Leaf shape varies margins as clustered crystals or as solitary crystals
from reniform to orbicular and petiolate in Galax in Galax. Internal glands are absent (Solereder
and Shortia to linear and petiolate in Berneuxia 1908; Metcalfe and Chalk 1950). A number of
to linear-lanceolate or linear-oblanceolate and members of the family undergo annual changes in
(nearly) sessile in Diapensia and Pyxidanthera. the coloration of their leaves. Diapensia lapponica
The size decreases in the series given above, and leaves are olive green in the summer and dark bur-
the reduction in size and shape is correlated with gundy in winter due to anthocyanin accumulation
adaptation to xeric or arctic-alpine habitats. Tri- (Day and Scott 1981). In the wood, primary
chomes are absent or infrequent except in Pyxi- medullary rays are observed only immediately
danthera. Diapensia forms domes or mats which above the nodes; secondary rays are absent, and
are adapted to windy habitats, and act as solar heat the vessels have small lumina (Solereder 1908).
traps, which restrict their distribution in temper- Pith is often heterogeneous, consisting of thin-
118 P.J. Scott
and thick-walled cells. Growth rings are poorly EMBRYOLOGY. The mature anther walls have a
defined in Diapensia lapponica (Metcalfe and well-developed fibrous endothecium. The tapetum
Chalk 1950). is binucleate and glandular. The ovules are gener-
The principal reserve foods in alpine and arctic ally anatropous, but vary from hemitropous to
plants are starches, sugars, and lipids. In evergreen campylotropous. In Shortia uniflora the ovules are
subshrubs like Diapensia lapponica, lipids and unitegmic and the nucellus is single-layered and
some carbohydrates are stored in the old stems disappears during megaspore development. The
and old leaves (Petersen 1912). inner layer of the integument constitutes an
The roots have ecto- and endotrophic mycor- endotheliaceous tissue. The nucellus consists of an
rhizae. In Diapensia lapponica, the tap and adven- epidermis and a few cells at the base. The epider-
titious roots are generally shallow (<30 em) and mis disintegrates so that the gametophyte is in
convoluted. Cork is poorly developed and phloem contact with the inner layer of the single integu-
shows collenchymatous thickening in Diapensia ment (Palser 1963). The pollen tube passes down
lapponica (Metcalfe and Chalk 1950). the style through the stylar canal and enters
the ovules porogamously. Seed production is a
FLOWER STRUCTURE. Flowers are actinomorphic, problem for many species. Examination of ovules
hypogynous, and pentamerous except for the shows comparatively few with endosperm or
ovary which is tricarpellate. The sepals are per- embryo development and considerable signs of
sistent and free or slightly connate. The petals disintegration, suggesting failure of pollination or
are free in Galax but connate in the other fertilization (Palser 1963).
genera. The family has a modified diplostemo- The endosperm is cellular and there are no
nous androecium. The outer whorl consists of 5 endosperm haustoria. The embryo is straight,
functional stamens and the inner whorl has 5 cylindrical, approximately two-thirds the length of
staminodia. the seed. Cotyledons are approximately one-third
The stamens are adnate to the corolla and alter- the total length. The seed coat consists of 2-3 cell
nate with the corolla lobes. The staminodia are layers (Palser 1963; Yamazaki 1966).
sometimes reduced, or absent in Diapensia and
Pyxidanthera. When present, the staminodia occur POLLEN MORPHOLOGY. Pollen grains are
opposite the corolla lobes and are adnate to monads, tricolp(oroid)ate, semitectate or intec-
varying degrees. The anthers vary in length, tate, per-reticulate, with at least some brochi more
orientation, number of pollen sacs, and plane of than ljlm wide. The sexine is as thick as the nexine
dehiscence. The anthers of Pyxidanthera and or thicker; colpi are tenuimarginate with an equa-
Galax have appendages. The relative size, number torial constriction (Fig. 39). The pollen grains are
of thecae and pollen chambers, orientation of not of a very distinctive type; they resemble those
dehiscence, and presence of appendages show con- of many families (Erdtman 1952; Xi and Tang
siderable variation. There are, however, a number 1990).
of features in common. Anthers are definitely or
obliquely introrse. The vascular strand of the fila- KARYOLOGY. A chromosome number of 2n = 12
ment extends to the apex of the connective where has been determined for the family except in
it tapers off in Diapensia and Shortia. It ends in a Galax urceolata where diploids and tetraploids
knob of vascular tissue below the horizontal exist that are indistinguishable morphologically.
pollen sacs in Galax and Pyxidanthera. The tetraploid cytotypes are considered an excel-
The ovary has three locules with several to many lent example of autopolyploidy in natural popula-
ovules. The placentae are axile, usually enlarged. tions (Nesom 1983).
There is a split in the central column in the upper
part of the ovary which, in some genera, extends POLLINATION AND REPRODUCTIVE SYSTEMS.
completely through the placentae so that the pla- Flowers of Diapensia lapponica are visited by a
centation becomes parietal. The ovary tapers into number of members of Hymenoptera and Diptera
the style which varies in relative length and has a (Fig. 40; Petersen 1912; Day and Scott 1981). Prob-
stylar canal. The stigma is 3-lobed. Ovules vary lems with reproduction have been noted in the lit-
from few to many. In some species, nectar is erature and few seedlings have been observed in
secreted by a few layers of cells on the outer populations. Although some species are abundant
surface of the ovary base. locally, their distributions are often limited. There
For the vascular supply of the flowers, the reader have been a number of reasons suggested: poor
is referred to the study by Palser (1963). seed dispersal, seed being retained in the capsule
Diapensiaceae 119
ponica (Bliss 1962). Seeds germinate in 21-28 days

(in 11 days in Diapensia lapponica) after 3 months
of chilling. There may be as many as 50 flowers per
year on very large plants of Diapensia lapponica,
with each flower giving rise to a mean production
of 121 seeds per capsule, with seed set averages of
70-75% and an average of240 ovules/capsule (Day
and Scott 1981).
DISPERSAL. The upright capsules of many species

would suggest dislodgement and limited dispersal
by wind. Capsules of Diapensia lapponica have
been observed projecting through the snow,
dehisced and with mature seed in mid-winter. This
suggests dispersal across the surface of the snow
(Day and Scott 1981).
PHYTOCHEMISTRY. The family is noted for

accumulating significant amounts of aluminium.
Shortia is surpassed by only a couple of other
plants in the amounts accumulated (Gibbs 1974).
AFFINITIES AND PHYLOGENY. Diapensiaceae are

Fig. 39. Diapensiaceae. Pollen of Galax urceolata, SEM photo, characterized by the presence of staminodia,
bar= lO~m (Palynological Laboratory, Stockholm)
single pollen grains, absence of a disk, 3-carpellate
gynoecium, !-layered testa, absence of integu-
mentary tapetum, absence of terminal and cha-
laza} haustoria, and aluminium accumulation
(Anderberg 1993). The family has often been
associated with the narrowly defined Ericales
(Takhtajan 1980, among others) and more recently
has been included in the broadly defined Ericales
sensu APG (1998). The five-gene sequence analy-
sis by Anderberg et al. (2002) places Diapensiaceae
in close association with Styracaceae, a position
also supported by embryological traits.
A phylogenetic analysis based on morphology
and four genes (Ronblom and Anderberg 2002)
showed that Galax is sister to all other genera, and
Pyxidanthera sister to the remaining genera at the
Fig. 40. Diapensiaceae.Diapensia lapponica flowers, visited by next higher node.
Diptera. Photo P.J. Scott
DISTRIBUTION AND HABITATS. The present dis-
tribution suggests that the family was formerly
where it germinates, inviability of seeds, and more widespread. Presently, it is represented by a
fungal attack of seeds (Ross 1936; Primack and few relict and well-defined species. Most species
Wyatt 1975). In localities at the southern limit of are confined to the two centres of diversity for
the range of Diapensia lapponica, some plants in the family, the Appalachian Mountain system of
each population bloom in late summer and dis- eastern North America and eastern Asia, except for
perse their seeds through the winter. This appears Diapensia lapponica which is circumpolar. These
to be an adaptation to avoid fungal infection (Day plants are probably relicts of the Arctotertiary
and Scott 1981). forest which had a circumboreal distribution
before the Pleistocene. The majority of species do
SEED. The seeds have a straight embryo with not compete well with other plants and are con-
ample endosperm, high in fat in Diapensia lap- fined to arctic/alpine habitats except Pyxidanthera
120 P.J. Scott
which grows in sandy pine barrens and Galax

which grows in open hardwood forests. Diapensia
lapponica is described as snow-fearing and is
usually found growing in areas blown clear of
snow during the winter (Tiffney 1972).
ECONOMIC IMPORTANCE. Members of the family

have little importance in commerce. The leaves of
Galax urceolata are preserved and used by florists.
Specialist growers cultivate some of the Diapensia,
Pyxidanthera, and Shortia.
KEY TO THE GENERA

1. Staminodia absent; flowers solitary; leaves narrow and
imbricate, sessile or very shortly petiolate 2
- Staminodia 5, opposite the petals (absent in some Asian
Shortia); flowers solitary or not; leaves orbicular to obovate
or spatulate, long-petiolate 3
2. Anther loculi dehiscing transversely, awned at base; flowers
sessile 4. Pyxidanthera
- Anther loculi dehiscing lengthwise, unawned; flowers
pedicellate 3. Diapensia
3. Corolla lobes toothed or fringed 2. Shortia
- Corolla lobes entire 4
4. Anthers 2-locular; style elongated 1. Berneuxia
- Anthers )-locular; style very short 5. Galax
i,l .
.. '
'
1. Berneuxia Decne. N
Berneuxia Decne., Bull. Soc. Bot. France 20: 159 (1873).
Perennial herb, subacaulescent; caudex thickened

and creeping. Leaves long-petiolate. Inflorescence
with slender peduncle about 18cm long (about
same length as leaves), 10-15-flowered; anthers
off
2-locular; stamens and staminodia fused and
forming a ring at base of petals; staminodia
reduced to scales; style elongated, inserted in
depression on top of ovary. A single species, B. Fig. 41. Diapensiaceae. A-H Shortia galacifolia. A Habit. B
thibetica Decne., in Himalayan Mts. (Li 1943). Lateral view of flower, corolla removed. C Flower. D Expanded
corolla. E Stamen. F Staminode. G Ovary, vertical section.
H Ovary, transverse section. 1-0 Galax urceolata. I Habit. J
2. Shortia Torr. & Gray Fig. 41A-H Flower. K Corolla and stamina) tube, expanded. L Stamen and
anther. M Ovary with 3 sepals and prophyll. N Fruit. 0 Seed.
Shortia Torr. & Gray, Am. J. Sci. Arts 42: 48 (1842), nom. cons. (From Wood and Channelll959)
Schizocodon Sieb. & Zucc. (1843).
Low perennial herbs forming dense clumps or 3. Diapensia L. Fig. 40

carpets, spreading by horizontal rhizomes. Leaves
long-petiolate. Flowers solitary or in racemes, Diapensia L., Sp. Pl. 1: 141 (1753).
nodding; peduncle bracted; petals united, the
lobes undulate-crenate notched; stamens 2- Cushion-like subshrubs. Leaves short-petiolate.
loculed, bent sharply inward; staminodia borne Flowers terminal on branchlets, solitary; pedun-
near base of petals; style elongated, filiform. cles elongated and scape-like after anthesis;
Five species in East Asia and one in E North stamens adnate to base of corolla; anthers 2-
America. loculed, with oblique, divergent loculi; staminodia
absent or vestigial. Four species, D. lapponica L.
circumboreal, three in the Himalayan Mts.
Diapensiaceae 121
4. Pyxidanthera Michx. Darlington, C.D., Wylie, A.P. 1955. Chromosome atlas of flow-
ering plants. London: Allen and Unwin.
Pyxidanthera Michx., Fl. Bor.-Am. 1: 152 (1803}. Day, R.T., Scott, P.J. 1981. Autecological aspects of Diapensia
lapponica L. in Newfoundland. Rhodora 83: 101-109.
Prostrate creeping evergreen subshrub. Leaves Diels, L. 1914. Diapensiaceen-Studien. Bot. Jahrb. Syst. Suppl.
50: 304-330.
awl-pointed, sessile. Flowers solitary and sessile on Erdtman, G. 1952. See general references.
short, densely leafy branches; petals united by Evans, W.E. 1927. A revision of the genus Diapensia with
broad stamen-filaments to form a tube; anther special reference to the Sino-Himalayan species. Notes Roy.
locules 2, awned at base and dehiscing trans- Bot. Gard. Edinburgh 15: 209-236.
Gibbs, R.D. 1974. Chemotaxonomy of flowering plants. Mon-
versely; pollen only discharged when awned tip treal: MeGill-Queen's University Press.
pushed downward; staminodia absent; style as Li, H.-L. 1943. On the Sino-Himalayan species of Shortia and
long as the corolla tube. A single species, P. barbu- Berneuxia. Rhodora 45: 333-337.
lata Michx., in eastern North America. Metcalfe, C.R., Chalk, L. 1950. See general references.
Murphy, H.T., Hardin, J.W. 1976. A new and unique venation
pattern in the Diapensiaceae. Torreya 103: 177-179.
5. Galax Sims Fig. 411-0 Nesom, G.L. 1983. Galax (Diapensiaceae): geographic varia-
tion in chromosome numbers. Syst. Bot. 8: 1-14.
Galax Sims, Curtis Bot. Mag. 5: t. 754 (1804}. Palser, B.F. 1963. Studies of floral morphology in the Ericales.
VI. The Diapensiaceae. Bot. Gaz. 124: 200-219.
Herbaceous perennial forming thick matted tufts; Petersen, H.E. 1912. The structure and biology of arctic flow-
ering plants. II. Diapensiaceae. Diapensia lapponica L.
rhizomes scaly-bracted, branched. Leaves petio- Meddel. Gronland 36: 141-154.
late. Inflorescence a raceme on a naked peduncle. Primack, R.B., Wyatt, R. 1975. Variation and taxonomy of Pyx-
Petals distinct to base; stamens and staminodia idanthera (Diapensiaceae). Brittonia 27: 115-118.
approximately equal, their filaments united and Ronblom, K., Anderberg, A.A. 2002. Phylogeny of Diapensi-
forming a tube adnate to petals at base and falling aceae based on molecular data and morphology. Syst. Bot.
27: 383-395.
with petals; anthers 1-loculed, dehiscing horizon- Ross, M.N. 1936. Seed reproduction of Shortia galacifolia. Bot.
tally across the top, lower part of anthers tapering Gdn. J. 37: 208-211.
to an obtuse point; style short. A single species, G. Samuelson, G. 1913. Studien iiber Entwicklungsgeschichte der
urceolata (Poir.) Brummitt, in eastern N America, Bliiten einiger Bicornestypen. Svensk Bot. Tidskr. 7: 97-
188.
with diploid and tetraploid races.
Scott, P.J., Day, R.T. 1983. Diapensiaceae: a review of the tax-
onomy. Taxon 32:417-423.
Solereder, H. 1908. Systematic anatomy of the dicotyledons.
Selected Bibliography Oxford: Clarendon Press.
Takhtajan, A.L. 1980. Outline of the classification of flowering
plants. Bot. Rev. 46: 225-359.
Anderberg, A.A. 1992. The circumscription of the Ericales and
Tiffney, W.N. Jr. 1972. Snow cover and the Diapensia lappon-
their cladistic relationships to other families of "higher"
ica habitat in the White Mountains, New Hampshire.
dicotyledons. Syst. Bot.!?: 660-675.
Rhodora 74: 358-377.
Anderberg, A.A. 1993. Cladistic interrelationships and major
Wood, C.E. Jr., Channell, R.B. 1959. The Empetraceae and
clades of the Ericales. Pl. Syst. Evol. 184: 207-231.
Diapensiaceae of the southeastern United States. J. Arnold
Arbor. 40: 161-171.
APG (Angiosperm Phylogeny Group) 1998. See general refer-
Xi, Y.-Z., Tang, Y.-C. 1990. Pollen morphology and phylogenetic
ences.
relationships in the Diapensiaceae. Cathaya 2: 89-112.
Bliss, L.C. 1962. Caloric and lipid content in alpine tundra
Yamazaki, T. 1966. The embryology of Shortia uniflora with
plants. Ecology 43: 481-529.
brief review of the systematic position of the Diapensiaceae.
Brummitt, R.K. 1972. Nomenclatural and historical considera-
J. Jap. Bot. 41: 245-251.
tions concerning the genus Galax. Taxon 21: 303-317.
122 C. Bayer
Dirachmaceae
C. BAYER
Dirachmaceae Hutch., Fam. Fl. Pl., ed. 2: 248 (1959). the margin of this cup. Between each petal and the
stamen in front of it, a pubescent projection of the
Trees or shrubs; hairs simple and glandular- petal is found. According to Link (1994), nectar is
peltate. Leaves alternate, simple, lobed to serrate, probably produced by epithelial glands under-
petiolate; venation pinnate; stipules linear-trian- neath these appendages. The cup-shaped cavity
gular. Flowers pedicellate, single in the axils of of the flower is apically closed by the petal
foliage leaves; pedicel provided with 4-8 lanceo- appendages and their indumentum, and may
late or linear-triangular bracts. Flowers actino- retain the nectar (Fig. 42).
morphic, hermaphrodite, tetracyclic, perigynous; The fruit valves spread when ripe, but their
sepals 5-6 or 8, with valvate aestivation, pubescent; apical portions remain attached to the style. They
petals white, free, imbricate, as many as petals, expose clusters of long unicellular hairs that
with ventral, basal appendages; stamens 5-6 or 8, enclose the seeds. Based on his observations on
opposite the petals and basally fused with them; flower and fruit structure of Dirachma, Link
filament bases deltoid; anthers basifixed, extrorse, (1994) suspected melittophily and anemochory.
tetrasporangiate, opening by longitudinal slits; Boesewinkel (1988, 1995) and Boesewinkel and
gynoecium syncarpous; ovary pubescent, 5-6- or Bouman ( 1997) described the seed as exotestal
8-locular, radially lobed; style distally glabrous; with one antiraphal integumentary bundle and an
stigma clavate to cylindrical; ovules 1 per locule, endotegmic pigment layer.
bitegmic, anatropous-apotropous, placentation
basal-axile. Fruits capsular, septicidal and sep- POLLEN MORPHOLOGY. According to the short
tifragal, opening from base to apex, with long hairs descriptions given by Erdtman (1952) and Link
on the inner surface; seeds smooth, brown; (1991, 1994), the pollen grains are prolate and tri-
endosperm scanty. colporate; the exine is finely reticulate.
A monogeneric family with two species,
Dirachma socotrana Schweinf. ex Balf. f. from EMBRYOLOGY. Some embryological features of
Socotra, and D. somalensis D.A. Link from Dirachma have been described by Boesewinkel
Somalia. and Bouman (1997). The tapetum is secretory and
includes binuclear cells. Mature pollen contains
VEGETATIVE STRUCTURES. The branching system starch grains. The crassinucellate ovules are later-
of Dirachma socotrana includes long and short ally flattened and have a zigzag-shaped micropyle.
shoots, the latter arising from the axils of foliage Embryo sac development probably corresponds to
leaves on long shoots. Yakovleva (1994) reported the Polygonum type; the antipodals degenerate
the presence of mucilage-containing cells in the after fertilisation.
leaf epidermis of Dirachma socotrana.
PHYTOCHEMISTRY. Comparing several genera
REPRODUCTIVE STRUCTURES. Apart from simple of Geraniaceae, Bate-Smith (1973) found trivial
hairs, peltate glandular trichomes occur on the flavonoids in Dirachma, similar to those of
pedicels and on the outside of the sepals of Geranium.
Dirachma somalensis, and on the ovary of D. soco-
trana (Link 1994). The pedicels are provided with AFFINITIES. Following the original description by
four or up to eight sterile bracts, which are some- Balfour (1888), who found most affinities with
times described as epicalyx or hypocalyx. The Vivianieae and Wendtieae (formerly included in
flowers are octamerous in D. socotrana, and pen- Geraniaceae), many subsequent authors placed
tamerous or hexamerous in D. somalensis. The Dirachma at the level of tribe or family in or close
ovary is enclosed by a cup of receptacular or sepa- to Geraniaceae. However, Balfour (1888) himself
line origin. Petals and stamens are inserted near had indicated a certain resemblance with Tili-
Dirachmaceae 123
(Ronse Decraene and Smets 1995). However, it is

unlikely that these characters justify the position
of Dirachma within this alliance, because Mal-
valean characters such as the presence of stellate
hairs, palmate venation, mucilage in tissues other
than the epidermis, and exotegmic seed coats,
are missing. Several other important characters
of Dirachma, including the shoot anatomy, are
unknown so that the systematic position of
Dirachma remains uncertain. Recent studies
exclude close affinities with both Geraniales and
Malvales but suggest a placement in a redefined
rhamnalean clade, even if the relationships to
Barbeya, another isolated genus placed in this
alliance, remain unresolved (Boesewinkel and
Bouman 1997; Thulin et al. 1998).
DISTRIBUTION, HABITATS AND CONSERVATION.

Dirachma socotrana is endemic to Socotra, occur-
ring together with Buxus hildebrandtii on lime-
stone boulders. D. somalensis was collected in
Central Somalia on lime as well as on sand. Both
species are very rare and highly endangered
(Bazara'a et al. 1991; Link 1994).
One bispecific genus:
Dirachma Schweinf. ex Balf. f. Fig. 42

Dirachma Schweinf. ex Balf. f., Trans. Roy. Soc. Edinburgh 31:
45, t. 8 (1888).
Characters as for family.
Balfour, B. 1888. Botany of Socotra: Dirachma. Trans. Roy. Soc.

Edinburgh 31: 45-46, pl. 8.
Fig. 42. Dirachmaceae. A Flower of Dirachma somalensis Bate-Smith, E.C. 1973. Chemotaxonomy of Geranium. Bot. J.
(perianth and androecium removed in part; asterisk: nectar Linn. Soc. 67: 347-359.
gland). B D. socotrana. Flower segment in vertical section (G: Bazara'a, M., Guarino, L., Miller, A., Obadi, N. 1991. Dirachma
gynoecium, St: stamen, P: petal, Pa: petal appendage, arrow: socotrana - back from the brink? Oryx 25: 229- 232.
nectary). C Floral diagram of D. somalensis. (Link 1994) Boesewinkel, F.D. 1988. The seed structure and taxonomic
relationships of Hypseocharis Remy. Acta Bot. Neerl. 37:
111-120.
Boesewinkel, F.D. 1995. Seed coat structure and the
aceae. Hutchinson (1959) considered Dirachma to delimitation of the Geraniales. Acta Bot. Neerl. 44: 491
represent a distinct family and included it in his (Abstract).
Tiliales. Characters of Dirachma that have been Boesewinkel, F.D., Bouman, F. 1997. Ovules and seeds of
Dirachma socotrana (Dirachmaceae). Plant Syst. Evol. 205:
used to support this placement include the pres- 195- 204.
ence of a hypocalyx, a valvate, amucronate calyx, Erdtman, G. 1952. See general references.
petals with ventral projection and ventral nectary Hutchinson, J.1959. The families of flowering plants, ed. 2, vol.
(albeit epithelial instead of trichomatic), hairs on 2. Oxford: Clarendon Press.
the ventral side of the fruit wall (Link 1991, 1994), Knuth, R. 1931. Geraniaceae. In: Engler & Prantl, Die nattir-
lichen Pflanzenfamilien, ed. 2, 19a. Leipzig: Engelmann, pp.
special features of the mucilage cells (Yakovleva 43-66.
1994), and the obhaplostemonous androecium
124 C. Bayer
Link, D.A. 1991. Dirachma somalensis D.A. Link sp. nov. A new Thulin, M., Bremer, B., Richardson, J., Niklasson, J., Fay, M.F.,
species of a remarkable and highly endangered mono- Chase, M.W. 1998. Family relationships of the enigmatic
generic family. Bull. Jard. Bot. Belg. 61: 3-13. rosid genera Barbeya and Dirachma from the Horn of Africa
Link, D.A. 1994. Dirachma Schweinf. (Dirachmaceae) a highly region. Plant Syst. Evol. 213: 103-119.
remarkable and endangered bispecific genus. In: Seyani, Yakovleva, O.V. 1994. The ultrastructure of mucilage cells in
J.H., Chikuni, A.C. (eds.) Proc. 13th Plenary Meet. the leaf epidermis of Dirachma socotrana (Dirachmaceae).
A.E.T.F.A.T., Zomba, Malawi, 2-11 April1991. Vol. 2. Plants Bot. Zhurn. 79: 52-58, pl. 2.
for the people. Zomba: National Herbarium and Botanic
Gardens of Malawi, pp. 1229-1238.
Ronse Decraene, L.P., Smets, E.F. 1995. The distribution and
systematic relevance of the androecial character oligomery.
Bot. J. Linn. Soc. 118: 193-247.
Ebenaceae 125
Ebenaceae
B. WALLNOFER
Ebenaceae Giirke in Engler & Prantl, Nat. Pflanzenfam. IV, 1: tenuinucellate. Fruit a berry usually with a pulpy
153 (1891), nom. cons. endocarp; seeds 1-16 per fruit, pendulous; hilum
small, apical, inconspicuous; seed coat exotestal,
Mostly evergreen dioecious, rarely monoecious or pigmented, thin, parenchymatous, soft to leathery;
polygamous trees or shrubs; bark, roots and heart- endosperm hard, abundant, oily, smooth to rumi-
wood often black. Unicellular trichomes simple or nate; embryo surrounded by endosperm, turned
two-armed; multicellular glandular or peltate upside-down (radicle superior), with two, oval,
hairs sometimes present. Leaves simple, mostly well-developed foliaceous cotyledons and a
alternate, spirally or distichously arranged, mostly strongly developed radicle.
petiolate, exstipulate; leaf-margins usually entire; A family comprising two genera and c. 500-600
extrafloral nectaries usually present on abaxial leaf species, distributed mainly in the tropical and sub-
surface. Inflorescences axillary, cymose, fascicu- tropical regions of both the Old and the New
late, raceme-like or paniculate. Flowers usually World.
unisexual, less frequently hermaphroditic, articu-
lated at base, regular, 3-5( -8)-merous; calyx VEGETATIVE MORPHOLOGY. Most species of
mostly gamosepalous, valvate or imbricate, Diospyros are small to medium-sized trees in the
spreading or reflexed, persistent and often accres- forest understorey. Only few species grow up to
cent in fruit; corolla gamopetalous, isomerous large canopy trees. Euclea species are subshrubs,
with calyx, shortly to deeply lobed; tube promi- shrubs or trees up to 12m high. The growth archi-
nent or very short in male flowers; male flowers: tecture of many species of Diospyros conforms to
stamens (3-) 12-20( -100), mostly unequal in size, Massart's model (Halle et al. 1978). Leaf arrange-
often setulose, commonly inserted at base of ment is distichous on the branches and spiral on
corolla tube, hidden or slightly exserted, often the trunk. In the latter case the leaves are often
arranged in two hardly discernible whorls; fila- much reduced in size or only scale-like. Some
ments usually short and flattened, free or united in species of the fire-adapted savannahs in South
pairs, triads or fascicles, or sometimes into a America and Africa are geoxylic subshrubs.
central cylinder; anthers usually linear, erect, basi- Certain species have been reported to develop
fixed, distally often apiculate, tetrasporangiate, the root-suckers and to establish often large popula-
outermost introrse, the innermost extrorse, and tions or coppices (Skallerup 1953; Rathore 1971;
those in-between latrorse, longitudinally dehisc- White 1983; for further details see also WallnOfer
ing; pistillode usually reduced, lacking ovules, 2001).
style and stylodia, rarely totally absent, or well-
formed in the terminal (structurally or rarely VEGETATIVE ANATOMY. The bark of many tropi-
functionally hermaphroditic) flower of a male cal species is black outside. In cross-section the
cyme; female flowers: staminodes arranged in a bark of twigs shows a characteristic thin layer of
single whorl, usually inserted at base of corolla black tissue. Stone cells are present in young twigs,
tube, often markedly reduced in shape, fewer than leaves, and fruits of various species. The indu-
stamens in male flowers, rarely absent; rudimen- mentum consists of four kinds of grey, brownish
tary anthers introrse; disk absent or less frequently or often characteristically rusty-brown hairs: uni-
well-developed; gynoecium syncarpous, 2-8- cellular, simple trichomes; unicellular, two-armed,
carpellate; ovary superior, sessile; style short, style sessile or rarely short -stalked trichomes; small,
branches usually longer than style, same number club-shaped, grey, multicellular, glandular hairs,
as carpels; stigmas often variously lobed; carpels and peltate, multicellular hairs (Busch 1913;
biovulate, mostly bilocular due to a false, second- Solereder 1899, 1908, 1914; Ng 1971). Extrafloral
ary septum or rarely unilocular; placentation nectaries of the specialized Benincasa type (Fig.
apical; ovules pendulous, anatropous, bitegmic, 43L) occur on abaxial leaf surfaces near the base
126 B. Wallnofer
of the lamina and often scattered along, but some- Euclea are mostly unbranched, multiflorous
what away from the midvein (Contreras and racemes.
Lersten 1984; Eisler 1907). Characteristic epider-
mal papillae are present on the abaxial leaf sur- FLOWER MORPHOLOGY. Ebenaceae have unisex-
faces of several species of Diospyros. Each hairless ual or rarely hermaphroditic flowers and are
epidermal cell yields on its surface a more or less mostly dioecious; less frequently they are monoe-
centrally placed, coronulate papilla, which is star- cious or even more rarely polygamous. In Euclea
like connected to the papillae of the surrounding the male flowers are usually larger than the female
cells by cuticular folds (Eisler 1907). The stomata but similar in shape. In Diospyros the male flowers
are usually confined to the abaxial leaf surfaces are smaller than the female and different in shape.
and are mostly anomocytic, less frequently Hermaphroditic flowers occur generally in
actinocytic (Ng 1971; Wilkinson 1979). Diospyros sect. Royena (the former genus Royena)
The wood is usually hard, heavy, fine-textured, and, according to Salter (1953) and White and
and pale or dark coloured. The heartwood of Barnes (1958), are only structurally but not func-
about 50 species is coloured black or dark brown tionally hermaphroditic. Among species of other
("ebonised") due to a dark substance (derivatives sections of Diospyros, hermaphroditic flowers are
and oxidative decomposition products of naph- sporadically found and occupy the terminal posi-
thoquinones) filling the lumina. The heartwood of tions in otherwise male inflorescences (Yasui 1915;
some species is irregularly composed of coloured Namikawa et al. 1932; Franceschi 1993).
and uncoloured parts and has a variegated, The number of floral parts is known to vary
banded, or marbled aspect. According to Wright within species and cannot be used for species
(1904) and Hillis and Soenardi (1994), the black delimitation or to segregate smaller genera, as had
deposits are formed as a response after injury and been done in the past. The carpels are biovulate
subsequent infection by fungi (for further details and in some species remain unilocular but mostly
see also WallnOfer 2001). are divided by a false longitudinal septum that
The structure of the wood is remarkably con- originates from the carpellary wall opposite the
stant throughout the family: vessels solitary and in placenta. In the first case, each locule is biovulate
radial multiples of 2, 3, 4 or sporadically more, and ovaries are 2-, 3- or 4-locular. In case of the
with simple perforations; pits small (to 8 Jlm uniovulate locules, ovaries are 4-, 6-, 8-, 10-, 12-,
diameter) between vessels as well as between 14- or 16-locular (Ng 1971, 1991b).
vessels and ray cells or parenchyma; parenchyma
predominantly apotracheal, scattered and in EMBRYOLOGY. The anther wall comprises the epi-
numerous uniseriate lines; sometimes forming dermis, the fibrous or non-fibrous endothecium,
vasicentric sheaths round the vessels; rays 1-2 two middle layers, and multinucleate tapetal cells.
cells, rarely to 3 or 4 cells wide, less than 1 mm The ovule is pendulous, oblong, anatropous,
high, heterogeneous; fibers with small pits, and bitegmic, and tenuinucellar. The archesporia!
with walls thinner than lumina, rarely thicker (Ng cell functions as megaspore mother cell, under-
1971, 1991b; Metcalfe and Chalk 1950). goes meiotic divisions, and produces a linear
tetrad. The chalazal megaspore develops into a
INFLORESCENCE STRUCTURE. The Ebenaceous Polygonum type of embryo sac. Endosperm devel-
inflorescence (Ng 1991b, Franceschi 1993) is opment is cellular. The embryogeny and organo-
conventionally referred to as a cyme. It is always genesis probably correspond to the Polygonad
axillary; the pedicels are articulated and, if type-Chenopodiad variation or Chenopodiad type
lateral, subtended by a bract. If there is one (Davis 1966; Johri et al. 1992). The seed coat is
flower only (especially in the female condition), formed only by the outer integument.
it occupies a terminal position on a bracteate
peduncle. Quite often, the inflorescence is a POLLEN MORPHOLOGY. The pollen grains are tri-
three-flowered dichasium. Numerous variations colporate and prolate-spheroidal to prolate. The
are seen ranging between three-flowered dichasia sexine is as thick as the nexine or thicker and
and multibranched paniculate inflorescences but, smooth or very finely warty. The ora are mostly
in nearly all cases, the terminal units are dichasia. well defined, usually lalongate, sometimes with
Usually within the same species the male inflores- indistinct lateral edges or rarely indistinguishable.
cences are many-flowered, whereas the female The pollen is remarkably constant throughout the
ones are only 1(3)-flowered. The inflorescences of family, and the main variations are in size and
Ebenaceae 127
shape of the grains and the apertures (Erdtman fruits of various cultivars of D. kaki are seedless
1952; Ng 1971; Franceschi 1993). (Yonemori et al. 2000).
Each fruit of Diospyros contains 1-16 seeds,
KARYOLOGY. In both genera the basic chromo- which are 8-40mm long. The seeds of Euclea are
some number is x = 15. An euploid series with 2n usually solitary and 3-10mm in diameter when
= 30, 60, 90, and 135 has been observed in Diospy- ripe, if more than one, then forming a sphere
ros, with 36 species reported to be diploid, and a together. During the development of the seed of
few others tetra- and hexaploid (Franceschi 1993; Euclea, the other (abortive) ovules and the axis are
Yonemori et al. 2000). Eight species of Euclea are pushed to one side of the fruit. The solitary seeds
diploid (White and Vosa 1980). are, therefore, encircled by a curved vascular loop
(like on a tennis ball), which is apically linked by
PoLLINATION. The flowers of various species are a shallow, vertical groove which represents the
reported to be visited by bees (Hague 1911; Ayala- impression of the displaced axis (Ng 1971; Winter
Nieto and Ludlow-Wiechers 1983). Beetles (con- 1963).
sidered to be the main pollinators), flies and wasps The rumination of the endosperm in Diospyros,
(all c. 3-6mm long) were observed visiting the if present, is of the Annona type (Periasamy 1966),
male and female flowers of Diospyros pentamera in whereas in Euclea it takes the peculiar form of
Australia (Irvine and Armstrong 1990). Diospyros ingrowths from the testa forming a cylinder round
hispida, a species of the Brazilian cerrado, flowers the whole radicle (Ng 1971; White 1983). The
during the night and attracts small nocturnal Lep- rumination in Diospyros sect. Royena forms only
idoptera (moths) (Silberbauer-Gottsberger and a shallow ingrowth round the distal part of the
Gottsberger 1988: 657). Also various rainforest- radicle. The embryo is turned upside-down
species are reported to flower during the night, but (radicle superior), possessing two oval, well-
nothing is known about the pollinators. developed foliaceous cotyledons and a strongly
developed radicle.
FRUIT AND SEED. The fruit is a multilocular,
usually indehiscent berry subtended by a persist- DISPERSAL. The fruits and seeds are dispersed by
ent, 3-8-lobed or rarely unlobed (truncate) calyx, various fruit -eating birds and mammals (Ridley
which is usually accrescent in Diospyros but not in 1930; White 1983; Pannell and White 1988). The
Euclea. The fruits of some African species of fruits of coastal and riverine species as well as
Diospyros sect. Royena become dry and occasion- those of species growing in periodically inundated
ally are tardily dehiscent (Winter 1963; White habitats may perhaps also be water dispersed (for
1983). In no species, however, is the fruit com- further details see also WallnOfer 2001).
pletely and spontaneously dehiscent. At fruit-
maturity the pericarp is either completely soft or GERMINATION AND SEEDLING. The germination
possesses a thinner or thicker, leathery or hard of Diospyros seeds is mostly epigeal, less fre-
outer layer. A hypodermal layer of stone-cells, very quently hypogeal or of the type "durian" (=crypto-
rarely of radial fibers or a mixture of fibers and epigeal). The cotyledons are emergent and
stone-cells is developed. The mesocarp is often photosynthetic or not, or are non-emergent, and
exiguous and sometimes two-layered. A layer of shed together with the seed body. The epicotyle-
pulpy, sometimes fibrous endocarp, which is donary leaves are usually alternate, but in a few
mostly present, adheres closely to the seeds and species the first two are subopposite (Ng 1971,
resembles a sarcotesta (Ng 1971, 1991a). Idioblasts 1991a; for further details see also Wallnofer 2001).
containing tannins occur generally in the pericarp
of the fruits (Corner 1976; Utsunomiya et al.1998). PHYTOCHEMISTRY. Naphthoquinones, terpen-
Especially in the immature fruits, tannins are oids, benzopyrones, and condensed tannins
responsible for the characteristic astringent taste, (mostly based on prodelphinidin) are widely dis-
which diminishes when the fruits ripen but rarely tributed and are characteristic of Ebenaceae
disappears completely as in some cultivars (Mallavadhani et al. 1998; Hegnauer 1966, 1989).
(Yonemori et al. 2000). The fruits of some species Naphthoquinones are frequent and their oxidation
are poisonous, presumably when raw, and are used products provide the dark colour of the bark,
in various parts of the world to catch fish. The heartwood, leaves and fruits. Idioblasts containing
diameter of mature fruit varies in the range 5-100 tannins occur in various organs, including fruits
mm (Ng 1971, 1991a). Due to parthenocarpy, the (Yonemori et al. 2000).
128 B. Wallnofer
AFFINITIES. Ebenaceae have traditionally been (zapote prieto, black sapote), D. blancoi (velvet
placed in the order Ebenales, together with all or apple or mabolo ), and D. lotus. The fruits of many
some of the following families: Sapotaceae, other species are of more regional importance.
Styracaceae, Symplocaceae and Lissocarpaceae The leaves of D. melanoxylon ("tendu") are uti-
(Morton et al. 1997). Recent molecular studies, lized in India as wrappers for cigarettes called
based upon rbcL sequence data, have demon- "bidi" (Hunter 1981). Ebony timbers are used
strated that Ebenaceae are monophyletic, whereas mainly for the production of musical instruments,
Ebenales in their traditional circumscription for carved work, cabinet making, and as decora-
are not. In these studies Ebenaceae appear as tive veneer for the production of furniture. The
the sister-group of the Primulales within the true ebony timbers are obtained from various
expanded Ericales as defined by APG (1998). species having a completely or partially black
Lissocarpa, the only genus of Lissocarpaceae, had heartwood, the economically most important of
been included in Ebenaceae by some authors and, which are Diospyros ebenum and D. melanoxylon
according to Ng (1971, 1991b) and Franceschi from India and Sri Lanka.
(1993), is the closest relative of Ebenaceae. This is
strongly endorsed by the molecular studies of KEY TO THE GENERA
Berry et al. (2001) and Anderberg et al. (2002).
1. Calyx usually accrescent on fruits; berry usually many-
seeded; seeds usually laterally flattened, encircled by a more
DISTRIBUTION, HABITAT AND ECOLOGY. Ebe- or less straight vascular loop round the periphery, which is
naceae occur mainly in the lowlands of the tropi- apically not linked by a vertical groove 1. Diospyros
cal and, to a lesser extent, subtropical regions of - Calyx not accrescent on fruits; berry usually 1(3)-seeded;
seeds usually subglobose, encircled by a curved vascular
the world. Only a few species are found at higher
loop, which is apically linked by a shallow, vertical groove
elevations or in warm-temperate regions. Euclea is 2.Euclea
restricted to Africa and Arabia, whereas Diospyros
is pan tropical. Many species of the latter genus are
small to medium-sized trees of the forest under- 1. Diospyros L. Fig. 43
storey and often have a remarkably low population Diospyros L., Sp. Pl., ed. 1: 1057 (1753); Hiern, Trans. Cam-
density. Several species grow along rivers; some bridge Philos. Soc. 12 (1 ): 27-300 (1873), rev.; Bakhuizen van
are rheophytes (White 1998), and others are char- den Brink, Bull. Jard. Bot. Buitenzorg, III, 15 (1-5): 1-515
acteristic elements of swamps, beach communities (1936-1955), SE Asian spp.; White, Bull. Jard. Bot. Belg. 48:
245-358 (1978), 58: 325-448 (1988), African spp.; Perrier de
and periodically flooded habitats. Some species Ia Bathie, Flore de Madagascar et des Comores (1952).
occur in dryer vegetation, in fire-prone savannahs, Royena L. (1753).
or in deciduous forests. Maba J.R. Forster & J.G.A. Forster (1775).
Tetraclis Hiern (1873).
PALAEOBOTANY. The fossil record has been Rhaphidanthe Hiern ex Giirke (1891).
compiled by Hiern (1873), Franceschi (1993),
Collinson et al. (1993), and Mai (1995). Most finds Leaves usually alternate and with entire margins.
are from the Tertiary. Fossil leaves, because of their Inflorescences usually cymose or fasciculate if
rather nondescript and very variable appearance, male and uniflorous (less frequently few-flowered)
can probably not be assigned to the family with if female. Male flowers usually smaller than the
absolute certainty. It seems that in the past not female ones, and differing in shape. Calyx variable,
much attention was paid to the presence of extra- 3-8-lobed, or cup-shaped and entire, persistent
floral nectaries and the coronulate papillae on the and usually accrescent on fruits. Stamens 2 to ca.
abaxial leaf surface. Probably many fossils which 100. Ovary 2-8-carpellate. Fruit a 1-16-seeded
have been assigned to Ebenaceae may have to be berry. 2n =30, 60, 90, 135. About 500-600 spp., with
reconsidered. 200-300 in Asia and the Pacific area, ca. 95 in
Madagascar, 94 on the African mainland, ca. 100 in
USES AND ECONOMIC IMPORTANCE. Ebenaceae the Americas, and 15 in Australia.
are the source of some economically important A comprehensive monograph is badly in need
products, the most valuable being the persimmon since the last one dates back more than a century
fruits and ebony timbers. Diospyros kaki is the (Hiern 1873). Besides Diospyros (with 15 sections),
most important fruit-yielding species and is culti- Hiern (1873) recognized also Maba (with 6 sec-
vated in many varieties on a large scale, especially tions), Royena and Tetraclis, which now are all
in temperate east Asia (Yonemori et al. 2000). included in Diospyros. Bakhuizen van den Brink
Other frequently cultivated species are D. digyna (1936-1955) grouped the SE Asiatic and Pacific
Ebenaceae 129
JM
cyathiform or patelliform, persistent but not
accrescent in fruit. Stamens 10-30. Ovary 2-3-
carpellate. Fruit usually a 1(3)-seeded berry. 2n =
~
B
30. About 12-20 spp. confined to Africa, Arabia,
Socotra and the Comoro Islands; many species
restricted to the Cape flora.
The species pertaining to subgen. Rymia (Endl.)
Verde. occur only in South Africa, Namibia and SW
Angola and have an urceolate to subglobose, shal-
lowly 5-8-lobed corolla. The species belonging to
subgen. Euclea are widespread and have a cam-
panulate, deeply 4-5-lobed corolla.
m
Anderberg, A.A. et a!. 2002. See general references.

references.
D Ayala-Nieto, M.L., Ludlow-Wiechers, B. 1983. Catalogo
Palinol6gico para Ia Flora de Veracruz. No. 13. Familia
Ebenaceae. Bi6tica 8: 215-226.
Bakhuizen van den Brink, R.C. 1936-1955. Revisio Ebe-
nacearum Malayensium. Bull. Jard. Bot. Buitenzorg III, 15
(1- 5): 1- 515.
Berry, P.E., Savolainen, V., Sytsma, K.J., Hall, J.C., Chase, M.W.
2001. Lissocarpa is sister to Diospyros (Ebenaceae). Kew
Bull. 56: 725-729.
Busch, P. 1913. Anatomisch-systematische Untersuchung der
Gattung Diospyros. Crefeld: Wilhelm Greven.
Collinson, M.E., Boulter, M.C., Holmes, P.L. 1993. Magnolio-
phyta ('Angiospermae'). In: Benton, M.J. (ed.) The fossil
record 2. London: Chapman & Hall, pp. 809-841.
Contreras, L.S., Lersten, N .R. 1984. Extrafioral nectaries in
Ebenaceae: anatomy, morphology, and distribution. Am. J.
Bot. 71: 865-872.
Fig. 43. Ebenaceae. A-K Diospyros curranii. A Fruiting Davis, G.L. 1966. See general references.
branch. B Calyx with ovary. C Ovary. D Transverse section of Eisler, E.1907. Das extrafiorale Nektarium und die Papillen der
ovary. E Transverse section of fruit. F Fruit. G Calyx of fruit, Blattunterseite bei Diospyros discolor Willd. Sitzungsber.
outside view. H Flower. I Same, vertical section. J Stamens. K Kaiser!. Akad. Wiss., Math.-Naturwiss. Cl., Abt. 1, 116:
Staminodes. L Diospyros koeboeensis, underside of leaf with 1563-1590.
nectaries. (Bakhuizen van den Brink 1936-1955) Erdtman, G. 1952. See general references.
Franceschi, D. de 1993. Phylogenie des Ebenales: analyse de
l'ordre et origine biogeographique des especes indiennes.
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virginiana. Bot. Gaz. (Crawfordsville) 52: 34-44, pl. 1- 3.
Halle, F., Oldeman, R.A.A., Tomlinson, P.B. 1978. Tropical trees
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Euclea Murray, Syst. Veg., ed. 13: 747 (1774); Hiern, Trans.
Hiern, W.P. 1873. A monograph of Ebenaceae. Trans.
Cambridge Philos. Soc. 12 (1): 27-300 (1873),rev.; de Winter,
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less frequently compound or occasionally flowers cal forests of Australia. In: Bawa, K.S., Hadley, M. (eds.)
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Johri, B.M. et al. 1992. See general references. Solereder, H. 1899. Systematische Anatomie der Dicotyledo-
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Elaeagnaceae 131
Elaeagnaceae
I.V. BARTISH and U. SwENSON
Elaeagnaceae Juss., Gen. Pl.: 74 (1789), nom. cons. tured pits are present, and nonvestured pits are
also common. Rays are heterogeneous to almost
Anemophilous or entomophilous small trees, homogeneous in some species of Elaeagnus. Axial
shrubs or rarely woody climbers; shoots often parenchyma is diffuse, sometimes very scanty.
reduced to spines; young branches, leaves, and Phloem is usually tangentially stratified into hard
calyx tube covered with peltate and stellate tri- and soft layers (Jansen et al. 2000). Sieve-element
chomes; nodules on the roots containing nitrogen- plastids are of Ss-type.
fixing bacteria. Leaves alternate, opposite, or rarely
in pseudowhorls, simple, petiolate, without stip- EMBRYOLOGY. Microsporogenesis is simultane-
ules; lamina entire, pinnately veined. Inflores- ous, and pollen grains are 2-celled or 3-celled
cences axillary, short, fasciculate, spicate or when shed. Embryo sac development is of the
racemose, or rarely flowers solitary. Flowers actin- Polygonum type or (Shepherdia) Allium type.
omorphic, apetalous, often fragrant, perfect, or Endosperm formation is nuclear. An endospermal
plants monoecious, dioecious, or rarely polyga- chalazal haustorium develops in Elaeagnus.
mous, the female flowers without staminodes, the
male without pistil; sepals 2 or 4( -6), joined into a POLLEN MORPHOLOGY. The reader is referred to
hypanthium; hypanthium free, constricted above Erdtman (1952), Leins (1967), and Sorsa (1971).
the gynoecium, white, cream, or yellow; receptacle Pollen is (2)3(4)-colporate, oblate or oblate-
tubular (bisexual and female flowers) or mostly spheroidal. In Elaeagnus, three areas with a com-
flat (male flowers); stamens in one (rarely two) paratively thin sexine are separated by three
whorl(s), (2-)4 or 8(-12), adnate to hypanthium, longitudinal, aperturiferous strands of incrassate
equal in length, all fertile, alternisepalous and/or sexine. Pollen grains of Shepherdia argentea and S.
oppositisepalous, erect in bud; filaments free, rotundifolia are angulaperturate or oblate. In con-
very short; anthers dorsifixed or basifixed, non- trast, grains of S. canadensis are longicolpate-
versatile, longitudinally dehiscing; gynoecium parvorate and, as to their shape, distinctly
one-carpellate (probably pseudomonomerous; different from other elaeagnaceous pollen grains.
Takhtajan 1997); ovary superior, 1-celled and 1- The pollen of Hippophae is suboblate or oblate-
ovulate; style elongate; stigma dry, non-papillate; spheroidal.
ovule with a short and broad funicle, anatropous,
bitegmic, crassinucellate; placentation basal. Fruit KARYOLOGY. The basic chromosome number in
drupe- or berry-like with thin, membranous peri- Hippophae is x = 12 (Rousi 1965) and in Elaeagnus
carp, enclosed by the persistent calyx tube, which x = 14 (Arohonka and Rousi 1980). Shepherdia
becomes fleshy. Seed solitary; testa hard; embryo canadensis has x = 11 and S. argentea x = 13
straight and achlorophyllous; endosperm scanty (Arohonka and Rousi 1980, and references
or absent; cotyledons fleshy, plano-convex. Germi- therein).
nation phanerocotylar.
A family with three genera and 30-50 species, PoLLINATION. Hippophae is fully anemophilous,
distributed mainly in the temperate regions of the while Elaeagnus and Shepherdia often have fra-
Northern Hemisphere, some in tropical SE Asia grant flowers and are entomophilous.
and eastern Australia.
SEED DISPERSAL. The fleshy fruits of all three
VEGETATIVE STRUCTURES. The stems and leaves genera are eaten by birds and may be carried for
are covered with peltate or scaly trichomes. The long distances. Fruits of Hippophae dried during
leaves are leathery, simple, entire and without winter are known to float and disperse by spring
stipules. Stomata are anomocytic. Vessels have floods along rivers.
simple perforations, and small to vestigial ves-
132 I. V. Bartish and U. Swenson
REPRODUCTIVE SYSTEMS. Hippophae and Shep- PALAEOBOTANY. The oldest fossil pollen attrib-
herdia are dioecious, Elaeagnus is monoecious or uted to Elaeagnaceae is recorded from the upper
polygamous. Eocene of Central Asia (Pulatova 1973 ). In Europe,
the oldest, well-documented fossil record for
PHYTOCHEMISTRY. Elaeagnaceae are strongly Elaeagnus or a related genus appears to be
tanniniferous; both ellagitannins and condensed Slowakipollis hippophaeoides from the Oligocene
tannins are present. Further compounds reported (Krutzsch 1962). Shepherdia pollen has been
by Hegnauer (1966, 1989) include indol alkaloids, reported from the Miocene of Oregon (Graham
sinapinic acid, flavonols, pentacyclic triterpenes 1963), and from the Miocene of Idaho (Smiley et
and L-quebrachit. al. 1975). Boehlensipollis hohli from the Oligocene
of Europe may represent an extinct genus ofElaea-
SYSTEMATICS. Phylogenetic studies of morpho- gnaceae (Muller 1981).
logical, karyological, and molecular characters
have identified three clades, corresponding to the MYCOTROPHY. Mycorhiza fungi are present in
genera Elaeagnus, Hippophae and Shepherdia the cortex of the rootlets of Hippophae and Shep-
(Hyvonen 1996; Bartish et al. 2002). Sequence herdia. All genera possess root nodules contain-
analysis of the chloroplast DNA rbcL fragment ing nitrogen-fixing actinomycetes (Frankia) with
indicates a close relationship between Elaeagnus nitrogenase activity (Gardner 1958; Vanstraten et
and Shepherdia, Hippophae being sister to that al. 1977). Frankia strains isolated from Elaeagnus
clade (Richardson et al. 2000). and Hippophae are able to cross the inoculation
barriers and infect Alnus species (Bosco et al.
AFFINITIES. Elaeagnaceae have been placed 1992).
either into Proteales (Cronquist 1988) or Rham-
nales (Thorne 1992), or in a monotypic order,
Elaeagnales, close to the two former (Takhtajan
1997). More recent molecular and morphological
analyses (Thulin et al. 1998; Richardson et al. 2000;
Soltis et al. 2000) place Eleaeagnaceae together
with Barbeyaceae, Cannabaceae, Cecropiaceae,
Celtidaceae, Dirachmaceae, Moraceae, Rham-
naceae, Rosaceae, Ulmaceae, and Urticaceae in a
group of families forming the expanded order
Rosales sensu APG (1998). Barbeyaceae attach as
sister to Elaeagnaceae with a moderate support;
Barbeya, a small dioecious tree with opposite
leaves and wind-pollinated flowers, is in many
respects similar to Hippophae. It is unclear,
however, whether Rhamnaceae or Ulmaceae
are most closely related to Elaeagnaceae and
Barbeyaceae.
DISTRIBUTION AND HABITATS. The family is

mostly deciduous and has a typical Northern
Hemisphere range; in SE Asia it extends to Malesia
' A
and NE Queensland. Elaeagnus occurs throughout
this range, whereas Hippophae is found through-
out Eurasia (Lian et al. 1998). Shepherdia is
restricted to North America. Elaeagnaceae occur
chiefly in steppes, river deltas and mountain ter-
races, valley slopes and along coasts at altitudes
from 0 to 5000m (Rousi 1971; Lian et al. 1998) but,
in SE and E Asia, Malesia and NE Queensland,
Fig. 44. Elaeagnaceae. Shepherdia canadensis. A Branch of
Elaeagnus is a lianaceous rain forest element female plant with young fruits. B Fascicle of male flowers. C
(Veldkamp 1986). Fascicle of female flowers. Drawn by Pollyanna von Knorring
(Orig.)
Elaeagnaceae 133
Fig. 45. Elaeagnaceae. Elaeagnus angustifolia. A Flowering

branch armed with thorns. B Hermaphrodite flower, dissected,
one stamen fallen off. CFruiting branch. Drawn by Pollyanna
von Knorring (Orig.)
Fig. 46. Elaeagnaceae. Hippophae rhamnoides. A Fruiting
branch armed with thorns. BShort shoot of male plant with
EcoNOMIC IMPORTANCE. A number of Elaeagnus flowers. C Male flower with two sepals and four stamens (one
species are grown as ornamental shrubs. Elaeag- of which removed). D Female flowers. Drawn by Pollyanna von
nus angustifolia, E. macrophylla, E. pungens, and E. Knorring (Orig.)
umbellata are mainly grown as deciduous or ever-
green shrubs for their attractive foliage. The fruits
of a number of species are edible, for example,
those of Hippophae rhamnoides (sea buckthorn), 2. Polygamous or monoecious; flowers often scented, sepals 4
Elaeagnus angustifolia (Russian olive, or oleaster), 2. Elaeagnus
and Shepherdia argentea (silver buffalo berry). - Dioecious; flowers odourless; sepals 2 (male flowers)
3. Hippophae
Fruits of sea buckthorn are rich in vitamins A, C,
E, and have been used for centuries in Europe and
Asia for the production of juices, jams, and bever-
ages. Seed oil of sea buckthorn was formally listed 1. Shepherdia Nutt. Fig. 44
in the Pharmacopoeia in 1977 and clinically tested
Shepherdia Nutt., Gen. N. Am. Pl. 2: 240 (1818); Servettaz, Beih.
in Russia and China (Xu 1994). The most impor- Bot. Centralbl. 25: 1-420 (1909).
tant pharmacological functions attributed to sea
buckthorn oil are: anti-inflammatory, bactericidal, Dioecious. Leaves opposite. Flowers scented, ento-
pain relief, and promotion of tissue regeneration. mophilous; in male flowers stamens 8, alternate to
Currently, Hippophae plantations are also used in and opposite with the 4 sepals. Three species in
China to prevent erosion of soil (Lu 1992). temperate North America.
KEY TO THE GENERA

1. Leaves exclusively opposite; stamens 8 (dioecious) 2. Elaeagnus L. Fig. 45
1. Shepherdia
- Leaves alternate, rarely subopposite or in pseudowhorls; Elaeagnus L., Sp. Pl.: 121 (1753); Servettaz, Beih. Bot. Centralbl.
stamens 4 2 25: 1-420 (1909).
134 LV. Bartish and U. Swenson
Polygamous or monoecious. Leaves alternate. Graham, A. 1963. Systematic revision of the Sucker Creek and
Flowers 4-merous, often scented and ento- Trout Creek Miocene floras of southeastern Oregon. Am. J.
Bot. 50: 921-936.
mophilous; perianth quadrangular; stamens equal Hegnauer, R. 1966, 1989. See general references.
in number and alternating with the sepals. Fruit Hyvonen, J. 1996. On phylogeny of Hippophae (Elaeagnaceae).
usually with eight longitudinal ribs. About 20-45 Nordic J. Bot. 16: 51-62.
species, most in Eurasia and from there to North Jansen, S., Piesschaert, F., Smets, E. 2000. Wood anatomy of
America (only one species, Elaeagnus commu- Elaeagnaceae, with comments on vestured pits, helical
thickenings, and systematic relationships. Am. J. Bot. 87:
tata), and through tropical SE Asia and Malaysia 20-28.
to N Queensland. The most widespread species is Krutzsch, W. 1962. Stratigraphisch bzw. botanisch wichtige
Elaeagnus angustifolia with discontinuous range neue Sporen- und Pollenformen aus dem deutschen Tertiar.
from China and C Asia to Europe. The genus is in Geologie 11: 265-307.
Leins, P. 1967. Morphologische Untersuchungen an
need of revision. Elaeagnaceen-Pollenkornern. Grana Palynologica 7: 390-
399.
Lian, Y.S., Chen, X., Lian, H. 1998. Systematic classification of
3. Hippophae L. Fig. 46 the genus Hippophae L. Seabuckthorn Res. 1: 13-23.
Lu, R. 1992. Seabuckthorn: A multipurpose plant species for
Hippophae L., Sp. Pl.: 1023 (1753); Rousi, Ann. Bot. Fennici 8: fragile mountains. Katmandu: ICIMOD Publication Unit.
177-227 (1971); Swenson and Bartish, Nord. J. Bot. 22: Muller, J. 1981. See general references.
369-374 (2002). Pulatova, M.Z. 1973. The upper Eocene flora of the Tadjik
depression by palynological data. Palinologiya kaynofita.
Moscow: Nauka, pp. 114-121 (in Russian)
Dioecious. Leaves usually alternate, rarely in Richardson, J.E., Fay, M.F., Cronk, Q.C.B., Bowman, D., Chase,
pseudowhorls or subopposite. Flowers odourless, M.W. 2000. A phylogenetic analysis of Rhamnaceae using
anemophilous; calyx with only 2 lobes, valvate; rbcL and trnL-F plastid DNA sequences. Am. J. Bot. 87:
male flowers with 4 stamens, alternate to and 1309-1324.
Rousi, A. 1965. Observations on the cytology and variation of
opposite with the sepals. Fruit with longitudinal European and Asiatic populations of Hippophae rham-
ribs in several species. Seven species, four of them noides. Ann. Bot. Fennici 2: 1-18.
recognized recently, in temperate areas of Eurasia. Rousi, A. 1971. The genus Hippophae L.: a taxonomic study.
Hippophae rhamnoides is the most widespread Ann. Bot. Fennici 8: 177-227.
species with fragmented range from C China to the Smiley, C.J., Gray, J., Huggins, L.M. 1975. Preservation of
Miocene fossils in unoxidized lake deposits, Clarkia, Idaho.
Atlantic coast of Europe. J. Paleontol. 49: 833-844.
Soltis, D.E. et al. 2000. See general references.
Sorsa, P. 1971. Pollen morphological study of the genus Hip-
Selected Bibliography pophae L., including the new taxa recognized by A. Rousi.
Ann. Bot. Fennici 8: 228-236.
Swenson, U., Bartish, LV. 2002. Taxonomic synopsis of
APG (Angiosperm Phylogeny Group) 1998. See general Hippophae (Elaeagnaceae). Nord. J. Bot. 22: 369-374.
references. Takhtajan, A. 1997. See general references.
Arohonka, T., Rousi,A.1980. Karyotypes and C-bands in Shep- Thorne, R.F. 1992. Classification and geography of flowering
herdia and Elaeagnus. Ann. Bot. Fennici 17: 258-263. plants. Bot. Rev. 58: 225-348.
Bartish, LV., Jeppsson, N., Nybom, H., Swenson, U. 2002. Phy- Thulin, M., Bremer, B., Richardson, J., Niklasson, J., Fay, M.F,
logeny of Hippophae (Elaeagnaceae) inferred from parsi- and Chase, M. W. 1998. Family relationships of the enigmatic
mony analysis of chloroplast DNA and morphology. Syst. rosid genera Barbeya and Dirachma from the Horn of
Bot. 27: 41-54. Africa region. Pl. Syst. Evol. 213: 103-119.
Bosco M., Fernandez, M.P., Simonet P., Materassi R., Normand, Vanstraten J., Akkermans, A.D.L., Roelofsen, W. 1977. Nitroge-
P. 1992. Evidence that some Frankia sp. strains are able to nase activity of endophyte suspensions derived from root-
cross boundaries between Alnus and Elaeagnus host speci- nodules of Alnus, Hippophae, Shepherdia and Myrica spp.
ficity groups. Appl. Env. Microbiol. 58: 1569-1576. Nature 266: 257-258.
Cronquist, A. 1988. The evolution and classification of flower- Veldkamp, J.F. 1986. Elaeagnaceae. In: Flora Malesiana I, 10:
ing plants, 2nd edn. New York: New York Bot. Garden. 151-156.
Erdtman, G. 1952. See general references. Xu, M. 1994. The medical research and exploitation of sea
Gardner, LC. 1958. Nitrogen fixation in Elaeagnus root buckthorn. Hippophae 7: 32-34.
nodules. Nature 181: 717-718.
Elaeocarpaceae 135
Elaeocarpaceae
M.J.E. COODE
Elaeocarpaceae Juss. ex DC., Essai Propr. Med. Pl., ed. 2: 87 A family comprising 12 genera and c. 550
{1816), 'Elaeocarpeae', nom. cons. species, widely distributed in tropica~ an~ war~
Tremandraceae R. Br. ex DC. (1824), nom. cons.
temperate southern regions but lackmg m conti-
nental Africa.
Trees to suffrutescent, sometimes ericoid; some-
times buttressed; bark without exudate; indumen-
VEGETATIVE MoRPHOLOGY. The family has some
tum of simple hairs, sometimes gland-tipped or
suffrutescent subshrubs, sometimes ericoid, and a
stellate. Leaves spirally arranged, distichous, oppo-
few shrubs; most species are small to large trees.
site or rarely whorled, simple (rarely pinnatis.ect
There are occasional reports of epiphytes or
or pinnate in juveniles), entire to ser~ate, venatwn
scrambling shrubs (Elaeocarpus bilobatus). 1'._1any
usually pinnate, ± trinerved at bas~ m Vallea and
can have buttresses or prop-roots, sometimes
in some Aristotelia and Sloanea; stipules present,
(Sloanea and Elaeocarpus) very large. Terminalia-
or 0 when colleters may occur. Flowers regular,
branching ('modeled' Aubreville': Halle :t al.l97~)
bisexual (unisexual in some Aristotelia and
occurs sporadically in Sloanea and particularly m
Elaeocarpus), solitary and axillary, in fascicles or
Elaeocarpus.
in terminal or axillary simple or compound
inflorescences, either cymose or racemose; sepals
VEGETATIVE ANATOMY. The reader is referred
usually free, valvate, rarely imbricate at tip; petals
to Metcalfe and Chalk (1950), Carlquist (1977),
present (0 in some Sloanea), free, valvate at mser-
and Gasson (1996). The leaf epidermis is often
tion and usually (induplicate-)valvate further up
mucilaginous; the indumentum consists of unicel-
but sometimes overlapping at apex (in some
lular and glandular hairs; stellate hairs are found
Sloanea variously fused even into a tube), usually
in the tremandraceous genera. Stomata are para-
expanded and variously toothed or lobed, ~orne
cytic or encyclocytic. Crystals are mostly. solitary.
times entire and sepaloid; disk a toothed nng or
Cork arises superficially. Nodes are tnlacunar.
separate lobes or pulvinate or flat-topped, or. 0;
Sieve element plastids are of the Ss type. In the
stamens 4-300, free, inserted on or above the disk
wood, growth rings are usu~lly discerna~le;
or around free disk-lobes, filaments shorter to
vessels are often in radial multiples. PerforatiOn
longer than the anthers, anthers basifixed,
plates are mainly simple b.ut sometimes. are
tetrasporangiate, opening at apex by 1-2 po~es
accompanied by some scalanform perforatw?s.
(sometimes extended into a tube) or short shts
Vessel ray pitting has reduced borders and vanes
(sometimes extending to base), connective some-
from circular to elongated (vertical, oblique, or
times extended ('awned'); ovary superior, syncar-
scalariform). Fibre-tracheids are absent, but libri-
pous (internal fusion sometimes incomplete at
form fibres are generally present; they are some-
apex); loculi 2-8(9); ovules 1-30/loculus, pen~u
times septate. Axial parenchyma is sparse and
lous, anatropous, attached to axis in 1-2 sene.s;
tends to be marginal. Rays are heterocellular,
style simple or apically branched, rarely stylo~Ia
mixed uniseriate and pluriseriate, the latter up to
free. Fruit indehiscent (a drupe or berry) or dehis-
7 cells wide, and varying strongly in height. Their
cent (walls thin, horny or woody, initially fleshy in
margins are composed of several ~ows of. upright
Vallea), splitting loculicidally or loculicidally and
cells, and uniseriate rays consist entirely of
septicidally, sometimes spiny or bristly (Sloanea
uprights. Prismatic crystals are found in ~1 genera
p.p.). Seeds 1-many, up to c. IS/loculus, variously
except Aristotelia and Vallea, located m cham-
'arillate' or with watery or fleshy sarcotesta or
bered or non-chambered ray cells or chambered
seed coat dry when sometimes with fine or rare!y
axial parenchyma cells.
stout (Platytheca) hairs; embryo embedded m
endosperm, usually straight with broad (some-
INFLORESCENCE STRUCTURE. Inflorescences are
times narrow) cotyledons.
terminal or usually axillary panicles, cymes,
136 M.J.E. Coode
thyrses or racemes, but are also solitary flowers or PoLLINATION. Very few observations have been
fascicles (all can be found in Sloanea). The most made on pollination of Elaeocarpaceae. The red-
floriferous are probably the panicles of Elaeocar- flowered Crinodendron hookerianum has all the
pus stipularis var. brevipes with up to 90 flowers. features of a bird-pollinated flower and seems to be
Bracts are usually early caducous; a few species of pollinated by a humming-bird (Bricker 1992). The
Elaeocarpus have prophylls on the pedicels as well. rest appear to be entomophilous; thrips have been
found in flowers. It is currently assumed that the
FLOWER STRUCTURE. Flowers are regular, mostly majority at least are adapted to 'buzz-pollination'
bisexual (some dioecious species in Aristotelia and (Matthews and Endress 2002) which tends to be
monoecious species in Elaeocarpus). The calyx associated with pendant flowers, short stamen
is nearly always of free valvate sepals, but is fused filaments, basifixed anthers withapical dehiscence
in Crinodendron and occasionally ± imbricate in and punctiform stigmas; the assumption is also
Sloanea. Petals vary greatly; absent in two made that the flowers are generally not nectarifer-
Australian and all but one New World species of ous, which is not supported by field experience
Sloanea. Disk usually present, variable, absent in (M.J.E. Coode) that flowers of most species of
Tetratheca and Platytheca. Stamens are sometimes Elaeocarpus at least are scented and some obvi-
twice the petals ('Tremandraceae'), more usually ously contain nectar.
not an exact number, from 4 in Aristotelia fruti-
cosa of New Zealand and 5 in Elaeocarpus bilong- REPRODUCTIVE SYSTEMS. Dioecy is known in the
vinas of New Guinea to 300 in some Sloanea; New Zealand species of Aristotelia, and perhaps
filaments various; anthers basifixed and opening also in A. chilensis of South America. Unisexual
initially by 1-2 apical pores or short slits, some- flowers are known in some species of the
times the connective extended into an awn Acronodia and Polystachyus groups of Elaeocarpus
(Sloanea p.p., Elaeocarpus p.p.) or tube. The ovary (W and C Malesia, Borneo: Coode 1996a, 1996b)
is superior, carpels 2-9-fused, style usually single, and in Madagascar (Tirel 1985). Tirel reports
although sometimes there are style branches as that some individuals have bisexual flowers while
many as carpels and even ± free stylodia (Aris- others have male flowers only. Corner (1939: 309)
totelia p.p., Sloanea p.p.); stylodia in Peripentade- said of the 'subgenus Acronodia' that there were
nia mearsii are free at base. For floral anatomy, see 'male and monoecious individuals' while in the
Payer (1857), van Heel (1966) and Matthews and Polystachyus group there are species with separate
Endress (2002). male and female inflorescences, but whether
always on separate trees is not established yet.
EMBRYOLOGY. Pollen grains are 2-celled when Otherwise, the flowers are bisexual, but see Coode
shed. The ovule is anatropous, bitegmic and crass- (1985) for possible unisexual flowers in Vallea. In
inucellate with a zigzag micropyle formed by both some species of Elaeocarpus, fruits which are
integuments. An endothelium and hypostase have apparently perfect may contain no seeds. Whether
been found in an Elaeocarpus species. The embryo there are barriers to self-pollination seems to be
sac is of the Polygonum type, endosperm forma- unknown.
tion is nuclear (Johri et al. 1992).
FRUIT AND SEED. There is wide variation in fruit
PoLLEN MORPHOLOGY. Pollen grains are unusu- type and consequently seed type. Of those with
ally small, prolate to suboblate (longest axis dehiscent fruits, the three 'tremandraceous' genera
8-25J.Lm) and tricolopor( oid)ate, sometimes syn- have small fruits with thin, flimsy walls; Crin-
colpate; the exine is often distinctly columellate; odendron, Peripentadenia and Dubouzetia have
the sexine is crassitectate and the tectum is much larger fruits with tough, horny walls often
obscurely to faintly psilate, rugulate or finely retic- separating into two layers and splitting loculici-
ulate and microperforate (Erdtman 1952; Sharma dally at first, sometimes septicidally subsequently
1969; Barth and Barbosa 1973; Lobreau-Callen in - the resultant open fruits can resemble fruits of
Tirel1985; Tang and Wu 1990). Euphorbiaceae at a similar stage; fruit of Sloanea
have woody walls generally much thicker than
KARYOLOGY. Several Aristotelia have been those of the three preceding genera and never sep-
counted with 2n = 28, two Elaeocarpus with 2n = arating into layers (Fig. 47E), whereas Vallea fruits
24 and 30 respectively (Fedorov 1969); Crinoden- open while the walls are still fleshy. The four
dron hookerianum has n = 21, and Dubouzetia genera with indehiscent fruits are few-seeded
elegans n = ca. 90. and have either berries (Sericolea, Aristotelia) or
Elaeocarpaceae 137
drupes (Elaeocarpus, Fig. 48E; Aceratium). Seeds from Aristotelia species, which are tryptamin-
may have dry coats or sarcotestas (fleshy in monoterpene derived. Elaeocarpus and Peripen-
Sloanea, thinner and ± watery in Crinodendron, tadenia species have furnished alkaloids of the
Sericolea and some Aristotelia, thick and spongy elaeocarpidin group, which are derived from
in Dubouzetia galorei). In Aristotelia, Dubouzetia, tryptamin or 1,3 diaminopropan or putrescin and
Tetratheca and Tremandra there may be ± waxy, a polyketid moiety. A bitter principle, elaeocarpid,
apical outgrowths (strophioles) of the chalaza, has been isolated from Elaeocarpus species.
often spirally coiled. 'Arils' (formed from the Halphen-positive seed oils and large amounts of
chalaza or raphe or even seed-coat or all of them) mucilage, both characteristic of Malvales families,
may partly or wholly cover seeds in some Sloanea have not been found in Elaeocarpaceae (Hegnauer
and Peripentadenia, while in Vallea there is a 1973, 1990).
lop-sided development from the raphe. Seeds of
Sericolea are usually curved and in Elaeocarpus a SUBDIVISION AND RELATIONSHIPS. Previous
range of shapes (Weibel 1968; Coode 1984), from attempts at subdividing the family have been only
straight through curved to hooked, is useful in partly successful (e.g. Schlechter 1916). However,
defining infrageneric groupings. Testa and tegmen it does seem possible to group the genera as fol-
are multiplicative; the exotesta is thick-walled, and lows: Aristotelia+ Vallea; Sloanea; Crinodendron+
in Sloanea and possibly in Aristotelia as well the Peripentadenia+Dubouzetia; Elaeocarpus+Acerat-
tegmen has many outer layers of fibres which, ium+Sericolea. Sloanea is isolated; it would seem
according to Corner (1976), are relictual and deny to be near the base if not basal. The three Tre-
a close affinity with Malvales. mandraceous genera ( Tremandra, Tetratheca and
Platytheca) have been found, on molecular evi-
DISPERSAL. Drupes of some Elaeocarpus and dence, to be nested in Elaeocarpaceae (Savolainen,
Aceratium are known to be dispersed by birds, Fay et al. 2000; Bradford and Barnes 2001). That
such as fruit-pigeons and cassowaries (Crome morphology which appears comparable (especially
1975a, 1975b). Elaeocarpus fruits are often blue; the 'anthers' opening by a single pore) supports this
this is due not to pigment but to diffraction (Lee placement, but in other respects it is difficult to
1991). Aceratium fruits are usually red. Presum- evaluate their true position. Thompson (1976)
ably the other indehiscent (fleshy) fruits are also provides a table of distinctions, and it is clear that
bird-dispersed. The colour of seeds (whether dry while Platytheca and Tremandra are very distinct,
seed coat, sarcotestas or arils) contrasts with the Tetratheca is less easy to define.
colour of the loculus interior in many species of
Sloanea and would suggest attraction to birds AFFINITIES. Elaeocarpaceae were generally
such as hornbills; the role of broad-based or bristle thought to be part of, or close to, Tiliaceae and
spines on the fruit exterior is not documented. placed in Malvales; more recently other positions
Strophioles (elaiosomes) are attractive to ants; were proposed (Dahlgren 1988; Takhtajan 1997;
Tetratheca is discussed in Berg (1976) in this Thorne 2000). Muntingia was often included and,
respect; whether the strophioles of the tree- more understandably, Petenaea (both currently
forming genera are also indicative of ant-dispersal treated in Malvales, see vol. V of this series).
is not known. Clifford and Drake (1985) report on Recently, Elaeocarpaceae have been placed on
finding endocarps in kangaroo dung. The seeds of molecular evidence in Oxalidales. The family,
Dubouzetia galorei float and are bright red; within as enlarged recently by the inclusion of Treman-
the spongy outer layers is a dense, hard sclerotesta draceae also on molecular grounds, is difficult to
which sinks. Cassowaries are known to feed on the delimit on visible characteristics. Surveys with less
fallen seeds of this species (label of Balgooy 6596 patchy sampling are desirable to confirm the con-
from Aru), but perhaps fish also play some part clusions already being widely followed.
along the flood-plains of southern New Guinea.
DISTRIBUTION AND HABITATS. Elaeocarpaceae
PHYTOCHEMISTRY. Both condensed and hydroly- are an essentially southern Gondwana family,
sable tannins are present, particularly in the bark, of which only Elaeocarpus and Sloanea reach the
the former based on procyanidin, the latter northern hemisphere in any numbers; they are not
containing gallic acid, ellagic acid and methylel- found in continental Africa. Sloanea is found in
lagic acid. Common triterpenes and saponins Central and South America, Madagascar, NE India
also occur; one Sloanea is known to be cyano- and Nepal through to Sand C China, throughout
genic. Many different alkaloids have been isolated Malesia to the Solomon Islands, New Caledonia
138 M.J.E. Coode
and N and E Australia; it is the only genus of the that are strung together to make malas or rosaries,
family to be reported from the London Clay, as important in eastern religious practice. Aceratium
seeds and abraded fruits. Elaeocarpus has a wider oppositifolium fruits are edible. A few species of
distribution in the Old World but is missing from Elaeocarpus have edible mesocarp and are some-
the New: Madagascar, Mauritius, Ceylon, E India times pickled (E. robustus) in SE Asia. The seeds
through to C China and S Japan, Hawai'i, through- of Elaeocarpus appear to be edible but only worth
out Malesia and the western Pacific, Australia and extracting in large-fruited species (e.g. E. schlech-
New Zealand; fossils are known from Australia. terianus). Some large Australian Sloanea and
The rest have a more limited distribution. Valle a is Elaeocarpus yield useful timber. Species of Crin-
known from northern South America to Peru and odendron, Vallea and Aristotelia are grown in
Bolivia; Aristotelia from Chile and Argentina, New north-temperate horticulture, while species of
Zealand, Tasmania and Australia. Crinodendron is other genera are found in tropical gardens. Several
known from the southern part of South America, local uses for Sloanea in the New World are listed
Peripentadenia is endemic to NE Queensland, by Earle Smith {1954).
while Dubouzetia is known from the Moluccas,
New Guinea, New Caledonia and Australia (North- CoNSERVATION. Many species of most, if not all
ern Territory and NE Queensland). Tremandra genera probably need listing as under threat of
and Platytheca are endemic to SW West Australia; some sort, e.g. the 1-2 individuals of Elaeocarpus
Tetratheca extends from there across the southern bojeri and hardly more numerous E. integrifolius,
part of the continent to E Australia. Aceratium is both of Mauritius (Coode 1987b; see also 1997
known from the Moluccas through New Guinea IUCN Red List of Threatened Plants, Walter and
and the Bismarck Archipelago, Solomon Islands, Gilletts 1998: pp. 242 and 584). Many well-
Vanuatu with different species from NE Queens- characterised species are known from the type
land. Sericolea is endemic to highland New collection only.
Guinea, once found on Fergusson Island. Mostly
they are species of high, or disturbed, forest; at KEY TO THE GENERA
mid-altitudes especially, forests can be found with
1. Suffrutescent; indumentum with some gland-tipped hairs,
several species co-existing, with large numbers of or with stellate hairs; stipules absent, sometimes replaced
individuals. Some reach altitudes of 3000 m or by colleters; ovary flattened or narrowed in flower, ovules
more, in Malesia at least (Sericolea calophylla up in a single row or 1; fruits dehiscent, fragile, ± flattened,
to 3500 m), where they tend to be shrubby. Some less than 1.5 em 2
are limited to seral habitats while a few are - Shrubs to trees; indumentum of simple non-glandular
hairs only; stipules usually present though sometimes
pioneers, e.g. Aristotelia serrata and Elaeocarpus early caducous; ovary terete in flower, rarely somewhat
murukkai, the latter forming even-age stands with angular, ovules in pairs or biseriate; fruits dehiscent with
a species of Macaranga and virtually nothing else tough walls, mostly exceeding 1.5 em, or indehiscent 4
on landslips at mid-latitude in Papua New Guinea. 2. Indumentum mostly of stellate hairs; disk of minute, free
lobes distant from each other; stamens with filaments
Aristotelia chilensis, after having been introduced about as long as anthers 10. Tremandra
to the Island of Juan Fernandez off the Chilean - Indumentum of simple (some gland-tipped) hairs, none
coast, has become a threat to the endemic flora at stellate; disk absent; stamens with filaments shorter than
lower and middle elevations. The two Australian anthers 3
Dubouzetia are shrubby species which inhabit 3. Petals with a central vein stronger than the rest and with
pinnate side-branches; stamens with the 4 pollen sacs
rocky gorges at low altitudes. The three 'treman- arranged in 1 series; fruit septum present; seed surface
draceous' genera are found in open 'bushland' with thick hairs, seeds without appendages
vegetation. 11. Platytheca
- Petals with net-venation; stamens with the four pollen sacs
PALAEOBOTANY. Rozefelds {1990) and Rozefelds in 2 series; fruit septum 0; seed surface glabrous or with
fine hairs, seeds with an appendage at apex
and Christophel ( 1996a, 1996b) report on en do- 12. Tetratheca
carps of Elaeocarpus from the Oligo/Miocene in 4. Stamens dehiscing at least initially by 2 separate apical
Australian deposits. Sloanea is recorded from the pores 5
London Clay; recent material in better condition - Stamens dehiscing at least initially by 1 apical pore or
slit 7
would be welcome.
5. Ovules 8-16/loculus (seeds usually many); petals variously
much-divided, or entire and sepaloid, or entire and
ECONOMIC IMPORTANCE. E[aeocarpus angusti- expanded at apex, or absent 3. Sloanea
folius (syn. E. sphaericus or Ganitrus sphaerica) is 5. Ovules 2/loculus (seeds few); petals 3-lobed, not further
the source of endocarps used as beads (rudraksha) divided, or merely 3-notched 6
Elaeocarpaceae 139
6. Leaves opposite or in whorls of 3; stipules not foliaceous; 1. ARISTOTELIA ALLIANCE (see Coode 1985).
stamen filaments terete; fruits indehiscent, seeds with or
without a short terminal appendage and sometimes with
a thin sarcotesta 2. Aristotelia Stipules foliaceous or reduced; petals free,
- Leaves spirally arranged; stipules when present foliaceous; expanded; stamen filaments ± equalling anthers;
stamen filaments flattened, broad-based and contiguous; anthers initially dehiscing by 2 apical pores, often
fruits dehiscent, initially ± fleshy, seeds with an obvious splitting to base; ovules 2/loculus, collateral but
asymmetric outgrowth of raphe 1. Vallea overlapping; styles branched or not; fruit septum
7. Fruits indehiscent 8
- Fruits dehiscent 10 0; seeds 1/loculus at most, glabrous. Two genera,
8. Leaves alternate to spirally arranged or very rarely oppo- New World and southern hemisphere.
site; fruit a drupe with± woody endocarp usually separa-
ble from mesocarp (rarely with persistent radial fibres)
9. Elaeocarpus 1. Vallea L. Fil.
- Leaves opposite, sometimes alternate in young shoots; fruit
a drupe with weakly woody'core' and fibrous mesocarp, or Vallea Mutis ex L. Fil., Suppl. Pl. Syst. Veg.: 42 (1782).
a berry 9
9. Petals variously much-divided; stamens with filaments Shrubs; leaves with ternate basal venation; stipules
longer than anthers and setae arranged around the apical foliaceous, often early caducous; inflorescence
pore or slit of anther; ovules 6-14/loculus; fruit a drupe,
seed surface often with fine hairs, embryos straight
cymose, rarely flowers solitary; petals imbricate at
8. Aceratium tip, 3-lobed; disk mostly replaced by spread ovary-
- Petals merely notched or ± entire; stamens with filaments base; stamens 15-60, inserted in a ± single ring;
as long as anthers or less, no setae at anther apex; ovules filaments, sigmoid, bases broad, contiguous; ovary
2/loculus, seeds glabrous with thin sarcotesta, embryos 3-5-locular; styles branched at apex; fruit fleshy at
curved 7. Sericolea
10. Sepals fused into a cup, finally splitting along 1-2 lines; first, loculicidally part -dehiscing, valves persistent;
seeds without appendages but sometimes with a watery or seeds with a lopsided outgrowth of raphe, testa
thin sarcotesta 4. Crinodendron dry. 1-2 species from Colombia and Venezuela to
- Sepals free; seeds with strophiole (sometimes spiral) at Peru and Bolivia, in the lower slopes of the Andes.
apex, or with complete arillode (an outgrowth of chalaza)
or with thick spongy sarcotesta 11
11. Disk of large free lobes, the stamens inserted around the 2. Aristotelia L'Her.
lobes; petals without pockets at base; stamens with fila-
ments as long as anthers or less, apical pores extending to Aristotelia L'Her., Stirp. Nov. 31 t. 16 (1786) nom. cons.
lateral slits; seeds with complete arillode, an outgrowth of Friesia DC.
chalaza 6. Peripentadenia
- Disk pulviniform to annular, stamens inserted mostly
between disk and ovary base; petals with pockets at base
Shrubs to medium trees; leaves opposite, rarely in
corresponding with disk-lobes; stamens with filaments 3s or alternate, juveniles sometimes pinnatisect,
longer than anthers, apical pores not extending laterally basal venation sometimes ternate; stipules
into slits; seeds with strophiole at apex or with thick sar- reduced (to 0?); inflorescence a panicule or thyrse,
cotesta 5. Dubouzetia rarely a 3-flowered cyme or flowers ± solitary;
flowers bisexual or plants dioecious; petals imbri-
cate at tip, 3-lobed, -notched or ± entire; disk
I. CORE ELAEOCARPS (genera 1-9) cushion-shaped to annular, rarely minutely lobed;
stamens 4-15; ovary 2-4-locular; styles notched at
Shrubs to trees; hairs simple, not gland-tipped; apex or with as many branches as loculi, some-
leaves spirally arranged or opposite, simple when times to base; fruit a berry; seeds sometimes with
mature, venation usually pinnate; pedicels contin- a thin sarcotesta, with or without a short terminal
uous with calyx; sepals usually free; petal veins, appendage. 5 species from Chile, New Zealand and
when visible, netted, petals with or without basal Australia (Tasmania, New South Wales).
pockets; stamen filaments usually terete; anthers
with 2 bilobed contiguous thecae; ovary terete to
angular; ovules hi-seriate; fruits indehiscent or 2. SLOANEA ALLIANCE
dehiscent with tough walls; septum 0; seeds
usually glabrous; embryo usually straight with 3. Sloanea L. Fig. 47
broad cotyledons.
Sloanea L., Sp. Pl. :512 (1753); Earle Smith, Contrib. Gray Herb.
Harv. Univ. 175: 1-114 (1954), New World spp.; Coode, Kew
Bull. 38: 347-427 (1983), conspectus of Old World spp.
Echinocarpus Blume
Antholoma Labill.
140 M.J.E. Coode
seeds 1-many, glabrous, with a coloured sarcotesta

or variable cover of hard seed-coat by often
coloured arillode. About 100 species from C &
tropical South America, and over 50 from the Old
World including Madagascar (3), NE India & Nepal
(4), Burma (7), China (12-15), Vietnam (6), Borneo
(2), New Guinea (18), New Caledonia (9), Australia
(4).
3. CRINODENDRON ALLIANCE (see Coode 1987).
Stipules minute, often early caducous; petals free,

expanded; stamen filaments straight to weakly
sinuous, at least equalling anthers; anthers without
conne_ctive ex.tension and apical setae, dehiscing
by a smgle ap1cal pore sometimes developing into
slits; ovules biseriate; styles unbranched above;
fruit without spines, walls horny often of 2 layers,
dehiscent loculicidally at first, sometimes also
finally septicidally; seeds few to several per fruit,
1-4 per loculus, usually glabrous. Three genera,
southern hemisphere.
4. Crinodendron Molina
Fig. 47. Elaeocarpaceae. Sloanea rhodantha. A Flowering
branchlet. B Flower, vertical section. C Stamens, inside and Crinodendron Molina, Sagg. Chi!. ed. 1: 179 (1782); Bricker,
outside view. D Ovary, cross section. E Dehisced fruit. F Syst. Bot. 16: 77-88 (1991), rev.
Arill~te seed. G s:ed, cross section, with adhering fringes Tricuspidaria Ruiz & Pavon
of ani. H Seed, ani removed, the area of adherence of aril
cross-hatched. I A ril, expanded. J Embryo. (Capuron around
1965)
Shrubs to medium trees; flowers solitary with
articulated pedicels; sepals fused at first, later
splitting along 1-2 lines; petals 3-lobed to 3( -5)-
toothed, 2-pocketed at base; disk tall, resembling
Small to large trees, often buttressed; occasionally a cogwheel; stamens 13-18, inserted between disk
shrubby; leaves sometimes subopposite, juveniles and ovary; ovary 3-5-locular; ovules 4-15/loculus;
sometimes pinnate, basal venation sometimes styles unbranched; fruit-valves not detaching, with
ternate; stipules linear to foliaceous, often columella sometimes persistent; seeds glabrous
caducous, rarely 0?; inflorescence cymose, thyr- with complete watery sarcotesta and without
soid, a fascicle or raceme, or flowers solitary; appendage. About 5 species from S South America;
sepals sometimes imbricate at tip; petals expanded one very distinct with red, bird-pollinated flowers
and divided or toothed (rarely entire) and im- (C. hookerianum).
bricate at tip or fused into a partial or complete
corona or sepaloid and acute or 0 (most New 5. Dubouzetia Brongn. & Gris
World, some Australia}; disk flat-topped to pul-
Dubouzetia Pancher ex Brongn. & Gris, Bull. Soc. Bot. Fr. 8: 199
viniform; stamens (8- }20- 300; filaments shorter
(1861).
than to exceeding anthers, sometimes ± 0; anthers
often awned, apical setae 0, dehiscing by 2 apical
Inflorescence short-racemose to fasciculate, rarely
pores to short apical slits sometimes extending flowers solitary with articulated pedicels; petals
down the sides; ovary 2-5-locular; ovules
rounded, ± entire to minutely serrulate, 2-pock-
8-30/loculus in 2 rows; styles unbranched or with
eted at base; disk pulviniform to slightly 5-lobed;
as many branches as loculi, sometimes to base;
stamens 15-45, inserted on disk surface and/or at
fruit woody, loculicidally dehiscent, valves gener-
ovary base; filaments longer than anthers; ovary
ally massive, persistent (often brightly coloured
3-5-locular; ovules 4-12/loculus; fruit valves not
inside), often with spines (broad based to bristly;
Elaeocarpaceae 141
detaching; seeds few to several, 1-3 per loculus, apical confluent slit; ovary 2-locular; ovules
glabrous or with fine spreading hairs, seed-coat 2/loculus; styles unbranched but notched at tip;
hard with an apical spiral waxy strophiole but D. fruit a berry; seeds 1-4 per fruit, glabrous, with
galorei with thick, spongy sarcotesta surrounding thin complete sarcotesta (not obvious when
a small sclerotesta. 6 species from New Caledonia, dried), without appendages. Embryo weakly
of which 1 is also in Moluccas to New Guinea, 3 curved; cotyledons fairly broad. About 16 species
more from New Guinea and 2 endemic to small from submontane to subalpine New Guinea.
areas of northern Australia.
8. Aceratium DC.
6. Peripentadenia L.S. Sm.
Aceratium DC., Prodr. 1: 519 (1824).
Peripentadenia L.S. Sm., Pro c. Roy. Soc. Queens!. 68:45 (1957).
Shrubs to small trees; leaves opposite, sometimes
Juvenile leaves sometimes pinnatisect; flowers alternate on young shoots; stipules minute or ?0;
grouped to fasciculate, perhaps condensed- inflorescence short-racemose to sub-umbellate;
solitary, pedicels not articulated; petals 3-lobed petals variously divided to laciniate, without
with lobes further few-divided, incurved but pockets at base; disk tall, resembling a cogwheel;
without pockets at base; disk with 5 separate stamens 12-20( -30), inserted between disk and
masses; stamens 50-75, inserted around the disk- ovary base; filaments straight in young bud, later
lobes; filaments sometimes exceeding anthers; usually sigmoid-inrolled, longer than anthers;
anthers eventually splitting down sides to base; anthers without connective extension, apical setae
ovary 3-locular; ovules 4-9/loculus; styles fused present and obvious, surrounding the apical pore
and unbranched above (in P. mearsii free at base); or short slit not extending down sides; ovary
fruit with valves detaching, columella persistent; 3-5-locular; ovules (5)6-14/loculus; styles un-
seeds 1(-2) per fruit, glabrous, with a hard seed- branched; fruit± drupoid, the meso carp with radi-
coat with complete aril attached only at chalaza. 2 ating fibres persistent on thin-walled or ± woody
species endemic to NE Queensland. endocarp; seeds 1-2 per fruit, glabrous or with
fine spreading hairs, lacking sarcotesta or other
appendage. About 20 species, 5 endemic to NE
4. ELAEOCARPUS ALLIANCE Queensland, the rest from Moluccas to New
Guinea, Solomon Islands and Vanuatu.
Stipules minute to foliaceous, often early
caducous; petals free, usually expanded; stamen
9. Elaeocarpus L. Fig. 48
filaments various; anthers dehiscing by a single
apical pore or slit, with or without connective Elaeocarpus L., Sp. Pl.: 515 (1753).
extension and/or apical setae; ovules biseriate; Many synonyms, listed in e.g. Tirel1985: 5.
styles unbranched above; fruit a berry or drupe;
seeds few to several per fruit, 1-2 per loculus, Shrubs to large trees; leaves spirally arranged to
usually glabrous. Three genera, (sub)tropical Asia distichous, very rarely opposite (Mauritius, New
and Pacific Islands to Australia and Madagascar. Guinea); stipules minute to large, linear to folia-
ceous, sometimes early caducous; inflorescence
a raceme, rarely short-racemose or paniculate;
7. Sericolea Schltr. flowers usually bisexual, sometimes unisexual (on
Sericolea Schltr., Bot. Jahrb. 54: 95 (1916); van Balgooy, Blumea monoecious plants?); petals expanded or sepaloid,
28: 103-141 (1982), rev. sometimes small, variously lobed and/or laciniate,
or entire and ± sepaloid, rarely entire and
Shrubs to small trees; leaves opposite; stipules expanded, with or without pockets at base; disk
linear, minute; inflorescence paniculate or race- variously tall or low, sometimes resembling a cog-
mose, branches and flowers opposite; petals small, wheel, or lobed, sometimes merely annular with
minutely lobed or notched, without pockets at minute teeth, rarely pulviniform; stamens 5-200,
base; disk weakly to distinctly lobed; stamens inserted on disk surface and/or at ovary base; fil-
(8-)10-15(-18), inserted between disk and ovary aments straight to sigmoid, shorter than, equalling
base; filaments straight to sinuous, shorter than or exceeding anthers, sometimes± 0; anther con-
to equalling anthers; anthers without connective nective extension often present as awn or 0, apical
extension or apical setae, dehiscing by a single setae sometimes present (but never surrounding
142 M.J.E. Coode
II. TREMANDRACEOUS GENERA (genera 10-12;

Thompson, Telopea 1: 139-215 (1976), and pers.
comm. March 2003.
Suffruticose to sometimes ericoid shrublets; hairs

simple (with some gland-tipped) or stellate; leaves
usually opposite to verticillate, simple, where
visible often penninerved; stipules absent or
replaced by colleters; flowers solitary; pedicels
discontinuous with calyx; sepals free, valvate;
petals free, expanded or linguiform, without basal
pockets, veins netted or pinnate; anthers with 4
thecae, without connective extension and apical
setae, dehiscing by a single apical pore; disk
usually 0; ovary somewhat to clearly flattened or
narrowed, 2-locular (rarely 3); ovules 1-2, rarely
more, uniseriate or single on a central axis; styles
unbranched; fruits somewhat to clearly com-
pressed, dehiscent, walls flimsy, septum present
or 0.
10. Tremandra R. Br. Fig. 49

Tremandra R. Br. in Flinders, Voy. Terra Austral. 2: app. 3: 544
(1814).
Subshrubs rarely to 2m; hairs mostly stellate

(simple on anthers and petals); leaves opposite,
simple, sometimes± lobed and/or± trinerved at
base; flowers scattered; petals entire and rounded,
Fig. 48. Elaeocarpaceae. Elaeocarpus sericoloides. A Flowering expanded,± net-veined; disk with small distinct
branchlets. B Flower, longitudinal section. C Pistil and disk, lobes; stamens 10, inserted at ovary base; alternate
some stamens removed. D Stamens. E Fruit. F Fruit stone.
Drawn by Lucy T. Smith. (Coode 2002)
stamens with thickened disk-like structure, fila-
ments ± straight, about equalling anthers, thecae
in 2 series; ovary flattened; fruit regularly dehis-
cent, valves detaching, septum 0; seeds 1-2 per
apical pore), dehiscing by a single apical confluent fruit, with fine slender-spreading to ± adpressed
slit often extending down sides sometimes to base; hairs, seedcoat hard with apical appendage. 2
ovary 2-9-locular; ovules 2-16/loculus; styles species endemic to SW West Australia.
unbranched; fruit a drupe with woody often sculp-
tured endocarp usually separable from the meso- 11. Tetratheca Sm.
carp, sometimes with ± persistent fibres as in
Aceratium; seeds 1-5(-7)/fruit, at most 1/loculus, Tetratheca Sm., Specimen. Bot. New Holland: 5, t. 2 (1793);
Thompson, Telopea 1: 139-215 (1976), rev.
glabrous and without sarcotesta or appendages.
Embryo straight with broad cotyledons to curved
with broad or narrow cotyledons. The largest Subshrubs, sometimes ericoid or trailing, usually
genus, with about 350 species altogether: Mada- to 1 m, rarely more; indumentum often mixed of
gascar 8, Ceylon & India c. 30, Thailand c.18, China fine hairs, thick-based bristles and gland-tipped
c. 20, Japan 2, Peninsular Malaysia c. 30, Java c. 15, hairs; colleters sometimes present in Western
Borneo c. 70, Sulawesi c. 40, Philippines c. 50, Australia; leaves alternate to opposite, sometimes
New Guinea c. 85, New Caledonia 29, Australia in verticels, often reduced or caducous; flowers
c. 25, Lord Howe Is. 1, New Zealand 2, Fiji 22, scattered or sometimes grouped; petals free, ligu-
Hawai'i 1. late to expanded, entire; disk 0; stamens 8-10,
inserted at ovary base, sometimes fused at base to
varying extents into pairs, filaments straight but
Elaeocarpaceae 143
often much expanded, shorter than anthers, thecae Fig. 49. Elaeocarpaceae. Tremandra stelligera. A Flowering
branch. B Stellate hairs. C Flower. D Outer and inner stamen.
in 2 series, apical pore usually extended into a
E Anther, inside view. F Pistil. G Ovary, vertical and transverse
tube, ovary often aberrantly 3-locular; ovules section. Drawn by B. Johnsen. (Thorne 1983)
1-2/loculus (1 sp. with 8-10 ovules), rarely more;
fruit dehiscent loculicidally (occasionally later
along septa), valves persistent, septum 0; seeds 1-4
per fruit, so 1-2 per loculus, usually with fine Selected Bibliography
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Ericaceae 145
Ericaceae
P.F. STEVENS
with J. LUTEYN (most Vaccinieae); E.G.H. OLIVER (Erica); T.L. BELL, E.A. BROWN, R.K. CROWDEN,
A.S. GEORGE, G.J. JoRDAN, P. LADD, K. LEMSON, C.B. McLEAN, Y. MENADUE, J.S. PATE, H.M. STACE,
C.M. WEILLER (Styphelioideae)
Ericaceae Juss., Gen. Pl.: 159 (1789) ("Ericae"), nom. cons. A mainly temperate, warm temperate and
Epacridaceae R. Brown (1810). montane tropical family of some 124 genera and
Monotropaceae Nuttall (1818).
Empetraceae S.F. Gray (1821). 4100 species.
Vacciniaceae DC. ex Gray (1821).
Pyrolaceae Dumort. (1829). CHARACTERS oF RARE OccuRRENCE. Cushion
plants in some species of Dracophyllum, and in
Evergreen or deciduous shrubs, rarely scandent, Disterigma empetrifolium and Rhododendron
lianes, or trees, epiphytic or not, or herbs, rarely saxifragoides. Leaves distichous in Vaccinium dis-
achlorophyllous and/or rhizomatous; hair roots tichum and perhaps some species of Disterigma.
present, with investing mycorrhizal fungal hyphae Leaf like the finger of a glove, or peltate, in some
forming a loose covering over hair roots and pen- Cassiope; Calluna with tails from the abaxially
etrating only the outer cortical cells; indumentum recurved portions of the lamina. Linear leaves rel-
unicellular and multicellular hairs, or unicellular atively common in Styphelioideae but otherwise
hairs only, or rarely none; terminal bud scaly, sporadic, e.g. Diplycosia, Rhododendron, Rusbya,
rarely naked, or aborting; leaves spiral, opposite or and Agarista (where they are V-shaped in trans-
whorled, entire or serrate, rarely margins strongly verse section). Leaves 60 em to 1m long in arbores-
revolute and leaves needle-like ("ericoid"), exstip- cent Dracophyllum and Richea pandanifolia.
ulate; inflorescences terminal or axillary, usually Whole inflorescence concolourous in Oligarrhena
racemose; prophylls paired, rarely 0; flowers rarely and some Vaccinieae. Calyx much longer than the
single, rarely multibracteolate, usually conspicu- corolla in a few Erica. Corolla in Vaccinium section
ous, hermaphroditic, rarely unisexual, poly- Polycodium open during development, fused into
symmetrical, rarely monosymmetrical, sepals a cap that falls at anthesis in Richea, scarcely
(2-)4-5( -7), fused at the very base, petals opening in Conostephium and Cosmelia, blue in a
(3)4-5( -7), fused, rarely free or fused as a cap; number of species of Andersonia. Most Styphe-
stamens (2-)5(-8), 10(-16), free from the corolla, lioideae lack glandular hairs, but corolla hairs
rarely adnate; anthers tetrasporangiate, rarely bis- glandular in Melichrus. Corolla tube of Utley a with
porangiate, inverting during development, with solid spurs; that of most Kalmia with pouches in
2( 4) apparently terminal or dorsal appendages or which anthers are held under tension. Antesepa-
not, dehiscence introrse or terminal, rarely latrorse line whorl of stamens in Diplarche adnate to the
or extrorse, with pores or short, rarely long slits or corolla, antepetaline whorl largely free; calyx free,
a slit; endothecium lacking, rarely present; pollen corolla adnate halfway up ovary in Erica serrata.
in tetrahedral tetrads or rarely in monads; nectary Flowers 4-merous, with 2 fertile stamens, in Oli-
present, rarely absent; ovary superior to inferior, garrhena. Filaments reflexed and anthers lying
(1- )4-5(-12)-carpellate, placentation axile to along corolla lobes in Acrotriche; anthers cohering
intruded parietal, rarely apical or basal; ovules at margins in Sprengelia incarnata, which has
(1-)numerous/carpel, anatropous to subcampy- bulbous hairs on the anther, S. monticola and some
lotropous, unitegmic, tenuinucellate; style usually Vaccinieae, connate in Coleanthera; filaments in
about as long as corolla, hollow, rarely expanded particular connate sporadically in Vaccinieae. The
at the apex; stigma punctate to lobed; fruit a berry, mesocarp of Pentachondra dehiscens splits to
drupe, or capsule, rarely calyx fleshy; seeds small reveal the pyrenes inside.
to minute, testa usually single-layered, variously
winged or not; embryo straight, fusiform, rarely VEGETATIVE MORPHOLOGY. Ericaceae are ever-
embryo minute, undifferentiated; endosperm cel- green, sometimes deciduous, often rather xero-
lular, fleshy, well developed, with haustoria at both morphic shrubs, rarely root climbers, Hanes, small
ends; germination epigeal. trees or herbs. Roots developing from the stem
146 P.F. Stevens et al.
come from immediately above axillary buds. surface seems to be obliterated developmentally
Pyroleae are herbs or slightly woody; Pyrola picta (Stevens 1970a). Styphelioideae have xeromor-
may have almost leafless stems (P. aphylla!; Haber phic, flat to recurved leaf blades that are often
1987). Pterosporeae and Monotropeae are herba- pungent at the apex. Variation in leaf type within
ceous and completely lack chlorophyll (Henderson genera such as Kalmia, Agarista, Diplycosia and
1919); shoots develop from roots (as also in Cassiope is high (Stevens 1970a), and variation
Moneses). The young shoots of many tropical Vac- in vegetative features provides much systematic
cinieae are dark red, and the whole plant is red in information; Philipson (1985) summarises varia-
some Monotropeae, the plant drying black. Many tion within Rhododendron.
Ericaceae are rhizomatous, or have lignotubers The leaf margin is usually entire to only slightly
from which they sprout after fire, while some epi- serrate; the serrations are often (but not always in
phytic and epilithic tropical Vaccinieae in particu- Arbutoideae and never in most Styphelioideae)
lar have root swellings. associated with multicellular hairs. Some tropical
Temple (1975) found infraspecific variation in Vaccinieae have large glands at the base of the
architectural models sensu Halle et al. (1978), and margin or on the petiole. Secondary veins are
it is difficult to place many taxa in models (Stevens pinnate, but most tropical Vaccinieae are strongly
1982; Temple in Oliver 1991). In many Vaccin- pH-nerved and with a clear midrib while most
ioideae the apical bud aborts, in many Rhododen- Styphelioideae are pli- or parallel-nerved and lack
dreae it produces an inflorescence, but in Ericeae an obvious midrib. Fine details of the venation
it may remain vegetative. Stems are commonly have been little studied, but taxa like Agarista are
orthotropic, but in Vaccinium section Cyanococcus notable in having all veins almost equally promi-
and in Oxydendrum they are mixed, being nent, so forming a dense, even reticulum (e.g.
orthotropic initially and more or less plagiotropic Lems 1962b; Judd 1984).
by the end of the season. Some tropical Vaccinieae Ericaceae have many hair types, variation within
have orthotropic axes that sprawl under their own Rhododendron and Lyonia being particularly
weight unless supported, or the axes may be pen- notable (see Seithe 1980; Judd 1981); unicellular
dulous. Buds are generally perulate, excepting and multicellular hairs usually occur together.
many Ericeae and a few Rhodoreae; the outer pair Multicellular hairs are eglandular and dendroid to
of perulae is prominent and acicular in several stellate, !epidote, or flattened-setose, or variously
neotropical Vaccinieae (incorrectly called stip- glandular. In glandular hairs secretion may accu-
ules). The relationship of vegetative branches to mulate on the outside of the hair (Gaylussacia),
inflorescences varies; vegetative branches in or within the hair, separating individual cells
Enkianthus and Rhododendron sect. Sciadorho- (Rhododendron). Multicellular hairs, when
dion arise within the terminal bud from axils of present, are likely to be found at least on the leaf
perulae; more normally such branches arise from margin or base of the shoot. Most Styphelioideae
buds in leaf axils. lack multicellular hairs.
The leaves are usually spirally arranged -
pseudoverticillate in Pyroleae and sometimes else- VEGETATIVE ANATOMY. The anatomy of the
where - or whorled to opposite, as in Ericeae; mycorrhizal associations in the root provides
almost all conditions occur even in small genera interesting systematic information (Cullings
like Kalmia and Pieris. Empetreae, particularly 1994). In some Monotropoideae lateral roots arise
Empetrum, are labile, with spiral and whorled exogenously (e.g. Copeland 1935).
leaves on the one plant. Distichy is very rare. The vascular anatomy of the family needs study,
Leaves may be convolute or revolute in bud. general surveys such as those by Cox (1948a,
Ericoid leaves, in which the lamina is more or less 1948b) being dated and recent work limited in
linear, the margins being sharply recurved and scope (e.g. Noshiro et al. 1995 and references
concealing the lower surface and its stomata therein). Clemants (1995) provided a detailed
(Hagerup 1953; cf. Hara 1958), are common. Here survey of the wood of several species of Bejaria;
the serrulate "margin" with its multicellular hairs Carlquist (1989) described woods of Empetreae.
probably represents the point of reflexion of the Vessels are numerous, scattered, and have scalari-
leaf surface, not the true margin. Such leaves are form to simple perforation plates, tracheids occur,
almost restricted to Ericoideae, especially Ericeae, axial parenchyma is sparsely vasicentric, and rays
and intergrade with leaves that have strongly rev- are often uniseriate, sometimes wider, especially
olute margins, as in Erica itself (Oliver 2000). Some in lignotubers (see below), and are comprised of
Cassiope have peltate leaves in which the abaxial upright cells. Pith can be homogeneous and more
Ericaceae 147
or less lignified, heterogeneous (small cells with toideae and perhaps Pyroleae (Pyykko 1968),
thick walls and large cells with thin walls inter- although it is rarely constant within genera. A 1-,
mingled), or Calluna-type (thick-walled cells 2- or more layered hypodermis is common, espe-
restricted to the periphery of the pith; Watson cially in tropical Vaccinieae.
1964a). Homogeneous pith is widespread. There The palisade mesophyll of some thick-leaved
are intermediates between these types, and infra- tropical epiphytic Vaccinieae is especially tall with
generic variation, as in Gaultheria and Leucothoe, transverse thickened bands on the walls (e.g. Dis-
but the character is valuable; variation is similar, terigma), it (and the hypodermis) perhaps being
but less extensive, in the cortex. Position of initia- involved in water retention during dry periods.
tion of the cork cambium provides important In several Vaccinieae and also in large-leaved
information in Vaccinieae, where it can be super- Richeeae, the spongy mesophyll adjacent to the
ficial, or deep-seated, being initiated inside the abaxial epidermis is thick-walled and lignified.
ring of pericyclic fibres (the basal condition for The thickening of the cellulose walls of the spongy
the family), or develop in a canker-like fashion mesophyll varies considerably within Vaccinium.
(Stevens 1974; Odell et al. 1989). Monotropoideae In Diplycosia and some species of Gaultheria,
lack a cork cambium (Henderson 1919). fibres wander through the mesophyll, and the dry
Whether fibres in the secondary phloem are in leaf is notably tough. The mesophyll is dorsiven-
bands or scattered separates groups of genera in tral or centric in symmetry, inverse dorsiventral in
Vaccinioideae (Stevens 1970b). Nodes in Ericaceae Cassiope and also in the bases of leaves of Cos-
are predominantly unilacunar, but Rhododendron melieae, where symmetry becomes isobilateral
may have several traces in the cortex arising from then centric along the leaf from base to tip. Veins
one or more gaps (Philipson and Philipson 1968), are deeply embedded within the mesophyll, some-
as also Therorhodion (inflorescence only), Bejaria times offset to the abaxial side. In Styphelioideae
and Lebetanthus have three traces coming from a they are usually all about equal in size and are
single gap, while Richieae are tri-or multilacunar associated with abaxial bands of sclerenchyma.
(Watson 1967b). Styphelioideae often have several These reach the abaxial epidermis in Styphelieae
arcuate vascular bundles in the petiole; in other and some Epacrideae; fibres form transcurrent
Ericaceae the single bundle varies from arcuate to bands in Richeeae and one species of Leucopogon
annular. There are sclereids of various types in the (e.g. Watson 1967b). The fibres link with the peri-
cortex, especially in the petiole, but their variation cyclic fibres of the stem. Young leaves especially of
and distribution are poorly understood. Fibres Styphelioideae are bordered abaxially by a layer of
occur in the pericyclic area in the stem (and leaf); unthickened cells that produce crystals within
they are less well developed in the basal clades. In their walls. These cells disintegrate with maturity,
Ericeae in particular the petiole bundle has broad leaving free crystals in mature leaves. Druses and
flanges of lignified tissue on either side, a feature sometimes crystals of calcium oxalate commonly
that needs more study. occur in the petiole, lamina and elsewhere in the
The leaves are usually bifacial, but in some Arc- plant.
tostaphylos they are held vertically; only rarely are
there stomata on the adaxial surface of the lamina, INFLORESCENCE STRUCTURE. Flowers with paired
as in Cassiope and Cosmelieae. The two main prophylls (bracteoles) are borne in the axils of
stomatal types are anomocytic and paracytic, bracts and arranged in a raceme, or spike in
rarely parallelocytic (Richeeae) or stephanocytic. several Styphelioideae. Racemes range from being
Stomatal type tends to be constant in groups of much elongated and with large, leafy bracts to
genera, but it varies within small genera like Elliot- short and with small bracts. They are often borne
tia and Enkianthus. Stomata are unoriented or on wood of the previous season or flush, or on
oriented parallel to the long axis of the lamina, but older wood (Lems 1962a), while in some Lyonia
in Ericeae, Cassiope, etc., they may be oriented (Judd 1981) and Styphelioideae all leaf axils of the
transversely. The adaxial epidermis is lignified in uppermost part of the previous season's innova-
Styphelioideae, some genera of the Lyonia-group tion may be occupied by inflorescences. Many taxa
(Stevens 1970b; Judd 1979) and Notopora. The have axillary fascicles, probably reduced racemes.
adaxial epidermis and/or hypodermis, rarely the In Ericeae inflorescences may be reduced to single
abaxial epidermis as well, may have "mucilagi- flowers, which in turn are aggregated and localised
nous" cells which bulge into the palisade meso- on the stem, so suggesting a raceme (Hansen 1950;
phyll (e.g. Lavier-George 1936; Copeland 1943); Oliver 1991, 2000). The terminal, corymbose or
this is known only in some Ericoideae, Arbu- capitate inflorescences of many Rhodoreae and
148 P.F. Stevens et a!.
the axillary panicles along defoliate branches in on the stamens. Genera ofVaccinieae in particular
some Thibaudia are all indeterminate. Single have been characterised by stamina! features (but
flowers with paired prophylls occur in the axils of see below).
leaves at the beginning of the current season's The flowers range from 1-60 mm long and are
growth in Vaccinium section Myrtillus and at the typically bisexual, polysymmetrical and 5-merous.
end in Chamaedaphne, while the panicles in Oxy- In Rhododendron and some related genera the
dendrum terminate the season's growth. The median sepal is abaxial, rather than the normal
single, axillary flowers of some Gaultheria and adaxial condition. Ericeae are 4-merous, apart
Styphelioideae are here called multibracteolate; from Erica subgenus Pentapera; some Erica are 3-
the "bracteoles" are probably reduced inflores- merous. The ovary is superior, except in Vaccinieae,
cence bracts. In genera like Cyathodes with such however, a few Gaultheria are semi-inferior, while
flowers, the basal bracteoles (which have been in some Vaccinium the ovary is not completely
described as bracts elsewhere) are strongly keeled, inferior; ovary condition was the main feature used
and the number of bracteoles varies considerably to justify the segregation ofVacciniaceae.
(see also Virot 1975). In Styphelia, Coleanthera and Calyx and corolla are usually distinct from each
some Astroloma, the terminal branch rudiment is other and from bracts or prophylls. The calyx con-
a flattened, expanded bract-like structure, and sists of free or almost free, imbricate (quincuncial)
what appear to be single, axillary flowers are sepals. In inferior ovaries, "calyx tube" describes
reduced inflorescences (Watson 1964b; see also the inferior portion, "calyx limb" the free part
Lemson 1996; Cherry et al. 2001). In the generic borne on top of the ovary. Bracts and calyx are
descriptions, it can be assumed that flowers in coloured in some taxa. In Empetreae calyx and
racemes or spikes are subtended by a bract and corolla are almost indistinguishable, in Monotro-
have two prophylls; multibracteolate flowers are peae both may be similar to the bracts, and in
probably incorrectly designated, but pending some Styphelioideae prophylls and calyx may be
more detailed comparative studies they describe similar (Fig. 58, p. 174). In Rhododendron the calyx
the single flowers mentioned above. Truly termi- is often obsolete. Both calyx and corolla are dry
nal flowers are uncommon, but are known from and marcescent in Ericeae and the calyx is scari-
Craibiodendron (Judd 1986), Elliottia (Bohm et al. ous in many Styphelioideae. Elsewhere they are
1978), Monotropa, Enkianthus, and perhaps more or less fleshy and wither after flowering,
Calluna (Oliver 2000) and Epacrideae. Inflores- although the calyx becomes fleshy in most
cences in Richeeae are particularly complex. Gaultheria and Diplycosia. The sepals are largely
Inflorescences of many Rhodoreae have large free, although joined at the very base, and are over
and usually deciduous perulae surrounding them lOmm long in some Ceratostema; in some tropi-
(Fig. 56, p. 171). Similar perulae in Cavendishia, ascal Vaccinieae they form a tube 12 mm long with
well as bracts, are brightly coloured and (sub)per- calyx lobes barely evident. Winging of the inferior
sistent (Luteyn 1983); they are known from a few ovary and of the calyx tube has been used as a
other Vaccinieae. Leaves near the flowers are some- generic character in Vaccinieae.
times brightly coloured in Erica (Oliver 1991). The corolla is usually sympetalous, forming a
The position of the prophylls on the pedicel cylindrical, campanulate or urceolate tube with
varies greatly, although they are usually towards or free lobes; the lobes are valvate or imbricate
at the base. In Diplycosia and Tepuia they are char- in bud. Empetreae, most Monotropoideae and
acteristically immediately under the calyx (see Fig. scattered genera elsewhere are polypetalous.
61B, p. 182) and in Disterigma they envelop the Polypetalous taxa have often been called primi-
inferior ovary. In many Ericeae both bracts and tive, but polypetaly is derived from sympetaly
prophylls are recaulescent and become part of the (Copeland 1943, 1947; Kron and Judd 1990; Judd
calycine whorl, perhaps even "replacing" the sepals and Kron 1993); the only exception could be
(Oliver 2000). Monotoca elliptica may lack bracts; Monotropoideae. The shape of the corolla is
prophylls are absent in Pyroleae and some broadly correlated with taxonomy - Rhodoreae
Monotropeae and Ericeae. often have a broadly open corolla; Arbutoideae are
overwhelmingly urceolate; Vaccinioideae and Sty-
FLORAL STRUCTURE. Ericaceae show extensive phelioideaeare often urceolate to tubular; Pyroleae
variation in meristicity and in the degree of fusion are rotate. However, in Phyllodoce (sympetalous)
of the corolla. The androecium is especially the corolla is campanulate or urceolate, in Bejaria
diverse, "Bicornes", the name early given to part of (polypetalous), rotate to tubular. The corolla of
the family, referring to the distinctive appendages Anthopterus in particular is strongly winged. Poly-
Ericaceae 149
symmetry is pervasive, but many Rhododendron

have monosymmetrical flowers often associated
with colour patterning on the adaxial and adjacent
corolla lobes, while in Elliottia and in several
Pyroleae bisymmetry is evident mainly in the
curvature of the style and, in the latter, in the
positioning of the anthers as well. Some Paphia,
Thibaudia and Erica have curved corollas. Indu-
c
D
mentum on the outside of the corolla is like that
elsewhere on the plant. On the inside there are
often unicellular hairs (uniseriate in Prionoteae)
that are especially conspicuous in Styphelioideae.
They may be in distinct tufts, as in Brachyloma
and Astroloma, sometimes with beautiful surface
sculpturing (e.g. WeiHer 1996b).
G
Most Ericaceae have twice as many stamens as
petals, but there are numerous exceptions. Sty-
phelioideae have only as many stamens as corolla
lobes. These are borne opposite the sepals and are
more or less strongly epipetalous, being inserted
at the base or throat of the corolla tube; there are
no staminodes. Epipetaly is rare elsewhere (but see
Diplarche). Filaments are free to fused, variously H
shaped, sometimes geniculate, and often hairy
(but almost never in Styphelioideae). Anther
dehiscence is predominantly by two, terminal
subintrorse pores (Figs 50, 51; but see below); these
may be confluent in some Vaccinieae and rarely
latrorse or extrorse. There are slits almost the
length of the anther in the basal Enkianthus
N
(Fig. 55, p. 166), scattered in Monotropoideae and
Ericoideae (including the wind-pollinated Empe-
1
treae, see Fig. 57, p. 172) and throughout Styphe-
lioideae. In Vaccinieae in particular the anther
thecae often have very long, apical tubules through
which the pollen exits by pores or short slits (Fig.
51). In one section of Leucopogon, the anthers are
sterile above the thecae, while in Melichrus there Fig. 50. Ericaceae. Stamens in Arbutoideae, Cassiopoideae,
is a sterile basal portion (Paterson 1961). Anthers Ericoideae and Harrimanelloideae. A Arbutus andrachne.
B Cassiope lycopodioides. C Bejaria racemosa. D Bryanthus
are sometimes connate, most notably in Colean- gmelinii. E Ledothamnus sessiliflorus. F Menziesia lasiophy lla.
thera and some Vaccinieae. The pollen in some G Rhododendron fictolacteum. H Rhododendron tanakae.
Styphelioideae is red or purple; the usual colour is I Elliottia pyroliflora. J Epigaea asiatica. K Rhodothamnus
white or cream. chamaecistus. L Calluna vulgaris. M Daboecia cantabrica. N
Erica australis. 0 Harrimanel/a stelleriana. (Stevens 1971)
There has been much argument over the nature
of the stamen appendages. They may be either
paired, varying in position from the tips of the
anthers to the filaments, or four, when they are tubules of many Vaccinieae; the apex of the anther
always on the tips of the anthers. Appendages on may bifurcate, hence the paired awns of some
the "backs" of anthers (see below), including the Gaultheria (Hermann and Palser 2000). The pres-
tubules, or on the filaments are called spurs; they ence of spurs is loosely correlated with the
are flattened to terete, and where they are borne stamens being included (Matthews and Knox
often varies infragenerically, as in Erica and Pieris. 1926; Hermann and Palser 2000).
Appendages on the tips of the anthers are called There is usually a nectary at the base of the ovary
awns (e.g. Enkianthus, Arbutoideae, Cassiope, etc.), (or on top, as in Vaccinieae ); in some Monotropeae
and may be morphologically comparable to the (Wallace 1977) and Ericeae it has prominent
projections. Palser (e.g. 1961) provides extensive

details on the morphology of the nectary.
The gynoecium is usually about as long as the
corolla. The ovary has 1-12locules with apical to
basal and axile to parietal placentation; there are
one to hundreds of ovules per carpel. The ovary is
glabrous to variously hairy; in some Arbutoideae
it is covered with multicellular papillae that are
fleshy in fruit. The style is long, slender, and
usually hollow, but it is relatively short and stout,
I
especially in some Ericeae and Monotropoideae. It
I may join the apex of the ovary smoothly (style
continuous) or be more or less deeply impressed.
When it is very deeply impressed (e.g. Agarista,
Prionotes), the placentae, which of course always
arise below where the style and ovary join, appear
to be basal. The stigma is punctate to variously
lobed, the apex of the style sometimes also being
F expanded, and is at least sometimes wet and
papillate.
FLORAL ANATOMY. Palser and collaborators and

Copeland have carried out extensive studies of
floral anatomy; for Styphelioideae information
may also be found in Etienne (1919), Copeland
(1954), Jackes (1968) and Lemson (unpublished).
Palser (1961) found that variation in floral
anatomy was not very highly correlated with that
of other features, and thus species with inferior
ovaries vary greatly in how much the whole vas-
cular supply is fused. Details of floral vasculature
are largely ignored below.
M There is extensive variation in the distribution
of stomata on both surfaces of the calyx and
corolla (Watson 1962, 1967a; Stevens 1971). The
stomata can be difficult todetect, and more exten-
sive surveys are needed, but there is a correlation
with phylogeny.
When there are twice as many stamens as petals,
the androecium is generally obdiplostemonous,
but the traces of the sepaline stamens may diverge
from the central stele before those of the petaline
stamens (e.g. Leins 1964; Palser and Murty 1967).
K 0 p If there is only one whorl of stamens, it is always
the antesepaline whorl. The anthers are tetraspo-
Fig. 51. Ericaceae. Stamens in Vaccinioideae. A Oxydendron
arboreum. B Pieris nana. C Lyonia foliosa. D Andromeda rangiate. In Styphelioideae, however, they are bis-
polifolia. E Zenobia pulverulenta. F Chamaedaphne calyculata. porangiate and dehisce by two (Prionoteae, a few
G Leucothoe keiskii. H Gaultheria suborbicularis. I Gaultheria other genera) or one (the rest) slits; the latter con-
procumbens. J Diplycosia malayana. K Vaccinium poasanum. dition is caused by the breakdown of the tissue
L Symphysia racemosa. M Notopora schomburgkii. N Orthaea separating the two sporangia. The anthers usually
apophysata. 0 Themistoclesia compacta. P Gaylussacia
braziliensis. (Stevens 1971) invert during development and the vascular
bundle forms an apical hook as it turns 180°.
Hence, the "apical" pores of many Ericaceae are
really basal and the "introrse" slits of, for example,
Styphelioideae (see above) are extrorse; we refer to
Ericaceae 151
the apparent position in the discussion here. absent in Monotropeae (Copeland 1941), or the
Inversion in Enkianthus, Monotropoideae and micropylar haustorium alone may be present, as in
Arbutoideae is late or may not occur at all (e.g. Elliottia (Copeland 1943).
Copeland 1941, 1947); note that anthers in Cyrilla
(Cyrillaceae, sister to Ericaceae) do not invert. The POLLEN MORPHOLOGY. Most woody Ericaceae
development of the anther wall is unexpectedly show little variation in pollen morphology (see
variable (Hermann and Palser 2000). There is Oliver 1987, 1991; various contributors in Luteyn
much variation in how the anthers dehisce. There 1995; Warner and Chinnappa 1986). The grains are
is a well-developed endothecium in Enkianthus usually in isopolar, radiosymmetric, triangular to
and Bejaria (Copeland 1943), although it is other- globular tetrahedral tetrads 23-82Jlm in diameter.
wise absent, or in Monotropeae and perhaps a few They are usually tricolporate, sometimes tricol-
Rhodoreae only poorly developed. Elsewhere an pate or tricolporoidate, and with definite trans-
exothecium only is found. Anthers in Ericoideae verse furrows. Aperture number varies from 3 to
open either by collapse tissue, or by resorbtion 5 at the species level in Enkianthus (Anderberg
tissue, the cells of the latter containing much oxalic 1994b). Some Monotropeae have porate pollen
acid (D'Arcy et al. 1996); resorbtion tissue is wide- (Wallace 1995). The exine is scabrate, verrucate,
spread, occurring in i.a. Styphelioideae (Paterson microrugulate or psilate (the latter especially by
1961). Vaccinioideae may have white disintegra- aperture margins). Calluna vulgaris has distinc-
tion tissue on the backs of the anthers that some- tive irregular tetrads that have only faint colpi and
times extends to the apices of the filaments. distinct ora (Foss and Doyle 1988). Viscin threads
In some Ericeae and Styphelioideae the carpels occur in most Rhodoreae, Phyllodoceae, and
are opposite the petals, rather than opposite the Bejarieae (Fig. 52). Enkianthus, some Erica (Oliver
sepals, as is common (Palser 1961; Palser and 1991), and many Monotropeae have pollen in
Murty 1967). Numerous species of Vaccinium in monads, as does Andersonia macranthera.
particular have inpushings of the outer walls of the Chimaphila has polyads, tetrads and monads, and
carpel, and are described as being pseudo-10- may even have very reduced colpi (Takahashi 1986,
locular. The ten carpels of Gaylussacia alternate 1987).
with both sepals and petals (Palser 1961); it is In Styphelioideae, pollen is in tetrads 23Jlm
unclear if they are derived from a pseudo-10- (Sprengelia) to 82Jlm (Epacris) across, or less com-
locular gynoecium or not. In Arbutoideae extra monly in reduced tetrads or monads. Monads are
carpels appear to develop in the plane of the septae rare outside Styphelieae, but occur in some species
(Palser 1954). Epigaea has carpellary bundles in of Richea. In Astroloma and some species of
the plane of the septae (Palser 1952). The style is genera that otherwise have normal tetrads, irreg-
hollow and the stylar canal is confluent with the
ovary chamber.
EMBRYOLOGY. The tapetum is secretory, and

microsporogenesis is simultaneous. Tapetal cells
may be uninucleate (Epacridoideae), binucleate
(Monotropoideae, Vaccinium), or multinucleate
(Ericoideae, ?some Vaccinioideae; see Johri et al.
1992). Pollen is binucleate.
The ovules are usually anatropous, unitegmic,
tenuinucellate, and have an endothelium. The
megagametophyte is 8-nuclear and develops from
a single archesporia! cell; in Styphelia the three
megaspores that do not form the embryo sac
may remain functional (Brough 1924). The mega-
gametophyte has sharp, ear-like processes in
Enkianthus and Epigaea (Palser 1952).
The embryo in Monotropa may consist of only
two cells (Olson 1980); all Monotropeae have
much reduced, undifferentiated embryos. There
are usually well-developed endosperm haustoria Fig. 52. Ericaceae. Pollen tetrad of Ledothamnus atroadenus
at both ends of the embryo, although they may be with adherent viscin threads (x2350). Photo J. Luteyn
ular tetrads with 0-4 viable grains are produced. POLLINATION AND REPRODUCTIVE SYSTEMS.
About half the genera of Styphelieae have pollen Little is known about pollination in Ericaceae,
grains that appear to be monads, and these range especially its tropical members, despite its popu-
from 10J..Lm (Oligarrhena) to 100J..Lm (Astroloma) larity among horticulturalists and its often large
across. They have been described as "pseudomon- or conspicuous flowers (although the corolla is
ads" (Smith-White 1948a, 1955), but in most cases only some 0.7mm long in Erica petricola). Most
no evidence of aborted grains remains at maturity. records of "pollinators" are simply of flower visi-
In Monotoca the pollen is a reduced tetrad (T- tors, and how effective these visitors may be in
monad) with three aborted grains attached to the pollination is unknown.
pole of the fertile grain. Meiosis in many temperate Ericaceae occurs in
the late summer of the year preceding anthesis.
KARYOLOGY. All chromosome numbers in the Inflorescences overwinter either enclosed in a
accounts are given as haploid (n) numbers. North perulate bud (Rhododendron) or completely
temperate Ericaceae and some Styphelioideae are exposed (Pieris); the flowers open in the early
moderately well-known cytologically, but the rest spring.
of the family is very poorly known. Janaki Ammal Flowers are usually hermaphroditic. Autogamy
et al. (1950) surveyed Rhododendron, finding is quite common (e.g. Reader 1977; Oliver 1991;
numbers as high as 2n = 156 (= 12x) in R. Kraemer 2001; see also Luteyn 2002). Preliminary
manipurense. Intrageneric polyploid series are studies of Cyathodes glauca indicate that it is an
quite common, and include some Styphelioideae obligate oucrosser (pers. comm. C.M. WeiHer), as
(Smith-White 1955; WeiHer 1996a, 1996c; Dawson is Pentachondra pumila (Godley 1966). Gynodioe-
2000 for numbers in that subfamily). For chromo- cism is known in Gaultheria in particular (e.g.
some numbers in Gaultheria and its relatives, see Middleton 1991; Hermann and Camhi 1992) and
Middleton and Wilcock (1990) and Luteyn (in in Styphelieae. Leptecophylla is functionally dioe-
Luteyn 1995); Atkinson et al. (1995) give counts cious with the flowers effectively unisexual: all
for tropical Vaccinieae which suggest that a more species produce larger and apparently hermaph-
extended survey will provide valuable systematic rodite but actually male flowers with apparently
information. In general, base numbers are con- functional gynoecia and male-sterile flowers. In
stant within genera or groups of genera, but the latter, anther development is aborted at an
Gaultheria in particular varies from n = 11, 12, or early stage, resulting in reduced anther size and no
13. The base number for the family is unknown, pollen formation, but fruit is set. The functionally
but in Enkianthoideae n = 11. dioecious L. juniperina has a prezygotic incom-
Many Styphelioideae are n = 13 or n = 12; no patability mechanism in the "male" flower, causing
counts are known from Prionoteae. It is likely that the arrest of pollen tube growth and associated
either n = 12 or n = 13 is the plesiomorphic char- pollen tube abnormalities at various stages from
acter state (Stace et al. 1997). Probable chromo- the stigma to the base of the style (pers. comm.
some number reduction series (including n = 13, C.M. WeiHer). All species are protandrous and the
12, 11, 10, 9, 8, 7, 6, 4) are especially marked in stigmas are normally coated with pollen at anthe-
Styphelieae (Smith-White 1948a, 1955); possible sis (as in Planocarpa and Cyathodes). In
infrageneric reduction series are Leucopogon Planocarpa two species (P. petiolaris and P.
sens. str. (n = 12, 11, 10), Cyathodes (n = 12, 10), sulcata) are assumed to be gynodioecious, with
Brachyloma (n = 9, 7), and Sphenotoma (n = 7, 6). both hermaphrodite and male-sterile flowers
Derived chromosome numbers may provide setting fruit. Monotoca, Pentachondra and Tro-
synapomorphies to mark certain problematic chocarpa all have some species with unisexual
lineages in Styphelieae (Stace et al. 1997). A flowers, while dioecy occurs in the wind-
remarkable case of stabilised triploidy was pollinated Empetreae. Monotoca exhibits a range
reported in "Leucopogon" juniperinus, in which in states of reduction of the anthers. Some species
haploid chromosome numbers (n = 4) are trans- of the genus have bisexual flowers, some are uni-
mitted to the pollen and diploid numbers (2n =8) sexual with reduced and barren anthers, while in
are transmitted in the megaspore. The maternal others the anthers are completely aborted with
and paternal chromosomes may be responding only bare filaments visible.
differentially to a cytoplasmic gradient that leads Flower colour is an important attractant,
to the formation of derived monad pollen by although flowers of some Acrotriche, Andersonia,
apoptosis of cells of the ericoid tetrad pollen grain Erica, Vaccinium, and Rhododendron, etc., smell,
(Smith-White 1948b). whether sweetly or unpleasantly (see also Knudsen
Ericaceae 153
and Olesen 1993). A common colour is bright corolla tube with a narrow orifice and attract
white or cream, often with a pinkish tinge, while moths, butterflies and long-tongued bees such as
many tropical Vaccinieae have red flowers. Orange, Amegilla species. Andersonia micrantha has a
pink, yellow, blue and mauve/purple are less foetid odour and attracts flies. Some Monotoca
common. In species with cryptic flowers, the species have a very short corolla tube forming an
colours may be dull cream/brownish (Monotoca open bowl (M. tamariscina) and are visited by
tamariscina) or greenish, in which case they may small dipterans (Keighery 1996).
have a strong odour (Acrotriche spp.). The corolla The corolla tube of Conostephium has a narrow
is usually single coloured. Simple bicolours are entrance, and the stigma, rather than blocking the
known in Rhododendron, Erica, tropical Vac- entrance, is exserted one to several millimetres
cinieae, Epacris, and large-flowered species of from the tube. Insect visitors are relatively large
Agapetes (where there is often horizontal and quite specialised (Leioproctus bees and
banding); many temperate species of Rhododen- Amegilla pulchra), but no nectar is produced and
dron have spots on their adaxial petal(s), presum- the flower is buzz-pollinated (T. Houston, pers.
ably honey guides. In Epacris impressa, there are comm., cf. Keighery 1996). Buzz pollination occurs
four races with short white corollas, long pink elsewhere, as in Vaccinium stamineum (Cane et al.
corollas, and broad white or pink corollas (Stace 1985; Knudsen and Oleson 1993).
and Fripp 1977). Many plants occur as pure-colour Hummingbirds visit the small, urceolate flowers
stands in different habitats and show some differ- of a number of Arbutoideae in eastern North
ences in flowering times, but some populations are America (Diggs 1995), although confirmation of
panmictic with pink- and white-flowered plants, pollination is needed. In general, species of Eri-
and at Wilson's Promontory there are heteroge- caceae known or suspected to be bird-pollinated
neous populations with red and white flowers. · lack scent and many, but by no means all, have
Flowers are usually animal-pollinated. Bees are long, tubular, red flowers (in Styphelioideae
common visitors and effective pollinators of many mostly Astroloma, Cosmelia, a few Andersonia,
Ericaceae with small, scented urceolate flowers several Styphelia); the ovary may be superior or
(e.g. Reader 1977; contributors to Luteyn 1995), inferior. Gullet-type flowers are almost restricted
and they also visit many larger-flowered species, to Rhododendron within Ericaceae. Humming-
e.g. Rhododendron, as well as many Monotropeae birds are pollinators in the New World (Luteyn and
(Wallace 1977). In Ericoideae that have viscin Sylva 1999; Luteyn 2002 and references therein)
threads mixed with the pollen, strands of tetrads and sunbirds, honeyeaters (Meliphagidae) and
become attached to the flower visitors. In some other groups in the Old World. Flower mites are
Kalmia pollination is explosive; bees release the dispersed by birds in both hemispheres (Colwell
anthers, held under tension in pockets in the 1973; Stevens 1976; Naskrecki and Colwell 1998).
corolla, and get dusted with pollen. Pollination by Nectar content in neotropical Vaccinieae averaged
hovering flies with long probosces is common in 20.4(7.1-32)% sucrose, largely appropriate for
southern Africa Erica (Rebelo et al. 1985), while hummingbirds (Luteyn and Sylva 1999). About 60
pollination by thrips is suspected both there species of Erica may be bird pollinated, but few
(Rebelo et al. 1985; cf. Oliver 1991) and in north- species of birds are involved, and only one, the
western Europe (Hagerup and Hagerup 1953). sunbird Nectarinia violacea, is largely restricted
Species pollinated by sphingids have a very sweet to the genus (Rebelo et al. 1985; Oliver 1991). In
and heavy scent, as have possibly bat-pollinated Papuasia bird pollination seems to predominate in
taxa (in Rhododendron: the flowers are very large species of Rhododendron living at high altitudes
and robust, and are often 6-merous or more); (over 3000 m), species probably pollinated by but-
flowers pollinated by bees and butterflies are also terflies, sphingids and even bats being found more
scented. commonly below 3000m (Stevens 1976). Cosmelia
Perhaps 90% of Western Australian Styphe- and Astroloma, Styphelioideae visited by hon-
lioideae are insect-pollinated (Keighery 1996). The eyeaters and spinebills, have a relatively long
corolla tube in Leucopogon is variable in length corolla in which the entrance to the tube is
but in most species is relatively short (1-3 mm). occluded by either adpression of the very short
Visitors include Tarsipes (seen on several species), lobes or stigma and corolla hairs blocking the
but more generally bees, including feral Apis mel- entrance; access to nectar is restricted to species
lifera, moths, butterflies and dipterans (Clifford that are reasonably forceful in probing the flower.
and Drake 1981; Keighery 1996; Brown et al. 1997). Of particular interest is the pollination of the
Sphenotoma and Lysinema have a relatively long functionally dioecious Leptecophylla divaricata by
several species of birds including honeyeaters tube (Leucopogon spp., Vaccinium spp., Brachy-
(Higham and McQuillan 2000). loma preissii). In many Rhodoreae, Styphelia and
The Honey Possum (Tarsipes rostratus) utilises a Cyathodes the anthers are well exserted, as they are
number of Styphelioideae (Turner 1982) although, in Richea sprengelioides and the "brush blossoms"
given their floral morphology, it is unclear how of Andersonia setifolia (Keighery 1996). The stick-
it would pollinate many species that it visits. iness of the pollen also varies, as in Styphelioideae,
Acrotriche species with small, cauliflorous flowers from very dry and loose in the buzz-pollinated
appear, on the basis of flower size, poorly adapted Coleanthera (Keighery 1996), Sprengelia and
to pollination by a comparatively large animal Conostephium to very sticky in Acrotriche and
(McConchie et al. 1986) but, on the New England species of Astroloma and Cosmelia visited by
plateau, flowers of A. aggregata form an important birds. In Leucopogon, species with pendulous
part of the diet of the marsupial mouse (Antechi- flowers tend to have enclosed anthers that contain
nus stuartii) and it may also be the pollinator dry, loose pollen, while species with erect flowers
(Fletcher, in McConchie et al. 1986). In Western have anthers in the corolla throat that have rela-
Australia, the only recorded visitor to Acrotriche tively sticky pollen. In buds of Brachyloma preissii
cordata is a muscid fly (Keighery 1996), so it is a globule of sticky fluid is exuded by the stigma
unclear if mammals are always involved in the pol- onto the unopened anthers. The globule is then
lination of the genus. spread across the anthers and pollen as the flower
Secondary pollen presentation in clumps at the opens and the anthers dehisce, ensuring that the
ends of the petals has been suggested for Elliottia pollen is well coated in sticky material.
(Copeland 1943). Many Ericaceae have various In many species the stigma is turgid and appears
forms of hairs associated with the corolla, and in very wet from the mature bud stage to anthesis
many cases these are related, either directly or (see also Heslop-Harrison and Shivanna 1977).
indirectly, to pollination. In Acrotriche the anthers In other species, especially a number of Styphe-
dehisce in bud and shed the sticky pollen onto lioideae, particularly those with an exserted style,
the hair tufts at the tips of the corolla lobes. At the stigma is not enlarged and appears virtually
anthesis the lobes recurve, taking the pollen with dry when mature.
them - again, secondary pollen presentation Barriers to hybridization are often poorly devel-
(McConchie et al. 1986; Ladd 1994). In Leucopogon oped. In Rhododendron hybridization between
and Styphelia hairs extend from the corolla lobes species in secondary (often anthropogenic) habi-
into the mouth of the tube. The situation is similar tats is common (for Erica, see Oliver 1991), and
in Astroloma. In many cases the dehiscing anthers introgression may occur (Kobayashi et al. 2000).
shed pollen among the hairs, which may assist Hybrids between several "genera'' have been
in retaining pollen within the corolla tube where reported. A hybrid between Gaultheria and
it more effectively contacts pollinators. (In Pernettya (xGaulnettya Marchant, Luteyn 1995
Astroloma, Brachyloma, Lissanthe and Melichrus for references), was one of the first synthesized.
the corolla hairs seem to assist in retaining nectar However, in Erica x Simocheilus (Oliver 1991),
at the base of the corolla tube.) Rhododendron x Ledum, and Gaultheria x
Empetreae are likely to be wind pollinated. Their Pernettya, the latter members of the pairs have
inconspicuous perianth has free parts, the anthers been synonymized, and this is a likely course with
are exserted, and the stigmatic surface relatively Phyllodoce, Rhodothamnus and Kalmiopsis, given
much enlarged; as is common in wind-pollinated the hybridization reported to occur between
taxa, they are largely dioecious and ovule number the genera (e.g. Dome 1999) and their underlying
is reduced. Anderberg {1994a) examined the cor- similarity. Purported hybrids between Kalmia
relation between ploidy level, breeding system and and Rhododendron are misidentified.
fruit colour in Empetrum. Some 100 species of
Erica, especially species previously placed in FRUIT AND SEED. The flowers are usually hori-
Philippia and other minor genera, are also proba- zontal to pendulous and the fruits are erect (Fig.
bly wind pollinated (Rebelo et al. 1985; Oliver 53). Rhododendron subsect. Ledum has erect
1991); there is no association with dioecy here. flowers and pendulous capsules opening from the
Richea sprengelioides and R. procera are candi- base first; some Vaccinium have pendulous flowers
dates for wind-pollination. and fruits. Capsule dehiscence is either septicidal
Anthers are commonly enclosed in the corolla or loculicidal, rarely septifragal, the two common
tube, but they may form a funnel-shaped structure types characterising large groups of genera.
that presents pollen at the mouth of the corolla Details of how the wall is thickened or the margins
Ericaceae 155
individually, or all seeds together in the drupes

that occur in Empetreae, Styphelieae, many Arbu-
toideae, and Gaylussacia.
The testa is usually only a single cell layer thick,
but several cells thick in the region of the raphe,
although only the walls of the outer layer are much
thickened; in Enkianthus vascular tissue persists
in this region (Netolitzky 1926). The testa of
Vaccinium sect. Oxycoccus and Andromedeae is
thicker. All cells may be papillate (Daboecia, rarely
in Erica), or a few cells may be more or less balloon-
like and in aggregate form a wing (Enkianthus,
many Leucothoe). Seeds also have elongated
"tails" (Rhododendron), outgrowths from one end
(Pterospora) or one side (Andromeda, where it is
many cells thick), Craibiodendron), or they may be
generally flattened (Elliottia racemosa). The cells
of the testa are more or less isodiametric in surface
view, or strongly elongated, at least 2x longer than
wide; intermediates and infrageneric variation
occur in Kalmia, Vaccinium, Erica and Pieris. The
anticlinal walls are usually straight, but sinuous in
some Erica (Oliver 1991). Thickening is best devel-
oped on the anticlinal and inner periclinal walls,
and varies from less than 3j..tm (especially in taxa
with elongated cells) to over 50 j.lm thick (in some
Costera). The outer periclinal walls are rarely much
thickened, although Gaultheria and its relatives in
particular are exceptions; in Erica these walls may
have surface ornamentations (Oliver 1991). The
other walls have fine plasmodesmata 2j..tm or less
in diameter, but pits are larger in some Pyroleae
(Takahashi 1993), Calluna and Elliottia. Some
tropical Vaccinieae have broad, scalariform bands
of thickening on the testal cells; the testa often
becomes mucilaginous on wetting. The walls of
Fig. 53. Ericaceae. Fruit. A Enkianthus pauciflorus. B Cassiope the testa cells are particularly thin when the
mertensiana. C Bejaria racemosa. D Ledothamnus sessilijlorus. endocarp is thick, as in Arbutoideae, while in
E Diplarche multiflora. F Rhododendron lapponicum. G Rhodo- some Erica (Oliver 2000) the ultrathin testa is
dendron tomentosum. H, I Elliottia (Tripetaleia) bracteata, in
I one valve removed. J Epigaea rep ens, ovary with top removed. transparent.
K Epigaea asiatica, note seeds on placentae. L Phyllodoce aleu- The straight, white and more or less terete, rarely
tica. M Kalmia (Leiophyllum) buxifolia. N Calluna vulgaris. 0 spatulate embryo is embedded in endosperm;
Daboecia azorica. (Stevens 1971) the cotyledons are relatively short. Some tropical
Vaccinieae have green embryos; in these taxa the
separate helps delimit individual genera (e.g. testa is mucilaginous.
Lyonia, Ledothamnus, Pyrola). Capsule shape
(elongated versus spherical) is of systematic DISPERSAL. There are two main dispersal syn-
significance in Rhodoreae and Erica. dromes in the family, by wind and animals. Cap-
Some fruits are baccate, developing from an sular fruits have small and quite often variously
inferior (Vaccinieae) or, rarely, a superior (some winged or tailed seeds which are presumably dis-
Diplycosia and Gaultheria) ovary; in both the persed by wind; the walls of the testa are usually
latter genera, the calyx alone is usually fleshy. The thin. The capsule in Erica is xerochastic, and seeds
endocarp surrounding the loculi often has one or of some Erica have an elaiosome, presumably for
two layers of fibres. These layers are much devel- ant dispersal (E.G.H. Oliver, pers. comm.). The
oped and form the stone surrounding the seeds flowers of Richea scoparia are visited by the snow
skink, Niveoscincus microlepidotus, which removes common in Ericoideae. In a survey of flavonoids

the corolla to get at the nectar and so later facili- in Monotropoideae, Pterospora and Hypopitys
tates seed release (Olsson et al. 2000). alone were found to lack flavonol glycosides
Baccate fruits are often dark purple or red when (Bohm and Averett 1989). Ellagic acid is known
ripe. The testa cells are rather thick-walled, and the mainly from arbutoids (Stevens 1971; Harborne
fruits are presumably eaten by birds, dispersal and Williams 1973) and methyl salicylate from
being endozoochorous. However, in Vaccinium Gaultheria and its relatives (cf. Luteyn et al. 1980).
section Oxycoccus the fruits float, and the seeds In Styphelioideae, eight types of compounds
can also (they have a multilayered testa); water dis- occur in leaf epicuticular waxes, viz. alkanes, alde-
persal seems an option. In some tropical Vac- hydes, triterpenes, primary alcohols, esters, 13-
cinieae the testa is mucilaginous and the embryo diketones, flavonoids and diterpenes. Most are
is green; exozoochorous dispersal is a possibility. widespread, but 13-diketones are confined to
Old World Vaccinieae with such seeds often have Richeeae, flavonoid waxes are restricted to Pri-
more or less translucent white, yellow or orange onotes, and diterpenes to Richeeae and Epacrideae
fruits; in the New World white or lilac fruits are (Salasoo 1983, 1985; Mihaich 1989). Both the mor-
common. Seeds with this syndrome of characters phology of the wax layer, which includes plates,
germinate better on moss than do seeds with a ribbons and tubes (Mihaich 1989; WeiHer et al.
thick-walled testa, which germinate better on soil 1994), and wax composition show more variation
(S. Vander Kloet, pers. comm.); taxa with a in Styphelieae, both at the level of the tribe and of
mucilaginous testa are often epiphytic or epilithic, individual species.
places where moss or humus accumulations are
high. Seeds in taxa which have capsules and fleshy RELATIONSHIPS WITHIN THE FAMILY. Ericaceae
calyces have thick-walled testa cells and a white are sister to the rather poorly-known Cyrillaceae,
embryo; again, endozoochory is likely, although and in turn these are sister to Clethraceae (Ander-
wind dispersal is also possible (Luteyn 1995, berget al. 2002). Detailed studies of these groups,
Gaultheria). In those taxa in which single seeds particularly their stamen development and the
individually or all the seeds together are sur- nature of any mycorrhizal associations, are inte-
rounded by endocarp, the unit of dispersal is the gral to understanding the evolution of Ericaceae.
pyrene (aggregate). Mammals eat the fruits of a Ericaceae were the subject of important phenetic
number of taxa, as the name bearberry (Arc- studies (e.g. Watson et al. 1967). Stevens (1995)
tostaphylos uva-ursi) suggests; bears are also very summarized some earlier classifications, while
fond of the fruits of Vaccinium spp. (McCloskey Copeland (1941) divided the Monotropoideae into
1948). The more or less indehiscent fruits of some tribes. A phenetic analysis of pollen characters by
Ericeae are barely fleshy; the development of Warner and Chinnappa (1986) resulted in the
pyrenes varies considerably, and dispersal mecha- break-up of most groups studied, but their taxo-
nisms are unknown (Oliver 1991). Drupes of nomic conclusions have mostly not been con-
Styphelieae are dispersed by animals, usually after firmed. Recent morphological and molecular work
ingestion and passing through in the faeces. Birds (e.g. Anderberg 1993; Judd and Kron 1993; Crayn
(in Australia, especially the emu, Dromaius novae- et al. 1996; Kron 1996; Kron et al. 2002a, 2000b) has
hollandiae) are the usual agents, but several mar- changed our ideas of higher-level relationships,
supials have been recorded eating the fruit. In and Kron et al. (2002a) provide a detailed review of
Epigaea the placentae alone are fleshy and seeds, the family and a new classification (Fig. 54). Earlier
which have thick walls, may be dispersed by ter- conventional classifications (Stevens 1971; Watson
restrial animals. 1976) are decidedly unsatisfactory, but the
informal classification of Watson (1967b) for
PHYTOCHEMISTRY. Harborneand Williams (1973) Epacridaceae has many good points.
surveyed the flavonoids and simple phenols of Enkianthus (Fig. 55, p. 166) is sister to the rest of
Ericaceae, while Moore et al. (1970) provided the family. Within Monotropoideae, the relation-
details for Empetreae; anthocyanins (Jarman and ships of Monotropeae and Pterosporeae have been
Crowden 1974) and flavones (Jarman 1975,Jarman controversial. Furman and Trappe (1971) thought
and Crowden 1977) have been surveyed for that they were derived from Pyroleae, Anderberg
Styphelioideae. The group has a fairly distinctive (1993) the reverse, and Copeland (1941) that they
chemical spectrum. were of polyphyletic origin from Arbutoideae; see
The flavonol gossypetin 3-galactoside links also Wallace (1975). Bidartondo and Bruns (2001)
Empetreae with other Ericaceae, being especially have recently begun to clarify relationships within
Ericaceae 157
Actinidia
Arbutus Arbutoideae
Bejaria
Call una Ericoideae
Ceratiola
73
Rhododendron
100 Cassiope Cassiopoideae
100 Chamaedaphne Early anther
Gaultheria Vaccinioideae inversion
Vaccinium
Cosmelia
64 Epacris
Pentachondra Styphelioideae
Dracophyllum
Prionotes
Pyrola Monotropoideae
Enkianthus Enkianthoideae
59
Cyrilla
Clethra
Diapensia
Symplocos
Diospyros- Outgroup
the group (cf. also Kron et al. 2002a); we follow Fig. 54. Ericaceae. Strict consensus of three trees found in the
their work in recognising Hypopitys, but the combined sequence analysis of 18 s, rbcL, and matK genes for
22 taxa of Ericaceae and their closest relatives. Bootstrap
number of monotypic genera is disconcerting. values greater than 50% are given above/below branches. Taxa
Variation in fruit type, inflorescence size, and in boldface type represent Ericaceae. Redrawn from Kron et
anther morphology do not correlate well (Wallace al. (2002a)
1987). Freudenstein (1999) looked at relationships
and character transformation in Pyroleae. Within
Arbutoideae, Hileman et al. (2001) suggest that fruit, although Drude (1889), Watson (1967b) and
Arbutus, plesiomorphic at least as regards some Powell ( 1983) separated Richea, Dracophyllum and
aspects of fruit anatomy, is paraphyletic. Sphenotoma as a separate subfamily, Richeoideae,
A major lineage has Cassiopoideae as sister to based on the presence of annular leaf scars. Sty-
the diverse and speciose Ericoideae that includes phelioideae were then divided into tribes. Styphe-
Bejaria, Erica, Rhododendron and Empetrum and lioideae s.l. form a monophyletic group within
their relatives. The inclusion of the Empetraceae in Ericaceae (Anderberg 1992; Crayn et al.1996, 1998;
Ericaceae is supported by chemical (Moore et al. Kron 1996; Kron et al. 1999b), and we recognize
1970) and molecular (e.g. Kron 1996) data, and by seven monophyletic tribes within them. Pri-
the distribution of hosts of the fungi Chrysomyxa onoteae (Prionotes and Lebetanthus) are basal, and
andExobasidium (Savile 1979).Anderberg (1994a) possess plesiomorphic features such as multicellu-
discussed relationships in Empetreae. lar hairs and filaments free from the corolla. The
Harrimanella, with Vaccinium and Epacris monotypic Archerieae are probably sister to the
and their numerous relatives, form another rest of the subfamily, but relationships between the
clade. Epacridaceae had traditionally been divided remaining five tribes are unclear (Kron et al.
into two subfamilies, Styphelioideae and 2002b; for morphological studies, see Powell et al.
Epacridoideae, based on features of the ovary and 1996, 1997). Styphelieae, the largest tribe, are in
some respects most derived, but generic limits HABITATS AND SYMBIONTIC RELATIONSHIPS.
here and in Epacrideae in particular need atten- Ericaceae are particularly prominent in arctic and
tion (e.g. Cherry et al. 2001). temperate regions, the montane tropics of South
Evolutionary studies had suggested two group- East Asia-Malesia and Central and South America,
ings of genera within the erstwhile Andromedeae, where Vaccinieae in particular are frequently
one including Lyonia and its relatives, the other Hanes and epiphytes. They are also common in
Gaultheria and its relatives (Stevens 1970b). Judd Mediterranean climates, including California and
(1979) clarified cladistic relationships within especially in heathland in South Africa (fynbos)
Lyonieae. Recent studies only partly confirm and Australia (kwongan of SW Australia, wallum
other tribal groupings in Vaccinioideae (Kron of SE Queensland). Ericaceae are heliophilous,
et al. 1999a, 2002b), and relationships around most members preferring acid conditions, notable
Gaultheria need much study (Powell and Kron exceptions being Rhododendron hirsutum in the
2001). Alps and some Ericeae in the Cape region (Oliver
Characters involved in pollination syndromes 1991). Such habitats are characteristically very low
and/or variation in the numbers of flower parts in available nutrients, contain mostly low levels of
have previously been considered very important organic matter, and are often seasonally dry, as in
in generic delimitation. Larger or smaller groups the Mediterranean climates. Some epiphytic and
of species distinguished by such features epilithic tropical Vaccinieae in particular, and
were removed from Rhododendron, Erica and rarely Rhododendron, have root tubers apparently
Vaccinium which have, hardly surprisingly, turned involved in water storage (Luteyn 2002).
out to be highly paraphyletic. Realignments in Ericaceae also grow successfully in acid bogs
Rhodoreae were suggested by Kron and Judd and as epiphytes, both habitats often being high in
(1990), while both molecular (Kron and King organic matter and the latter also liable to dry out
1996) and morphological (N. Gift, pers. comm.) intermittently. They are a major component of
data suggest major changes in generic limits in the vegetation in north temperate barrens and other
Phyllodoceae. We adopt a broad circumscription rocky outcrops. Some Vaccinium are epilithic in
of Kalmia below (see Kron et al. 2002a for the Malesia, while Monotropeae and Pterosporeae are
formal transfers), but more changes may be mycotrophic herbs of the forest floor.
needed. Oliver (2000, for a summary) showed that Ericaceae are noted for being mycorrhizal. These
meristic and fruit differences used to segregate fungal associations develop on specialised, very
genera from Erica are unsatisfactory; thus, genera fine (40-70 f.!m diameter) "hair roots" that effec-
like Salaxia and Simocheilus (= Thoracospermum), tively replace ordinary root hairs which may even
previously placed in the Salaxidae, are probably be absent (Wood 1961). Hair roots are ephemeral,
related to separate sections of Erica, hence the particularly in species growing in habitats sub-
broad limits of that genus below. jected to summer drought (Bell and Pate 1996a).
Luteyn (e.g. 1997) suggested some changes to The cortex of the hair root is reduced to two
generic limits in neotropical Vaccinieae but, as single-celled layers, an endodermis and an exo-
Vander Kloet (1985) noted, the limits of Vaccinium dermis, both with Casparian strips on their radial
itself were unsatisfactory. Indeed, Kron et al. walls (Allaway and Ashford 1996); the exodermis
(2002b, see also 1999a) indicate the magnitude of in some Styphelioideae may have thickened walls
the problem. Major clades they note include and a suberised lamella which constitutes the out-
Asian Agapetes, perhaps with many other SE Asian ermost layer in older hair roots once the epider-
species of Vaccinium (see the description of mis collapses or is sloughed off (Briggs and
Agapetes below), Oceanic Agapetes (= Paphia) + Ashford 2001). The vasculature of a hair root is
Dimorphanthera, and a large, only weakly sup- minimal - a single xylem tracheid and a single
ported New World group that in turn includes sieve element plus companion cell (McLennan
well-supported Andean and Meso-American/ 1935).
Caribbean clades. Many genera described below In ericoid mycorrhizae (see Read 1996) the outer
are para- or polyphyletic. Other smaller and cortical cells of hair roots are almost filled by
perhaps independent lineages include mainly branching fungal hyphae. The hyphae first develop
species of Vaccinium. Unfortunately, it is too early a loose network around each hair root, from the
to perceive the limits of the major groupings; cyto- region behind the root apex where epidermal cells
logical data (Atkinson et al. 1995) may help clarify are differentiating to the older regions of the same
the limits of the clades. Formal intergeneric trans- root where epidermal cells have enlarged and
fers should be made only with great caution. matured (Duddridge and Read 1982). Hyphae
Ericaceae 159
positioned alongside the outermost epidermal The fungal association may (a) improve the
cells branch at right angles and eventually pene- mobilization of nutrients from the soil rhizos-
trate the wall, usually at a single specific point phere, (b) increase the exploitation of a larger
per cell (Hutton et al. 1994), presumably using volume of soil and (c) make otherwise intractable
enzymes. Despite earlier reports, there is no lateral sources of nutrients available. Mobilization of
or internal spread of infection laterally via adja- nutrients is the key issue in improving competitive
cent epidermal cells (see Duddridge and Read effectiveness of ericaceous hosts whose preferred
1982; Hutton et al. 1994). The hyphae in the epi- habitats are often high in organic matter and/or
dermal cells branch, forming prolific intracellular low in available nutrients, especially nitrogen.
coils which retain their structural integrity after Facilitating the uptake of nitrogen from substrates
the infected root cell senesces (Duddridge and such as proteins, peptides and other high-molecu-
Read 1982). In Lysinema ciliatum certain thick- lar weight insoluble forms is an important func-
walled epidermal cells containing hyphal coils tion of the fungal component, not making
separate at the middle lamella and may then phosphorus available, as was originally thought
be released in the soil, whereas others remain (Read 1983; Perotto et al.1995). Inorganic nitrogen
attached to their parent roots (Ashford et al.1996). in the form of ammonium and nitrate is readily
Either may be a source for infection of other hair taken up by non-infected and mycorrhizally-
roots formed in the same or subsequent seasons. infected roots, and the absorbed N is incorporated
Arbutoid mycorrhizae, which also occur in the into the insoluble fraction of the plant shoot (Read
Pyroleae, show the same penetration of the cells, 1996; Bell and Pate 1996a). Nitrate is the greatest
but the root is invested by a Hartig net, the hyphae component of xylem sap when inorganic N is in
following the cell walls or even forming a complete good supply in the soil, as it is after fire or during
sheath. In monotropoid mycorrhizae, the root is major nitrification events following initial winter
invested in the same way, but the hyphae form peg- rains. Ammonium is dominant when N is limited,
like intrusions of the cell wall. This categorization as during the greater part of the growing season
may be something of an oversimplification (Bell and Pate 1996a). Ammonium is likely to be
(Largent et al. 1980), and the mycorrhizal status the form in which N passes from fungus to host
of Enkianthus (and Cyrillaceae) appears to be root, and it moves to the shoot in organic form,
unknown. principally as glutamine, but in some species argi-
Several species of ascomycete fungi form ericoid nine predominates (Bell and Pate 1996a; Smith
mycorrhizae and some of these fungi associate and Read 1997). The endophyte receives carbohy-
with ectomycorrhizal roots as well (Bergero et al. drate from the host. Finally, the achlorophyllous
2000). Eight species of fungus have been recorded Monotropa, at least, apparently receives both 14C-
from Gaultheria shallon alone (Xiao and Berch and 32 P-labelled compounds from trees by way of
1995). There is little evidence of host specificity at its mycorrhizal associate (Bjorkman 1960; Leake
both the genus and species level (Pearson and 1994).
Read 1973; Hutton 1997). Morphologically and Fire is a major formative influence on vegetation
culturally distinct groups of fungi have been of the Mediterranean-type ecosystems in Australia
isolated from hair root material of Styphelioideae and South Africa in which many Styphelioideae
across southern Australia, but most had little in and Erica occur. Growth, morphology, carbohy-
common with Hymenoscyphus ericae and Oidia- drate storage and accumulation, and partitioning
dendron spp. that infect other Ericaceae (Hutton of biomass differ between fire-sensitive (killed by
1997; McLean 1999). Basidiomycetes have been fire) and fire-tolerant (surviving fire) plants (Bell
implicated in the formation of arbutoid and and Pate 1996b; Bell and Ojeda 1999). Although
monotropoid mycorrhizae (Read 1983; Leake most species are fire-sensitive or "seeders", the
1994; Cullings et al. 1996), and Bidartondo and remainder, the "resprouters", survive the fire. A
Bruns {2001; see also Kretzer et al. 2000) found seeder typically has a single main stem associated
strong specificity in the relationship between with a compact canopy and limited lateral root
fungus and host in Monotropeae and Pleuri- extension, while a resprouter generally shows a
cosporeae, at least. One (or more, but similar) spreading, multi-stemmed canopy with large-
species of fungus is associated with the host diameter roots or lignotuberous root stocks (Bell
throughout its range, different hosts being associ- and Ojeda 1999). However, there are intermediate
ated with different basidiomycetes. When different forms of fire response types (facultative seeder
hosts grow together, they maintain their associa- sprouters) and a great deal of infraspecific varia-
tion with their "own" species of fungus. tion, particularly in southern African ericas.
Seeder species have much smaller starch reserves Tristan da Cunha and the south of South America.
in roots than resprouters. This starch is mostly A few Ericaceae ( Vaccinium spp., related to the
confined to rays of xylem parenchyma and scat- widespread north temperate/ Arctic section Myr-
tered groups of inter-ray parenchyma, and the tillus, some Styphelioideae) grow in Hawaii, and
usually much greater storage capacity of roots of several Styphelioideae and two Vaccinieae in New
resprouters is associated with their broader rays Caledonia. There are few dates for the events
(Bell and Pate 1996b; Bell and Ojeda 1999). In causing these patterns, but relationships within
Styphelioideae, amounts of starch in shoots are Arbutoideae suggest a vicariant event between
similarly low in both seeders and resprouters, western North America and the Mediterranean at
but seeders tend to begin flowering earlier (within
the Palaeogene/Neogene boundary; western North
one to three years after germination) and achieve American species of Arbutus were not involved
at least a three-fold or greater shoot: root dry (Hileman et al. 2001).
weight ratio (Bell and Pate 1996b ). Other Ericaceae Within Vaccinieae, the clade of Paphia and
have lignotubers that also sprout after fire or inDimorphanthera, with some 100 species, is
response to damage to the above-ground parts restricted to Papuasia and the west Pacific
(e.g. Judd 1981). (Dimorphanthera is not immediately related to the
South-western Australian New World Satyria, cf. Stevens 1974), while
Styphelioideae
resprout in several ways. The shoot bud bank can another probable clade including many species
be exhausted while not fully depleting the starch currently placed in Vaccinium and also Agapetes s.
reserves held in the root stock - "bud-limited" str. (= Agapetes below), some 400+ species, grows
sprouting, as in Leucopogon verticillatus. Alterna-
from China and the Himalayas to eastern Malesia.
tively, root starch may be totally depleted before the
Many tropical American Vaccinieae are related
shoot bud bank is exhausted - "energy reserve- with Andean and Central America-Caribbean sub-
limited" sprouting, as in Conostephium pendulum groups (Kron et al. 2002b).
(Bell and Pate 1996b). Leucopogon striatus, com- Massings of rather closely related species are
monly found in banksia woodlands of Western common. These include the ca. 400 species of
Australia, shows a third resprouting pattern, the Agapetes s.l. from Southeast Asia and Malesia, and
potential for limited resprouting during the firstthe ca. 285 species of Rhododendron sect. Vireya
(monophyletic) in Malesia, with only a few else-
few years after seedling establishment, followed by
a loss of this ability as the plant matures (Bell1995).
where. Styphelioideae are especially diverse in
Ericaceae often tolerate heavy metals. In Australia, where nine genera containing about 60
California they are notable components of ser- species are endemic in the south-west; Luteyn
pentine vegetation, while in Papua New Guinea (2002) discusses diversification and diversity in
Rhododendron beyerinckianum can grow directly neotropical Vaccinieae. An even more remarkable
on ultramafic rock. In New Caledonia, Dracophyl- aggregation is the 207 species of Erica found in less
lum and Styphelia (Leucopogon) are prominent on than 625km2 of the south-western Cape (Oliver
maquis on cuirasse derived from serpentine rocks. et al. 1983), with over 450 species in the south-
Ericaceous shrubs are often found in active vol- western Cape as a whole (Rebelo et al. 1985).
canic areas in both Central America and Malesia. Species of Agarista and Gaylussacia sect. Gaylus-
sacia form smaller aggregations in eastern Brasil,
Lateritic soils are the habitat of some Lyonia in the
Caribbean (Judd 1981) and of Styphelioideae in while Lyonia is diverse in the Antilles.
Australia. In general, the greatest density and presumably
also diversification of Ericaceae is in Mediter-
DISTRIBUTION. Ericaceae show a diversity of ranean climates or in high-rainfall temperate to
distributional patterns. These include an amphitropical climates in geologically young and active
south Atlantic disjunction within Agarista areas with high relief. The great majority of
(Stevens 1970b; Judd 1984), an amphi-north species in the family is found in these areas.
Atlantic disjunction within Corema, a disjunction Heads (2003) has produced a lengthy panbio-
between North and South America (Gaylussacia), geographic analysis of Malesian Ericaceae which
and numerous north-temperate disjunctions appeared too late for incorporation into the
within Rhododendron, Lyonia, Elliottia, and account.
Epigaea. Within Styphelioideae, there are several
vicariant relationships between taxa growing in FossiL RECORD. Much of the fossil record of
the SW and SE of Australia (Smith-White 1948a). Ericaceae is of leaf impressions and of doubtful
Empetrum is widely disjunct, growing in the utility (e.g. Stevens 1970b). However, reports of
northern hemisphere and the Falkland Islands, large-leaved species of Kalmia like K. cuneata
Ericaceae 161
from Europe (Fischer 1992) are of great biogeo- A few Styphelioideae are grown as container
graphical significance, such species currently plants. However, there are propagation problems
being restricted to North America. There are seeds (McLean 1999); seeds are difficult to germinate
and/or fruits of several other taxa no longer and cutting success is variable (Thompson 1986;
growing there, e.g. Zenobia, Lyonia, Eubotrys, Williams 1986), although the latter is improved by
?Enkianthus; Epacricarpidium from the Middle mycorrhizas (McLean 1995).
Miocene from Denmark (Friis 1985); Leucothoe- Blueberries ( Vaccinium sect. Cyanococcus ),
like seeds from the Maastrichtian from central bilberries (Vaccinium sect. Myrtillus), cranberries
Europe (Knobloch and Mai 1986); and seeds of (Vaccinium sect. Oxycoccus, see Eck 1990) and, to
Rhododendron from Palaeocene deposits in a lesser extent, cowberries or lingonberries (V.
England (Collinson and Crane 1978). Styphe- vitis-idaea) are cultivated or gathered in the wild
lioideae have a sparse fossil record (see Jordan and and used for jams and jellies, and even Christmas
Hill 1996). The record extends back to the Early ornaments (cranberries). Astroloma humifusum
Tertiary and possibly the Late Cretaceous and is has potential for the "bush food" industry; its
concentrated in south-eastern Australia, particu- fruits have been used for jam. The fruits of the
larly Tasmania. Generalised ericalean pollen from Andean Gaultheria myrsinoides are reported to be
the Late Cretaceous may indicate the first appear- variously hallucinogenic, poisonous, or harmless
ance of Styphelioideae, and more distinctive (Luteyn 1995, under Pernettya), and those of
epacrid pollen appears in the Middle Eocene. Comarostaphylos discolor (Diggs 1995) may also
Fossil leaves of Richeeae and Epacrideae are be poisonous. Andromedotoxins can cause serious
known from the early Oligocene, and of Styphe- poisoning or even death (Ewan-Nyambi et al.1993
lieae from the latest Oligocene-early Miocene. and references therein), while the honey of Rhodo-
Trochocarpa and Cosmelieae (probably Spren- dendron can be fatal, as Xenophon found in his
gelia) are at least as old as the Early Pleistocene, retreat across Asia Minor.
the former possibly the Late Oligocene/Early There are more minor uses of the family, and
Miocene. Monotoca pollen is known from the mid- there are many records of its use in local pharma-
Miocene. Endocarps identified as Epacridaceae copeias. For instance, tea has been made from the
from the Eocene in England should be studied leaves of Rhododendron sect. Ledum and from
further. Pollen probably of Ericeae is found Vaccinium arctostaphylos. Gaultheria procumbens
between 71 and 64Ma in Namaqualand, although was an important source of salicylic acid before it
similar pollen is known elsewhere as early as the was made by synthesis; G. shallon and Kalmia
middle Cretaceous (Scholtz 1985). latifolia are much used as "greens" by florists in
The discovery of the minute (<2 mm long) North America; the root burls of Erica arborea and
flower buds of Paleoenkianthus from the Turonian E. scoparia are used for briar (from "bruyere",
(90Ma B.P.) is more tantalizing than anything else. heather) pipes (Oliver 1991). Calluna vulgaris
The plant has anthers suggestive of those of ("Scotch heather"), grazed by sheep and grouse
Enkianthus, but it also has viscin threads; features alike, is an important element in the economic and
of the pollen surface and the stigma-style complex social fabric of much of highland England and
are not known from any extant Ericaceae (Nixon Scotland.
and Crepet 1993). The Cretaceous Ericalean flora
of the Atlantic Coastal Plain is very diverse (Crepet
CoNSERVATION. In Australia, more than 100
et al. 2001), and we may need to change our ideas species of Styphelioideae are considered under
on the evolution of the family when we understand threat from various sources (Briggs and Leigh
this flora better. 1989). Most occur in the south-west of Western
Australia (Keighery 1996 lists five species as
ECONOMIC IMPORTANCE. Rhododendron is one of Declared Rare Flora). Pathogenic Phytophthora
the more popular ornamental shrubs in north spp. are now widespread in southern Australia and
temperate gardens. Many cultivars have been are a threat to Styphelioideae and many other
developed and hybridization is common; kurume species. In Western Australia, Coleanthera virgata
azealeas (R. kaempferi x R. kiusianum) have been and Leucopogon cryptanthus have not been seen
developed in Japan. Tropical species of Rhododen- since the types were collected in the 1840s and
dron sect. Vireya also have their devotees. are considered extinct. In Victoria Choristemon
However, naturalised R. ponticum forms almost humilis, now assigned to Leucopogon, has not been
impenetrable thickets over substantial areas of the collected since 1923 and is also thought to be
western British Isles. Genera like Pieris, Kalmia extinct. Several Erica spp. in South Africa are very
and Erica are grown for their flowers and foliage. localized and are in danger of extinction.
CONSPECTUS OF ERICACEAE 3. Plant lacking multicellular hairs; axes nodding

12. Monotropopsis
I. Subfamily Enkianthioideae - Plant with multicellular hairs; axes erect 4
Genus 1. 4. Axis slender, bracts not overlapping; corolla urceolate;
II. Subfamily Monotropoideae anthers with long spurs; seeds winged 6. Pterospora
1. Tribe Pyroleae - Axis stout, bracts overlapping; corolla campanulate;
Genera 2-5. anthers with very short spurs; seeds not winged
2. Tribe Pterosporeae 7. Sarcodes
Genera 6-7. 5. Axes nodding 6
3. Tribe Monotropeae - Axes erect 8
Genera 8-16. 6. Style continuous; fruit baccate 11. Monotropastrum
- Style impressed; fruit a capsule 7
III. Subfamily Arbutoideae 7. Flowers single, white 10. Monotropa
Genera 17-20. - Flowers 2-4, purplish 14. Hypopitys
IV. Subfamily Cassiopoideae 8. Style impressed 8. Allotropa
Genus 21. - Style continuous 9
9. Petals fused IS. Hemitomes
V. Subfamily Ericoideae - Petals free 10
1. Tribe Bejarieae 10. Flowers 3-merous; filaments flattened 9. Cheilotheca
Genera 22-24. - Flowers 4-5-merous; filaments terete 11
2. Tribe Rhodoreae 11. Anthers elongated, not inverting, with two long slits
Genera 25-28. 16. Pleuricospora
3. Tribe Phyllodoceae - Anthers hippocrepiform, inverting, with a single apical
Genera 29-34. slit 13. Pityopus
4. Tribe Empetreae 12. Plants ± herbaceous; petals free 13
Genera 35-37. - Plants woody; petals usually fused 16
5. Tribe Ericeae 13. Plant with erect, leafy stems; inflorescence umbellate
Genera 38-40. 4. Chimaphila
VI. Subfamily Harrimanelloideae - Plant with rosette of leaves ± at ground level; inflores-
Genus 41. cence not umbellate 14
14. Inflorescence with >6 flowers, racemose 15
VII. Subfamily Styphelioideae - Inflorescence of a single flower 3. Moneses
1. Tribe Prionoteae 15. Lamina finely serrate; inflorescence secund 5. Orthilia
Genera 42-43. - Lamina at most obscurely crenate; inflorescence with
2. Tribe Archerieae flowers all around the axis 2. Pyrola
Genus 44. 16. Ovary superior (if inferior, then anthers with awns) 17
3. Tribe Cosmelieae - Ovary inferior 91
Genera 45-47. 17. Flowers nearly always 4-merous; corolla ± scarious,
4. Tribe Epacrideae persistent; leaves opposite or whorled, nearly always
Genera 48-52. ericoid 18
5. Tribe Oligarrheneae - Flowers rarely 4-merous; corolla not scarious; leaves
Genera 53-54. usually spiral, rarely ericoid 19
6. Tribe Richeeae 18. Leaves sessile, tailed 38. Calluna
Genera SS-57. - Leaves petiolate, not tailed 40. Erica
7. Tribe Styphelieae 19. Fruit septicidal; pollen usually with viscin threads 20
Genera 58-76. - Fruit loculicidal, or indehiscent; pollen lacking viscin
VIII. Subfamily Vaccinioidaee threads 34
1. Tribe Oxydendreae 20. Leaves opposite or whorled; anthers usually with slits
Genus 77. their entire length 21
2. Tribe Lyonieae - Leaves scattered to pseudo-verticillate; anthers rarely
Genera 78-81. with slits their entire length 22
3. Tribe Andromedeae 21. Lamina ericoid 24. Ledothamnus
Genera 82-83. - Lamina with recurved margins, but not ericoid
4. Tribe Gaultherieae 35. Kalmia (part)
Genera 84-89. 22. Inflorescence bud with large, usually brown perulae;
5. Tribe Vaccinieae bracts and bracteoles usually deciduous 23
Genera 90-124. - Inflorescence bud lacking such perulae; bracts and
bracteoles usually persistent 25
23. Inflorescence axis elongated, with leaves below the
KEY TO THE GENERA (to those of Styphelioideae
flowers; bracts and prophylls leafy 25. Therorhodion
partly based on Powell 1983) - Inflorescence axis not elongated, lacking leaves below the
flowers; bracts and prophylls not leafy 24
1. Plants without chlorophyll 2 24. Capsule subspherical 27. Menziesia
- Plants with chlorophyll 12 Capsule longer than wide 28. Rhododendron
2. Sepals very different from petals, the latter fused 3 25. Petals free 26
- Two perianth whorls not clearly distinguishable, inner Petals fused 28
members usually free 5
Ericaceae 163
26. Leaves ericoid; vegetative plant prostrate 23. Bryanthus - Corolla >5 mm long, with distinctly revolute lobes 48
- Leaves rarely ericoid; plant erect 27 48. Leaves scattered along stem 75. Styphelia
27. Filaments± terete, anthers with terminal pores - Leaves distinctly clustered in false whorls
22. Bejaria 65. Cyathodes
- Filaments flattened, anthers with elongated slits 49. Corolla tube conical in upper part, lobes very small, erect;
29. Elliottia anthers 2-lobed 63. Conostephium
28. Anthers with slits at least half their length 29 - Corolla tube cylindrical, campanulate or rotate, lobes
- Anthers with terminal pores or short slits 30 spreading or recurved; anthers rounded SO
29. Lamina <2 em long; antesepalous stamens strongly SO. Corolla tube with tufted hairs or scales below middle
epipetalous 26. Diplarche inside; filaments usually flat 51
- Lamina >5 em long; all stamens free from petals - Corolla tube glabrous below middle inside, rarely puberu-
30. Epigaea lent or pubescent; filaments terete, usually thin 52
30. Corolla with 10 pouches 34. Kalmia (part) 51. Corolla tube short, broad, 5 tufts of hairs or ring of glands
- Corolla lacking pouches 31 at base, lobes spreading 71. Melichrus
31. Lamina ericoid, or with strongly recurved margins 32 - Corolla tube elongated, cylindrical, 5 tufts of hairs or
- Lamina plane 33 fringed scales in tube, lobes erect around anthers, spread-
32. Lamina ericoid; style impressed 31. Phyllodoce ing or recurved at top 60. Astroloma
- Lamina with strongly recurved margins; style and ovary 52. Locules often more than 5; fruit with separate stones
continuous 39. Daboecia 53
33. Lamina with ciliate margins; plant lacking sessile glan- - Locules usually 5 or fewer; fruit with a single stone 55
dular hairs 32. Rhodothamnus 53. Corolla valvate in bud; filaments thin, terete
- Lamina lacking ciliate margins; plants with sessile glan- 73. Pentachondra
dular hairs 33. Kalmiopsis - Corolla ± imbricate in bud; filaments short, thick and
34. Lamina rarely xeromorphic, margin often with gland- tapered 54
tipped teeth, venation pinnate, midrib usually evident; 54. Adaxial surfaces of corolla lobes with hairs towards the
plant nearly always with multicellular hairs 67 base 74. Trochocarpa
- Lamina xeromorphic, with entire margins, if serrulate, - Adaxial surfaces of corolla lobes with widespread hairs
not gland-tipped, venation closely parallel- or pH-nerved, 67. Decatoca
midrib not evident; plant lacking multicellular hairs 35 55. Flowers pedicellate, prophylls basal 70. Lissanthe
35. Stems with annular scars after leaf fall 36 - Flowers sessile, prophylls inserted immediately below
- Stems without annular scars after leaf fall 38 calyx 56
36. Corolla ovoid or conical, forming a cap that falls at anthe- 56. Ovary and style with hairs 64. Croninia
sis, the lobes not opening 56. Richea - Ovary and style glabrous 57
- Corolla cylindrical or campanulate, not shed at anthesis, 57. Corolla lobes imbricate in bud 61. Brachyloma
the lobes opening 37 - Corolla lobes valvate in bud 58
37. Flowers in short spikes; bracts persistent; corolla tube 58. Corolla tube elongate 59
narrow 57. Sphenotoma - Corolla tube short; filaments terete or filiform
- Flowers in compound racemes or panicles, rarely solitary; 68. Leptecophylla
bracts falling; corolla broadly cylindrical or campanulate 59. Corolla lobes glabrous or papillose; ovary 1-2-locular
55. Dracophyllum 72. Monotoca
38. Leaves sheathing 39 - Corolla lobes hairy, rarely glabrous; ovary (2-)5(-10)-
- Leaves not sheathing 41 locular 60
39. Filaments adnate to corolla tube 45. Cosmelia 60. Corolla lobes with erect tufts of hairs near apex and fine
- Filaments free 40 hairs at throat; flowers often yellow-green to green
40. Corolla glabrous; tube very short; lobes imbricate; 58. Acrotriche
anthers connivent or cohering 46. Sprengelia - Corolla lobes sparsely to densely bearded, rarely glabrous;
- Corolla hairy inside; tube cylindrical; lobes imbricate; flowers white, cream or red 61
anthers free 47. Andersonia 61. Drupe± spherical; mesocarp fleshy or thin
41. Flowers 4-merous 42 69. Leucopogon
- Flowers 5-merous 43 - Drupe depressed; mesocarp thick, pulpy
42. Stamens 4 66. Cyathopsis 74. Planocarpa
- Stamens 2 54. Oligarrhena 62. Flowers pedicellate, bracteoles two, basal, quite different
43. Fruit drupaceous, indehiscent, rarely splitting irregularly from sepals 44. Archeria
and transversely 44 - Flowers sessile, bracteoles several, imbricate, passing into
- Fruit a loculicidal capsule 62 ~ili ~
44. Corolla lobes incurved at the apex, style impressed into 63. Corolla lobes contorted in bud 64
ovary 53. Needhamiella - Corolla lobes imbricate in bud 65
- Corolla lobes rarely incurved; style and ovary continuous 64. Lamina ovate, acuminate, pungent, erect above base then
45 spreading widely 52. Woollsia
45. Anthers exserted above corolla tube 46 - Lamina linear to ovate-lanceolate, erect, appressed to
- Anthers included within corolla tube, or lying between stem 50. Lysinema
spreading lobes 49 65. Filaments long, inserted at base of corolla tube; anthers
46. Anthers connate in a cone around style adnate, connivent around style 51. Rupicola
62. Coleanthera - Filaments short, inserted in corolla throat; anthers
- Anthers free 47 attached above middle 66
47. Corolla <1.6 mm long, the lobes spreading 66. Filaments terete, shorter than anthers; anthers dorsally
59. Androstoma attached to filament, included 49. Epacris
- Filaments compressed, about as long as anthers; anthers - Inflorescence fasciculate; lamina with fibres free in
adnate to filament for most of their length, exserted mesophyll (break lamina!), lacking marginal glands
48. Budawangia 88. Diplycosia
67. Lamina ericoid and <2cm long, fruit a drupe 68 88. Fruit fleshy, or fruit dry but calyx fleshy
- Lamina rarely ericoid-like, if so, >3 em long, or fruit a 87. Gaultheria
capsule 70 - Fruit and calyx dry 89
68. Inflorescence capitulate 36. Corema 89. Corolla >1 em long, campanulate 83. Zenobia
- Inflorescence not capitulate 69 - Corolla <1 em long,± urceolate 90
69. Lamina with wooly hairs in abaxial groove; style longer 90. Prophylls basal 85. Leucothoi!
than stigmatic lobes 37. Ceratiola - Prophylls apical 86. Eubotrys
- Lamina lacking wooly hairs in abaxial groove; style 91. Filaments and/or anthers of alternating stamens strongly
shorter than stigmatic lobes 35. Empetrum unequal 92
70. Inflorescence terminal; ovary superior; fruit fleshy, - Filaments and anthers of alternating stamens similar to
usually indehiscent 71 somewhat unequal (but spurs may differ considerably)
- Inflorescence usually axillary; fruit proper (i.e. not 95
including calyx) fleshy only if ovary is inferior 74 92. Filaments unequal; anthers not notably robust 93
71. Leaves in whorls of three 19. Ornithostaphylis - Filaments ± equal; anthers rather woody, the tubules
- Leaves scattered 72 widening distally 94
72. Ovary smooth 20. Arctostaphylos 93. Anthers equal; stamens <1/2 the length of the corolla;
- Ovary papillate 73 bracts small or early deciduous 102. Orthaea
73. Ovary with several ovules/loculus; fruit a berry; lamina - Anthers unequal; stamens as long as corolla (-1/2 as
usually <3 times longer than broad 17. Arbutus long); bracts usually large and persistent
- Ovary with 1 ovule/loculus; fruit a drupe; lamina usually 101. Cavendishia
>3 times longer than broad 18. Comarostaphylis 94. Filaments connate their entire length (New World)
74. Lamina <1 em long 75 105. Satyria
- Lamina > 1 em long 76 - Filaments free (Old World) 98. Dimorphanthera
75. Leaves decussate; flowers axillary 21. Cassiope 95. Fruit pyrenoid 124. Gaylussacia
- Leaves spiral; flowers terminal 41. Harrimanella - Fruit baccate 96
76. Stamens with geniculate filaments and/or with spurs 77 96. Anthers with slits the length of the thecae, tubules minute
- Stamens with ± straight filaments, awned or not 80 94. Lateropora
77. Corolla urceolate to campanulate; seeds with wing on one - Anthers with pores or short slits, tubules usually well
side 78. Craibiodendron developed 97
- Corolla urceolate to tubular; seeds unwinged, spindle- 97. Anther tubules with other than terminal or introrse pores
shaped or rounded 78 or short clefts, much shorter than the thecae 98
78. Capsule with much thickened, often whitish ribs, or - Tubules with terminal or introrse pores or short clefts
filaments with delicate spurs that have disintegration (if lateral, then tubules longer than thecae) 99
tissue 81. Lyonia 98. Anther tubules with short, extrorse clefts; pedicels stout
- Capsule lacking thickened ribs; stamen spurs lacking 95. Notopora
disintegration tissue 79 - Anther tubules with short, latrorse clefts; pedicels slender
79. Lamina with veins on the abaxial surface graded in to stout 93. Didonica
prominence; stamens spurred 79. Pieris 99. Prophylls at apex of pedicel enveloping ovary (often with
- Lamina with veins on the abaxial surface about equal in inflorescence bracts); flowers (3)4(5)-merous
prominence; stamens without spurs 80. Agarista 114. Disterigma
80. Inflorescence terminal; anthers with a single pair of - Prophylls rarely at apex of pedicel, not enveloping ovary;
awns 81 flowers usually 5-merous 100
- Inflorescence rarely terminal; if so, then anthers not with 100. Ovary 5-locular, with 5 additional inpushings; cork-
awns 84 cambium superficial (surface of leafy stems lacking
81. Inflorescence perulate; anthers with slits for much of their longitudinal cracks) 92. Agapetes
length 1. Enkianthus - Ovary usually simply 5-locular; cork cambium deep-
- Inflorescence not perulate; anthers with short, terminal seated, or more superficial, but localised (surface of leafy
slits or pores 82 stems often developing longitudinal cracks) 101
82. Indumentum of scales; inflorescence a foliaceous raceme 101. Stamens 5-10; calyx lobes free, 5-9mm long
84.Chamaedaphne 111. Oreanthes
- Indumentum never of scales; inflorescence racemose, not - Stamens (4, 5, 8) 10; calyx lobes usually smaller, connate
foliaceous 83 or not 102
83. Inflorescence >10cm long, branched 77. Oxydendrum 102. Calyx continuous with pedicel 103
- Inflorescence <3 em long, unbranched 82. Andromeda - Calyx articulated with pedicel 117
84. Anthers with slits their entire length 85 103. Anther tubules elongate, half as wide (or less) than thecae
- Anthers with pores or short slits 86 104
85. Corolla <5 mm long 42. Lebetanthus - Anther tubules at most slightly longer than thecae, rarely
- Corolla >10mm long 43. Prionotes thin, if elongate, more than half as wide as thecae 105
86. Anthers with terminal tubules, lacking awns 87 104. Calyx lobes >2mm long; filaments with tuft of retrorse
- Anthers usually lacking terminal tubules, frequently hairs on back 113. Pellegrinia
awned 88 - Calyx lobes <1 mm long; filaments without tuft of retrorse
87. Inflorescence racemose, rarely paniculate; lamina lacking hairs on back 110. Semiramisia
fibres in mesophyll, with one or more pairs of marginal
glands near base 89. Tepuia
Ericaceae 165
105. Calyx and corolla conspicuously winged or rarely angled 122. Anthers spurred, if not, connective thickened; tubules dis-
106 tinct; corolla never angled, often abruptly constricted
- Calyx terete or angled, rarely winged; corolla terete 108 104. Psammisia
106. Leaves usually opposite or pseudoverticillate; corolla - Anthers not spurred; tubules often connate; corolla
wings quite broad distally 122. Anthopterus (part) angled or not, gradually narrowed distally
- Leaves scattered; corolla wings at most narrow distally 103. Macleania
107 123. Corolla ca. 4mm long, tube with solid spurs opposite the
107. Corolla terete 118. Themistoclesia (part) lobes; calyx with wings alternating with the lobes
- Corolla narrowly winged, wings broadest at the base 96. Utleya
122. Anthopterus (part) - Corolla often longer, tube lacking spurs; calyx usually not
108. Calyx angled, rarely winged 109 winged 124
- Calyx terete 112 124. Corolla urceolate to campanulate, <12 mm long (-22 mm
109. Calyx angles opposite the lobes 107. Polyclita long, tubular), the lobes often imbricate, thin and drying
- Calyx angles alternating with the lobes 110 membranaceous, or flowers polypetalous; filaments often
110. Calyx limb l<cm long 100. Agapetes scortechinii longer than the anthers 90. Vaccinium (part)
- Calyx limb <4 mm long Ill - Corolla± tubular, (8-)12mm long, the lobes thick, drying
111. Lamina linear, with midrib alone evident 117. Rusbya coriaceous, valvate; filaments shorter than the anthers
- Lamina not linear, with more complex venation 125
118. Themistoclesia (part) 125. Anther tubules usually 2-5 times longer than thecae;
112. Corolla (I0-)14mm long, carnose to coriaceous; embryo calyx limb often 2-4 times longer than tube
white 113 119. Plutarchia
- Corolla <lOmm long (if longer, then filaments - Tubules up to 2 times longer than thecae; calyx limb as
longer than anthers), thinly fleshy; embryo green (white) long as or shorter than tube 126
114 126. Calyx with wings longer than lobes; anthers alternately
113. Lamina 8 em long; inflorescence racemose, flowers spurred; stamens about half as long as corolla
numerous; anther tubules at most 2 x length of thecae 108. Anthopteropsis
120. Thibaudia (part) - Calyx with at most obscure wings; anthers lacking spurs;
- Lamina <3( -6) em long; inflorescence fasciculate, flowers stamens about as long as corolla 127
1-3 (racemose, flowers numerous); an,tber tubules 2-5 x 127. Flowers 6-8-merous; corolla flaring 91. Symphysia
longer than thecae 121. Demosthenesia - Flowers 5-merous; corolla urceolate to tubular 128
114. Pedicels usually long and filiform, cernuous; corolla urce- 128. Calyx limb to as long as tube; filaments much shorter to
olate; filaments usually longer than anthers longer than anthers; tubules to 2.5x longer than thecae
115. Sphyrospermum 99. Paphia
- Pedicels at most thin, not cernuous; corolla various; - Calyx limb often longer than tube; filaments much
filaments usually shorter than anthers 115 shorter than anthers; tubules about as long as thecae
115. Lamina pli-nerved, with prominent marginal glands near 120. Thibaudia (part)
base, lacking surface glandular hairs; inflorescence fasci-
culate 123. Costera GENERA OF ERICACEAE
- Lamina pli- or penni-nerved, lacking marginal glands
near base, usually with surface glandular hairs; inflores- Basic features
cence various 116 Plant woody; hair roots invested by fungal hyphae
116. Lamina ± pli-nerved; corolla tubular, rarely urceolate;
embryo green 116. Diogenesia that form a Hartig net; pericyclic fibres at most
- Lamina ± penni-nerved; corolla urceolate, rarely cam- poorly developed; node unilacunar; pith homoge-
panulate; embryo white 90. Vaccinium (part) neous; bud scales present; leaves spiral, convolute
117. Anther tubules elongate, half as wide (or less) than thecae in bud, scattered, with midrib and reticulate vena-
118
- Anther tubules at most slightly longer than thecae, rarely
tion; inflorescence terminal; flowers 5-merous,
thin, if elongate, more than half as wide as thecae 120 pedicels present, not articulated with calyx, pro-
118. Corolla with lobes at least 4mm long, the base often phylls basal; corolla sympetalous; anthers with
ventricose 109. Ceratostema slits, endothecium present; nectary at base of
- Corolla with lobes <2 mm long, the base never ventricose ovary; placentation axile, numerous ovules/locule;
119
119. Filaments connate; anther pores terminal
capsule loculicidal; raphe with vascular bundle;
112. Siphonandra embryo white, terete.
- Filaments free or connate; anther pores oblique, or Note: these features can be understood to be
tubules with short slits 97. Gonocalyx applicable to all genera of the family, unless mod-
120. Thecae conspicuously papillate; tubules rigid, connate or ified in the subfamilial or tribal characterizations,
not; stamens often <2/5 the length of corolla 121
- Thecae smooth or papillate; tubules flexible; stamens and the characters in these behave similarly.
often as long as corolla 123
121. Calyx turbinate, with wings or angles opposite the lobes;
connective extending dorsally up each anther tubule I. SUBFAMILY ENKIANTHOIDEAE Kron, Judd &
106. Mycerinus Anderberg (2002).
- Calyx with wings (if present) alternating with the lobes;
connective terminating at the bases of anther tubules Enkiantheae P.F. Stevens (1971).
122
Fig. 55. Ericaceae. A-F Enkianthus campanulatus. A Flower- of capsule more or less thickened; seeds winged or
ing branchlet. B Flower. C Same, opened, showing awned not; n = 11. 16 spp., four sections, E Asia; low to
anthers. D Stamen with inverted, awned anther; one of the
thecae dehiscing with longitudinal slit. E Capsules. F Seed. G
mid altitude.
Enkianthus perulatus, stamen, the anther with protruding
connective. Drawing by Polyanna von Kni:irring. (Anderberg All other Ericaceae
1994b)
Raphe lacking vascular bundle; exothecium
present.
Pith heterogeneous; inflorescence perulate; calyx
valvate; anthers with paired awns, megagameto-
phyte with ears; testa cells not elongated, rather II. SUBFAMILY MONOTROPOIDEAE Arnott
tall. (1832).
1. Enkianthus Loureiro Fig. 55 Pyroloideae Kostel. (1834).
Enkianthus Loureiro, Fl. Coch. 1: 339 (1793); Anderberg, Herbs; multicellular hairs absent; inflorescences
Nordic J. Bot. 14: 385-401 (1994).
racemes, eperulate.
(Sub )deciduous shrubs; leaves pseudoverticillate;
inflorescences perulate, corymbose to umbellate, 1. TRIBE PYROLEAE Dumortier (1829).
rarely paniculate, with terminal flower, bracts and
prophylls poorly developed; corolla campanulate Rarely subshrubs; fungal hyphae with complex,
to urceolate, lobed at most ca. 1/3; anthers smooth coiled intrusions into exodermal cells of hair
apart from awns; style impressed or not; sutures roots; multicellular hairs absent; leaves pseudover-
Ericaceae 167
ticillate, ± serrate; flowers weakly monosymmetri- 2. TRIBE PTEROSPOREAE Baillon (1891).

cal or not, prophylls absent; petals free; filaments
thin, anthers with short terminal tubules (rarely £chlorophyllous; glandular multicellular hairs
absent) and pores; testa cells moderately elon- present; leaves entire; axes pink to reddish, many-
gated, little thickened; embryo small, hardly flowered; petals fused; anthers with 2 spurs; style
differentiated. impressed, rather short.
2. Pyrola L. 6. Pterospora Nuttall

Pyrola L., Sp. Plant. 1: 396 (1753); Krisa, Bot. Jahrb. Syst. 90: Pterospora Nuttall, Gen. N. Am. Pl. 1: 269 (1818); Wallace,
476-508; Haber, Syst. Bot. 12: 324-335 (1987), Can. J. Bot. 66: Wasmann J. Bioi. 33: 63-68 (1975).
1993-2000 (1988).
Rootballs tight; axes pink to reddish; corolla urce-
Rhizomatous herb; leaves in rosette, thick; pollen olate; stamens 1/2 as long as corolla, anthers long-
in tetrads; no disc; style with collar at the spurred, inverting late?, with apical and lateral
apex; capsule valves with cobwebby edges; testa slits; nectary not obvious; fruit dehiscing from the
cells with moderate-sized to no plasmodesmata; base first; seeds winged, cells somewhat elongated,
n = 23, 46. Ca. 30 spp., north temperate, Sumatra, thin-walled; n = 8. One sp., P. andromedea Nutt., W
Guatemala. Infrageneric classification and species North America including Mexico, the Great Lakes
limits difficult; the anthers of P. minor L. lack region; 60-3675 m alt.
tubules, the buds of P. chlorantha lack scales.
7. Sarcodes Torrey
3. Moneses S.F. Gray
Sarcodes Torrey, Smithson. Contrib. Know. 6: 18, pl. 10 (1853);
Moneses S.F. Gray, Nat. Arr. Brit. Pl. 2: 396,403 (1821). Wallace, Wasmann J. Bioi. 33: 68-72 (1975).
Herb, shoots arising from roots; leaves in rosette, Roots fibrous; axes red to orange; corolla campan-
thick; flower single, on long stalk, bract somewhat ulate, slightly saccate at base; stamens ca. 2/3
below flower; pollen in tetrads; no disc; style with length corolla, anthers elongated, with very small
collar at the apex; testa cells with thin scalariform spurs, with terminal extrorse slits, nectaries with
thickenings; n = 13. 2 spp., north temperate. low lobes; fruit with irregular dehiscence; seeds
subovoid, cells not elongated, thick-walled; n = 32.
One sp., S. sanguinea Torrey, USA (California and
4. Chimaphila Pursh
adjacent Oregon) and Mexico (Baja California);
Chimaphila Pursh, Fl. Am. Sept. 1: 279 (1814). 1070-3050m alt.
Rhizomatous subshrub; leaves coriaceous; inflo-

rescence a corymb, bracts ± borne up pedicel; 3. TRIBE MONOTROPEAE Dumortier (1829).
pollen in tetrads to monads; style short, with collar
at the apex; testa cells with small or no plasmod- £chlorophyllous; shoots arising from roots; fungal
esmata; n = 13. Five spp., boreal and north-tem- hyphae forming peg-like inpushings into exoder-
perate, S to Guatemala and Hispaniola. Pollen in mal cells of hair roots; roots rarely endogenous;
tetrads, monads or polyads. leaves entire; bracts (sub )persistent, as large or
larger than and as conspicuous as sepals; prophylls
usually absent; sepals usually as large as
5. Orthilia Rafinesque
petals/corolla; style rather short, stigma expanded;
Orthilia Rafinesque, Aut. bot.: 103 (1840). embryo minute, undifferentiated.
Rhizomatous herb; leaves thin; inflorescence

8. Allotropa A. Gray
secund; anthers lacking tubules, pollen in monads;
disc lobed; style long, no collar at the apex; capsule Allotropa A. Gray in Newberry, Rep. Expl. Surv. Railroad 6(3):
valves with cobwebby edges; cells of testa with 80 (1858); Wallace, Wasmann J. Bioi. 33: 13-18 (1975).
moderate to small plasmodesmata; n = 19. One sp.,
0. secunda (L.) House, north-temperate. Roots fibrous; axes erect, red and white striped,
many-flowered; perianth a single whorl, 5, free,
campanulate; stamens (9)10(11), as long as peri- Laos and Vietman, Sumatra (Indonesia); 1000-
anth, anthers inverting late, with pore-like slits 2700m alt.
ca. 1/3 the length of anther; style impressed; seeds
linear, cells elongated, thin-walled; n = 13. One sp.,
A virgata A. Gray, W USA; 80-3000m alt. 12. Monotropsis Schweinitz
Monotropsis Schweinitz in Elliott, Sketch Bot. S. Carol. Georgia
1: 478 (1817); Wallace, Wasmann J. Bioi. 33: 49-53 (1975).
9. Cheilotheca J.D. Hooker
Roots coralloid; axes nodding, violet to purple,
Cheilotheca Bentham & J.D. Hooker, Gen. Plant. 2: 607 (1876);
Wallace, Wasmann J. Bioi. 33: 18-23 (1975). several-flowered; flowers (4)5(6)-merous; sepals
Andresia Sleumer, Fl. Males. I, 6: 669 (1967). present, different in texture; corolla subtubular,
petals fused, slightly saccate at the base; stamens
Root mass loose; axes erect, white to purple, !-few- ca. 3/5 the length of the corolla, with extrorse
flowered; sepals 2-5; corolla tubular-campanulate, pores; nectary crenate; ovary with largely parietal
petals 3, free, concave; stamens 6, 1/2-2/3length of placentation, style continuous; fruit baccate; seeds
corolla; anthers linear, thecae free, with long slits, ovoid, cells little elongated, thick-walled. One sp.,
or fused, hippocrepiform, with a single terminal M. odorata Schweinitz, SE USA, 600-1350 m alt.
slit; ovary 6-carpellate, placentation parietal, style
continuous; fruit baccate; seeds? Two spp., Assam
(India), the Malay Peninsula (Malaysia) and 13. Pityopus Small
Sumatra (Indonesia); 600-1500m alt. Pityopus Small, N. Am. Fl. 29: 161 (1914); Wallace, Wasmann J.
Bioi. 33: 53-58 (1975).
10. Monotropa L.
Rootballs tight; axes erect, white to yellowish,
Monotropa L., Sp. Plant. 1: 387 (1753); Wallace, Wasmann J. (1-)several-flowered, branched or not; flowers 4-
Bioi. 33: 29-42 (1975). (5- )merous; sepals present; corolla tubular, petals
free; stamens 2/3 the length of the anthers, thecae
Rootballs ± tight; axes nodding, white to reddish, horizontal, connate, hippocrepiform, with a single
!-few-flowered; flowers 3-8-merous; sepals absent apical slit; ovary with parietal placentation, style
or not; corolla tubular, petals free, saccate at base; continuous, stigma umbiculate; fruit baccate;
stamens as long as corolla, anthers horizontally seeds ellipsoid, cells not elongated, thick-walled.
reniform, with terminal slit or slits; nectary with One sp., P. californicus (Eastwood) H.F. Copeland:
paired lobes between antepetalous stamens; ovary W USA; 30-2000m alt.
with axile placentation, style impressed, stigma
umbilicate to funneliform; fruit capsular; seeds
spindle-shaped, cells elongated, thin-walled; 14. Hypopitys Crantz
n = 16, 24. 1 sp., M. uniflora L., north temperate; Hypopitys Crantz, Inst. Rei Herb. 2: 467 (1766).
0-2000m alt.
Rootballs ± tight; axes nodding, cream to red or
11. Monotropastrum H. Andres purple, (1)2- to several-flowered; flowers 4-5-
merous, subcampanulate; sepals (0- )4-5; corolla
Monotropastrum H. Andres, Notizbl. Berl.-Dahl. 12: 696, fig. 8 free, saccate at the base; stamens almost as long as
(1935); Wallace, Wasmann J. Bioi. 33: 42-48 (1975).
the sepals, anthers horizontal, thecae connate,
Eremotropa H. Andres, Bot. Jahrb. Syst.76: 103, pl. 6 (1953);
Wallace, Taxon 36: 128-130 (1987). hippocrepiform, with single terrminal slit;
nectary with paired lobes between opposi-
Root mass dense; axes nodding, white, !-few-flow- tipetalous stamens; style impressed, stigma umbil-
ered; flowers 3-5-merous; sepals absent or not, icate; seeds spindle-shaped, cells elongated,
more or less distinct or not; corolla tubular, petals thin-walled; n = 8, 16, 24. 1 sp., H. monotropa
free, more or less saccate at base; stamens as long Crantz, north temperate region, Central America;
as corolla, anthers horizontally reniform, with ter- to 400 m alt. Possibly to be divided (see Bidartondo
minal slit or slits; nectary with paired lobes and Bruns 2001).
between antepetalous stamens; ovary with parietal
placentation, style continuous, stout to slender, 15. Hemitomes A. Gray
stigma funneliform; fruit baccate; seeds ovoid,
cells reticulate. Two spp., Punjab (India) to Korea Hemitomes A. Gray in Newberry, Rep. Expl. Surv. Railroad 6(3):
80, pl. 12 (1858); Wallace, Wasmann J. Bioi. 33: 24-29 (1975).
Ericaceae 169
Roots fibrous; axes erect, cream to reddish, 1- to Trees or large shrubs; lamina often serrate, smooth
several-flowered, branched or not; flowers 4( -6)- abaxially; anthers with apical projection; ovary
merous; sepals and prophylls apparently present; papillate, ovules 2-several/carpel; fruit baccate,
corolla tubular-campanulate, petals connate; papillate; seeds few; embryo spatulate. 10 spp.,
stamens ca. 2/3 the length of the corolla, anthers British Columbia S to Nicaragua, circum-Mediter-
not inverting, with long slits; ovary 8-carpellate, ranean; seasonal climates, s.l.-3100m alt. Possibly
with parietal placentation, style continuous; fruit paraphyletic (Hileman et al. 2001).
baccate; seeds ellipsoid, testa cells little elongated,
?thick-walled. One sp., H. congesta A. Gray, W USA;
18. Comarostaphylos Zuccarini
30-2700m alt.
Comarostaphylos Zuccarini, Abh. Math.-Phys. Cl. Kiinigl.
Bayer. Acad. Wiss. 2: 331 (1837); G.M. Diggs Jr., Fl. Neotrop.
16. Pleuricospora A. Gray 66: 146-193 (1995).
Pleuricospora A. Gray, Proc. Am. Acad. Arts Sci. 7: 369 (1868);
Wallace, Wasmann J. Biol. 33: 58-63 (1975). Shrubs to trees; lamina often serrate, papillate
abaxially; anthers lacking apical projection; ovary
Roots coralloid; axes erect, cream to yellowish, ( 1) papillate, carpels 4-6, 1 ovule/carpel; fruit drupa-
to several-flowered; flowers usually 4-merous, ceous, papillate, pyrenes not separating; embryo
tubular; sepals present; petals free; stamens as spatulate. 10 spp., S California to Panama;
long as corolla, anthers not inverting, with long montane.
slits; nectary not seen; ovary with parietal placen-
tation, style continuous, stigma barely expanded; 19. Ornithostaphylos Small
fruit indehiscent; seeds ovoid, testa cells not elon-
gated, thick-walled; n = 26. One sp., P. jim brio lata Ornithostaphylos Small, N. Am. Fl. 24: 101 (1914).
A. Gray; W North America; 150-2750 malt. As with
several other Montropeae, the flower can also be Shrub; leaves whorled, lamina entire, smooth
described as lacking a corolla, but with large and but densely hairy abaxially; anthers with apical
closely-associated prophylls. projection; ovary smooth, 2 ovules/carpel; fruit
drupaceous; embryo spatulate; n = ? One sp.,
All other Ericaceae 0. oppositifolia (C. Parry) Small, S California,
Baja California.
Evergreen; fungal hyphae with complex, coiled
intrusions into exodermal cells of hair roots;
anthers without endothecium, pollen in tetrads. 20. Arctostaphylos Adanson
Arctostaphylos Adanson, Pam. Pl. 2: 165,520 (1763), nom. cons.
Xylococcus Nuttall (1889).
III. SuBFAMILY ARBUTOIDEAE (Meisner) Arctoiis Niedenzu (1890).
Niedenzu (1890).
Evergreen, rarely deciduous shrubs to small trees;
Arbuteae Meisner (1839). lamina entire to serrate, smooth abaxially; anthers
without apical projections; ovary smooth, carpels
Bark usually flaking, the surface smooth; leaves 5-10, ovules 1(2)/carpel; fruit drupaceous, 4-10
convolute in bud, often serrulate; inflorescences seeded, segments separating or not; embryo allan-
eperulate; corolla urceolate, lobes short; filaments toid, rarely spatulate. To 60 spp., mostly W USA S
swollen at the base, anthers with paired, reflexed to Guatemala, two spp. circumboreal, s.l. to alpine.
awns, almost smooth, with terminal pores or short Species limits difficult; Arctous (perulate inflores-
slits; placentae apical, style continuous, stigma cence, deciduous) and Xylococcus (spatulate
slightly lobed; fruit fleshy; testa cells moderately embryo, barely swollen filaments) often segre-
elongated, thick-walled; n = 13. Ellagic acid is gated (see also Hileman et al. 2001). The leaves
common. may be held edge on.
17. Arbutus L. All other Ericaceae
Arbutus L., Sp. Plant. 1: 395 (1753); P.D. Sorenson, Fl. Neotrop. Hartig net absent; pericyclic fibres in stem and
66: 194-221 (1995). fibres associated with leaf bundles; anthers with
pores, or slits, inverting early; style impressed.
Cassiopoideae + Ericoideae Prostrate shrub; leaves ericoid, inflorescences

Leaves entire{!). racemes, pedicels articulated; anthers with short
terminal slits, resorbtion tissue?; style continuous,
stigma subtruncate. One sp., B. gmelinii D. Don, NE
Asia. The petals may be connate basally.
IV. SuBFAMILY CASSIOPOIDEAE Kron & Judd
{2002).
24. Ledothamnus Meisner
Cassiopeae P.F. Stevens (1971).
Ledothamnus Meisner, Fl. Bras. 7: 171 (1863); Luteyn, Fl.
Neotrop. 66: 107-122 (1995), in P.E. Berry et a!. (eds) Fl.
Leaves a variant of ericoid; flowers single, axillary; Venez. Guyana 4: 745-748 (1998).
corolla campanulate, 1/3lobed; anthers with pores
and paired, reflexed awns; stigma truncate; testa Shrubs; leaves ericoid, decussate to whorled;
cells elongated, thin-walled. flowers (5)6( -9)-merous; stamens as many as
petals, rarely twice as many, filaments flattened,
21. Cassiope D. Don anthers with slits their entire length; ovary
strongly verrucose, 5{ -7)-carpellate; stigma
Cassiope D. Don, Edinburgh New Phil. J. 17: 157 (1834). subtruncate; seeds irregular, testa cells little
thickened. Seven spp., Guayana Highlands of S
Small shrubs, pith Calluna-type; leaves opposite, Venezuela and adjacent Brasil; 1000-2800m alt.
acicular, tubular or channelled and with auricles,
sessile; flowers single, axillary, with 4-6 basal
bracteoles; n = 13. Ca. 12 spp., circumboreal, most 2. TRIBE RHODOREAE DC. ex Duby {1828).
spp. Himalayas-East Asia; s.l. to alpine.
Diplarcheae Airy Shaw (1964).
V. SUBFAMILY ERICOIDEAE Link {1829) Inflorescences perulate; median petal adaxial;

("Ericeae") anthers with collapse tissue; testa cells elongated,
thin-walled. Gossypetin is present.
Subfamily Rhododendroideae (Juss.) Sweet (1828).
Anthers lacking appendages, pollen with viscin 25. Therorhodion Small

threads; fruit a septicidal capsule. Therorhodion Small, N. Am Fl. 29: 45 (1914).
Prostrate deciduous shrub; leaves revolute in bud;

1. TRIBE BEJARIEAE Copeland {1943). inflorescence elongated, borne above leaves of
current flush, bracts and prophylls leafy; flowers
Leaves revolute in bud; petals free, corolla rotate; monosymmetrical; corolla subrotate; anthers
anthers with resorbtion tissue; testa cells ca. 2x with pores; seeds?; n = 12. Two spp., W Alaska, NE
longer than broad, thin-walled. Asia.
22. Bejaria L. 26. Diplarche J.D. Hooker & Thompson

Bejaria L., Mant. Plant.: 152, 242 (1771) ("Befaria"); S.
Diplarche J.D. Hooker & Thompson, J. Bot. Kew Misc. 6: 380
Clemants, Fl. Neotrop. 66: 54-106 (1995). (1854).
Shrubs to trees; stomata tetracytic; flowers 5-7- Small evergreen shrubs; pith?; bud scales?; leaves
merous; corolla also tubular; anthers with short convolute in bud; inflorescences eperulate,
apical slits, with endothecium; fruit partly loculi- corymbs, elongating greatly in fruit; corolla
cidal; seeds elongated; n = ? 15 spp., sect. Race- hypocrateriform; antesepalous stamens epi-
mosae Fedtsch. & Basil. one sp. in SE USA, sect. petalous, antepetalous stamens less so, anthers
Bejaria Cuba, central Mexico to N Peru and the with slits their entire length, resorbtion tissue?;
Guianas; s.l.-3900 m alt. capsule septifragal, the outer walls separating into
two layers periclinally. Two spp., E Himalayas;
23. Bryanthus J.G. Gmelin 3300-4500 m alt.
Bryanthus J.G. Gmelin, Fl. Sibir. 4: 132 (1769).
Ericaceae 171
27. Menziesia J.E. Smith

Menziesia J.E. Smith, Plant. Icon. Hact. Ined. 3: t. 56 (1791).
Deciduous shrubs; leaves revolute in bud; indo-

mentum often of flattened hairs; flowers 4- or 5-
merous, tubular or urceolate, lobes short; stamens
5, 8 or 10, anthers with short slits; seeds spindle-
shaped. 7 spp., temperate East Asia and North
America, low alt. Viscin threads absent, but cf.
Copeland (1943).
28. Rhododendron L. Fig. 56

Rhododendron L., Sp. Plant. 1: 392 (1753); Sleumer, Fl. Males.
I, 6: 474-668 (1967); W.R. Philipson, Notes Roy. Bot. Gard.
Edinburgh 34: 1-71 ( 1975 ); Luteyn & O'Brien, Contributions
towards a classification of Rhododendron (1980); Cullen,
Notes Roy. Bot Gard. Edinburgh 39: 1-207 (1980); Cham-
berlain, Notes Roy. Bot. Gard. Edinburgh 39: 209-486 ( 1982);
Davidian, The Rhododendron species, vol. 1 (1982), vol. 2 Fig. 56. Ericaceae. Rhododendron scabribracteatum. A Flow-
(1989), vol. 3 (1992); W.R. Philipson & M.N. Philipson, Notes ering branchlet. B Pistil. C Stamen. D, E 1\vo inner perulae
Roy. Bot. Gard. Edinburgh 44: 1-23 (1986); Kron, Edinburgh from outside, their scars at tip ofbranchlet. F Prophyll. Drawn
J. Bot. 50: 249-364 (1993); Judd & Kron, Edinburgh J. Bot. 52: by C. van Creve!. (Sleumer 1967)
1-54 (1995).
Azalea L. (1753).
Ledum L. (1753).
Tsusiophyllum Maxim. (1870).
29. Elliottia Muhlenberg
Evergreen, less often deciduous shrubs or more Elliottia Muhlenberg in Elliott, Bot. S. Carol. Georgia 1: 448
rarely trees; indumentum diverse, often glandular (1817); Bohm eta!., J. Arnold Arbor. 59: 311-341 (1978).
or eglandular scales, or branched eglandular hairs; Cladothamnus Bongard (1833).
leaves convolute or revolute in bud; inflorescences Tripetaleia Siebold & Zuccarini (1873).
rarely axillary; corolla variable in shape, shallowly
lobed to polypetalous, monosymmetrical or not; Deciduous to evergreen shrubs; multicellular hairs
stamens 5-20, anthers with pores or rarely slits; absent; rarely pith almost Calluna-type; leaves
ovary 5-12-locular; capsule ca. 2x longer than convolute in bud, thin; inflorescences panicles to
broad; seeds spindle-shaped to ellipsoid or flat- racemes or cymes; flowers 3-5-merous; petals free;
tened, variously winged, testa cells rarely polygo- anthers with long slits; style curved, stigma much
nal; n = 13(-12x). 8 subgenera, ca. 850 spp., north expanded; seeds ovoid to flattened or winged; testa
temperate, esp. SE Asia, Malesia (very large section cells with large pitted areas; embryo with small
Vireya), two spp. in NE Australia. R. tanakae chalaza! haustorium; n = 11. Four spp., SE and W
(Maxim.) has anthers that dehisce by slits and a North America, Japan; s.l. to tree line.
3-carpellate ovary.
30. Epigaea L.
3. TRIBE PHYLLODOCEAE Drude (1897). Epigaea L., Sp. Plant. 1: 395 (1753).
Epigaeae Britton & Brown (1913). Procumbent shrublets; stomata tetracytic; leaves
Cladothamneae Copeland (1943). revolute in bud; calyx lobes large, green; flowers
with uniseriate hairs; corolla hypocrateriform to
Pith heterogeneous; inflorescences eperulate; widely campanulate; anthers with slits at least half
pedicels ± articulated; median sepal abaxial; their length; stigma expanded; capsule spherical,
anthers with resorbtion tissue; style impressed; thin-walled, with fleshy placentae; testa cells
testa cells not elongated, little thickened. not elongated, rather thick-walled. Three spp, SE
Black Sea region, Japan, E North America; s.l.
to montane.
31. Phyllodoce Salisbury 4. TRIBE EMPETREAE Pax (1891).

Phyllodoce Salisbury, Parad. Lond.: t. 36 (1805).
Shrubs; leaves basically whorled, ericoid; plants
dioecious, or rarely monoecious, or rarely flowers
Small shrubs; bud scales?; leaves revolute in bud,
hermaphroditic; perianth parts free; anthers
ericoid; inflorescences corymbs; corolla urceolate
without appendages, with elongated slits, pollen
or campanulate, lobed less than 1/3; anthers with
lacking viscin threads; disc absent; 1 basal
short slits; stigma subcapitate; testa cells at most
ovule/loculus; fruit a drupe, stones separate;
slightly elongated, not thickened; n = 12. 7 spp.,
n= 13.
north temperate; s.l. to alpine.
35. Empetrum L. Fig. 57G-N

32. Rhodothamnus Reichenb.
Empetrum L., Sp. Plant. 2: 1022 (1753); Vasilyev, Rod [the
Rhodothamnus Reichenb., Fl. Germ. Exc. 1: 417 (1830). genus] Empetrum (1961).
Small shrubs; corolla subrotate, lobed ca. 2/3; Prostrate shrubs; leaves also spiral, without woolly
anthers with short slits or pores; stigma capitate; hairs in abaxial channel; flowers axillary, solitary;
capsule often slightly loculicidal; testa cells not perianth segments 6, 2-seriate; stamens 3; ovary
elongated; n = 12. Two spp., Europe, Turkey; 6-9-locular; fruit red or black. Two spp., north
mid elevation. Hybridizes with Phyllodoce
(xPhyllothamnus Schneider).
33. Kalmiopsis Rehder

Kalmiopsis Rehder, J. Arnold Arbor. 13: 30 (1934).
Small shrubs; inflorescences corymbs; corolla

broadly campanulate, lobed 1/3-1/2; anthers with
short flaring slits; stigma slightly capitate; testa .
~
cells little elongated. One sp., K. leachiana (L.F.
Henderson) Rehder, W USA (Oregon); montane.
Doubtfully distinct from Phy llodoce, with which it J~
hybridizes.
34. Kalmia L.
Kalmia L., Sp. Plant. 1: 391 (1753); Jaynes, Kalmia: The Laurel
Book II (1988).
Leiophy llum Hedwig f., Gen. Plant.: 313 (1806).
Loiseleuria Desvaux, J. Bot. 1: 35 (1813).
Small to large, rarely deciduous shrubs; pith rarely

homogeneous; leaves alternate, opposite or
pseudoverticillate, revolute in bud; inflorescences
often corymbs, rarely axillary; corolla broadly
campanulate, with 10 pouches, or polypetalous,
rotate; stamens 10(5), anthers with short to long
slits; ovary 2-5-locular; style rarely continuous,
stigma more or less capitate; capsule often with
short loculicidal slits; testa cells moderately
elongated or not, fairly thick to thin-walled; n = 12, Fig. 57. Ericaceae. A-F Corema conradi. A Male flower, two
24. 11 spp., Kalmia (= Loiseleuria) procumbens L. perianth segments removed. B F emale flower. C M ale flower.
± circumboreal, the rest North America, Cuba; s.l. D Ovary. E Fruit. FSame, transverse section showing pyrenes.
G- N Empetrum nigrum. G Habit. H Male flower. I Pistil.
to above timberline. J Ovary, transverse section. K Fruit, vertical section. L
Pyrene. M, NSame, transverse and vertical section. (Schneider
1912)
Ericaceae 173
temperate, extreme S South America, Falkland growth; prophylls absent; flowers 4-5(6)-merous;
Islands, Tristan da Cunha; s.l. to alpine. Species corolla urceolate; anthers lacking appendages,
limits uncertain. papillate, with short, terminal slit, resorbtion
tissue present; ovary continuous; testa cells thick-
walled, papillate. One or two spp., SW Europe,
36. Corema D. Don Fig. 57A-F Azores; s.l.-moderate alt.
Corema D. Don, Edinburgh New Phil. J. 2: 63 (1826).
40. Erica L.
Erect shrubs; leaf without woolly hairs; inflores-
cence terminal, subcapitate; perianth segments Erica L., Sp. Pl. 1: 352 (1753); Schumann & Kirsten, Ericas of
3-4, 1-seriate; stamens 3(4); ovary 3-locular; fruit South Africa (1992); E.G.H. Oliver, Contrib. Bolus Herb. 19:
1-483 (2000).
white. Two spp., E USA, SW Europe, Azores; low alt. Blaeria L. (1753).
Sa/axis Salisbury (1802).
Sympieza Roem. & Schultes (1818).
37. Ceratiola Michaux Scyphogyne Brongn. in Duperry (1829).
Ceratiola Michaux, Fl. Bor.-Am. 2: 221 (1803). Bruckenthalia H.G.L. Reichenbach (1831).
Eremia D. Don (1834).
Thamnus Klotzsch (1834) ("1835").
Erect shrubs, leaves with woolly hairs; inflores- Thoracosperma Klotzsch (1834) ("1835").
cences axillary, fasciculate; perianth segments Philippia Klotzsch, Linnaea (1834) ("1835").
4( -6), 2-seriate; stamens 2; ovary 2-carpellate; fruit Coccosperma Klotzsch (1838).
red. One sp., C. ericoides Michaux, SE USA, low alt. Ericinella Klotzsch (1838).
Grisebachia Klotzsch ( 1838).
Acrostemon Klotzsch (1838).
Coilostigma Klotzsch, Linnaea 12: 234 (1838); E.G.H. Oliver,
5. TRIBE ERICEAE DC. ex Duby (1829). Bothalia 17: 163-170 (1987).
Simocheilus Klotzsch (1838).
Salaxidae Drude (1889). Anomalanthus Klotzsch (1838).
Calluneae P.F. Stevens (1971). Syndesmanthus Klotzsch (1838).
Daboecieae P.F. Stevens (1971 ). Platycalyx N.E. Brown (1905).
Arachnocalyx Compton (1934).
Eremiella R.H. Compton (1953).
Evergreen shrubs; leaves ericoid or very strongly Nagelocarpus Bullock (1954).
revolute, rarely recurved, whorled, rarely opposite Stokeanthus E.G.H. Oliver (1976).
or scattered; flowers nearly always 4-merous; calyx
and corolla marcescent; anthers often spurred, Prostrate shrubs to trees to 10( -30) m tall; pith
pores more or less elongated, pollen lacking viscin homogeneous; bud scales sometimes absent;
threads. bracts and prophylls rarely recaulescent; sepals
(1-3)4(5), small and bract-like to large and
38. Calluna R.A. Salisbury coloured; stamens (4-6)8( -10), filaments straight
to S-shaped, spurs on the backs of anthers or apex
Calluna R.A. Salisbury, Trans. Linn. Soc. Lond. 6: 317 (1802). of filaments, flattened to terete; pollen rarely in
monads; carpels (1- )4( -8), rarely 1 ovule/carpel;
Pith Calluna-type; leaves sessile, tailed; flowers style much longer than ovary or rarely almost
single in leaf axils, with 3-5(+) pairs of bracteoles; absent, rarely continuous, slightly expanded or
sepals large, coloured; corolla tubular-capanulate, rarely much expanded, cup-shaped, or reflexed;
lobed halfway; capsule septifragal; testa cells stigmatic lobes none to well developed; fruit
barely elongated, thick-walled, with large pits; loculicidal or indehiscent, ± drupaceous; cells of
n = 8. One sp., C. vulgaris (L.) Hull, Europe, Turkey; testa variable, not to much elongated, thin to thick,
s.l. to montane. pits, if present, small; n = 12, rarely 18. 860 spp.,
southern Africa (90%), also Europe, Middle East,
39. Daboecia D. Don south-west Arabian peninsula, Africa, Madagas-
car; s.l. to afroalpine. Infrageneric classification is
Daboecia D. Don, Edinburgh New Phil. J. 17: 150 (1834). unresolved; Bruckenthalia has monads, n = 18, but
cannot otherwise be separated.
Pith very heterogeneous; leaves strongly revolute;
inflorescences racemes terminating season's
VI. SUBFAMILY HARRIMANELLOIDEAE

Kron & Judd (2002).
41. Harrimanella Coville

Harrimanella Coville, Proc. Wash. Acad. Sci. 3: 570 (1901).
Small shrubs; bud scales? absent; leaves acicular,

petiolate; flower single, terminal, prophylls absent; c
corolla subcampanulate; anthers with long spurs;
n = 8. 2 spp., interruptedly circumboreal.
Styphelioideae + Vaccinioideae
Leaves with gland-hairs on the margins; inflores-
cences axillary.
VII. SuBFAMILY STYPHELIOIDEAE Sweet (1828).
Subfamily Epacridoideae Link (1829).
Epidermis lignified; adaxial fibre caps of leaf

veins absent, abaxial fibre caps massive, not con-
tacting the abaxial epidermis; leaves xeromorphic,
pungent, usually discolorous, secondary veins
palmate; inflorescence axillary; sepals free, imbri-
cate, persistent; stamens equal to and alternate Fig. 58. Ericaceae. Lebetanthus americanus. A Flowering
branch. B Same, detail. C Flower. D Same, petals removed. E
with corolla lobes, filaments glabrous, anthers bis- Stamen. F Pistil. G Ovary, transverse section. H Dehisced fruit.
porangiate, with two long introrse slits. Drawn by N. Raspini. (Dimitri 1972)
1. TRIBE PRIONOTEAE Drude (1889). impressed, stigma small. One sp., P. cerinthoides R.
Br., Australia (Tas.), endemic.
Scrambling or climbing; leaves spreading, subses-
sile; flowers solitary, pendulous; bracteoles several; All other Styphelioideae
corolla glabrous, lobes imbricate in bud, later ±
spreading; anthers included. Multicellular hairs absent; leaf teeth not glandular,
midrib not evident, veins commonly evident on
abaxial surface only; anthers with a single slit.
42. Lebetanthus Endl.
Lebetanthus Endl., Gen. Pl. 10: 749 (1839).
2. TRIBE ARCHERIEAE Crayn & Quinn (1998).
Scrambling shrubs; pedicel short; corolla subur-
ceolate; nectary scales 5; ovules ca. 10 per locule, Ovary deeply 5-lobed, style inserted between the
stigma discoid. One sp., Lebetanthus americanus lobes.
Endl., Patagonia and Tierra del Fuego.
44. Archeria J.D. Hooker
43. Prionotes R. Br. Archeria J.D. Hooker, Fl. Tasmania 1: 262, t. 80, 81 (1857).
Prionotes R. Br., Prodr.: 552 (1810).
Shrubs, erect or spreading; leaves distichous or
Climbing or epiphytic shrubs; leaves not crowded; imbricate, shortly stalked or sessile; veins parallel;
pedicel long, slender; corolla cylindrical, some- inflorescences short terminal racemes, or flowers
what constricted at throat; nectary scales very solitary, axillary towards stem apex, bracteoles
small; ovules many per locule, style strongly several, basal, caducous; corolla cylindrical or
campanulate, glabrous, lobes valvate, imbricate or
Ericaceae 175
contorted in bud, later ± recurved, pilose or 47. Andersonia R. Br.

glabrous; stamens inserted towards top of tube, fil-
Andersonia R. Br., Prodr.: 553 (1810); L. Watson, Kew Bull. 16:
aments short, anthers at throat; nectary annular 85-128 (1962).
disk or scales free; ovules several per locule, style
included or shortly exserted, stigma capitate. Shrubs, usually slender, glabrous or hairy; flowers
Seven spp., five endemic in Australia (Tasmania), also solitary in axils of bracts; sepals linear to nar-
two in New Zealand. rowly ovate, glabrous to hairy and usually ciliate,
if hyaline then only below the middle, usually
Remaining Styphelioideae equal to corolla; corolla tubular to campanulate,
Pedicels short. hairy inside, tube at least half as long as lobes,
lobes valvate in bud, later recurved, rarely
incurved; stamens free, anthers included, or par-
3. TRIBE COSMELIEAE Crayn & Quinn (2002). tially or fully exserted; nectary scales free, or par-
tially to fully fused; placentation basal, ovules 1-25
Stomata often cyclocytic; leaves amphistomatic, per locule, style exserted or not, stigma clavate or
spirally twisted, crowded, imbricate, sessile, capitate, 5-lobed; seeds small, reticulate; n = 13.
sheathing at base, scars absent, veins campylodro- About 35 spp., two sections, endemic in SW
mus; flowers single, terminal; bracteoles several, Australia.
imbricate, foliose, grading upwards in size to
sepals, persistent.
4. TRIBE EPACRIDEAE Dumortier (1829)
("Epacreae").
45. Cosmelia R. Br.
Cosmelia R. Br., Prodr.: 553 (1810). Leaves crowded; bracts many, persistent, imbri-
cate, grading to sepals; flowers single in upper
Stiff, erect shrubs; lamina erect to spreading; axils; corolla lobes later spreading; ovules at least
flowers on short branches; sepals not petaloid, several per locule.
ciliolate, otherwise glabrous; corolla much
longer than sepals, 5-lobed, red, tube cylindrical, 48. Budawangia I. Telford Fig. 59
glabrous; lobes elliptic, short, imbricate in bud,
later suberect; stamens inserted near throat, fila- Budawangia I. Telford, Telopea 5: 231 (1992).
ments flat, hirsute, anthers included, linear, upper
half dorsally adnate to filament; nectary annular; Decumbent, rhizomatous shrubs; leaves shortly
ovules 20-30 per locule, style long, not exserted, petiolate, with 3 sub-parallel veins below; flowers
stigma capitate, lobed; seeds numerous, reticulate; "pedunculate"; bracts ciliate; corolla broad-
n = 13. One sp., C. rubra R. Br., SW Western campanulate, short, glabrous, lobes imbricate in
Australia. bud, mainly glabrous; stamens inserted in throat,
exserted, spreading; filaments filiform, anthers
dorsally adnate to filament, oblong; nectary 0;
46. Sprengelia J.E. Sm. ovules many per locule, style well-exserted, stigma
Sprengelia J.E. Sm., Kongl. Vetensk. Acad. Nya Hamil. 15: 260 small, truncate; seeds numerous, reticulate. One
(1794). sp., B. gnidioides (Summerh.) Telford, endemic in
Australia (N.S.W.).
Slender glabrous shrubs; flowers also crowded in
"heads"; sepals with wholly hyaline margins,
49. Epacris Cav.
glabrous or ciliate; corolla tube short, or petals free
or shortly cohering, glabrous, lobes imbricate to Epacris Cav., Icon. 4: 25, t. 344 (1797).
almost valvate in bud, later spreading widely,
glabrous; stamens free, anthers with two slits, Shrubs, usually erect; leaves sometimes stem-
cohering around style, rarely hairy on outer clasping, also scattered, parallel-veined, com-
surface, exposed when corolla lobes spread; monly concolorous; sepals usually ciliate; corolla
nectary 0; ovules several per locule, style shortly cylindical or campanulate, glabrous, lobes imbri-
exserted, stigma small, lobed or truncate; n = 12. cate in bud, glabrous; stamens inserted in throat,
Four spp., Australia, all States except W.A., N.T.; filaments short, anthers included or shortly
one sp. in New Zealand but possibly introduced.
oblong, exserted, fused to filament from midpoint

or below, connivent, slit short, apical; nectary
absent; style well exserted, stigma small, truncate;
seeds many, reticulate. Four spp., endemic in
Australia (N.S.W.).
52. Woollsia F. Muell.

Woollsia F. Muell., Fragm. 8: 52,55 (1873).
Erect shrubs; leaves crowded; finely parallel-

veined abaxially; sepals ciliate; corolla cylindrical,
lobes contorted in bud, later spreading widely,
glabrous; stamens free, anthers included; nectary
scales distinct, acuminate; style exserted, stigma
truncate; n = 13. One sp., W. pungens (Cav.) F.
Muell., endemic in Australia (Qld and N.S.W.).
5. TRIBE OLIGARRHENEAE Crayn & Quinn

(2002).
Very small shrubs without lignotuber; leaves

Fig. 59. Ericaceae-Styphelioideae. Budawangia gnidioides. A crowded, appressed, veins parallel, obscure; pro-
Flowering stem. BFlower. C Stamens. D Pistil. E Fruit. FSeed.
Drawing by I. Telford. (Telford 1992)
phylls 2; corolla glabrous; stamens episepalous,
anthers included; ovary 2-locular, 1 apical ovule
per locule, style short; fruit a drupe.
exserted; nectary scales 5, occasionally united;
style short to exserted, stigma small or capitate; n 53. Needhamiella L. Watson
= 13.40-50 spp., mainly Australia (all states except
W.A. and N.T.), several in New Zealand and one, Needhamie/la L. Watson, Kew Bull. 18: 272 (1965).
Needhamia R. Br. (1810) non Scop. (1777).
probably introduced, in New Caledonia.
Leaves opposite and spiral; flowers in upper axils;
50. Lysinema R. Br. prophylls densely ciliate; sepals ciliate; corolla
Lysinema R. Br., Prodr.: 552 (1810).
tube cylindrical, sometimes sparsely hairy outside,
lobes induplicate-valvate in bud, later widely
Slender shrubs; leaves also scattered, 1 vein spreading, apex incurved, hairy around throat;
abaxially; inflorescences spike-like; corolla slender, stamens inserted in tube; nectary scales united in
tubular, lobes contorted in bud, glabrous; stamens a disc; style short, stigma capitate, 5-lobed, below
free, anthers versatile, included or barely exserted; anthers; fruit a dry drupe. One sp., N. pumilio (R.
nectary scales 5; ovules many, stigma capitate; Br.) L. Watson, SW Australia.
n = 13. Five spp., endemic in SW Australia.
54. Oligarrhena R. Br.
51. Rupicola Maiden & Betche Oligarrhena R. Br., Prodr.: 549 (1810).
Rupicola Maiden & Betche, Proc. Linn. Soc. New South Wales
23: 774 (1898); Telford, Telopea 5: 234- 239 (1992).Inflorescence a spike; prophylls thin, ciliate;
flowers 4-merous; sepals thin, ciliate; corolla cam-
Erect or decumbent shrubs; leaves subsessile, panulate, glabrous, lobes valvate in bud, later
with 3-5 sub-parallel veins; flowers "pedunculate"; spreading; stamens 2, inserted near apex of tube,
bracteoles usually glabrous except ciliate margin; 2 staminodes sometimes present; nectary scales 4,
corolla broad campanulate, glabrous, lobes imbri- distinct; stigma minute; fruit falling with calyx and
cate in bud, glabrous; stamens inserted at base of corolla as a unit. One sp., 0. micrantha R. Br., SW
corolla-tube, filaments thick, anthers with one slit, Australia.
Ericaceae 177
6. TRIBE RICHEEAE Crayn & Quinn (2002). 7. TRIBE STYPHELIEAE Bartling (1830).
Nodes tri- or multilacunar; pith heterogeneous; Veins parallel; stamens epipetalous; one apical
stomata paracytic; leaves crowded, bases sheath- ovule per locule, style continuous; fruit a drupe.
ing stem, scars annular; veins parallel, with adaxial
and abaxial fibre caps extending to the epidermis;
58. Acrotriche R. Br.
inflorescences terminal.
Acrotriche R. Br., Prodr.: 547 (1810).
55. Dracophyllum Labill.

Erect to spreading, rarely procumbent shrubs;
Dracophyllum Labill., Voy. Rech. Perouse 2: 211, t. 40 (1800). leaves crowded, sessile or shortly petiolate, entire
or toothed, mucronate or aristate, glabrous or
Erect or prostrate shrubs, rarely small trees; leaves hirsute, veins also radiating, glaucous abaxially;
minutely serrate; inflorescences also axillary, inflorescences spikes, rarely cauliflorous; sepals
panicles, racemes or spikes, or flowers solitary; obtuse, ciliolate; corolla tube cylindrical or cam-
pedicels often present, bracts usually caducous; panulate, with tufts of hair spreading across
corolla cylindrical or campanulate,lobes imbricate throat, glabrous below, lobes valvate in bud, later
in bud, later spreading to recurved; stamens free spreading, with a tuft oflong hairs at the thickened
or inserted on tube, included, anthers dorsifixed; and commonly recurved apex; stamens inserted in
nectary scales free; ovules many per locule, stigma throat, filaments short, anthers exserted, oblong or
5-lobed; n = 13. About 50 spp., E Australia (Qld, orbicular, versatile, lying between the corolla
N.S.W., Tas.), Lord Howe Is., New Caledonia, New lobes; nectary annular; ovary 2-10-locular, style
Zealand. conical or cylindrical, not exserted, stigma small,
lobed; mesocarp dry or pulpy; endocarp some-
times splitting into nutlets; n = 9. 15 spp., endemic
56. Richea R. Br.
in Australia, all states except N. T. The pollen is pre-
Richea R. Br., Prodr.: 555 (1810). sented on the tuft of hairs on the corolla lobes.
Shrubs or small trees; flowers crowded in a spike

59. Androstoma J.D. Hooker
or panicle; bracts and bracteoles up to 6; sepals
glabrous; corolla ovoid or conical, glabrous, almost Androstoma J.D. Hooker, Fl. Antarctica: 44 (1844).
closed at apex, splitting transversely near base and
falling early; stamens 4 or 5, free, exserted, fila- Straggling shrubs; leaves crowded, linear, coria-
ments persistent, anthers versatile; nectary scales ceous, with callus tip, slightly discolorous, with 3
large, obscure or absent; ovules many per locule; veins below, shortly petiolate; inflorescences ter-
stigma capitate or small, lobed; n = 13 (Smith- minal and axillary spikes of 1-3 flowers; corolla
White 1955).11 spp., endemic in Australia (N.S.W., campanulate, glabrous, lobes valvate in bud, later
Vic., Tas.). spreading; stamens inserted in throat, filaments
filiform, anthers exserted; ovary 3-4-locular, style
equal to or longer than tube; mesocarp pulpy; 2n
57. Sphenotoma (R. Br.) Sweet
= 24. One sp., A. empetrifolia J.D. Hooker, endemic
Sphenotoma (R. Br.) Sweet, Fl. Australas. t. 44 (1828). in New Zealand.
Slender erect shrubs; inflorescences dense ovate to

60. Astroloma R. Br.
oblong spikes; bracts persistent; sepals glabrous or
hairy, ciliolate; corolla hypocrateriform, glabrous, Astroloma R. Br., Prodr.: 538 (1810).
tube shorter or longer than sepals, lobes valvate in
bud, later spreading, with prominent longitudinal Low shrubs, commonly mat-like; leaves crowded,
ridges at base constricting corolla throat; stamens sessile to shortly petiolate, veins also radiating;
inserted on tube, anthers included, dorsally flowers solitary with several bracteoles or 2 or 3 on
attached at apex or midpoint to filament; nectary a short peduncle; bracts ciliolate; sepals ciliate;
annular or of distinct scales; ovules several per corolla cylindrical, with tufts of hairs in throat or
locule, style included in tube, stigma 5-lobed; near base, lobes valvate in bud, later revolute,
n = 6, 7. Six spp., endemic in Western Australia. commonly bearded inside; stamens inserted in
throat, filaments flat or terete, anthers included or
exserted; nectary annular; style not or shortly lar, thick, fleshy, valvate in bud, later spreading,
exserted, stigma capitate, 5-lobed; n = 4, 7, 8, 16.28 shortly bearded inside; stamens inserted just
spp., endemic in Australia, in all states except N.T. below throat, filaments terete, anthers attached
close to apex, bifurcate; nectary annular; ovary
villous, style equal to or exceeding tube, hirsute,
61. Brachyloma Sonder
stigma truncate; mesocarp thin. One sp., C.
Brachyloma Sonder,in J.G.C. Lehmann, Pl. Preiss.1: 304 (1845). kingiana (F. Muell.) J. Powell, endemic in SW
Western Australia.
Shrubs; leaves crowded, veins also radiating;
flowers solitary or in small spikes; corolla tubular,
65. Cyathodes Labill.
with 5 tufts of reflexed hairs in throat, lobes ±
imbricate in bud, later spreading, papillate Cyathodes Labill., Nov. Holl. Pl. Spec. 1: 57, t. 81 (1805); C.M.
adaxially; stamens inserted near top of tube, WeiHer, Aust. Syst. Bot. 9: 491-507 (1996).
anthers not exserted; nectary annular; ovary 4-10-
locular, style short, stigma lobed. Seven spp., Low, diffuse to erect compact shrubs; leaves
endemic in Australia, all states except N.T. pseudoverticillate, spreading; lower surface glau-
cous; flowers solitary, terminal or axillary, erect,
subsessile; bracteoles many, closely imbricate;
62. Coleanthera Stschegl. corolla tube suburceolate or cupular, exceeding
Coleanthera Stschegl., Bull. Soc. Imp. Nat. Moscou 32 (1): 4 calyx, sparsely pubescent inside; lobes valvate in
(1859). bud, later revolute, shorter than tube, sparsely
pubescent; stamens inserted near top of tube, fila-
Shrubs; leaves crowded; inflorescences 1-3- ments thick and tapered, anthers exserted; nectary
flowered spikes; sepals ciliate; corolla tube short, annular, truncate; ovary (5)6-10-locular, stigma
hairy in throat, lobes valvate in bud, later strongly small, lobed; mesocarp pulpy; n = 12 or 18. Three
revolute, hairy inside; stamens inserted in throat, spp.: endemic in Australia (Tasmania).
anthers exserted and forming a cone around the Cyathodes dealbata R. Br. is an anomalous
style, with two slits; nectary scales none or united species from Tasmania and New Zealand. It is an
in an obscure disc; stigma small. Three spp., alpine mat plant with small, silver-backed leaves, a
endemic in SW Australia. narrow, cylindrical corolla with short, densely-
bearded lobes, and a succulent orange drupe.
63. Conostephium Benth.
66. Cyathopsis Brongn. & Gris
Conostephium Benth. in Endl., Enum. Pl. Huegel.: 76 (1837).
Conostephiopsis Stschegl., Bull. Soc. Imp. Nat. Moscou 32: 5 Cyathopsis Brongn. & Gris, Bull. Soc. Bot. France 11:66 (1864);
(1859). Virot, Fl. Nouv. Caled. 6: 24-29 (1975), as Styphelia.
Small shrubs; leaves crowded; flowers solitary, Compact shrubs; leaves crowded, subsessile, veins
pedicellate, usually pendulous; bracteoles several; radiating; inflorescence a spike, crowded in upper
corolla tubular, conical towards apex, hairy in axils, basal bracts 8-12; sepals 4; corolla tube
throat; lobes very small, valvate in bud, later not short, glabrous inside and out, lobes twice as long
spreading; stamens inserted at base of cone of as tube, valvate in bud, later spreading, hirsute
corolla, anthers included, with two slits; nectary inside; stamens inserted between lobes and almost
scales 5, distinct, or absent; style exserted; stigma as long, the anthers exserted; nectary scarcely
small; ovary 5-locular; n = 8. Six spp., endemic in lobed; style short, stigma small. One sp., C. flori-
south-western Australia. bunda Brongn. & Gris, New Caledonia.
64. Croninia J. Powell 67. Decatoca F. Muell.

Croninia J. Powell, Nuytsia 9: 125 (1993). Decatoca F. Muell., Trans Roy. Soc. Victoria n.s. 1: 25 (1889).
Small shrubs; leaves strongly veined; flowers soli- Shrubs or small trees; leaves petiolate, veins radi-
tary; bracteoles several, grading into sepals, scari- ating; inflorescence a terminal or axillary spike;
ous, ciliolate; sepals scarious, ciliolate; corolla sepals 4 or 5; corolla tube cylindrical, 8-10-ribbed
tube cylindrical, pubescent inside, lobes triangu- in lower half, hairy inside in upper two-thirds,
Ericaceae 179
lobes ± imbricate in bud, later spreading, glabrous 70. Lissanthe R. Br.

inside above hairy base; stamens attached below
Lissanthe R. Br., Prodr.: 540 (1810).
lobes, filaments short, thick and tapered, anthers
scarcely exserted; nectary 5-lobed; ovary 10-
Erect to spreading or bushy shrubs; leaves petio-
locular, style thick, tapering, stigma small; meso-
late, entire, tip mucronate or aristate; inflores-
carp thick, fruit with multiple pyrenes. One sp., D.
cences racemes, terminal or in upper axils, flowers
spenceri F. Muell., Papua-New Guinea.
pedicellate; sepals ciliolate~ corolla tube cylind~i
cal, hairy inside above middle, lob~s val~at~ m
68. Leptecophylla C.M. Weiller bud, later recurved, glabrous to villous msid~;
stamens inserted at throat, filaments short, fib-
Leptecophylla C.M. WeiHer, Muelleria 12: 196 (1999).
form anthers half exserted or obvious at throat;
nect;ry annular; ovary 3-9-locular, style c~lindri
Erect shrubs, rarely prostrate or a tree to 6 m tall;
cal, not exserted, stigma lobed; n = 7, 14. SIX spp.,
leaves spreading or suberect, usually pungent,
endemic in Australia, all states except W.A. and
lower surface glaucous, striate; flowers solitary,
N.T.
terminal and axillary; bracteoles many, usually
closely imbricate; flowers effectively unisexual, the
plants dioecious; corolla campanulate. or subur- 71. Melichrus R. Br.
ceolate, exceeding or about equalhng calyx,
Melichrus R. Br., Prodr.: 539 (1810).
glabrous or pubescent inside, lobes valvate in bud,
later spreading, glabrous to densely bearded
Prostrate to erect shrubs; leaves crowded towards
inside; stamens inserted in throat, filaments terete
stem apex; flowers solitary, ± sessile, commonly
or filiform, anthers wholly or partly included;
below current season's growth; bracteoles several,
nectary annular, truncate, or lobed and toothed;
ciliolate; sepals ciliate; corolla urceolate or alm~st
ovary 5-7-locular, style usually continuous, short,
rotate 5 tufts of hairs or a ring of glandular hairs
stigma at or below level of anther~, or long and
near base, lobes valvate in bud, later spreading,
with a conspicuous bend and the stigma exserted;
sparsely hairy inside; stamens inserted ~ear base,
mesocarp pulpy; n = 10, 12 (Smith-White 1955, as
filaments short, flat, anthers usually mcluded;
Cyathodes).13 spp.,Australia:Ne~ Zealand, Papua
nectary annular; ovary 4-6-locular, style s~o~t,
New Guinea, and several PaCific Islands.
thick, stigma small; n = 8. Four spp., endemic m
Australia (Qld, N.S.W., Victoria).
69. Leucopogon R. Br.
Leucopogon R. Br., Prodr.: 541 (1810). 72. Monotoca R. Br.
Monotoca R. Br., Prodr.: 546 (1810).
Shrubs or small trees; leaves rarely opposite or
pseudoverticillate, petiolate or se~sile, commonly
Shrubs or small trees; leaves petiolate, commonly
crowded, entire, ciliate or denticulate; flowers
glaucous abaxially; inflor:scences (te~minal)
solitary, multibracteolate, or in spikes; sepals
spikes or racemes; flowers bisexual or umsexual;
commonly ciliolate; corolla tube commonly
sepals 4 or 5, ciliate; corolla campanulate; lob~s
cylindrical, glabrous inside or h~iry in throat,
valvate in bud, later recurved, glabrous or papil-
lobes valvate in bud, later spreadmg, commonly
lose; stamens inserted at top of corolla tube, fila-
bearded inside; stamens inserted at top of tube,
ments short, anthers enclosed or half-exserted;
anthers enclosed or very shortly exserted, com-
nectary scales distinct or united in a disk; ovary l-
monly with sterile tips; nectary scales 5 or u~ited
or 2-locular, style short, stigma small, lobed; n =12.
as a disc; ovary 2-5-locular, style slender, stigma
17 spp., endemic in Australia, all states except N. T.
capitate to truncate, strongly to slightly lobed.
About 230 spp., most in Australia (especially tem-
perate regions), a few in New Zealand, New Cale- 73. Pentachondra R. Br.
donia, SE Asia and some Pacific islands. Probably
Pentachondra R. Br., Prodr.: 549 (1810).
to be split; includes taxa often placed in Styphelia
in the past.
Small shrubs, sometimes prostrate; leaves sessile or
shortly petiolate, crowded; spike (2)3-5( -7)-flow-
ered, or flowers solitary, terminal and in upper
axils, multibracteolate, bisexual or pistillate; corolla and tapered, anthers shortly exserted; nectary
cylindrical, sparsely hairy inside, lobes valvate in annular; ovary 8-11-locular, style long, rarely very
bud, later recurved, bearded adaxially; stamens short, stigma lobed; fruit with multiple pyrenes;
inserted in throat, anthers almost sessile, enclosed n = 10.About 12 spp.,Malesia (Papua-New Guinea,
or ± exserted; nectary scales distinct or united in a Borneo, Celebes) and Australia (all states except
disk; ovary 5-11-locular, style included or exserted, N.T.).
stigma lobed; fruit with multiple pyrenes; n = 14.
Four or five spp., Australia (N.S.W., Vic., Tas.) and
New Zealand. VIII. SUBFAMILY VACCINIOIDEAE Arnott
(1832).
74. Planocarpa C.M. Weiller
Apical bud of vegetative shoot aborting; stigma
Planocarpa C.M. Weiller, Austral. Syst. Bot. 9: 510 (1996). truncate; n = 12.
Small shrubs; leaves crowded towards tips of inno-

vations, erect to spreading, oblong to ovate, fiat or 1. TRIBE 0XYDENDREAE Cox (1948).
slightly convex, the margin sometimes recurved;
plants hermaphrodite or gynodioecious; flowers Pith Calluna-type; stomata paracytic; anthers
erect, solitary and multibracteolate, or in 1-3- lacking appendages, with slits half their length.
fiowered spikes near branch tips, on short, erect
peduncle; corolla tube campanulate or cylindrical,
77. Oxydendrum DC.
glabrous or puberulent inside, lobes valvate in
bud, later spreading, glabrous or densely bearded, Oxydendrum DC., Prodr. 7: 601 (1839).
shorter than tube; stamens inserted in throat,
anthers included; nectary annular, glabrous; Deciduous trees; hairs not glandular; inflores-
style short, stigma small; ovary 5-8-locular; fruit cences terminating the flush, panicles; corolla
depressed globose, mesocarp thick, pulpy; n = 9. subtubular; filaments stout, anthers ?with disinte-
Three spp., endemic in Australia (Tasmania). The gration tissue dorsally; seeds elongated, testa cells
perulae of leaf buds persist, becoming grey. elongated, moderately thick walled. One sp., 0.
arboreum (L.) DC., SE USA; low alt.
75. Styphelia J.E. Sm.
Styphelia J.E. Sm., Spec. Bot. New Holland: 45 (1795). 2. TRIBE LYONIEAE Kron & Judd (1999).
Shrubs, occasionally prostrate, with or without Epidermis lignified; filaments geniculate.

lignotuber; leaves crowded; flowers pedicellate,
solitary, multibracteolate, or rarely a 1-3-fiowered
78. Craibiodendron W.W. Smith
raceme; corolla tubular, hairy in throat and at base,
lobes valvate in bud, later revolute, hairy inside; Craibiodendron W.W. Smith, Rec. Bot. Surv. India 4: 276 (1911);
stamens inserted at base, anthers exserted, versa- Judd, J. Arnold Arbor. 67: 441-469 (1986).
tile; nectary scales distinct or united in a shallow
cup; stigma small, exserted; n = 4. 15 spp., endemic Trees; leaves entire; inflorescences racemes or
in southern Australia. panicles; corolla urceolate to campanulate, thick;
anthers lacking appendages; seeds with wing on
one side, testa cells much elongated, thin-walled;
76. Trochocarpa R. Br. n = ? Five spp., SE Asia, 200-3200 m alt.
Trochocarpa R. Br., Prodr. 548 (1810).
79. Pieris D. Don.
Shrubs or small trees; leaves crowded, shortly
petiolate; inflorescences terminal and axillary Pieris D. Don, Edinburgh New Phil. J. 17: 159 (1834); Judd, J.
Arnold Arbor. 63: 103-144 (1982).
spikes; flowers unisexual or bisexual; sepals ciliate;
Arcterica Coville (1901).
corolla cylindrical or campanulate, with refiexed
hairs in throat, lobes ± imbricate in bud, later
Shrubs, rarely vines; pith heterogeneous, rarely
recurved, glabrous or sparsely hairy adaxially;
homogeneous; leaves rarely whorled, serrate,
stamens inserted in throat, filaments short, thick
Ericaceae 181
rarely entire; inflorescence terminal or axillary

panicles or racemes; filaments expanded at the
base, rarely straight, anther spurs at the filament
junction; disintegration tissue rarely absent; seeds
angular, rarely spindle-shaped, testa cells not to
much elongated, thin- to moderately thick-walled.
7 spp., subgenus Arcterica (Coville) Judd mono-
typic; subgenus Pieris with two sections: East Asia,
E USA, Cuba; low to moderate alt.
80. Agarista G. Don

Agarista G. Don, Gen. Syst. 3:837 (1834); Judd, J.Arnold Arbor.
65: 255-352 (1984), Fl. Neotrop. 66: 295-344 (1995).
Agauria (DC.) Bentham & J.D. Hooker, Gen. Plant. 2: 586
(1876); Sleumer, Bot. Jahrb. 69: 374-394 (1938).
Shrubs; pith Calluna-type (± heterogeneous);

leaves revolute, rarely convolute in bud, entire,
rarely serrate; inflorescences racemes, rarely ter-
minal, rarely panicles; corolla urceolate to tubular;
anthers lacking appendages; style slightly (rarely
not) expanded near apex; seeds spindle-shaped,
Fig. 60. Ericaceae. Lyonia ovalifolia. A Habit. B Flower. C Pistil.
testa cells much elongated, thin-walled; n = 12. 30
D Ovary. E vertical section of ovary. F Transverse section of
spp., SE USA (one sp.) to South America, esp. E ovary. G Stamens. Drawn by R. van Crevel. (Sieumer 1697)
Brasil, Africa (scattered), Madagascar, and
Reunion; s.l.-3400 m alt. Only one sp. with several
variants reported (Sleumer) in the Old World, but 82. Andromeda L.
variation in Madagascar is very great.
Andromeda L., Sp. Plant 1: 393 (1753).
81. Lyonia Nuttall Fig. 60 Shrub; leaves revolute in bud, entire; inflores-
cences terminating the flush, subcorymbose;
Lyonia Nuttall, Gen. N. Am. Pl. 1: 266 (1818), nom cons.; Judd,
J. Arnold Arbor. 62: 63-209, 315-436 (1981), Fl. Neotrop. 66: anthers with one pair of suberect awns, disinte-
222-294 (1995). gration tissue absent; seeds angular. One to two
spp., north temperate.
Evergreen to deciduous shrubs, rarely trees; pith
heterogeneous or homogeneous; leaves entire to 83. Zenobia D. Don
serrate; inflorescences racemes to fascicles, rarely
panicles; flower 4-7(8)-merous; anthers with Zenobia D. Don, Edinburgh New Phil. J. 17: 158 (1834).
spurs on filament, rarely none; fruit with pale,
thickened sutures down lines of dehiscence; seeds Deciduous shrub; leaves serrulate; inflorescences
spindle-shaped, testa cells much elongated, thin- fasciculate; anthers with two pairs of erect awns; n
walled; n = 12. 35 spp., four sections: SE Asia (sect. = 33. One sp., Z. pulverulenta (Willd.) Pollard, SE
Pieridopsis (Rehder) Airy Shaw) and SE USA USA; low alt.
(sects. Arsenococcus (Small) Judd, Lyonia and
Maria (DC.) C.E. Wood) to Mexico and the Greater
Antilles; low alt. to montane. 4. TRIBE GAULTHERIEAE Niedenzu {1889).
Stomata usually paracytic; corolla often urceolate;

3. TRIBE ANDROMEDEAE Klotzsch (1838). filaments swollen, anthers with short slits, with
disintegration tissue; n = ?
No multicellular hairs except on leaf margin; pith
heterogeneous; anthers with awns; testa several- 84. Chamaedaphne Moench
layered, cells not elongated, all walls moderately
thickened. Chamaedaphne Moench, Meth. 457 (1 794).
Subevergreen shrubs; pith heterogeneous; indu- 11 and 12. 130 spp., circum-Pacific, extending to
mentum !epidote; leaves serrate?; inflorescences the Himalayas, E USA and E Brasil; s.l. to sub-
terminating the flush, racemes or panicles, bracts alpine. Sometimes smelling of wintergreen.
foliaceous; filaments flattened, anthers lacking
appendages, disintegration tissue?, tubules ca. 1/2
their length; testa cells ca. 2x longer than broad, 88. Diplycosia Bl. Fig. 61
rather thick-walled; n = ?. One sp., C. calyculata Diplycosia Bl., Bijd.: 857 ( 1856); Sleumer, Fl. Males. I, 6: 696-740
(L.) Moench, circumboreal, low to medium alt. (1967).
Pernettyopsis King & Gamble, J. Asiat. Soc. Bengal 74: 79
(1905); Sleumer, Fl. Males. I, 6: 675-677 (1967).
85. Leucothoe D. Don
Shrubs, sometimes the roots or stem bases
Leucothoe D. Don, Edinburgh New Phil. J. 17: 159 (1834).
swollen, often with ± setose indumentum; leaf
margins entire, rarely serrulate; inflorescences
Evergreen, rarely deciduous shrubs, rarely small
fascicles, rarely 1-flowered; flowers rarely 4-
trees; pith heterogeneous or almost Calluna-type;
leaves ± serrulate; inflorescences racemes; anthers
with 1 or 2 pairs of erect awns, or muticous; seeds
rounded or winged by fringe of enlarged cells,
testa cells little elongated or thickened; n = 11. 6
spp., rather diverse (five sections!), Southeast Asia,
SE USA; low alt.
86. Eubotrys Nutt.

Eubotrys Nutt., Trans. Am. Phil. Soc. 8: 269 (1842).
Deciduous shrubs; pith ± homogeneous; leaves

serrulate; inflorescences racemes; prophylls apical;
anthers with 1 or 2 pairs of erect awns; seeds
rounded or winged by fringe of enlarged cells,
testa cells little elongated or thickened; n = 11. 2
spp., SE USA; low alt.
87. Gaultheria L.
Gaultheria Kalm ex L., Gen. Pl. ed. 5, 187 (1754); Sleumer,
Fl. Males. I, 6: 677-696 (1967); Middleton & Wilcock,
Edinburgh J. Bot. 47: 291- 301 (1990); Middleton, Bot. J.
Linn. Soc. 106: 229- 258 (1991); Luteyn, Fl. Neotrop. 66:
384-488 (1995).
Pernettya Gaudichaud, Ann. Sci. Nat. 5: 102 (1825) (as
Pernettia); Luteyn, Fl. Neotrop. 66: 365- 383 (1995).
Chiogenes Salisbury (1817).
Shrubs, sometimes procumbent; pith homoge-

neous to heterogeneous; leaves serrate or not;
flowers in racemes, rarely terminal, or rarely
panicles, or rarely flowers solitary and multib-
racteolate; corolla rarely tubular or campanulate;
anthers with (1)2 pairs of awns, these sometimes
muticous, tubules rarely present, rarely with D
slits almost the length of the anther; capsule sur-
rounded by fleshy calyx, or rarely a berry with per-
sistent calyx, or not fleshy; seeds angular, testa cells Fig. 61. Ericaceae. Diplycosia pinifolia. A Habit. B Flower at
beginning of anthesis. C Flower at full anthesis. DStamens in
not elongated, all walls moderately thick to thick; front and side view. E Pistil and disk. F Young fruit. G Seed.
n = 11 (most common), 12, 13, polyploidy on base Drawn by R. van Creve!. (Sleumer 1967)
Ericaceae 183
merous; anthers lacking appendages, with tubules

ca. 1/2 their length, rarely with pores; capsule sur-
rounded by fleshy calyx, rarely a berry; seeds
angular, testa cells elongated, somewhat thickened;
n = 18. Ca. 100 spp., but poorly known, Malesia;
often at moderate altitudes. Sometimes smelling of
wintergreen; mesophyll with fibres.
89. Tepuia Camp

Tepuia Camp in Gleason & Killipp, Brittonia 3: 178, figs 4-5
(1939); Luteyn, Fl. Neotrop. 66: 351-364 (1995), in Fl. Venez.
Guyana 4: 760-763 (1998).
Shrubs; lamina often strongly recurved, entire,

with marginal glands at or near base; inflores-
cences racemes, rarely panicles; anthers lacking
appendages, tubules ca. 1/2 their length; fruit
baccate, rarely with persistent calyx; seeds angular,
testa cells isodiametric. 7 spp., Guayana Highlands
of SE Venezuela; 1500-2500m alt. Sometimes
smelling of wintergreen.
5. TRIBE VACCINIEAE Reichenbach (1831).

Fig. 62. Vaccinium oxycoccus. A Flowering and fruiting plants.
Thibaudieae Bentham & J.D. Hooker (1876). B Flower. C Calyx and gynoecium. D Stamens, left back view,
right front view. (Ross-Craig 1979)
Stomata paracytic; leaves usually coriaceous,
pli-nerved; inflorescences axillary; pedicel articu-
lated; calyx and corolla usually valvate, calyx
limb as long or shorter than tube; stamens present, thecae smooth or papillate, tubules ± long
about equalling corolla, isomorphic or alternately and slender, with terminal pores, rarely with slits;
slightly dimorphic, anthers with tubules, rarely ovary 4-5(rarely falsely 10)-locular; fruit purple to
with spurs; ovary inferior; fruit fleshy, many- black, rarely red or white; n = 12, 24. Ca. 140 spp.,
seeded; testa hard; embryo white. Generic limits in arctic and north temperate (few spp.), Central
Vaccinieae are particularly unsatisfactory. and South America, SE Asia, Pacific, Africa and
Madagascar. Rarely epiphytic. Wildly paraphyletic,
limits unclear.
90. Vaccinium L. Fig. 62
Vaccinium L., Sp. Pl.: 349 (1753); Sleumer, Bot. Jahrb. Syst. 71: 91. Symphysia K.B. Presl
423- 493 (1941); Vander Kloet, The genus Vaccinium in
North America (1988), Pacific Sci. 47: 76-85 (1993); Vander Symphysia K.B. Pres!, Epist. Symphysia (1827); Vander Kloet,
Kloet & Dickinson, Bot. Mag. Tokyo 105: 601- 614 (1992). Taxon 34: 440-447 (1985).
Oxycoccus J. Hill (1756). Hornemannia Vahl, Skrift. Naturh.-Selsk. Ki0ben. 6: 120
(1827), non Willd. (1809).
Shrubs, rarely trees, lianes, or rhizomatous; leaves
rarely thin, deciduous, penni-nerved; inflores- Flowers (5)6-8-merous; inflorescence racemes;
cences racemes, rarely terminal, or rarely fascicles, pedicels expanded at the apex; calyx limb shorter
or flowers single, axillary; flowers (4)5(6)-merous; than the massive tube, erect, lobes minute; corolla
rarely pedicel continuous; calyx limb erect, rarely <1 em long, flaring tubular, thick, sub-bistratose;
spreading, often shorter than tube, lobes variable; stamens shorter than corolla, filaments long,
corolla <1.5 em long, cylindric, urceolate, or cam- anther thecae papillate, tubules shorter, narrower,
panulate, lobes imbricate, rarely valvate, rarely with oblique pores; fruit black. n = 24?. One sp., S.
parted to base; filaments short to long, anthers racemosa (Vahl) Stearn, N Caribbean, Jamaica and
with or without spurs, disintegration tissue rarely E Cuba to Martinique.
92. Agapetes G. Don length of the anther; fruit blue-black; seed? Three
Agapetes G. Don, Gen. Hist. 3: 862 (1834); Sleumer, Bot. Jahrb.
spp., Costa Rica and Panama; medium alt.
Syst. 71: 423-493 (1941) (Vaccinium, in part), Fl. Males. I, 6:
746-878 (1967) (Vaccinium, excl. sect. Pachyantha)
95. Notopora J.D. Hooker
Shrubs, rarely trees, trailing vines, sometimes with Notopora J.D. Hooker, Icon. Pl. 12: 53, pl. 1159 (1876);
lignotubers; phellogen superficial; leaves rarely Steyermark, Acta Bot. Venez. 2: 288-298 (1967); Luteyn,
Fl. Venez. Guyana 4: 750-752 (1998).
pen?i-nerved; inflorescences racemes, rarely
fascicles; flowers (4)5-merous, pedicels articulated,
Terre~trial shrubs; leaves congested, pinniveined,
rarely continuous, calyx limb rarely longer than
tube, erect, rarely spreading, lobes variable; corolla margms often strongly recurved; inflorescences
<5 em long, cylindric or urceolate, rarely campan- fascicles or racemes; calyx limb slightly longer
ulate,lobes imbricate (to 1/3 tube); filaments short than tube, erect, lobed halfway; corolla 2-4 em
long, carnose, cylindric to urceolate-subcylindric;
~o long, .anth.ers with or without spurs, rarely dis-
filaments longer than anthers, weakly adherent to
mtegratwn tissue present, thecae papillate, tubules
short and broad to long and slender, with termi- base of corolla, thecae papillate, tubules slightly
nal pores or slits; ovary falsely 10-locular; fruit narrower and shorter, with wide, extrorse clefts;
purple to black, orange; testa hard or mucilagi- fruit unknown. Five spp: Guayana highland region
nous; embryo white or green; n = 12, 24. Ca. 400 of Venezuela and adjacent Guyana; 400-2500 malt.
spp., SE Asia and Malesia, montane, perhaps to
New Caledonia. Sometimes epiphytic or even 96. Utleya Wilbur & Luteyn
epilithic. Most Indo-Malesian species of Vac-
Utleya Wilbur & Luteyn, Brittonia 29: 267 (1977).
cinium are provisionally included here (see Kron
et al. 2002b: the Agapetes and Bracteata-Oarianthe
clades). Epiphytic shrub; leaves pinniveined; inflores-
cences short racemes; calyx tube strongly 5-
wi.nged to sinuses, li~b erect; corolla <1 em long,
93. Didonica Luteyn & Wilbur thick, urceolate, conspiCuously 5-spurred opposite
Didonica Luteyn & Wilbur, Brittonia 29: 255 (1977); Luteyn,
lobes; filaments short, thecae smooth, curved
Syst. Bot. 16: 587-597 (1991). inwards at the base, tubules as long and wide,
flexible, with elongate, oval slits; fruit unknown.
Epiphytic shrubs, leaves flat to involute; inflores- One sp., U. costaricensis Wilbur & Luteyn, endemic
cences fascicles or racemes; flowers 5-6-merous; to Costa Rica; medium alt.
calyx limb usually campanulate; corolla 0.7-2.2 em
long, thin or succulent, cylindric-campanulate; 97. Gonocalyx Planchon & Lindley
filaments short, anthers finely papillate, incurved
at the base, tubules slightly shorter, as wide, rigid, Gonocalyx Planchon & Lindley, Gard. Chron. 1856: 152 (1856);
Luteyn, Syst. Bot. 15: 747 (1990), key.
somewhat spreading, with short, oval, latrorse slits
which do not extend to the actual tip; fruit ?white;
seed? Four spp., Costa Rica and Panama; 500- Small epiphytic, rarely terrestrial shrubs; leaf
margin flat to revolute; inflorescences one- to few-
1400m. alt.
flowered fascicles or loose racemes; flowers (4)5-
merous; calyx limb spreading, the tube rarely
94. Lateropora A.C. Smith narrowly winged; corolla (0.6- )1 em long, tubular
or cylindric-campanulate to globose-urceolate
Lateropora A. C. Smith, Contrib. U.S. Nat!. Herb. 28: 333 (1932);
Wilbur & Luteyn, Ann. Miss. Bot. Gard. 68: 162-163 (1981). bistratose at the apex; filaments short, free o;
connate, thecae papillate, tubules ca. 3x as long,
Terrestrial or epiphytic shrubs; inflorescences narrower, with small, subterminal, oblique pores
racemes, often umbelliform; pedicel short, stout; or short slits; fruit (red) purple to black; n = 23/24.
calyx limb erect; corolla <1 em long, thick, broadly 9 spp., Costa Rica to N Colombia, Caribbean; low
urceolate or squatly campanulate; filaments short, to medium alt.
weakly adherent to the base of corolla, thecae
papillate, strongly incurved at base with the lower
1/3 protruding inwardly or even turning upward,
tubules short, with lateral slits almost the entire
Ericaceae 185
98. Dimorphanthera F. Muell. Fig. 63

Dimorphanthera F. Muel!., Wing's South. Sci. Rec. n.s. 2 (1886);
Sleumer, Fl. Males. I, 6: 747-753 (as Vaccinium sect.
Pachyanthum), & 885-914 (1967); P.F. Stevens, Contrib.
Herb. Austral. 8: 1-34 (1974).
Trees to shrubs or Hanes, rarely epiphytes; stem

base rarely swollen; inflorescences racemes to fas-
cicles; pedicel rarely continuous; calyx tube rarely
winged, limb to as long, more or less erect, rarely
deeply lobed; corolla (0.5-) 1.2-4.5 em long, thick,
tubular or campanulate, ± bistratose lobes rarely
to 1/3; stamens ca. 1/2 the corolla length, filaments
short, anthers spurred or not, usually woody,
dimorphic, thecae papillate or not, tubules as wide,
ca. as long, broadened and flaring in larger
anthers, erect in smaller anthers, with elongate
pores, or rarely smaller anthers with connate
tubules and single pore; fruit purplish black; n =
24, 36.80 spp., most New Guinea (also Philippines,
Moluccas, New Britain and New Ireland);
(s.l.-)750-4000m alt. Section Pachyantha (6 spp.)
probably in the Paphia lineage.
99. Paphia Seemann

Paphia Seemann, J. Bot. 2: 77 (1864); Sleumer, Fl. Males. I, 8: Fig. 63. Ericaceae. Dimorphanthera amblyornidis. A Habit. B
878-885 (1967), as Agapetes; P.F. Stevens, Edinburgh J, Bot. Inflorescence. C Ovary, disk and style. D Major stamens, front
60 (2003 ), in press. and back view. E Minor stamens, front and back view. Drawn
by R. van Creve!. (Sleumer 1967)
Small tree to rhizomatous shrublet, Hanes, rarely
epiphytes; leaves rarely penni-nerved; inflores-
cences fascicles; pedicel rarely continuous; calyx
tube rarely shortly winged, limb to as long, erect
to subspreading; corolla 1.3 em long, thick, tubular, 101. Cavendishia Lindley, nom. cons.
terete to angled; filaments much shorter to longer Cavendishia Lindley, Bot. Reg. 21: pl. 1791 (1835); Luteyn, Fl.
than anthers, thecae slightly papillate, tubules Neotrop. Monogr. 35: 1-290 (1983).
shorter to 2.5x as long, broad to narrow, with elon-
gated slits; fruit blackish; n = 36. 20 spp., E New Epiphytic or terrestrial shrubs; leaves penni- or
Guinea (16 spp.), Bougainville, Australia (Queens- pli-nerved; inflorescences racemes or fascicles,
land), New Caledonia and Fiji; 1000-4000m alt. rarely flowers solitary; bracts usually large, showy,
persistent; calyx limb erect or spreading, lobed
100. "Agapetes scortechinii" to halfway; corolla (0.6- ) 1-4.6 em long, usually
thick, tubular; stamens rarely shorter than corolla,
Climbing (epiphytic) shrub; inflorescences fasci- dimorphic, filaments short, thecae smooth or
cles; pedicel continuous; calyx tube winged, limb slightly papillate, tubules twice as long, ca. as wide,
narrower, longer, erect, lobes long; corolla ca. 2 em with elongate slits; fruit dark blue-black; n =24/ca.
long, more or less tubular; stamens rather shorter 26. About 130 spp., Mexico (Oaxaca) to Bolivia, E
than corolla, filaments as long as anthers, thecae to Brazil (Amapa); lowland to high alt.
papillate, tubules as long, wide, with much elon-
gated slits; fruit colour?; n = 12. A. scortechinii
(King & Gamble) Sleumer, Malay Peninsula;
1200- 2100m alt.
102. Orthaea Klotzsch rarely winged, lobes (2-4)5; corolla 0.5-4 em long,
thick, subcylindric, elongate-urceolate or subglo-
Orthaea Klotzsch, Linnaea 24:23 (1851); Luteyn, Nordic J. Bot.
7: 31-37 (1987). bose, bistratose at apex; stamens (8)10(-12), from
Lysiclesia A.C. Smith, Contrib. U.S. Nat!. Herb. 28: 517 (1932). ca. 1/3 to often nearly equalling corolla, filaments
Empedoclesia Sleumer, Notizbl. Bot. Gart. Mus. Berlin-Dahlem free or connate, thecae all or alternately 2-spurred,
12: 124 (1934). rarely spurs lacking, strongly papillate, tubules 1/4
to as long, narrower, with elongate clefts; fruit
Epiphytic shrubs; leaves rarely subcoriaceous, usually green at maturity; testa mucilaginous;
penni-nerved; inflorescences racemes to fascicles, embryo green. 60 spp., Costa Rica to Bolivia, E to
rarely flowers solitary; bracts rarely large, showy French Guiana and Trinidad; lowland rainforest to
and caducous; calyx tube 3-winged, limb ca. 2x high alt.
as long, erect to suberect, lobes rarely greatly
enlarged; corolla >1 em long, tubular to subcylin-
dric; stamens usually ca. 1/3 the corolla length, 105. Satyria Klotzsch
filaments free or connate, long, dimorphic, Satyria Klotzsch, Linnaea 24: 14 (1851).
anthers equal, rarely slightly dimorphic; disinte-
gration tissue rarely present, thecae smooth, Epiphytic or terrestrial shrubs; inflorescences
tubules shorter, ca. as wide, with terminal or racemes to fascicles, often ramiflorous; calyx
slightly oblique pores; fruit blue-black. 35 spp., limb flaring or spreading, rarely longer then tube,
subgen. Orthaea and Lysiclesia (A.C. Smith) lobes (3-4)5; corolla (0.7-) 1 em long, cylindric to
Luteyn, Mexico to Bolivia, E to Guyana and slightly flaring, or rarely globose-urceolate, rarely
Trinidad. 0. crinita A.C. Smith may have minute lobed ca. 1/3; stamens up to 1/3 corolla length,
spurs on the filaments. filaments short, connate, anthers dimorphic,
thecae ± woody, slightly papillate to smooth,
103. Macleania W.J. Hooker tubules ca. as long, as wide or wider, longer
anthers: tubules often flaring distally and with
Macleania W.J. Hooker, Icon. Pl. 2: pl. 109 (1837); Yeo, Baileya in curved and ornately decorated tips, with latrorse
15: 45-59 (1967); Luteyn, BioLlania 6: 455-465 (1997).
pores, shorter anthers: tubules usually not flaring
distally, with introrse pores; fruit dark blue-black;
Epiphytic or terrestrial shrubs; leaves pinni- or n = 24. 25 spp., S Mexico to Bolivia, E to French
pH-nerved; inflorescences racemes, rarely termi- Guiana; lowland rainforest to cloud forest.
nal, or fascicles, rarely flowers solitary; pedicel
rarely continuous, often angled to conspicuously
winged, wings rarely protruding beyond the calyx 106. Mycerinus A.C. Smith
limb as a spur; calyx erect to spreading, lobes Mycerinus A.C. Smith, Bull. Torrey Bot. Club 58: 441 (1931);
(3-4)5; corolla >1 em long, rarely shorter, sub- Luteyn, Fl. Venez. Guyana 4: 749-750 (1998).
cylindric or elongate-urceolate, bistratose at apex;
stamens ca. 1/4 to nearly equalling corolla, fila- Terrestrial or epiphytic shrub; leaves thick-
ments free or connate, anther thecae strongly coriaceous, brittle, the margin flat or strongly
papillate, tubules ca. as long, somewhat narrower, recurved and with large glands at lamina base;
completely connate or not, rigid, broadly conical, inflorescences ± fascicles; prophylls apical;
with elongate, distinct or fused clefts; fruit blue- calyx limb dilated, narrowly winged to the tips of
black or white; seeds often mucilaginous; embryo lobes, lobes long; corolla 1-2 em long, carnose,
sometimes green; n = 48. 40 spp., S Mexico to Peru; cylindric to campanulate-subinfundibuliform;
most diverse in W Ecuador; s.l. to high alt. stamens shorter than corolla, filaments short,
stout, anther thecae finely papillate, connective
104. Psammisia Klotzsch continuing to the apex of tubules, tubules equal-
ing thecae, membranous, with oval clefts more
Psammisia Klotzsch, Linnaea 24:42 (1851); Luteyn, Opera Bot. than half their length, style stout; fruit reddish
92: 120-121 (1987), spp. with globose corollas.
purple; seed? 3 spp., Guayana Highland region of
Venezuela; 1300-2700 m alt. Related to Macleania
Epiphytic or terrestrial shrubs; leaves pinni- or and Psammisia.
pH-nerved; inflorescences,± fascicles to racemes,
rarely terminal, rarely flowers solitary; calyx limb
equal or ca. 2x as long as tube, erect or spreading,
Ericaceae 187
107. Polyclita A.C. Smith sometimes flaring pores; fruit dark blue-black,
rarely translucent white; testa mucilaginous;
Polyclita A.C. Smith, Bull Torrey Bot. Club 63: 314 (1936).
embryo green. 32 spp., Venezuela to N Peru,
26 spp. endemic to eastern (Oriente) Ecuador;
Terrestrial or epiphytic shrub; leaves penni-
(600-)800-3700(-1950)m alt.
nerved; inflorescence short racemes; pedicel
continuous; calyx tube angled or narrowly winged
opposite the lobes, limb erect, lobed ca. halfway; 110. Semiramisia Klotzsch
corolla ca. 1.5 em long, thick, tubular, bistratose at
Semiramisia Klotzsch, Linnaea 24:25 (1851); Luteyn, Syst. Bot.
the apex; filaments short, anthers shortly spurred 9: 359-367 (1984).
or not, thecae slightly papillate, rigid, tubules
slightly shorter, ca. as wide, flexible, with elongated Terrestrial or epiphytic shrubs, rarely lianoid;
slits; fruit blackish; seed? One sp., P. turbinata (0. leaves coricaeous, pli-nerved; inflorescences short
Kuntze) A.C. Smith, Bolivia; mid alt. racemes or 1-flowered; pedicel continuous; calyx
tube angled or winged, limb to 2x longer, erect to
108. Anthopteropsis A.C. Smith spreading, lobed to halfway; corolla > 1 em long,
thick, tubular, bistratose; stamens slightly shorter
Anthopteropsis A. C. Smith, Ann. Miss. Bot. Gard. 28:441 (1941). than corolla, filaments short, connate or free,
anther thecae strongly papillate, tubules ca. 112 as
Epiphytic shrub; leaves subcoriaceous; inflores- broad, 5-7x longer, with terminal or subterminal,
cences terminal, rarely axillary racemes; calyx oblique pores, sometimes flaring; fruit unknown.
tube conspicuously and broadly 5-winged to the Four spp., Venezuela to Peru; mid alt.
sinuses, limb rather longer, erect, apiculate; corolla
1.5-2 em long, thick, cylindric, inconspicuously 5-
angled; stamens ca. 112 corolla length, filaments 111. Oreanthes Bentham
short, anthers alternately and inconspicuously Oreanthes Bentham, Pl. Hartweg.: 140 (1844); Luteyn,
spurred, thecae slightly to coarsely papillate, Brittonia 29: 173-176 (1977); Fl. Ecuad. 54: 329-337 (1996).
basally shortly mucronate, tubules shorter, little
narrower, flexible, with slits their entire length; Terrestrial or epiphytic shrubs; leaves thick-coria-
fruit ?red; seed? One sp., A. insignis A.C. Smith, ceous, obscurely pli-nerved, usually succulent;
central Panama; medium alt. Close to Macleania inflorescences 1-flowered to fascicles; pedicel con-
and Psammisia. tinuous; calyx tube rarely bluntly 5-angled, limb to
3x longer, lobed to base; corolla> 1 em long, cylin-
109. Ceratostema Jussieu dric, terete to angled (lobes ca. 114); stamens 5 or
10, filaments free or connate, anther thecae essen-
Ceratostema Jussieu, Gen. Pl.: 163 (1789); Luteyn, J. Arnold tially smooth, tubules ca. 112 as broad, slightly to
Arbor. 67: 485-492 (1986); Luteyn, Fl. Ecuad. 54: 210-253 up to 6x longer, with terminal or subterminal and
(1998).
Periclesia A.C. Smith, Contrib. U.S. Nat!. Herb. 28: 357 (1932). oblique, usually flaring pores or rarely elongate
clefts; fruit unknown. 7 spp., Ecuador and adjacent
Terrestrial or epiphytic, rarely scandent shrubs, Peru; 1340-3290 m alt.
lignotubers common; bud scales rarely subulate;
leaves penni- or pli-nerved; inflorescences 112. Siphonandra Klotzsch
fascicles to racemes, rarely flowers solitary;
flowers (4)5(6)-merous, bracts small to large and Siphonandra Klotzsch, Linnaea 24: 24 (1851).
showy, pedicel rarely continuous; calyx tube rarely
winged, limb to 4x as long, suberect or spreading, Terrestrial or epiphytic; leaves penni-nerved,
lobes short to long, often distinctly veined; corolla rarely crenulate; inflorescence racemes; calyx limb
thick, >1 em long, rarely less, subcylindric, often spreading; corolla ca. 2.5 em long, cylindrical; fila-
ventricose at base, bluntly angled, rarely terete, ments short, connate, thecae papillate, tubules very
lobed to halfway; filaments short, distinct or narrow, 4-5x longer, flexible, with flaring terminal
connate, slightly dimorphic or not, anthers often pores; fruit black, testa ?mucilaginous; embryo
slightly dimorphic, thecae usually strongly papil- green. 1-2 spp., S Peru toN Bolivia; high alt.
late, tubules much narrower, 2-5x longer or rarely
shorter, flexible, separate to connate nearly to
apex, with short, oblique clefts or nearly terminal,
113. Pellegrinia Sleumer aments usually longer than anthers, somewhat

sigmoid at the base, anthers membranous, thecae
Pellegrinia Sleumer, Notizbl. Bot. Gart. Mus. Berlin-Dahlem 12:
287 (1935). smooth, rarely papillate, tubules to 3x longer,
usually little narrowed, with oval clefts; fruit
Usually epiphytic; bud scales rarely subulate; translucent white to lavender or purplish; testa
leaves pli- or penni-nerved; inflorescences fasci- mucilaginous; embryo green; n = 24, 48, ca. 75. 22
cles or !-flowered; pedicel continuous; calyx limb spp., S Mexico to Bolivia, esp. Ecuador, E to French
suberect or spreading, lobes long; corolla 2 em Guiana, and Haiti to Trinidad;
long, subcylindrical, sub inflated; filaments shorter
than anthers, with dense retrorse hairs abaxially, 116. Diogenesia Sleumer
anther thecae smooth, tubules narrow, 2-4x
longer, flexible, with small flaring or oblique pores; Diogenesia Sleumer, Notizbl. Bot. Gart. Berlin-Dahlem 12: 121
(1934); Notes Roy. Bot. Gard. Edinburgh 36: 251-258 (1978).
fruit blue-black; seed? Five spp., Peru; high alt. Eleutherostemon Herzog (1915), non Klotzsch (1838).
114. Disterigma (Klotzsch) Niedenzu Usually epiphytic shrubs, branches slender, often
pendent, rarely climbing; bud scales subulate;
Disterigma (Klotzsch) Niedenzu, Bot. Jahrb. Syst. 11: 160, 209 inflorescences fascicles or racemes, rarely flowers
(1889); A. C. Smith, Brittonia 1:203-232 (1933); Wilbur, Bull.
Torrey Bot. Club 119: 280-288 (1992 ); Luteyn, Fl. Ecuad. 54: solitary; flowers 4-5-merous, pedicel slender, con-
253-286 (1996). tinuous; calyx sub erect; corolla usually <1 em long,
Killipiella A.C. Smith, J. Wash. Acad. Sci. 33: 242 (1943). cylindric, urceolate or campanulate (lobed 1/2
way); stamens 4-6, 8 or 10, filaments longer or
Terrestrial or epiphytic shrubs; leaves often con- shorter than anthers, anther thecae smooth or
gested, usually less than 3 em long and obscurely faintly papillate, tubules equal or longer and
pH-nerved, rarely crenate; inflorescences fascicles, slightly narrowed, with oval pores or clefts; fruit
flowers rarely solitary; flowers (3)4(5)-merous, purple-black or rarely white; testa mucilaginous;
pedicel with apical prophylls investing ovary, very embryo green; n = 12. 13 spp., Venezuela to N
short, obscurely articulated; calyx limb somewhat Bolivia; mid alt. Close to Sphyrospermum.
longer than the tube, erect, lobed to base, rarely
imbricate; corolla <1 em long, subcylindric or
117. Rusbya Britton
campanulate-cylindric; stamens rarely same
number as corolla lobes, ca. 2/3 corolla length, Rusbya Britton, Bull. Torrey Bot. Club 20: 68 (1893).
filaments longer or shorter than anthers, thecae
smooth, tubules to 2x longer, little narrower, rarely Epiphytic shrub; bud scales subulate; leaves linear,
narrower, rarely fused, with elongate, elliptical, one-nerved; inflorescences 1-flowered; pedicel
introrse clefts or subterminal pores; fruit blue- continuous; calyx tube narrowly angled, limb
black or translucent white; testa mucilaginous (or sub erect; corolla <1 em long, cylindrical-urceolate;
not?); embryo green; n = >40. 35 spp., Guatemala S filaments dimorphic, anthers dimorphic, almost
to Bolivia and E to Guyana, 2/3 in Ecuador; low to smooth, tubules not narrowed and slightly longer,
high alt. with elongated slits; fruit not known. One sp., R.
taxifolia Britton, N Bolivia; mid alt.
115. Sphyrospermum Poeppig & Endlicher
118. Themistoclesia Klotzsch
Sphyrospermum Poeppig & Endlicher, Nov. Gen. Sp. Pl. 1: 4
(1835); A.C. Smith, Brittonia 1: 203-232 (1933); Luteyn, Fl. Themistoclesia Klotzsch, Linnaea 24: 41 (1851); Sleumer,
Ecuad. 54: 341-363 (1996). Notizbl. Bot. Gart. Mus. Berlin-Dahlem 13: 108-111(1936).
Epiphytic or terrestrial shrubs with pendent Often epiphytic shrubs; leaves subcoriaceous to
branches; leaves coriaceous, sometimes very coriaceous, rarely thick-fleshy, obscurely pH-
thickly, small, obscurely pH-nerved; inflorescences nerved, rarely pinnate; inflorescences fascicles or
fascicles, rarely flowers solitary; flowers 4-5- racemes, rarely flowers solitary; flowers (4)5-
merous, pedicels slender, cernuous, distally merous; pedicel continuous (articulated?); calyx
swollen; calyx limb ± spreading, lobes short to tube bluntly angled, rarely strongly winged, limb
long; corolla to 1.2( -2.0) em long, campanulate or erect to spreading, lobes rarely long; corolla ca. 1
cylindrical to infundibuliform; stamens 4-5(-10), em long, ovoid to cylindric, terete to angled;
ca. 3/4 corolla length, slightly dimorphic or not, fil- stamens 4, 5, 8 or 10, more than 1/2 to as long as
Ericaceae 189
corolla, filaments to as long as the anthers, anther slightly winged, limb to 2x as long, sub erect, often
thecae smooth, tubules nearly equal to 4x as long, deeply lobed; corolla >1 em long, cylindrical, rarely
rarely fused, little to moderately narrowed, with urceolate; filaments very short, often slightly
short clefts; fruit purple, black or cream; testa dimorphic, thecae submembranous, slightly papil-
mucilaginous; embryo green. 25 spp., Costa Rica to late, tubules flexible, not narrowed, to 6x longer,
Venezuela and Peru; mid to high alt. with elongate slits or oblique flaring pores; fruit
blue-black; seed? 11 spp., C Peru to N Bolivia; high
alt.
119. Plutarchia A.C. Smith
Plutarchia A.C. Smith, Bull. Torrey Bot. Cl. 63: 311 (1936).
122. Anthopterus Hooker
Terrestrial shrubs; leaves also opposite; inflores- Anthopterus Hooker, Icon. Pl. 3: pl. 243 (1840); Luteyn, Opera
cences fascicles; flowers 4-5-merous, bracts often Bot. 92: 109-113 (1987); Brittonia 48: 605-610 (1997).
conspicuous, pedicel articulated; calyx tube rarely
winged, limb erect, to 2-4 times longer, usually Shrubs or small trees, rarely epiphytes; bud
lobed to base; corolla >1 em long, cylindric to scales rarely subulate; leaves rarely subopposite or
urceolate-cylindric, terete or angled; stamens pseudoverticillate, penni- or pH-nerved; inflores-
(5- )8-10, filaments short, slightly dimorphic or cences racemes; bracts sometimes conspicuous;
not, thecae slightly papillate, tubules 2-4 times pedicel continuous; calyx tube strongly winged,
longer, rather narrower, flexible, with elongate limb also longer, suberect to spreading, moder-
clefts; fruit blue-black, rarely white; testa mucilagi- ately lobed; corolla ca. 1(-2) em long, subcylindric
nous; embryo green. 11 spp., most in Colombia to subglobose, narrowly to broadly 5-winged,
and Ecuador; high-elevation montane forest and lobes rarely long; filaments shorter than anthers
paramo. (rarely same length), often connate, anthers
membranous, thecae smooth, tubules slightly nar-
rowed, 1-3x longer, with short to elongate clefts;
120. Thibaudia Jaume Saint-Hilaire fruit blue; seed? 11 spp., SE Panama to NE Peru;
Thibaudia Jaume Saint-Hilaire, Expos. Fam. 1: 362 (1805). s.l.-2500m alt. Most species poorly known; close
Calopteryx A.C. Smith, J. Arnold Arbor. 27: 100 (1946). to Thibaudia and Themistoclesia.
Epiphytic or terrestrial shrubs; bud scales rareiy

123. Costera J.J. Smith
subulate; leaves rarely penni-nerved, crenate;
inflorescences axillary or terminal, rarely rami- Costera J.J. Smith, Icon. Bog. 4: 77, tab. 324 (1910); Sleumer, Fl.
florous fascicles, racemes or panicles; pedicel Males. I, 6: 740-746 (1967).
rarely continuous; calyx tube rarely narrowly 5-
angled, or rarely strongly winged to sinuses, limb Shrubs, climbing or epiphytic; inflorescences
often longer, erect to slightly spreading, lobed to fascicles; flowers 4-5-merous; pedicel continuous;
halfway; corolla >1 em long, cylindric to subcylin- calyx limb erect to spreading, lobed; corolla <1 em
dric, rarely 5-angled to strongly winged; filaments long, urceolate to campanulate, lobed to halfway or
short, separate or connate, anthers firm or mem- less; stamens 5, 8, or 10, filaments to as long as
branous, thecae smooth or slightly papillate, anthers, anther thecae papillate, tubules much to
tubules ca. as long and wide, with elongate clefts; slightly narrower and shorter to 1.5x longer,
fruit purple to blackish; testa mucilaginous or with oblique pores or short slits; fruit white to
hard; embryo white, rarely green. 60 spp., Panama dark purple; testa hard or mucilaginous; embryo
and Costa Rica, mostly from Colombia to Bolivia white or green; n = 12. 9 spp., W Malesia; mostly
and E to Guayana and Brazil; low to high alt. 500-1300m alt. Circumscription and relationships
uncertain.
121. Demosthenesia A.C. Smith
124. Gaylussacia Kunth, nom. cons.
Demosthenesia A.C. Smith, Bull. Torrey Bot. Club 63: 310
(1936). Gaylussacia Kunth, Nov. Gen. Sp. Pl. 3, ed. fol.: 215, ed. qu.: 275,
t. 275 (1819); Sleumer, Bot. Jahrb. Syst. 86: 309-384 (1967).
Usually epiphytic shrubs, rarely lianes, lignotubers
frequent; bud scales often subulate, leaves rarely Shrubs or subshrubs; indumentum sometimes
penni-nerved; inflorescences fasciculate or race- glandular; leaves coriaceous to membranous,
mose; pedicel continuous; calyx tube rarely evergreen to deciduous, pinni-nerved, apex glan-
dular; inflorescences racemes, flowers rarely Bidartondo, M.I., Bruns, T.D. 2001. Extreme specificity in epi-
solitary; flowers 4-5-merous; calyx limb deeply parasitic Montropoideae (Ericaceae): widespread phyloge-
netic and geographic structure. Molec. Ecol. 10: 2285-2295.
lobed; corolla <1 em long, cylindric, urceolate, or Bjorkman, E. 1960. Monotropa hypopitys L.- an epiparasite on
campanulate, often angled; filaments short to tree roots. Physiol. Plant. (Copenhagen) 13: 308-327.
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Bohm, B.A., Brim, S.W., Hebda, R.J., Stevens, P.F.1978. Generic
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limits in the tribe Cladothamneae (Ericaceae) and its posi-
locular; fruit drupaceous, with 10 pyrenes; n = 24. tion in the Rhododendroideae. J. Arnold Arbor. 59: 311-341.
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Luteyn, J.L. (ed.) Flora Neotropica. Monograph 66. Ericaceae
Part II, the superior-ovaried genera. New York: New York Generic concepts in Styphelieae are currently being revised.
Botanical Garden, pp. 13-27. The following genera do not appear to be monophyletic, and
Warner, B. G., Chinnappa, C.C.1986. The implications and evo- will undergo significant change: Astroloma R.Br., Brachyloma
lutionary trends in pollen of Canadian Ericales. Can. J. Bot. Sond., Leucopogon R. Br. sensu lato, Lissanthe R.Br., Monotoca
64: 3113-3126. R.Br. and Styphelia Sm. sensu Benth.
Fouquieriaceae 195
Fouquieriaceae
K. KuBITZKI
Fouquieriaceae DC., Prodr. 3: 349 (1828), nom. cons. shrubs or small trees to columnar stem succulents.
The woody habit is found in six species. The
Shrubs to small trees with woody or succulent remaining species are stem succulents with a
trunks, bearing simple to branched, outwardly highly parenchymatized, central xylary water-
arched to horizontal, spinose, heteroblastic storage tissue. In Fouquieria fasciculata and F. pur-
branches; periderm translucent, often exfoliating pusii, the succulent tissue is restricted to the basal
and associated with the persistent epidermis; stem or central portion of the stem, whereas the distal
cortex with peripheral groups of sclereids and parts are woody. Fouquieria columnaris is the
assimilatory tissue and inner starch- and anasto- tallest species and has conical, tapering trunks,
mosingly arranged water-storage tissue; nodal which are succulent from the base to the tip.
spines 2-45 mm long, subtended by tapering, con- Mature individuals are commonly 14, exception-
tinuous decurrent ridges laterally separated by ally 25m high and may consist of a single
distinct or shallow furrows, or by recurrent ridges unbranched or forked central trunk. In these
widely separated; nodes unilacunar, 1-trace. stems, in which a solid xylem forms only a small
Leaves alternate, simple, entire, exstipulate, petio- portion of the stem, mechanical strength is pro-
late to nearly sessile, glabrous or slightly pubescent vided by the turgescence of the parenchymatous
below, those of long-shoots elliptical, long- water-storing tissue, and upon drought the trunks
petiolate; spines rigid, separating abaxially from may collapse and bend downward (Fig. 64).
petioles of long-shoot leaves and continuing into The long-shoots have simple petiolate leaves,
the decurrent ridges of the cortex; short-shoot which tend to wilt at dry periods. The short-
leaves in axillary fascicles, shortly petiolate to shoots, produced in the axils of long-shoot leaves,
nearly sessile. Inflorescence determinate, terminal bear fascicles of shortly petiolate or sessile leaves,
or axillary, spicate, racemose to paniculate or which are retained even after prolonged drought.
corymbosely paniculate; flowers with 2 prophylls, The spines are associated with long-shoot leaves
hypogynous, perfect; sepals 5, distinct, strongly in a most characteristic fashion (Fig. 65). The
imbricate, quincuncial, persistent; petals 5, abaxial side of the petioles develops into a rigid,
connate into a tube, lobes imbricate; stamens conical spine composed of narrow, elongate fibres,
hypogynous, 10(-23 ), unevenly exserted; filaments which continue from the spine through the under-
slightly adnate to corolla base, occasionally with lying stem cortex to form the decurrent ridge
basal ligulate spur; anthers tetrasporangiate, characteristic of the stems. Upon maturation of
introrse, acuminate at apex, 2-lobed at base, the long-shoot, an abscission layer forms between
longitudinally dehiscent; gynoecium fused of 3 the leaf and spine tissue and eventually the leaf
carpels; ovary superior, with septiform parietal is shed. The development of the spines suggests
but basally axile placentation; style in upper half that they are of cortical origin (Humphrey 1931;
usually branched into 3 style branches; ovules Henrickson 1969).
anatropous, bitegmic, 6-20. Fruit a loculicidal The decurrent ridges cover young stems and are
capsule with columnar, axile placentae; seeds 6-15, separated only by deep or shallow furrows. With
oblong-elliptical with membranous margins of the enlargement of the stems, the ridges become
unicellular trichomes; endosperm scanty, oily and tangentially separated and eventually detach from
proteinaceous; embryo 3-9 mm long; cotyledons the stem. The underlying, translucent periderm
flat. x = 12. consists of alternating fibrous and suberinous
A monogeneric family with 11 xerophytic layers and usually exfoliates from older stems in
species from the southern U.S.A. to south Mexico. thin papery sheets. A rigid periderm forms only
under the ridges, whereas the intervening areas
VEGETATIVE STRUCTURES. The growth habit of are covered by a persistent epiderms. The cortex
Fouquieria species varies from small woody consists of an outer, thin, chlorenchymatous layer,
196 K. Kubitzki
Fig. 64. Fouquieriaceae. Fouquieria splendens, Baja California. which follows the pattern of translucent furrows
Photo K. Kubitzki and series of tightly packed sclereid nests. The
inner cortex consists of starch-storing tissue,
except for a network of water-storage tissue, which
also follows the anastomosing pattern of translu-
cent furrows (Humphrey 1935; Henrickson 1969,
1972).
In the wood, the vessel elements have simple
perforations; imperforate tracheary elements have
bordered pits; and wood rays are mostly homo-
cellular and 1-8 cells wide; wood parenchyma is
diffuse.
A detailed description of the vegetative anatomy
of Fouquieria has been given, among others, by
Henrickson (1969, 1972).
INFLORESCENCES. Inflorescences range from

elongate or congested panicles to racemoids or
spicoids and seem to be consistently determinate;
in some species such as F. columnaris the sequence
of flowering is basipetalous but acropetalous
sequence predominates [the latter, in spite of the
presence of terminal flowers in the schemes
of Henrickson (1972), interpreted by him as
Fig. 65. Fouquieriaceae. Fouquieria splendens. A Long-shoot indeterminate].
ending in an inflorescence. B Long-shoot with axillary, leaf-
bearing short-shoots, the long shoot leaves fallen off. C Long- FLOWER STRUCTURE. The histology and vascular-
shoot, the leaves abscising from the spines, which appear as
extensions of the cortical ridges of the axis. D Flower. E
ization of the flowers has been described by Hen-
Dehiscing capsule. F Seed. (Takhtajan 1981, from Henrickson rickson (1972). Stamen number is variable in
1969) Fouquieria but in all species stamens are borne
Fouquieriaceae 197
from the receptacle in a single whorl. Ten is the The flattened seeds contain a small embryo and
predominating and apparently basic number. a thin layer of endosperm. The seed wing is
formed from unicellular trichomes derived
EMBRYOLOGY. The ovules are anatropous, from the epidermis of the outer integument
bitegmic, and tenuinucellar with an integumen- (Henrickson 1972).
tary tapetum; the micropyle is formed by the inner
integument. The embryo sac is of Polygonum type, PHYTOCHEMISTRY. Presence of ellagic acid,
and endosperm development is ab initio cellular, caffeic acid, flavonols, glycosides of cyanidin and
with a chalazal haustorium (Johansen 1936; Khan (in red-flowered spp.) pelargonidin, dammaran
1943). (Hegnauer 1989) and seco-iridoid glucosides
(Dahlgren et al. 1976) have been reported. Ocotillo
POLLEN MORPHOLOGY. (Henrickson 1973). The wax from the cortex of Fouquieria splendens is a
pollen grains are 3-colporate, more or less oblate- dammaran derivative.
spheroidal and eureticulate; distinguishing fea-
tures include: ( 1) lumina diminishing in size AFFINITIES. Fouquieriaceae are notable for
towards the poles and colpus margins; (2) a ten- exhibiting several basal traits: their ovules are
dency to form variable, often striate patterns; (3) bitegmic, and their wood has tracheids and diffuse
thickening of the nexine towards the poles; and axial parenchyma. The family has often been com-
(4) absence of nexine thickening around the pared with Styracaceae and Ericaceae; Nash (1903)
apertures. Most distinctive are the grains of F. and Thorne (1968), among others, have suggested
columnaris, which have a crassitectate nexine with a relationship with Polemoniaceae. Molecular
large brochi. studies have resolved all these families as forming
part of a major clade, Ericales s.l., although the
KARYOLOGY. Chromosome numbers are known precise position of Fouquieria within this clade
for all but one species, seven of which are diploid remains uncertain in most recent comprehensive
(2n = 24), one tetraploid, and two hexaploid studies (for example, Soltis et al. 2000; Albach et al.
(Henrickson 1972). 2001). The relationship between Fouquieriaceae
and Polemoniaceae has been supported in the
POLLINATION. The family exhibits a great diver- combined tree of a five-gene study (Anderberg et
sity in floral features and modes of pollination al. 2002), but not by the combined plastid genes or
(Henrickson 1972). In all species, sweet nectar is the mitochondrial gene alone.
produced from nectaries around the ovary base.
Whitish flowers with relatively short floral tubes, DISTRIBUTION AND HABITATS. Fouquieria ranges
sometimes producing a sweet smell, are visited by from western Baja California to northern Arizona
various kinds of insects. Hummingbird visitation and from eastern Texas to south-eastern Oaxaca.
has been observed to the red tubular flowers borne Six species are narrow endemics. Species of the
in terminal inflorescences of species such as F. southern United States and northern Mexico
splendens and F. formosa. Some species have been occur mainly in desert habitats receiving 50-350
shown to be self-compatible, as isolated cultivated mm rainfall. Species in southern Mexico grow in
plants set seeds, but this may be irrelevant in the deciduous tropical forest and arid tropical scrub
wild because floral structure does not seem to averaging mostly 400-700mm precipitation.
favour self-pollination (Henrickson 1972). Between desert and forest species, no difference in
ecological preferences can be recognised.
FRUIT AND SEED. The fruit is a loculicidal capsule
containing a large, parenchymatous central axis A single genus:
and 5-18 broad, winged seeds; its histology has
been described by Henrickson (1972). Placenta-
Fouquieria Kunth in HBK. Figs. 64,65
tion changes characteristically during the matura-
tion of the fruit: at anthesis, the ovary has three Fouquieria Kunth in HBK., Nov. Gen. Sp. 6: 81 (1821);
parietal placentae borne on thin septa extending Henrickson, Aliso 7: 439-537 (1972), rev.
into the locule but, in the process of growth of the Idria Kellogg ( 1860).
ovary, the three septa are broken and the placen-
tae fuse into a common, central parenchymatous Description as for family.
column. Eleven spp. in the arid regions of Mexico and
adjacent south-western U.S.A. Three subgenera
were distinguished by Henrickson (1972): subgen.
198 K. Kubitzki
Fouquieria, woody throughout; periderm exfoliat- Hegnauer, R. 1989. See general references.
ing in thin sheets; style exserted, 8 spp.; subgen. Henrickson, J. 1969. Anatomy of periderm and cortex of
Fouquieriaceae. Aliso 7: 97-126.
Bronnia (H.B.K.) Henrickson, stems succulent, Henrickson, J. 1971. Anatomy of periderm and cortex of
epidermis persistent; style exserted, 2 spp.; Fouquieriaceae. Aliso 7: 97-126.
subgen. Idria (Kellogg) Henrickson, tall, succu- Henrickson, J. 1972. A taxonomic revision of the Fouquieri-
lent, periderm not exfoliating, style included, only aceae. Aliso 7: 439-537.
F. columnaris (Kellog) Kellog ex Curran. Henrickson, J. 1973. Fouquieriaceae DC. World Pollen Spore
Flora 1: 1-12.
Humphrey, R.R. 1931. Thorn formation in Fouqieria splendens
and Idria columnaris. Bull. Torrey Bot. Club 58: 263-264.
Selected Bibliography Humphrey, R.R. 1935. A study of Idria columnaris and
Fouquieria splendens. Am. J. Bot. 22: 184-207.
Albach, D.C., Soltis, P.S., Soltis, D.E., Olmstead, R.G. 2001. Phy- Johansen, D.A. 1936. Morphology and embryology of
Fouquieria. Am. J. Bot. 23: 95-99.
logenetic analysis of asterids based on sequences of four
genes. Ann. Missouri Bot. Gard. 88: 163-212. Khan, R. 1943. The ovule and embryo sac of Fouquieria. Proc.
Anderberg, A.A. et al. 2002. See general references. Natl. Inst. Sci. India 9: 253-256.
Bate-Smith, E. C. 1964. Chemistry and taxonomy of Fouquieria Nash, G.V. 1903. A revision of the family Fouquieriaceae. Bull.
splendens: a new member of the asperuloside group. Phyto- Torrey Bot. Club 30: 449-459.
chemistry 3: 623-625. Soltis, D.E. et al. 2000. Angiosperm phylogeny inferred from
Behnke, H.-D. 1976. Sieve-element plastids of Fouquieria, 18 S DNA, rbcL, and atpB sequences. Bot. J. Linn. Soc. 133:
381-461.
Frankenia (Tamaricales), and Rhabdodendron (Rutaceae),
taxa sometimes allied with Centrospermae (Caryophyl- Takhtajan, A. (ed.) 1981. See general references.
lales). Taxon 25: 265-268. Thorne, R.F. 1968. Synopsis of a putative phylogenetic classifi-
Dahlgren, R., Jensen, S.R., Nielsen, B.J. 1976. Iridoid cation of the flowering plants. Aliso 6: 57-66.
compounds in Fouquieriaceae and notes on its possible
affinities. Bot. Not. 129: 207-212.
Grubbiaceae 199
Grubbiaceae
K. KUBITZKI
Grubbiaceae Endl., Gen. Pl.: 327 (1837), nom. cons. lanceolate, and usually have strongly revolute
margins (Fig. 66A, B). The leaf bases are connected
Shrubs; leaves decussate, simple, entire, petiolate by a transverse, long-pubescent ridge or wing on
to nearly sessile; stipules wanting, but the leaf each side across the stem.
bases connected by a transversal ridge. Hairs The anthers are 2-sporangiate (Endress and
unicellular. Inflorescences small, axillary, 3(2)- Stumpff 1990); Hieronymus (1889) and Carlquist
flowered dichasia or many-flowered, cone-like (1977a) observed that one microsporangium in
compound dichasia; the flowers of one inflores- each theca is sterile. Cronquist ( 1983) stated that
cence with coherent or connate inferior ovaries. "the orientation of the anthers of Grubbia is very
Flowers subtended by bracts and provided with 2 suggestive of the Ericales, although they are not
prophylls, tiny, hermaphroditic, actinomorphic, exactly like those of any other family: they are
epigynous, monochlamydeous, tetramerous, dip- minute, inverted and adnate for their whole length
lostemonous; tepals distinct, valvate, sepaloid, to the distal part of the filament, thus appearing
canescent-hairy on outer surface, pink to red on to be extrorse". I have been unable to confirm
inner surface; stamens 8, 4 of these longer, oppo- Cronquist's observation, but an ontogenetic study
site the tepals and basally attached to them, and 4 may clarify the situation; inverted anthers often
alternating with the tepals; filaments linear, ending have a characteristic vascular hook where the
as a blunt tip beyond the thecae; anthers dorsi- vascular bundle from the filament enters the
fixed, 2-thecate, 2-sporangiate; dehiscence length- connective (Anderberg 1992).
wise extending over the upper and lower shoulders
of the anther, with 2 valves opening towards the ANATOMY. The vegetative anatomy, including
ventral side; disk epigynous, papillate to shortly wood anatomy, has been treated by Carlquist
hairy; gynoecium 2-carpellate, transversal; style (1977b, 1978). The nodes are trilacunar. The upper
unbranched, with 2 short apical lobes but appear- leaf epidermis has a thick cuticle or a cutinized
ing simple; ovary inferior, initially 2-locular, thick wall; the lower epidermis bears anomocytic
becoming 1-locular by disintegration of upper stomata and non-glandular unicellular trichomes.
part of septum above insertion of the ovules, these Rhombic crystals and crystal druses of calcium
solitary per carpel, pendulous from the upper part oxalate are found in the leaf tissue, and crystals
of the septum, anatropous, unitegmic, tenuinu- also occur in the ray cells of the wood. In the sec-
cellate. Fruits 1-seeded drupes, those of one ondary xylem the vessels are long and provided
infructescence becoming laterally compressed and with oblique perforation plates with many (16-43)
fused, forming a coenocarp suggesting a small bars. Lateral wall pitting is alternate on vessel-
cupressaceous cone; seeds with thin testa; tracheid contacts, and scalariform on vessel-ray
endosperm copious, oily, its outer surface heavily contacts. Tracheids are of almost the same length
cutinized; embryo straight, central, the cotyledons as the vessels. Rays are narrow multiseriate plus
shorter than hypocotyl and radicle. uniseriate, with procumbent cells only in central
A single genus of 3 spp., restricted to the Cape portions of multiseriate rays. Axial parenchyma
Province of South Africa. is sparse and diffuse. The wood of Bruniaceae
and Geissolomataceae is very similar.
MoRPHOLOGY. The species of Grubbia are ever-
green shrubs; they branch from a single stem EMBRYOLOGY. The ovules are provided with an
and a taproot, or ascend with numerous shoots endothelium and a long micropyle. The sporo-
from a lignotuber at the surface of the ground (G. genous cell functions as megaspore mother cell,
tomentosa). Young branches are longitudinally and the chalaza! megaspore develops into a
ridged and covered with a tomentum. The leaves Polygonum-type embryo sac. Endosperm
are leathery and linear or narrowly triangular- formation seems to be cellular (Dahlgren and van
200 K. Kubitzki
Fig. 66. Grubbiaceae. Grubbia rosmarinifolia. A Leaf, upper more (6-20) fruits fuse to form the syncarps of
side, with position of floral triad indicated. B Leaf, base of G. tomentosa and G. rourkei, which are 5 mm or
blade seen from below. C Floriferous branchlet. D Floral triad.
E Prophylls supporting the lateral flowers in the triad. F Lateral more in length (sect. Strobilocarpus Klotzsch); at
flower of triad and its bract. G Flower, vertical section. H maturity, they turn reddish or purple and are
Stamen and tepa! opposite this. I Stamen alternating with probably dispersed by birds (Carlquist 1977a).
tepals, anther locules opened. (Dahlgren and van Wyk 1988)
PHYTOCHEMISTRY. The leaves are tanninniferous,
but there are negative reports of fiavonoids or
Wyk 1988). The chalaza! end has a conspicuous, iridoids.
globose hypostase. At the cotyledonary stage the
massive cellular endosperm shows chalaza! and PHYLOGENY. At one time or another, families
micropylar haustoria (Fagerlind 1947). such as Santalaceae, Ericaceae, Hamamelidaceae
and Bruniaceae have been considered as the
PoLLEN MoRPHOLOGY. Pollen grains of G. ros- closest relatives of Grubbiaceae (Carlquist 1977b;
marinifolia are prolate (17 x 12J..Lm), tricolporate Dahlgren and van Wyk 1988). (The santalalean
with narrow colpi and lalongate ora, and tectate- affinity of Grubbia was based on its alleged
baculate. The tectum is psilate and not or incon- ategmic condition, which has proven to be wrong.)
spicuously pitted; tectal perforations seem to be Carlquist ( 1977b) was unable to definitely exclude
lacking (Dahlgren and van Wyk 1988). either of these possibilities but stressed the simi-
larity in wood anatomy between Grubbiaceae,
FRUIT AND SEED. The individual fruits are drupes Geissolomataceae and Bruniaceae. Supported by
with a thin, fleshy exocarp and a thick, bony endo- the presence of allegedly inverted anthers, endo-
carp. The syncarps of G. rosmarinifolia are com- sperm haustoria and unitegmic, tenuinucellate
posed of 2- 3 fused ovaries and do not exceed ovules, Cronquist (1981) and Takhtajan (1987)
1 mm in diameter; they may be barochorous or included Grubbiaceae in their narrowly circum-
wind-dispersed (sect. Ophira Burm. & L.). Many scribed Ericales. This was also supported by
Grubbiaceae 201
various cladistic (Anderberg 1992) and molecular Description as for family; 3 spp., S Africa.
(summarised by APG 1998) analyses, in which
Grubbiaceae together with Actinidiaceae appeared
among the "lower" Ericales. More recent mole- Selected Bibliography
cular studies, based on a broader sampling and
carefully selected outgroups and employing both Anderberg, A. 1992. The circumscription of the Ericales, and
chloroplast and nuclear genes (Xiang 1999; Xiang their cladistic relationships to other families of "higher"
et al. 2002), have resolved Grubbia with high sta- dicotyledons. Syst. Bot. 17: 660-675.
tistical support in the Cornales clade sister to Cur- references.
tisia. The close relationship between these two APG II (Angiosperm Phylogeny Group) 2003. See general
genera is also corroborated by a uniquely shared, references.
short insertion in the matK sequences (Xiang et al. Carlquist, S. 1977a. A revision of Grubbiaceae. J. S. Afr. Bot. 43:
115-128.
2002). Xiang et al. (2002) suggest to merge Curtisia Carlquist, S. 1977b. Wood anatomy of Grubbiaceae. J. S. Afr.
and Grubbia in a single family and adduce about Bot. 43: 129-144.
11 morphological characters common to them, Carlquist, S. 1978. Vegetative anatomy and systematics of
most of which, perhaps with the exception of the Grubbiaceae. Bot. Notiser 131: 117-126.
ridge connecting the bases of the opposite leaves, Cronquist, A. 1981. See general references.
Cronquist, A. 1983. Some realignments in the dicotyledons.
appear to be generalised cornalean characters, Nord. J. Bot. 3: 75-83.
whereas the fusion of the ovaries and resulting Dahlgren, R., van Wyk, A.E. 1988. Structures and relationships
syncarp structure in Grubbia and other differ- of families endemic to or centered in southern Africa. Ann.
ences (e.g. fruit structure) to my mind forbid such Missouri Bot. Gard. 25: 1-94.
a shotgun marriage. Endress, P.K., Stumpff, S. 1990. Non-tetrasporangiate stamens
in the angiosperms: structure, systematic distribution and
evolutionary aspects. Bot. Jahrb. Syst. 112: 193-240.
DISTRIBUTION AND HABITATS. On the basis of Fagerlind, F. 1947. Die systematische Stellung der Familie
their relatively inefficient water conducting system Grubbiaceae. Svensk Bot. Tidskr. 41: 315-320.
and the extensive infraspecific differentiation of Harms, H. 1935. Grubbiaceae. In: Engler, A., Harms, H. (eds.)
G. rosmarinifolia, Carlquist (1977a, 1977b) re- Die natlirlichen Pflanzenfamilien, ed. 2, 16b. Leipzig: W.
Engelmann, pp. 46-51.
garded the genus as a relictual group that has Hieronymus, G. 1889. Grubbiaceae. In: Engler, A., Prantl, K. Die
persisted in mesic pockets within the otherwise natiirlichen Pflanzenfamilien III, 1. Leipzig: W. Engelmann,
xeromorphic Mediterranean-type flora of the pp. 228-230.
Table Mountain sandstone of the Cape Province. Takhtajan, A.L. 1987. See general references.
van Tieghem, P. 1897. Sur les caracteres et les affinitees des
However, each of the three species occupies a Grubbiacees. J. Bot., Paris 11: 127-138.
different mesic habitat. Xiang, Q.-Y. 1999. Systematic affinities of Grubbiaceae and
Hydrostachyaceae within Cornales - insights from rbcL
A single genus: sequences. Harvard Pap. Bot. 4: 527-542.
Xiang, Q.- Y., Moody, M.L., Soltis, D.E., Fan, C.-Z., Soltis, P.S.
2002. Relationships within Cornales and circumscription of
Grubbia Bergius Fig. 66 Cornaceae - matK and rbcL sequence data and effects of
outgroups and long branches. Molec. Phylog. Evol. 24:
Grubbia Bergius, Kong. Vet. Acad. Handl. 28: 34 (1767); 35-57.
Carlquist, J. S. Afr. Bot. 43: 115-128 (1977), rev.
202 L. Hufford
Hydrangeaceae
L. HUFFORD
Hydrangeaceae Dumort., Anal. Fam.: 36, 38 ( 1829), nom. cons., among shoot axes. Cardiandra, Deinanthe, and
excluding Kania and Pottingeria. Kirengeshoma have only subterranean peren-
nating organs, and their aerial branches are
Perennial shrubs, vines, or herbs from woody rhi- herbaceous and annual. Populations of these three
zomes; bark exfoliating in strips or sheets. Leaves genera often consist of dense clusters of aerial
evergreen or deciduous, generally opposite, less stems. Cardiandra and Deinanthe evolved her-
commonly whorled or alternate, exstipulate; peti- baceous aerial stems independent of those in
oles prominent to inconspicuous; lamina simple, Kirengeshoma. Scrambling and vining shoots that
entire or toothed, uncommonly lobed. Inflores- climb by shoot-borne roots evolved separately
cences cymose, sometimes in corymbs, thyrses, or at least twice in Hydrangeae: (1) in Hydrangea
panicles; flowers few and large to numerous and anomala and Hydrangea section Cornidia, and (2)
small; sterile flowers with enlarged, showy calyces in Schizophragma, Pileostegia, and Decumaria.
in some. Flowers bisexual or unisexual, rarely Shoots of Whipplea are scandent, and scandent
dioecious; calyx imbricate or valvate; sepals 4-12, branches occur also in Platycrater and Philadel-
free or basally united; corolla imbricate, valvate, or phus. Barykina and Kapranova {1983) reported
convolute; petals 4-12, basally fused, or petals asexual reproduction through the rooting of scan-
completely united and calyptrate; androecium dent shoots in Philadelphus.
haplostemonous, diplostemonous, or polystemo- Leaves of most species are opposite, although
nous, stamens 4-numerous, free or basally united; some are whorled or alternate. Leaves are simple
filaments flat and linear, subulate, or filiform, dis- and are lobed only in Hydrangea quercifolia,
tal forks present or absent; anthers basifixed, H. sikokiana, and Kirengeshoma. Most leaves of
tetrasporangiate, with distal connective protru- Deinanthe bifida have a bilobed apex. Laminas
sion in some; gynoecium 2-12-carpellate, syn- of most species are largely ovate or elliptical.
carpous; ovary partially to completely inferior; Their margins range from prominently to incon-
placentation usually axile at the base and parietal spicuously toothed or entire. The laminas of
above, rarely strictly axile or strictly parietal; Carpenteria, Fendlera, Fendlerella, and various
ovules 2-many, anatropous; stylodia free, or a Philadelphus from the arid North American West
single style, sometimes with distal style branches; have entire margins. Entire margins evolved in
stigmas usually papillate, less commonly smooth. parallel in the two clades of climbing species,
Fruits capsules or berries. Seed length 10mm or notably in Pileostegia and in Hydrangea section
less; testa sculpture reticulate. Cornidia.
A family of 17 genera and about 220 species
distributed in temperate and subtropical regions ANATOMY. Gregory (1998) provided a compre-
of the Americas, Pacific islands, Asia, and hensive review of vegetative anatomy. All Hydran-
Europe. geaceae, except Hydrangea section Cornidia,
Kirengeshoma, and Pileostegia, have erect
VEGETATIVE MORPHOLOGY. Shoots are sympo- unicellular hairs that typically have a tuberculate
dial (Zhou and Hara 1988, 1989). Inflorescences surface and usually a pointed apex, and that
form at the apices of branches and the distal nodes are raised on multicellular bases (Hardin and
of branches. Most species are shrubs that have Pilatowski 1981; Gregory 1998). Similar trichomes
numerous branches at ground level, although are present also in the sister family Loasaceae.
these appear also to form subterranean rhizomes Two-armed, unicellular trichomes are present in
that affect asexual reproduction. Ishii and Takeda Kirengeshoma, whereas two-armed, multicellular
(1997) suggested that the strong basitonic branch- trichomes are present in Deinanthe. Stellate, uni-
ing and density-dependent turnover of stems lead cellular trichomes characterize Deutzia (Fig. 68F),
to intense intra-clonal competition for resources whereas stellate, multicellular trichomes are found
Hydrangeaceae 203
in both Pileostegia and Hydrangea section Corni- Bark that exfoliates in strips or sheets is charac-
dia. Trichomes that have a multicellular stalk and teristic of the family (absent only in a few species
a single apical cell occur in some species of of Hydrangea) and is possibly a derived state
Philadelphus. shared with the Loasaceae, in which the condition
Druse crystals have been reported in Carpente- is also common.
ria, Fendlera,]amesia, Philadelphus, and Whipp lea. Stomates are usually anomocytic, varying to
Many species of Deutzia and Kirengeshoma share paracytic in Deutzia, but are strictly paracytic in
small, spherical or lenticular crystals. Crystallifer- Dichroa and some species of Hydrangea (Gregory
ous idioblasts that bear raphides embedded in 1998).
mucilage are characteristic of tribe Hydrangeae Leaf venation is acrodromous in Fendlera,
(Umemoto 1974; Gregory 1998). Excluding the Fendlerella, Philadelphus, and Whipplea, but
wood, crystals have not been observed in pinnate in all other genera (Gregory 1998). Sec-
Fendlerella but have been reported for all other ondary veins in the pinnately veined taxa vary
genera. slightly: Carpenteria, Deutzia, and ]amesia have
Nodes are generally trilacunar and three-trace, basal secondary veins that are eucamptodromous,
although additional traces have been reported but most others are brochidodromous (or semi-
in Broussaisia, Kirengeshoma, Schizophragma craspedodromous in Deutzia). Kirengeshoma is
(Dhillon 1975; Kapranova 1976; Gregory 1998). craspedodromous, with transitions to actinodro-
Some Hydrangea are 5- or 7-lacunar and 5- or mous. Among Hydrangeae, Cardiandra, Dein-
7-trace, respectively. anthe, and Platycrater are eucamptodromous;
Wood anatomy has been described by Burkett Decumaria and Pileostegia are brochidodromous;
(1932), Quibell (1972), Stern (1978), Styer and Schizophragma is cladodromous, whereas Hy-
Stern (1979a, 1979b), and Snezhkova (1986, 1990). drangea s.s. is brochidodromous, although tending
Growth rings are generally present. The wood to semicraspedodromous in section Cornidia
ranges from diffuse-porous to ring-porous. Vessels (Watari 1939; Stern 1978; Hao and Hu 1996a;
are usually solitary, in radial multiples, and clus- Gregory 1998).
ters of a few cells, but Broussaisia is reported to
have strictly solitary vessels. Perforation plates FLOWER STRUCTURE. The floral dimorphism cen-
are oblique and exclusively scalariform in most tered on perfect fertile flowers and sterile display
species. Whipplea and some species of Dichroa flowers that have an enlarged calyx (Fig. 70) is
and Philadelphus have both scalariform and a derived feature characteristic of Hydrangeae
simple perforation plates. Perforation plates (but lost in Broussaisia, Dichroa, Pileostegia,
are generally simple in Fendlerella. Imperforate Decumaria, and some species of Hydrangea
elements consist of tracheids and fiber tracheids section Cornidia) (Hufford 1997). Floral dimor-
in Broussaisia, Carpenteria, Deutzia, Dichroa, phism associated with dioecy is characteristic of
Fendlerella, some species of Philadelphus, the Hawaiian island endemic Broussaisia.
and Hydrangea arborescens; only tracheids in The fertile flowers are actinomorphic and epig-
Fendlera, Fendlerella, ]amesia, some species of ynous. Ovaries are largely completely inferior in
Philadelphus, H. heteromalla, H. paniculata, and genera such as Deutzia, Decumaria, some Dichroa
H. quercifolia; only fiber tracheids in Decumaria, and most Hydrangea; closer to half inferior in
Schizophragma, most species of Hydrangea, and in Carpenteria, most Philadelphus, Broussaisia
the late wood of Whipp lea; and fiber tracheids and (female flowers), some Hydrangea and Dichroa;
septate fibers in Platycrater. Axial parenchyma and less than half inferior for instance in ]amesia,
is present in most genera, although absent in Kirengeshoma and Broussaisia (male flowers).
Platycrater, some species of Deutzia, and most Among most Hydrangeae, perianth merosity
Hydrangea. Rays are both uniseriate and multise- varies between tetramerous and pentamerous.
riate in most genera, except bi- and multiseriate in ]amesia, Fendlerella, and Whipplea typically have
Schizophragma, uni- and biseriate in Fendlerella, a pentamerous perianth, whereas that of Fendlera
and only uniseriate in Whipplea. Uniseriate rays and Philadelphus is typically tetramerous. In Car-
are generally heterocellular, but consist only penteria and Deinanthe, the perianth is pentamer-
of upright cells in Deutzia, Philadelphus, and ous to octamerous. The Decumaria perianth is
Whipplea. Multiseriate rays are heterocellular most commonly octamerous (Fig. 71B), although
(also the biseriate rays of Fendlerella), except in it ranges from hexamerous to dodecamerous.
Platycrater and some species of Hydrangea in Sepals are usually basally connate (most promi-
which cells are upright and square. nent in Kirengeshoma), but they appear more-or-
204 L. Hufford
less free in Carpenteria, Deinanthe, and Philadel- only parietal placentation. Examined Fendlerella
phus. The petals are free or tenuously connate at and Whipplea have axile placentation.
the base in all except Pileostegia and Hydrangea Ovules are pendant in Decumaria, Fendlera,
a no mala, in which the petals can be entirely united Fendlerella, Kirengeshoma, Pileostegia, Philadel-
and abscise as a unit at anthesis. The corolla is phus, Schizophragma, and Whipplea; oriented
funnel- to bowl-shaped, rotate, erect, or reflexed. upward in examined Deutzia, and H. heteromalla;
Both diplostemonous and polystemonous and- and horizontal in Broussaisia, Carpenteria,
roecia are common (Wettstein 1893; Gelius 1967; Dichroa, H. anomala subsp. petiolaris, H. hirta,
Hufford 1998). Haplostemony is present only in and H. macrophylla. In Cardiandra, Deinanthe,
Dichroa pentandra and D. platyphylla. Stamen ]amesia, and Platycrater ovule orientation varies
filaments of Fendlera and various species of with location.
Deutzia are distally flanged, and these form a Taxa that have style branches or stylodia have a
corona in the center of the flower. Filaments are stigma at the apex of each; the stigmatic zone
usually bifacial, at least basally, and strongly so in extends down the furrow on the ventral side in
Fendlerella, ]amesia, Whipplea, and especially in Fendlera, Fendlerella, and Whipplea. The com-
Fendlera and some Deutzia, although they are uni- pletely synstylous genera, including Deinanthe,
facial over most of their length in Carpenteria, have stigmas positioned along furrows on the
some Deutzia, Kirengeshoma, Philadelphus, and outer surface of the stylar apex. Stigmas are papil-
Hydrangeae. Anthers are basifixed, bithecate and late, except in Deinanthe. Stigmas of Deinanthe
tetrasporangiate. The connectives have distal pro- have a smooth surface that becomes coated by
trusions that project beyond the thecal junction in secretions.
Fendlera,Fendlerella,]amesia, Whipplea and some The functionally dioecious Broussaisia has
species of Deutzia. Fendlerella,]amesia, Whipplea, male flowers that have a sterile pistillode, and the
and Decumaria sinensis have a fleshy protrusion female flowers lack stamens or have rudimentary
on the abaxial side of the connective. The endothe- staminodes.
cium consists of a single cell layer, except at the
ends adjacent to the connective on the abaxial EMBRYOLOGY. Pollen is released at the binucleate
side of the anther at which 2-4 layers are present stage. The unitegmic, tenuinucellate ovules are
for a short distance. Endothecium is limited to anatropous. Embryo sac development is Poly-
the thecal regions. Two to three layers of thin, gonum type. The lower half of the embryo sac is
elongate cells are positioned between the endothe- enclosed by an endothelium. The embryo sac
cium and tapetum. The tapetum may be present extends into or beyond the micropyle in various
or absent at dehiscence (Saxena 1971). Anther genera. These genera have micropylar haustoria.
dehiscence is slightly introrse in ]amesia, The egg apparatus of examined Deutzia and
Fendlera, Fendlerella, Whipplea, Carpenteria, Philadelphus is positioned outside of the ovule.
Philadelphus, Kirengeshoma, and some Deutzia, The antipodal cells of Kirengeshoma become haus-
whereas Hydrangeae have evolved latrorse dehis- toria!. The polar nuclei fuse before fertilization.
cence, but Broussaisia and Decumaria sinensis Endosperm formation is ab initio cellular, except
have a reversal to introrse dehiscence. in Fendlera (Mauritzon 1933, 1939; Davis 1966;
Ovaries are connate over their entire length Saxena 1971).
(Morf 1950). They are synascidiate at the base
(characterized by continuous septa), but sympli- POLLEN MoRPHOLOGY. Pollen is relatively uni-
cate in the mid and upper regions (characterized form among the genera (Agababyan 1961;
by discontinuous septa) (Klopfer 1971, 1973). Wakabayashi 1970; Pastre and Pons 1973; Hideux
The genera differ in the extent of stylar conna- and Ferguson 1976; Hao and Hu 1996b). The
tion: Cardiandra, Deutzia, Fendlera, Hydrangea, tricolporate grains are spheroidal to slightly
Kirengeshoma, Platycrater, and some Dichroa have prolate or oblate. The exine is tectate to semi-
completely separate stylodia; genera such as tectate. The tectum is perforate to reticulate. The
Fendlerella and Philadelphus have styles with free exine can have a secondary sculpture of minute
style branches; and others (e.g., Broussaisia and granulations. Pollen grains are small {11.2-22.4 x
Schizophragma) have simple styles. Deinanthe also 9.8-21.0 J.tm).
has a simple style, but this arises through the post-
genital fusion of initially free stylodia. KARYOLOGY. Base chromosome numbers among
Placentae of most genera are axile at the base the genera include x = 11, 13, 14, 15, 16, 17, and 18.
and parietal distally. Some species of Deutzia have Sax (1931) reported the regular formation of poly-
Hydrangeaceae 205
ploid series among species of Deutzia. Tetraploids Deutzia fruits have two regions of dehiscence.
have also been reported in Hydrangea (Sax 1931), They are septicidal but, unlike the above taxa, split
although they are uncommon. Chromosome acropetally along ovarian septa from the base of
numbers do not vary among species of Philadel- the fruit. The individual carpels spread slightly at
phus, but morphological differences among the the base, creating a broad gap through which most
chromosomes of different species interfere with seeds appear to be dispersed. In addition, the fruits
their pairing in interspecific crosses and limit also dehisce apically, creating an opening between
natural hybridizations (Ammal 1951). Hamel the stylodia.
(1951) demonstrated size differentiation among Most Hydrangeae, including Cardiandra,
chromosomes in Deinanthe, Kirengeshoma, and Deinanthe, Platycrater, and Hydrangea, in which
Schizophragma. Chromosome complements of seed sizes are relatively small (Hufford 1995), have
Deutzia are reported to be little differentiated in dehiscence that creates a gap between the stylodia.
either size or shape (Ohba and Akiyama 1992). This has been lost in association with an increase
in seed length in Schizophragma, Pileostegia, and
REPRODUCTIVE SYSTEMS. The reproductive bio- Decumaria, in which intercostal portions of the
logy of Hydrangeaceae remains largely unex- lateral walls of the ovary separate from the costal
amined. Flower size and number as well as ovule ribs. After the intercostal pieces have fallen from
size and number vary among the clades. Trade-offs the fruits, they have a cage-like appearance.
in these features, which are associated with Among Hydrangeae, the tropical genera Brous-
changes in stamen number and pollen:ovule saisia and Dichroa shifted from dry, dehiscent
ratios, appear to be central to reproductive strate- fruits to berries. Those of Dichroa are bright pink
gies (Hufford, unpublished data). to bluish, and of Broussaisia purplish, red, or
In Hydrangea, Pilatowski (1981) reported that pinkish. Seeds of Broussaisia are surrounded by
the densely flowered inflorescences serve as a finger-like ingrowths of the fruit wall.
platform on which various unspecialized pollin- The inner wall layers surrounding the locules
ators land and forage. Robertson (1892) found differentiate during late ovary development as a
Hydrangea arborescens was visited primarily by mechanical region that has lignified cell walls. The
many kinds of bees (including various species of cells of this region are oriented perpendicular to
Halictus) and flies as well as by a few coleopterans the long axis of ovary. These layers oflignified cells
and lepidoterans. are lacking in Dichroa.
The tropical Broussaisia is the only member of
the family known to be dioecious. Degener (1945) SEEDS. Seeds are 5 mm or less in length, except in
reported that the flowers were visited by beetles Kirengeshoma in which seeds are up to 10mm. The
and wasps. embryo is straight and embedded in endosperm
(Corner 1976). Seeds are mostly ellipsoidal, ovate,
FRUIT. Fruits are capsular, except in the tropical or spindle-shaped, but funnel-form in Philadel-
Broussaisia and Dichroa. Capsular fruits open phus, urceolate in Broussaisia, Dichroa, and their
variously. Septicidal dehiscence that proceeds close relatives Hydrangea hirta and H. scandens,
basipetally from the base of the stylodia is obovate, disc-like in H. anomala, horn-shaped in
plesiomorphic and characteristic of Fendlera, Decumaria barbara, and bottle-form in Pileostegia
]amesia, Fendlerella, Whipplea, and Carpenteria. (Hufford 1995). All have a reticulate testa sculp-
The basipetal splits extend to mid fruit in Fendlera ture, although secondary sculptures vary, in-
and ]amesia. Dehiscence proceeds to near the base cluding papillae, granulations, rugae, striations,
of the fruit in Fendlerella and Carpenteria, al- and smooth. Exotesta cell walls are lignified
though the style of Carpenteria restricts the (Nemirovich-Danchenko and Lobova 1998). Seeds
apical separation of the fruit, creating window-like are typically winged, and the wing is usually
openings maintained until the styles break. In restricted to the chalaza! end but extends entirely
Whipplea, the individual, single-seeded carpels around the seed in Kirengeshoma and Hydrangea
separate entirely. Philadelphus has loculicidal cap- a no mala. Schizophragma, most Philadelphus, most
sules that dehisce basipetally from the base of the Deutzia, and Decumaria sinensis have a micropy-
style to the lower half of the fruit. Kirengeshoma lar flange that may function as wing. Fendlerella
has both septicidal and loculicidal dehiscence that and Whipplea have a funicular appendage.
progresses basipetally but extends little from the
apex of the ovary, mimicking the dehiscence of DISPERSAL. The small, winged seeds of most
Hydrangeae (especially Deinanthe). Hydrangeaceae are undoubtedly wind dispersed.
206 L. Hufford
They are readily thrown from the dehisced fruits AFFINITIES. Hydrangeaceae are the sister clade of
when branches move in wind. A seed wing is Loasaceae (Downie and Palmer 1992; Hempel et al.
lacking in Broussaisia, Dichroa, and their close rel- 1995; Soltis et al. 1995; Xiang et al. 1998; Hufford
atives H. hirta and H. scandens, which have urce- et al. 2001). They are well supported as a clade of
olate seeds. This is associated with a shift to Cornales (sensu APG 1998).
indehiscent, fleshy fruits and presumably animal Hydrostachys has been affiliated with
dispersal in Broussaisia and Dichroa. Hydrangeaceae in various molecular systematic
studies (Hempel et al. 1995; Albach et al. 2001);
PHYTOCHEMISTRY. Most genera of Hydran- however, its exceptional branch length makes this
geaceae contain secoiridoids and loganin (Jensen placement dubious (Hufford 1997; Hufford et al.
et al. 1975). Deutzia is unique among examined 2001).
members of the family in having iridoids that lack
C-10 (Frederiksen et al. 1999). Kaempferol and DISTRIBUTION AND HABITATS. Soltis et al. (1995)
quercetin are widespread; myricetin was found suggested that Hydrangeaceae had a New World
in Decumaria and ]amesia (Bohm et al. 1985). origin. Hydrangeaceae and their sister clade
Procyanidin and prodelphinidin have repeatedly Loasaceae may have originated in mesophytic or
been recorded (Bate-Smith 1978), and some xerophytic environments of Central America or
Hydrangea were found to contain ellagic and gallic southwestern North America. The three largest
acids (Hegnauer 1973). The alkaloids febrifugin genera, Deutzia, Hydrangea (also the mono-
and isofebrifugin (see Economic Importance phyletic Hydrangea s.l.) and Philadelphus, are dis-
below) have been isolated from Dichroa and junct between the Old and New Worlds. The two
Hydrangea (Hegnauer 1973). Hydrangea has fatty species of Decumaria, which are part of Hydrangea
acid compounds dominated by linoleic acid, as is s.l., are disjunct between eastern Asia and the
common among most other Cornales (Breuer et al. southeastern United States. Hu (1954) suggested
1987). that the most primitive species of Philadelphus are
in mesophytic areas of Central America, and that
SUBDIVISION AND RELATIONSHIPS IN THE major clades of the genus continue to have extant
FAMILY. Phylogenetic analyses (Soltis et al. 1995; representatives in mountainous areas of Mexico,
Hufford et al. 2001) indicate that ]amesia in which the genus radiated. Deutzia is disjunct
and Fendlera form a monophyletic group (sub- between Mexico and Central and eastern Asia. It is
fam. Jamesioideae) that is the sister clade to the unclear whether the Mexican species of Deutzia
rest of the family (subfam. Hydrangeoideae). (sect. Neodeutzia), which are uniquely polystemo-
Hydrangeoideae consist of the sister tribes nous in the genus, are a relictual group of an initial
Philadelpheae and Hydrangeae (sensu Hufford et radiation of the genus in Central America or a
al. 2001). In the Philadelpheae, current phy- derived clade. Hydrangea sect. Cornidia, which is
logenetic sampling indicates that Carpenteria limited to Central and South America, except for
and Philadelphus are sister clades, Deutzia and the disjunct H. integrifolia Hayata of the Philip-
Kirengeshoma are sister clades, and Fendlerella pine Islands and Taiwan, is nested as a derived
and Whipplea are sister clades. The sister of the clade of Hydrangea s.l. and, thus, may represent
Fendlerella- Whipplea clade within Philadelpheae a secondary expansion into Central America.
remains unclear. Relationships are less well As discussed below (Paleobotany), clades of
resolved in Hydrangeae (Hufford 1997; Hufford et Hydrangeoideae were widespread in the Northern
al. 2001). Cardiandra and Deinanthe are mono- Hemisphere during the Tertiary, and the fragmen-
phyletic, and there is some support for their place- tation and migration of Tertiary forest elements
ment as the sister of the rest of the Hydrangeae may account for the current distribution of
(=Hydrangea s.l.). Hydrangea s.str. is paraphyletic Hydrangea sect. Cornidia.
(Soltis et al. 1995; Hufford 1997; Hufford et al. Fendlera, Fendlerella, ]amesia, and Whipplea
2001). Broussaisia, Decumaria, Dichroa, Pileoste- are restricted to northern Mexico and western
gia, Platycrater, and Schizophragma evolved North America. Axelrod and Raven (1985) sug-
among the clades of Hydrangea s.str. Broussaisia gested ]amesia originated in the Southern Rocky
and Dichroa form a monophyletic group with H. Mountains (U.S.A) in the Eocene. Holmgren and
hirta, H. macrophylla, and H. scandens. The vines Holmgren (1989) suggested ]amesia spread to the
Decumaria, Pileostegia, and Schizophragma form west after Miocene uplifts.
a monophyletic group in Hydrangea s.l. (Soltis Various smaller genera of Hydrangeae are
et al. 1995; Hufford 1997; Hufford et al. 2001). limited in distribution to Asia. The vining Pileoste-
Hydrangeaceae 207
gia and Schizophragma are found on Taiwan and polonica from Europe. Dichroa bornensis,
in eastern Asia and the Himalayan region. Deinan- described from the Middle Oligocene of Europe by
the and Platycrater are distributed in southern Mai (1985), lacks character states sufficient to
Japan and China in moist, temperate forests. Car- include it in the extant genus Dichroa (Hufford
diandra, the sister clade of Deinanthe, is also 1995) but can be reasonably assigned to the
found in Japan and China, and also on Taiwan. "macrophylla'' clade of Hydrangea (Hufford et al.
Dichroa, which is distributed through southern 2001).
China, the Himalayas, and Southeast Asia (includ- Philadelphus fossils similar to the extant P.
ing the Malay Archipelago, Philippine Islands, and lewisii have been found from the Pliocene of
New Guinea), is nested within Hydrangea s.l. as California, U.S.A. (Condit 1944) and Upper Creta-
part of the "macrophylla'' clade (Hufford et al. ceous of Vancouver Island, Canada (Bell 1957),
2001). The "macrophylla'' clade also includes H. which are close to the current range of the extant
macrophylla, H. scandens, and H. hirta, and cur- species. Subfam. Jamesioideae is represented by
rently extends from the Himalayas through China fossil leaves of ]amesia caplanii Axelrod from
to Taiwan and Japan. The Hawaiian endemic the Oligocene of southwestern Colorado, U.S.A.
Broussaisia is also associated with the "macro- (Axelrod 1987). These fossil leaves are very similar
phylla" clade, and its present distribution may to those of the extant ]. americana, which is cur-
represent a long distance disperal from Asia. rently part of the flora of Colorado and the south-
]amesia and Fendlera, which compose sub- western U.S.A.
fam. Jamesioideae, grow in rocky, often steep
habitats that are exposed and relatively arid. In ECONOMIC IMPORTANCE. Roots of Dichroa febri-
tribe Philadelpheae (subfam. Hydrangeoideae), fuga have been used as a traditional source of anti-
Fendlerella and some species of Philadelphus grow malarial drugs in Asia (Fairbairn and Lou 1950;
in environments very similar to those of ]amesia Steinmetz 1972; Murata et al.1998; Kim et al. 2000).
and Fendlera. Carpenteria, Whipplea, and many Compounds from these roots are a potential
other species of Philadelphus are found in only source of anti-inflammatory drugs (Kim et al.
slightly less exposed, arid habitats. Kirengeshoma 2000). Dried roots of Hydrangea arborescens
and members of tribe Hydrangeae (subfam. have been used as a diuretic and diaphoretic
Hydrangeoideae) are found in more mesophytic (Spongberg 1972). Deutzia has been used in China
habitats. Many Hydrangeae are found in shaded to to treat enuresis, malaria, and scabies (He 1990).
open forests or forest margins, especially on slopes, The Japanese prepare the sweet beverage
cliffs, riparian banks. Broussiasia, Dichroa, and amacha from steamed and dried leaves of
Deinanthe are found in more moist, shaded habi- Hydrangea macrophylla (Spongberg 1972).
tats. The shifts of Broussaisia and Dichroa into wet Many genera are important ornamentals, espe-
tropical forests are clearly derived in the family. cially Deutzia (Za'ikonnikova 1966), Hydrangea
(McClintock 1956), Kirengeshoma (Cannon 1981),
PALEOBOTANY. Hydrangeaceae have a rich fossil Philadelphus (Wyman 1965), and Schizophragma
record that includes seeds, leaves, sterile flowers, (Nevling 1964).
and fruits. Most fossil Hydrangeaceae are from
the Tertiary when members of the family appear CoNSERVATION. Carpenteria is of conservation
to have been common elements of mesophytic concern in California, U.S.A. because of habitat
forests in the northern hemisphere (Hu and destruction. Kirengeshoma is rare in Japan.
Chaney 1940; Ozaki 1991; Manchester 1994; Mai
1998). SUBFAMILIAL CLASSIFICATION
Tribe Hydrangeae was more widespread in the I. Subfam. Jamesioideae Hufford (2001).
Northern Hemisphere during the Tertiary than Genera 1-2.
currently. Hydrangea fossils are known variously II. Subfam. Hydrangeoideae A. Br. (1864).
from the Oligocene to Pliocene in Europe and 1. Tribe Philadelpheae Reichenbach (1828).
western North American where they are not Genera 3-8.
members of the extant flora (Chaney and Sanborn 2. Tribe Hydrangeae DC. (1830).
Genera 9-17.
1933; MacGinitie 1941; Gorbunov 1970; Lancucka-
Srodoniowa 1975; Mai 1985; Manchester 1994).
Similarly, Schizophragma is known in extant floras KEY TO THE GENERA
only from eastern Asia, but Mai (1985) has 1. Aerial shoots strictly herbaceous and annual 2
described Pliocene fossils of Schizophragma - Aerial shoots woody and perennial 4
208 L. Hufford
2. All flowers fertile (sterile display flowers lacking); sepals - Stellate trichomes lacking; stylodia free, or style simple or
united for over half of their length; petals largely erect, branched; fruits either loculicidal or septicidal but not
curving outward only at their tips 6. Kirengeshoma both septicidal and apical between stylodia 15
- Both fertile and sterile flowers (the latter with enlarged 15. Stamens> 15; fruits loculidical 7. Philadelphus
calyces); sepals only slightly united at the base; petals - Stamens <15; fruits septicidal 16
reflexed from the base 3 16. Leaves entire; stamen filaments have two distal flanges;
3. Flowers less than 1 em wide; stamens 12-20; stylodia seeds >4 mm long 2. Fendlera
separate, stigmas terminal and papillate - Leaves toothed; stamen filaments lack distal flanges; seeds
10. Cardiandra <2mm long l.]amesia
- Flowers more than 1.5cm wide; stamens >100; style bottle-
brush-like because of protrusive stigmatic zone, stigmas GENERA OF HYDRANGEACEAE
smooth 9. Deinanthe
4. Style simple; stigma either capitate or with bottle-brush
furm 5 I. SUBFAM. JAMESIOIDEAE Hufford (2001).
- Stylodia free or style branched into style branches, lacking
capitate or bottle-brush stigmas 9 Shrubs. Sterile flowers absent. Perianth 4-5-
5. Shrubs with numerous basal branches; fruits septicidal
capsules or indehiscent berries 6
merous, petals imbricate, androecium diplostemo-
- Vines; fruits dehisce through loss of lateral intercostal nous, carpels 3-5. Fruit capsular, seeds <6 per
walls 7 locule.
6. Leaves deciduous; flowers bisexual; flowers more than 3 em
wide; stamens >200; fruits septicidally dehiscent; seeds
elliptical with a thin wing 8. Carpenteria 1. ]amesia Torr. & Gray Fig. 67
- Leaves evergreen; flowers unisexual, plants dioecious;
flowers less than 1.5 em wide, male flowers with 8-12 ]amesia Torr. & Gray, Fl. N. Am. 1: 593 (1840); Holmgren and
stamens; fruits fleshy, indehiscent (berry-like); seeds Holmgren, Brittonia 41: 335-350 (1989), rev.
urceolate, lacking a wing 13. Broussaisia
7. Leaves deciduous; petals separate or tenuously connate at Shrubs. Leaves deciduous, petiolate, lamina
their apices; seeds ellipsoidal with micropylar flange and toothed. Inflorescence cymose panicle. Sterile
chalaza) wing or horn-shaped with broadened micropylar
pole and pointed chalaza) wing 8 flowers absent. Sepals 4-5, valvate. Petals 4-5,
- Leaves evergreen; petal margins connate and corolla calyp- imbricate, spathulate, white to pinkish. Stamens
trate (abscising at anthesis); seeds bottle-shaped with 8-10, filaments subulate, anthers with abaxial
neck-like micropylar pole and rounded chalaza) pole
15. Pileostegia
8. Inflorescences of both sterile and fertile flowers; petals 4-5;
stamens 8- 10; seeds ellipsoidal 14. Schizophragma
- Inflorescences of only fertile flowers; petals usually 6-12;
stamens >12; seeds horn-shaped 16. Decumaria
9. Ovules 1 per locule; seeds with funicular appendage (easily
detached in Fendlerella) 10
Ovules at least 4 per locule; seeds without funicular
appendage 11
10. Erect shrubs; leaves entire; capsules ovoid to ellipsoidal;
seeds straight 3. Fendlerella
- Trailing shrubs; leaves toothed; capsules globose; seeds
slightly crescent-shaped 4. Whipplea
11. Inflorescences generally with both sterile and fertile
flowers; petals reflexed or absent; fruits dehiscing only
apically, forming an opening between the stylodia 12
- Inflorescences only with fertile flowers; petals funnel-form
to rotate or reflexed; fruits apically or basally septicidal (if
the latter then also with apical interstylar opening), or
loculicidal, or berries 13
12. Flowers less than 1 em wide; stamens fewer than 30
11. Hydrangea
- Flowers greater than 1 em wide; more than 200 stamens
17. Platycrater
13. Corolla valvate; petals reflexed; fruits berries; seeds
urceolate 12. Dichroa
- Corolla imbricate or valvate; petals funnel-form to rotate;
fruits capsular; seeds ellipsoidal to funnel-form 14
14. Stellate trichomes on vegetative and reproductive struc-
Fig. 67. Hydrangeaceae-Jamesioideae. ]amesia americana. A
tures; stylodia completely separate; fruits dehiscing both
Flowering branch. B Floral bud. C Flower. D Petal and stamens.
apically between the stylodia and septicidally from base
E Pistil with free stylodia. FFruit, after dehiscence. (Holmgren
acropetally 5. Deutzia
and Holmgren 1989)
Hydrangeaceae 209
and distal connective protrusion. Ovary 3-5- absent. Sepals (4)5, apert. Petals (4)5, apert,
carpellate, partly inferior, style short with long spathulate, white. Stamens (8)10, filaments dor-
style branches, or stylodia free; ovules oriented siventrally flattened and subulate, anthers have
variously. Fruit septicidal capsule. Seeds -20, abaxial and distal connective protrusion. Ovary
0.5-1 mm long, winged, without micropylar flange. (2)3( 4-5)-carpellate, partly inferior; short simple
n = 16 (Sax 1931; Hamel 1953; Love 1969). Two style with style branches, ovules pendant. Fruit
species. Most common in the Southern Rocky septicidal capsule. One seed per carpel, 1-2 mm
Mountains, U.S.A., but scattered in various moun- long, wing and micropylar flange lacking, funicu-
tain ranges of the American Great Basin and lar appendage present. Four species, in northern
Southwest and northern Mexico. Mexico and American Southwest in shrub or
pinyon pine-juniper communities on steep slopes
and canyon walls. Revision needed.
2. Fendlera Engelm. & Gray
Fend/era Engelm. & Gray, Smithson. Contrib. 3: 77 (1852).
4. Whipplea Torr.
Shrubs, bark of twigs longitudinally ridged. Leaves Whipplea Torr., Pacif. Rail. Rep. 4: 90 (1856).
deciduous, petiole inconspicuous, lamina entire.
Inflorescence dichasial cyme. Sterile flowers Shrubs, trailing or low growing, bark of twigs
absent. Sepals 4, valvate. Petals 4, imbricate, longitudinally ridged. Leaves deciduous, petiolate,
spathulate, white. Stamens 8, filaments broad, lamina toothed. Cymose panicle. Sterile flowers
linear, dorsiventrally flattened with forked apex, absent. Sepals (4)5( 6), apert. Petals (4)5( 6), apert,
anther with distal connective protrusion. Ovary spathulate, white. Stamens (8)10(12), filaments
4-carpellate, partly inferior, stylodia free, ovules dorsiventrally flattened and subulate, anthers with
pendant. Fruit septicidal capsule. Seeds 15-25, abaxial and distal connective protrusion. Ovary
4-5.5 mm long, wing and micropylar flange (2-3 )4-5-carpellate, partly inferior, style short
lacking. n = 11 (Sax 1931; Hamel1953). Two to four with style branches; ovules pendant. Fruit septici-
species, in the American Southwest and northern dal capsule. One seed per locule, 1-1.5 mm long,
Mexico in shrub or pinyon pine-juniper commu- wing and micropylar flange lacking, funicular
nities on steep slopes and canyon walls of high appendage present. One species, Whipplea mode-
deserts. Revision needed. sta Torr. In California, Oregon, and Washington,
U.S.A., in dry, rocky areas of open forests of coast
ranges.
II. SUBFAM. HYDRANGEOIDEAE A. Br. (1864).
5. Deutzia Thunb. Fig. 68
Shrubs or herbaceous perennials. Sterile flowers
absent or present. Perianth 4-12-merous, petals Deutzia Thunb., Nov. Gen.: 19 (1781); Zaikonnikova, Deutzias
imbricate or valvate, androecium diplostemonous, - ornamental shrubs - a monograph of the genus Deutzia
haplostemonous, or polystemonous, carpels 2 or Thunb. (1966), rev.; Zaikonnikova, Baileya 19: 133-144
(1975), key; He, Phytologia 69: 332-339 (1990), rev. Sichuan,
more. Fruit capsule or berry, seeds 1-many_per China spp.
locule.
Shrubs. Leaves deciduous, petiolate, lamina
toothed. Infloresence cymose panicle, dichasial
1. TRIBE PHILADELPHEAE Reichenbach (1828). cymes. Sterile flowers absent. Sepals 5, valvate.
Petals 5, imbricate or valvate, spathulate to ovate,
Shrubs or herbaceous perennials. Sterile flowers white to pinkish. Stamens 10-15, stamen filaments
absent. Capsular fruits either loculicidal or septi- subulate to linear with forked apex. Ovary 3-5-
cidal or both. carpellate, completely inferior, stylodia free, ovules
horizontal. Fruit capsular, dehisces both apically
3. Fendlerella A.A. Heller between styles and acropetally septicidal from the
base of the fruit. Seeds numerous, 0.5-3 mm long,
Fendlerella A.A. Heller, Bull. Torrey Bot. Club 1828: 626 (1828). winged. n = 13, 26, 39, 52,65 (Sax 1931; Love 1969;
Ohba and Akiyama 1992). Approximately 60
Shrubs, bark of twigs longitudinally ridged. Leaves species. Zaikonnikova (1966) recognized the sec-
deciduous, petiole inconspicuous, lamina entire. tions Neodeutzia Engl., Mesodeutzia C.K. Schn.,
Inflorescence cymose panicle. Sterile flowers and Deutzia as well as six subsections and 16
210 L. Hufford
Fig. 69. Hydrangeaceae-Hydrangeoideae. A-D Philadelphus

inodorus. A Flowering branch with dehisced capsules from
previous season. B Flower and bud with valvate calyx lobes. C
Style with style branches. D Transverse section of ovary. E-H
P. hirsutus. E Ovary in vertical section showing auriculate
placentae and pendulous ovules. F Young fruit soon after dis-
Fig. 68. Hydrangeaceae-Hydrangioideae. Deutzia paniculata. articulation of corolla and androecium showing the ovarian
A Flowering branch. B Flower. C Flower, seen from beneath. D disk. G Mature, dehisced capsule. H Seed. (Spongberg 1972)
Ovary, transverse section. E Fruiting twig. F Underside of leaf.
Drawn by T. Yamada. (Nakai 1926)
winged. 2n = 52, 54 (Hamel 1951; Funamoto and
series. The genus is disjunct between Mexico (only Nakamura 1989). Two species. Japan and Korea. In
the approximately four species of sect. Neodeutzia) mountain forests.
and Asia. Species of sects. Mesodeutzia and
Deutzia are distributed from Korea, Japan, Taiwan, 7. Philadelphus L. Fig. 69
and the Philippine Islands throughout most of
China and the Himalayan region. Most species are Philadelphus L., Sp. Pl.: 1: 470 (1753); Hu S.-Y., J.Arnold Arbor.
35: 275- 333 (1954), 36:52-109 (1955), 37: 15-90 (1956), rev.;
Asian (Zaikonnikova 1975; He 1990). Hwang, Acta Bot. Austro Sinica 7: 1-12 (1991), synopsis of
Chinese species.
6. Kirengeshoma Yatabe
Erect or semi-scandent shrubs. Leaves deciduous
Kirengeshoma Yatabe, Bot. Mag. (Tokyo): 5: 1 (1890). or persistent, petioles often inconspicuous, lamina
entire or toothed. Flowers solitary, in dichasial
Herbaceous perennial from rhizome. Leaves cymes, or cymes aggregated in panicles. Sterile
opposite-subopposite, petiolate, palmately lobed, flowers absent. Flowers often strongly fragrant.
lamina toothed. Inflorescence cymose panicle. Sepals 4{5), valvate. Petals 4{5), imbricate, spathu-
Sterile flowers absent. Sepals 5, valvate, prominent late to ovate, white. Stamens >15, filaments subu-
calyx tube. Petals 5, imbricate, spathulate, yellow to late to filiform. Ovary {3)4{5)-carpellate, partly to
creamy white. Stamens - 15, stamen filaments completely inferior, style with style branches;
subulate. Ovary 3- 4-carpellate, partly inferior, ovules pendant. Fruit loculicidal capsule. Seeds
stylodia free, ovules pendant. Fruit basipetally numerous, 1- 4 mm long, chalaza! wing, micro-
septicidal and loculicidal capsule. Seeds numer- pylar flange (rarely absent). n = 13 (Bangham 1929;
ous, 4-10 mm long, strongly flattened laterally, Sax 1931). Approximately 80 species. Asia, Europe,
Hydrangeaceae 211
North America and Central America. Tropical, and Kyushu) and D. caerulea from China. Both in
temperate, and arid environments in open forests, moist forests of mountains (Lancaster 1996).
scrublands, rocky slopes, and riparian banks.
Hu (1954) recognized four subgenera: Gemmatus
10. Cardiandra Sieb. & Zucc.
(Koehne) Hu, Macrothyrsus Hu, Deutzioides Hu,
and Philadelphus. Hu (1954) suggested that the Cardiandra Sieb. & Zucc. Fl. Jap.: 119 (1835).
genus originated in the New World, and pointed to
tropical and subtropical species of subgenus Gem- Herbaceous perennial from rhizome. Leaves
matus in Central America as the most primitive alternate, petiolate, lamina toothed. Inflorescence
extant species. cymose panicle. Sterile flowers present. Sepals
4-5(6), valvate. Petals 4-5(6), imbricate, ovate,
white, reflex at anthesis. Stamens 12-25, filaments
8. Carpenteria Torr.
filiform. Ovary 2-3(4-5)-carpellate, partly infe-
Carpenteria Torr., Smithson. Contrib. 6: 12 (1854). rior, stylodia free, most ovules erect. Fruit
capsular with apical dehiscence. Seeds numerous,
Shrubs. Leaves deciduous, entire, petioles in- 0.5-1.0 mm long, wing and micropylar flange
conspicuous. Inflorescence terminal and axillary present. About six species. Eastern Asia. Moist
dichasial cymes. Sterile flowers absent. Flowers forests in mountains. Revision needed.
often strongly fragrant. Sepals 5-7, valvate. Petals
5-7, imbricate, largely ovate with very short claw,
11. Hydrangea L. Fig. 70
white. Stamens >200, filaments filiform. Ovary
5-7 -carpellate, partly inferior, style simple; ovules Hydrangea L. Sp. Pl.: 397 ( 1754); McClintock, Proc. Calif. Acad.
pendant. Fruit septicidal capsule. Seeds numerous, Sci.29: 147-256(1957),rev.;Wei,Guihaia 14:101- 121 (1994),
0.5-1 mm long, winged, no micropylar flange. One rev. Chinese species.
species, Carpenteria californica Torr. Restricted
to dry ridges and slopes of the Sierra Nevada
foothills in California, U.S.A, between the San
Joaquin and King rivers. A conservation concern
because of habitat loss.
2. TRIBE HYDRANGEAE DC. (1830).
Shrubs or herbaceous perennials. Sterile flowers

usually present, rarely absent. Fruits either berries
or capsules with apical dehiscence or fragmenting
lateral walls.
9. Deinanthe Maxim.
Deinanthe Maxim. Mem. Acad. Petersb. VII, 16: 2 (1867).
Herbaceous perennial from rhizome. Leaves peti-

olate, lamina apex acute or bifid, margin toothed.
Inflorescence cymose panicle. Sterile flowers
present. Sepals S- 6, imbricate. Petals S- 8, imbri-
cate, ovate, white, pinkish purple to blue, reflex at
anthesis. Stamens >200, stamen filaments fila-
mentous. Ovary 4-6-carpellate, partly inferior,
stylodia connate postgenitally, ovules horizontal
and pendant. Fruit capsule with apical dehiscence
pulling apart the connate stylodia. Seeds numer-
ous, 0.5- 1.5 mm long, wing and micropylar flange Fig. 70. Hydrangeaceae-Hydrangioideae. Hydrangea serrata.
present. 2n = 34 (Hamel 1951). Two species: D. A Flowering branchlet. B Flower bud. C Flower. D Fruiting
bifida from Japan (southwest Honshu, Shikoku, branch. E Fruit. Drawn by T. Yamada. (Nakai 1926)
212 L. Hufford
Erect or scandent shrubs or vines, climbing by 13. ·Broussaisia Gaudich.

sho.ot-borne roots. Leaves deciduous or persistent,
Broussaisia Gaudich., Voy. Bot.: 479 (1826).
petiolate, margins rarely lobed, toothed or entire.
Inflorescence cymose panicle, often flat-topped
Shrub, stems hirsute (rarely glabrous), especially
or pyramidal. Sterile flowers present usually or
when young; leaf scars prominent (internodes
a?s~nt. Sep.als 4-:-5., valvate. Petals 4-5, valvate, per-
short). Leaves persistent at ends of branches
sistmg. until frmtmg and reflexed, abscising after
opposite or whorled, petiolate, lamina obovate t~
anthesis, or calyptrate and abscising at anthesis,
ovate or elliptical, margins toothed. Inflorescence
usually ovate, rarely spathulate, white, pink, blue,
cymose panicle. Sterile flowers absent. Dioecious.
or yellow. Stamens 8-20, stamen filaments filiform
Male flowers: sepals (4)5(6), valvate; petals
or subulate. Ovary 2-4(5)-carpellate, partly to
(4)5( 6), f:ee, val~ate, ovate, greenish blue, pink,
completely inferior, stylodia free, ovules pendant,
or greemsh white, reflex at anthesis; stamens
erect, or horizontal. Fruit capsular with apical
(8-) 1.0( -12), stamen filaments taper to apex;
interstylar dehiscence. Seeds numerous, 0.5-3 mm
~vanes (4)5(6)-carpellate, shallowly inferior, style
long; ellipsoidal, spindle-shaped, obovoid, or urce-
simple; ovules horizontal, sterile. Female flowers:
olate, W:inged and.some with micropylar flange or pink to dark blue; sepals (4)5-6, valvate; corolla
both wmg and micropylar flange lacking. n = 18,
rudimentary or lacking; stamens lacking; carpels
36 (Sax 1931; Schoenagel 1931; Love 1966, 1969).
(4}5( 6)-car~ellate, completely inferior, single style,
About 29 species. Eastern Asia (extending west-
stigma capitate, ovules horizontal. Fruit a fleshy
ward through the Himalayan region), including
berry. Seeds numerous and surrounded by projec-
Sumat~a, Java, Philippine Islands, Ryukyu Islands
tions from inner wall of ovary, about 0.5 mm long,
and Taiwan, southeastern North America, Central
urceolate. 2n =32 (Wagner et al.1990). One species
America, and western South America. In meso-
(B. arguta Gaud.), although various varieties and
phytic or moist forests and often on slopes, cliffs,
forma ?ave been described to account for poly-
or riparian banks. Five sections recognized, but
morphism (Fosberg 1939). Endemic to Hawaiian
their monophyly, except Cornidia, is dubious.
Is~~nds, where loca~ed on all main islands, except
Various temperate deciduous species that have
NIIhau and Kaho olawe, in dense, wet forests
sterile flowers are important ornamentals,
(Degener 1945).
especially cultivars of H. macrophylla and H.
paniculata.
14. Schizophragma Sieb. & Zucc.
12. Dichroa Lour. Schizophragma Sieb. & Zucc., Fl. Jap.: 58: t. 26 (1835).
Dichroa. L?ur., Fl. Cochinch.: 301 (1790); Hwang, Acta Phyto-

tax. Stmca 25: 384-389 (1987), synopsis and key.
Vines, climbing by shoot-borne roots. Leaves
deciduous, petiolate, margins toothed or entire.
Shrubs. Leaves persistent, petiolate, lamina Inflorescence cymose panicle. Sterile flowers
toothed. Inflorescence cymose panicle. Sterile present, one-lobed. Sepals (4)5, valvate. Petals (4)5,
flowers absent. Sepals (4)5(6), valvate. Petals valvate, abscising after anthesis, ovate, white.
(4)5(6), valvate, ovate, blue or pinkish; reflex at Stamens 8-10, stamen filaments filiform. Ovaries
anthesis. Stamens 4-5 or 8-12, filaments filiform. (4)5-carpell~te, deeply inferior, style simple;
st~gm~s capitate, ovules pendant. Fruits capsules
?vary (4)5( 6)~carpellate, partly to completely
mfenor, stylodia free or style with style branches; With mtercostal portions of the lateral walls
ovules horizontal. Fruit a berry. Seeds numerous, separating from costal ribs. Seeds numerous,
about ?·5 mm long; urceolate, lacking both wing 2.5-4.5 mm long, spindle-shaped, winged and with
micropylar flange. n = 14, 36 (Sax 1931; Hamel
and micropylar flange. n = 18 (Love 1969). Twelve
19~1, Nevling 1964). About 10 species. Eastern
species in two sections (sect. Dichroa, 10 species;
sect. Silvicola S.M. Hwang, 2 species). Distributed Asia. In mesophytic forests. Revision needed.
through mainland SE Asia southward to Pacific
islands. Three species endemic to New Guinea, 15. Pileostegia Hook. f. & Thoms.
two species endemic to Philippine Islands. The
Pileostegia Hook. f. & Thoms., J. Linn. Soc. 2: 57 (1858).
plants have been poorly collected, patterns of
character variation are not known, and a revision
is needed. Perennial vines, climbing by shoot-borne roots.
Leaves persistent, margins entire. Inflorescence
Hydrangeaceae 213
cymose panicle. Sterile flowers present, one-lobed. fruiting, ovate, white. Stamens >15, stamen fila-
Sepals (4)5, valvate, persist as rim on fruit. Petals ments filiform. Ovaries 6-12-carpellate, deeply
(4)5, valvate, triangular, white, calyptrate, abscis- inferior; style simple; stigmas capitate, ovules
ing at anthesis. Stamens (8)10, stamen filaments pendant. Fruits capsules with intercostal portions
filiform. Ovaries (4)5-carpellate, deeply inferior; of the lateral walls separating from costal ribs.
style simple; stigmas capitate, ovules pendant. Seeds numerous, 1.5-3 mm long, horn-shaped,
Fruits capsules with intercostal portions of the winged and with micropylar flange. n = 14 (Sax
lateral walls separating from costal ribs. Seeds 1931). Two species: D. barbara L., southeastern
numerous, 1-2 mm long, bottle-shaped, winged. North America; D. sinensis, China. In mesophytic
n = 18. Four species. Eastern Asia (China, Taiwan, forests.
Ryukyu Islands, India, and Himalayan region).
Mesophytic forests at mid elevations in moun-
17. Platycrater Sieb. et Zucc.
tains. Revision needed.
Platycrater Sieb. et Zucc., Fl. Jap. 62: t. 27 (1835).
16. Decumaria L. Fig. 71

Shrubs. Leaves deciduous, petiolate, toothed.
Decumaria L., Sp. Pl. ed. 2, 2: 1663 (1763). Inflorescence dichasial cymes or cymose panicle.
Sterile flowers present, with (2)3-4(5) lobes. Sepals
Perennial vines, climbing by shoot-borne roots. 4, valvate. Petals 4, valvate, ovate, white, abscising
Leaves deciduous, margins entire. Inflorescence at fruiting. Stamens >200, filaments filiform.
cymose panicle. Sterile flowers absent. Sepals Ovaries 2-4-carpellate, deeply inferior, stylodia
6-12, valvate. Petals 6-12, valvate, abscising at free, stigmas terminal, ovule variable. Fruit
capsular with apical interstylar dehiscence.
Seeds numerous, 1.0-1.5 mm long; spindle-
shaped, winged, micropylar flange lacking. One
species: Platycrater arguta Sieb. et Zucc. Warm
temperate areas of Japan and China.
Agababyan, V.S. 1961. A contribution to the palynomorphol-

ogy of the family Hydrangeaceae Dum. Izv. Akad. Nauk
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216 C. Erbar and P. Leins
Hydrostachyaceae
C. ERBAR and P. LEINS
Hydrostachyaceae Engler, Syllabus, ed. 2: 215 (1898), nom.

only at the initiation of inflorescences. This
cons. happens already during the submerged stage
whereas flowering starts only when the plants fall
Submerged aquatic perennial rosette plants with a dry.
short disk-like stem, attached to rocks in fast- At the top of the main axis, reduced leaves (scale-
flowing water; vascular system reduced. Leaves like hypsophylls) are formed and the axis termi-
alternate, entire to 2-3 times pinnatifid, leaf nates with a spike. Additional inflorescences are
base enlarged with intrapetiolar stipule; stomata formed in the axils of the uppermost pinnate
wanting; stem, leaves, and adventitious roots pro- leaves (polytelic synfiorescence). At the onset of
vided with numerous small, rounded, scale-like or flowering the plants have a Plantago-like appear-
fringed emergences. Plants dioecious, very seldom ance (Warming 1891a, 1891b; Rauh and Jager-
monoecious; unisexual flowers small, without Ziirn 1966; Cusset 1973; Jager-Ziirn 1998).
perianth, sessile in bracts of dense spikes; stami-
nate flower interpreted as consisting of a single VEGETATIVE ANATOMY. The vascular system is
stamen which is deeply divided into two thecae or reduced. In the disk-like stem, procambial tissue
of two independent bisporangiate stamens; with annular- and spiral vessels is present. In the
pistillate flower consisting of a single superior, peduncle of the spike, proto- and metaxylem and
bicarpellate, unilocular ovary with two elongate, phloem groups are diffusely scattered in the vas-
persistent, filiform stylodia; ovules numerous on cular tissue. The single bundle is closed. Some
two parietal placentas, anatropous, unitegmic, vessels with annular or spiral thickenings are close
tenuinucellar. Fruit a septicidal capsule with together and surrounded by thin-walled cells in
numerous tiny seeds. younger stages. Later these cells between the
A monogeneric family of 22 species native to vessels become sclerified at the base and col-
Madagascar and tropical and southern Africa. lenchymatous in the upper part of the inflores-
cence axis. The single vascular bundles differ from
VEGETATIVE MORPHOLOGY. The perennial hapa- the usual mode in that the proto-xylem and proto-
xanthous rosette plants start growing in fresh, phloem are not compressed in older stages but
fast-flowing water. The shoot of the water plants continue functioning. Due to the later-on strong
is fiat and disk-like and attached to rocks and elongation of the peduncle of the spike, the vessels
stones. Its diameter is usually a few centimetres. get disrupted. In older stages no entire vessels are
A primary root is not developed but numerous found. Adult plants have only schizogenic vascular
adventitious roots fasten the plants to the rocks. spaces with remnants of vessels.
The leaves are arranged spirally in tufts (Fig. 73A). The leaves have neither stomata nor special
The floating leaves are 30-50( -100) em long. aerenchyma. The petiole has a central main
They have an enlarged leaf base with a small, bundle and unusually numerous, small bundles
membranous, median stipule (intrapetiolar supplying the emergences. The main bundle is
stipule), a stout petiole and a flexible lamina .. The surrounded by thin-walled parenchymatous cells
leaves are simple or more frequently 2-3 times which later become collenchymatous. Annular and
pinnatifid. The rhachis, the pinnae as well as spiral vessels can be found only in young leaves
the petiole are densely covered with numerous whereas in older ones schizogenous vascular
rounded, tuberculate, scale-like, thread-like or spaces occur. Sieve tubes with simple sieve plates
fringed emergences. Their size varies between 1.5 have been observed only sporadically, e.g. in
and 5 mm. Emergences may be condensed at the Hydrostachys maxima (Warming 1891a; Rauh and
top of the leaflets. Jager-Ziirn 1966; Jager-Ziirn 1998).
The plants remain unbranched during the vege-
tative phase. The shoot system becomes sympodial
Hydrostachyaceae 217
is laterally flanked by a group of silky hairs each

(Cusset 1973).
Floral developmental studies support the
existence of two androecial organs. In the
earliest developmental stage of the male flower,
two androecial primordia arise separately on the
lateral edges of a transversally elongated, low
apex. The common base of the two bisporangiate
organs, which is formed by the floral apex,
remains rather short (Leins and Erbar 1988, 1990).
Because of their fully separate initiation, we can
interpret the male flower as consisting of two
A bisporangiate stamens. These may derive from
originally tetrasporangiate stamens in which
the sporangia of the neighbouring thecae have
fused apically. In a second phylogenetic process,
the synthecal sporangia shifted into a lateral
position.
The female flower consists only of a dimerous
and unilocular gynoecium with parietal placenta-
tion and the carpels arranged in a transversal posi-
tion. Silky hairs are formed on the lateral flanks
of the floral apex. Abaxially the ovary is wholly
enclosed by its subtending bract so that only the
two subulate stylodia with long stigmatic areas of
short papillae are visible in flowering inflores-
Fig. 72. Hydrostachyaceae. Hydrostachys multifida. A Male cences (Fig. 72B, C). Arrangement and shape of the
flower in the axil of its bract, adaxial view. B Part of female emergences on the dorsal side of the female bracts
inflorescence; the flowers covered by bracts, only stylodia are are of taxonomical value, distinguishing the
visible. CAdaxial view of adult female flower with subtending species (Cusset 1973).
bract. D Side view of female flower with silky hairs on lateral
flank. (Orig.)
The early development of the gynoecium is
remarkable. When the stylodia become visible at
the top of the two carpel primordia, plication takes
place in that the carpels start forming margins. It
FLOWER STRUCTURE. Nearly all species of Hydro- is very unusual that the stylodia are unaffected by
stachys are dioecious. Only Hydrostachys monoica this process. Below the insertion of the stylodia,
is monoecious and Hydrostachys stolonifera is the broad margins are inversely U -shaped. In early
facultatively monoecious (Cusset 1973; Jager-Ziirn stages the ovary becomes zygomorphic: coeno-
1998 ). Long, dense spikes of unisexual flowers (Fig. carpy is more pronounced on the abaxial than on
73B, D) terminate the main axis as well as the axil- the adaxial side. Later on, a suture extends to the
lary branches arising in the axils of the uppermost base of the ovary on its adaxial side (along this
pinnate leaves. All spikes are open (no terminal suture the ripe fruit opens septicidally). Only the
flower developed) and the uppermost bracts are base of the ovary bears a low septum. Most of the
sterile. ovary is unilocular, with many ovules on parietal
The simple male flower lies in the deepening of placentae. The base of the stylodia becomes over-
its subtending, shell-like bract (Fig. 72A). The male topped by growth of the ovary on the dorsal side
flower is either interpreted as consisting of a single of its upper choricarpous region (Fig. 72D; Leins
tetrasporangiate stamen which is deeply divided and Erbar 1988, 1990).
into two theca (e.g. Rauh and Jager-Ziirn 1966;
Cronquist 1981; Jager-Ziirn 1998) or as consisting EMBRYOLOGY. The anther can be interpreted as
of two independent, bisporangiate stamens tetrasporangiate (Rauh and Jager-Ziirn 1966;
(Warming 1891a, 1891b; Leins and Erbar 1988, Jager-Ziirn 1998) or bisporangiate (Warming
1990; Takhtajan 1997). The microsporangia, which 1891a, 1891b; Leins and Erbar 1988, 1990). The
face the bracts, open longitudinally by slits. In tapetum is secretory and its cells become 2-
many members of Hydrostachys, the androecium nucleate. Cell wall formation after meiosis is
simultaneous and the pollen grains are 2-celled FRUIT AND SEED. The fruit of Hydrostachys is a
when shed as permanent tetrads. septicidal capsule. Adaxially it opens in full length
The ovule is anatropous, unitegmic and tenuin- along a preformed suture, abaxially only in the
ucellate. The nucellus is weakly developed and short choricarpous region. The capsule releases
short, whereas the chalaza is conspicuous. One numerous, tiny seeds (0.25 x 0.5mm) lacking
subepidermal cell of the apex becomes the arche- endosperm (only negligible remains of the
sporia! cell which functions directly as the endosperm are present). The outer epidermis of
megaspore mother cell. The development of the the single integument develops into the testa
chalaza! megaspore follows the Polygonum type. whose outer cell walls are thickened and contain
Endosperm formation is ab initio cellular and a pectin. Getting wet, they swell up and the seed
micropylar haustorium is formed. Embryogeny surface becomes mucilaginous (Rauh and Jiiger-
follows the Onagrad type (Rauh and Jiiger-Ziirn Ziirn 1966; Jiiger-Ziirn 1998).
1966; Jiiger-Ziirn 1998).
DISPERSAL. Since the fruit ripens and opens
POLLEN MORPHOLOGY. The pollen grains are above the water level (Rauh and Jiiger-Ziirn 1966),
arranged in permanent tetrahedral or hexahedral the tiny seeds may be released from the septicidal
to rhomboidal tetrads. The individual pollen capsule by wind. A further transport by water also
grains are heteropolar trisymmetrical and inaper- seems possible, since seeds may easily be trans-
turate. The sexine is one-layered with a thick ported by currents. Contact with water, however,
tectum and as thick as the nexine. The tectum causes swelling of the epidermis which thus
partly exhibits microfurrows and suprategillar becomes mucilaginous. The mucilage fastens
microspinules. The exine has irregularly formed, the seed to the rocks during germination (myx-
thinner areas (Erdtman 1952; Straka 1988). These ospermy; Rauh and Jiiger-Ziirn 1966; Jager-Ziirn
are reduced and unspecific palynological features. 1998).
KARYOLOGY. The only count is 2n = 20-24 for PHYTOCHEMISTRY. Apart from kaempferol glyco-
Hydrostachys imbricata (Palm 1915). sides (Scogin 1992), caffeic acid has been recorded
from Hydrostachys (R0nsted et al., 2002). Iridoids
PoLLINATION. Flowering sets in when the water or alkaloids have not been found in Hydrostachys
level drops. The flowers elevated above the water up to the present.
surface are wind-pollinated (flowers apetalous and
stylodia with a long, papillate stigmatic area). AFFINITIES. The placement of the aquatic family
Although the yellow anthers in the male spikes and in angiosperm classifications is difficult due to the
the gay red or (seldom) yellow stylodia in the reductions in vegetative and floral morphology.
female spikes are somewhat striking, visit by In addition, in this taxon pollen characters and
insects was never observed (Rauh and Jiiger-Ziirn chemical compounds are not relevant for system-
1966). atics. The family has been associated with the
Podostemaceae by many authors for a long time,
REPRODUCTIVE SYSTEMS. The plants grow totally although Warming (1891 b, 1891 c) separated
submerged during the rainy season. Only in the Hydrostachyaceae as a different family and sug-
dry season, when the water level in the rivers sinks, gested a position near Saxifragaceae (Warming
do the vegetative organs emerge from the water, 1891a; but see Les et al. 1997). Podostemaceae are
the peduncles of the spikes enlarge, and flowering quite different in having bisexual flowers with cen-
and seed ripening take place. The inflorescences trifugal stamen inception on a ring primordium
have already been initiated during the submerged (Rutishauser and Grubert 1999,2000). On the basis
phase. Only a few species, such as Hydrostachys of their floral morphological and embryological
distichophylla and H. verruculosa, grow perma- studies, Rauh and Jager-Ziirn (1966) assigned
nently submerged in some places. They never the family a position within Asteridae near
come in bloom and propagate vegetatively by Scrophulariaceae-Plantaginaceae. Subsequently,
sprouts of the adventitious roots. Also deep-water most authors accepted a close relationship to the
species (e.g., Hydrostachys verruculosa) may prop- Scrophulariales (e.g., Cronquist 1981; Leins and
agate vegetatively during some years if conditions Erbar 1988, 1990; Wagenitz 1992; Takhtajan 1997).
for flowering are not given (Rauh and Jiiger-Ziirn Comparative DNA (rbcL) sequence analyses pro-
1966). vided the suggestion that Hydrostachyaceae are
allied with the Hydrangeaceae in the Cornales
Hydrostachyaceae 219
One genus:
Hydrostachys Thouars Fig. 73

Hydrostachys Thouars, Nova Madag. Gen. 2 (1806); Cusset,
Adansonia II, 13:75-119 (1973). About 22 spp., Madagascar,
tropical and southern Africa.
Characters of the family.
Albach, D.C. eta!. 2001b. See general references.

references.
Cronquist, A. 1981. See general references.
Cusset, C. 1973. Revision des Hydrostachyaceae. Adansonia II,
13: 75-119.
Engler, A. 1895. Hydrostachydaceae africanae. Bot. Jahrb. Syst.
20: 136-137.
Engler, A. 1898. Syllabus der Pflanzenfamilien. 2. ed. Berlin:
Borntraeger.
Erdtman, G. 1952. See general references.
Hempel, A.L., Reeves, P.A., Olmstead, R.G., Jansen, R.K. 1995.
Implications of rbcL sequence data for higher order rela-
tionships of the Loasaceae and the anomalous aquatic plant
Fig. 73. Hydrostachyaceae. Hydrostachys imbricata. A Habit. B Hydrostachys (Hydrostachyaceae). Pl. Syst. Evol. 194: 25-37.
Male inflorescence. C Female inflorescence. (Warming 1891c) Jager-Ztirn, I. 1998. Anatomy of the Hydrostachyaceae. In:
Handbuch der Pflanzenanatomie 13, 4. Landolt, E. (ed.)
Extreme adaptations in angiospermous hydrophytes.
Berlin: Borntraeger, pp. 129-196.
(Hempel et al. 1995; APG 1998). Further molecular Leins, P., Erbar, C. 1988. Einige Bemerkungen zur Bltitenen-
studies of rbcL sequences (Xiang 1999; Savolainen, twicklung und systematischen Stellung der Wasserpflanzen
Fay et al. 2000) and of combined sequences from Callitriche, Hippuris und Hydrostachys. Beitr. Bioi. Pflanzen
the rbcL, 18 S rDNA, and atpB genes (Albach et al. 63: 157-178.
2001b; Soltis et al. 2000) confirm a relationship of Leins, P., Erbar, C. 1990. The possible relationship of Hydro-
stachyaceae based on comparative ontogenetical flower
Hydrostachyaceae to Hydrangeaceae, although the studies. Mitt. Inst. Allg. Bot. Hamburg 23b: 723-729.
remarkably long branch leading to Hydrostachys is Les, D.H., Philbrick, C.T., Novelo R.A. 1997. The phylogenetic
problematic (Xiang 1999). In the analysis of the position of river-weeds (Podostemaceae): insights from
ndhF gene, Hydrostachys is sister to Escalloniaceae rbcL sequence data. Aquat. Bot. 57: 5-27.
Palm, B. 1915. Studien tiber Konstruktionstypen und Entwick-
(Albach et al. 2001b).
lungswege des Embryosacks der Angiospermen. Akad.
Afhandl. Stockholm 1915: 1-259.
DISTRIBUTION AND HABITATS. The habitats of Rauh, W., Jager-Ztirn, I. 1966. Zur Kenntnis der Hydrostachy-
Hydrostachyaceae are rapids and waterfalls of aceae. I. Bltitenmorphologische und embryologische
tropical and subtropical rivers. Most of the species Untersuchungen an Hydrostachyaceen unter besonderer
Berticksichtigung ihrer systematischen Stellung. Sitzungs-
(15) are endemic to Madagascar, a few (7) are ber. Heidelb. Akad. Wiss.-Math.-Naturwiss. Kl. 1966: 1-177.
found in southern Africa, hardly transgressing the R0nsted, N., Strandgaard, H., Jensen, S.R., M0lgaard, P. 2002.
equator. In Madagascar the species are found in Chlorogenic acid from three species of Hydrostachys.
large populations in rocky rapids of the central Biochem. Syst. Ecol. 30: 1105- 1108.
highlands (between 800 and 2000 m altitude). In Rutishauser, R., Grubert, M. 1999. The architecture of Mourera
fluviatilis (Podostemaceae): developmental morphology of
Africa Hydrostachyaceae are widely spread in inflorescences, flowers, and seedlings. Am. J. Bot. 86:
rivers throughout the mountain regions of Angola, 907-922.
Congo, Zaire, Tanzania, Malawi, Zimbabwe, Rutishauser, R., Grubert, M. 2000. Developmental morphology
Mozambique and South Africa (Jager-Ziirn 1998). of Apinagia multibranchiata (Podostemaceae) from the
They are submerged during the rainy season. In Venezuelan Guyanas. Bot. J. Linn. Soc. 132: 299- 323.
Savolainen, V., Fay, M.F. et a!. 2000. See general references.
the dry season at low water level, the plants Scogin, R. 1992. Phytochemical profile of Hydrostachys insig-
become emerged and flowering and seed ripening nis (Hydrostachyaceae). Aliso 13: 471-474.
take place. Soltis, D.E. et a!. 2000. See general references.
Straka, H. 1988. Die Pollenmorphologie von Hydrostachys, Warming, E. 1891b. Note sur le genre Hydrostachys. Overs.
Hydrostachyaceae, und ihre Bedeutung fiir die systematis- Kongel. Danske Vidensk. Selsk. Forh. Medlemmers Arbeider
che Einreihung der Familie. Beitr. Bioi. Pflanzen 63: 1891: 37-43.
413-419. Warming, E. 1891c. Familien Podostemaceae IV. Kongel.
Takhtajan, A. 1997. See general references. Danske Vidensk. Selsk. Naturvidensk. Math. Afh. Ser. 6, VII
Wagenitz, G. 1992. The Asteridae: evolution of a concept 4: 135-179.
and its present status. Ann. Missouri Bot. Gard. 79: 209- Xiang, Q.-Y. 1999. Systematic affinities of Grubbiaceae and
217. Hydrostachyaceae within Cornales - insights from rbcL
Warming, E. 1891a. Podostemaceae. In: Engler, A., Prantl, K. sequences. Harvard Pap. Bot. 4: 527-542.
Die natiirlichen Pflanzenfamilien III, 2a. Leipzig: Engel-
mann, pp.1-22.
Lecythidaceae 221
Lecythidaceae
G.T. PRANCE and S.A. MoRI
Lecythidaceae Poit., Mem. Mus. Hist. Nat. Paris 13: 141-165 up to 55 m tall. The bark of Neotropical Lecythi-
(1925). daceae (Mori et al. 1987a) is easily recognized in
Barringtoniaceae F. Rudolphi (1830).
Foetidiaceae (Nied.) Airy Shaw (1956). the field because of its fibrous nature and the
presence of files of unilaterally thickened, crystal-
Small to large trees; rarely shrubs; axis with cor- bearing cells in the phloem. Crystalliferous
tical vascular bundles. Leaves alternate, simple, phloem can be observed at low magnifications by
margins usually entire (opposite and clustered in making a tangential peel of the bark with a sharp
one species of Abdulmajidia), pinnately nerved; knife. The bark can often be peeled from the trunk
stipules absent or minute and caducous. Inflores- in long strips, and then the inner bark can be
cences terminal, axillary or cauline, simple beaten into thin sheets. It has been used for caulk-
racemes, panicles with 2 or 3 orders of racemose ing boats, cigarette paper, native clothing, and
or spicate branches or fascicles. Flowers actin- cordage. An excellent field feature is that the bark
omorphic or zygomorphic, hermaphrodite; sepals can be peeled from the twigs in long strips; this is
2-6 or rarely unlobed; petals 3-6(8), infrequently shared by species of Annonaceae and therefore
12 or 18, imbricate, free (absent in Foetidia); sterile collections of these two families are some-
stamens numerous, 10-1210, connate at base into times confused. External bark morphology of
a short or long stamina! ring (free to base in Foe- Neotropical Lecythidaceae is useful in differenti-
tidia), the ring actinomorphic or prolonged on ating species. Some species possess essentially
one side into a strap-like structure which arches smooth bark (e.g. Eschweilera collina), the bark of
over the summit of the ovary; anthers bilocular, others is scalloped (e.g. E. apiculata, E. laevicarpa,
latrorse, introrse or rarely poricidal; ovary inferior and E. micrantha) and still others have very
or semi-inferior, usually 2-, 3-,4- or 6-locular, with deeply furrowed bark (e.g. Corythophora rimosa
2-115 anatropous ovules in each loculus, the axile and Lecythis zabucajo ). The presence of cortical
placenta at the apex, base or throughout the length bundles helps characterize the Lecythidaceae, and
of the locule; ovules bitegmic, tenuinucellate; the orientation of these bundles (inversely ori-
style short or more rarely long, undivided. Fruits ented in the Planchonioideae and Foetidioideae
indehiscent, then dry, fleshy or woody or dehis- vs. normally oriented in the Lecythidoideae) has
cent by a circumscissile operculum, then woody, been used to distinguish subfamilies within the
sometimes very large; seeds exotestal/mesotestal, family (Prance and Mori 1979). However, the valid-
winged (Cariniana and Couroupita) or without ity of cortical bundle orientation needs further
wings in remaining genera; endosperm lacking investigation. The stem-node-leaf continuum of
or very scanty; arils present or absent; embryos Lecythidaceae has been accurately described by
undifferentiated or with fleshy plano-convex or Lignier (1890). Its principal features, at least in the
foliaceous cotyledons. New World Lecythidaceae, are (1) numerous cor-
Seventeen genera in three subfamilies, the tical bundles that enter the petiole and are indi-
Lecythidoideae exclusively neotropical with 210 rectly connected to the stele via fusion with stelar
species, the Planchonioideae with 55 species in traces; and (2) three traces which depart from the
Africa, Madagascar and tropical Asia to N Aus- stele at different levels, the median trace usually
tralia, and the unigeneric Foetidioideae with 17 departs at the same node as the leaf it enters
species in Madagascar, Mauritius and E Africa. whereas the lateral traces depart from below that
node (Mori and Black 1987). The leaves of most
VEGETATIVE MORPHOLOGY. Almost all species of species of Planchonioideae and of the more
Lecy~hidaceae are trees with only three species primitive genera of Lecythidoideae (Gustavia,
growmg as shrubs. Some species, such as Berthol- Grias, Couroupita) tend to be large and clustered
letia excelsa, are extremely large, emergent trees towards the ends of the branches. These comply to
Corner's model of architecture (e.g. Gustavia
222 G.T. Prance and S.A. Mori
grandibracteata, G. monocaulis, and Barringtonia cells associated with parenchyma bands in all
calyptrocalyx), Schoute's model (all species of genera, crystal cell walls unilaterally or uniformly
Grias, Gustavia grandibracteata, and G. superba), thickened, the latter always occurring within unis-
and Rauh's model (the cauliflorous Couroupita eriate parenchyma bands; crystal cell wall thick-
guianensis ). Most other species have smaller leaves ening either exclusively one or the other type, or
and a much branched crown with typically lepta- in mixture in part of the genera; gum often
caul form. In all, eight types of architecture have present. Fibers medium to moderately long,
been reported in Lecythidaceae, the others of pitting small to very small, clearly bordered in
which are: - Leeuwenberg's model: Barringtonia Allantoma, Bertholletia, Cariniana, Couratari, and
edulis; - Koriba's model: Petersianthus africanus; Couroupita; gelatinous fibers present in most
- Massart's model: Couratari stellata; - Roux's genera".
model: Bertholletia excelsa; and - Troll's model: Before his death, de Zeeuw had completed a
Lecythis pisonis, Eschweilera sp. series of manuscripts on the wood anatomy of
Stipules are usually absent but, in a few species, Foetidia (de Zeeuw, manuscript), Careya and Chy-
minute caducous stipules occur. Several species of denanthus (de Zeeuw, manuscript), Petersianthus
Lecythidoideae and Planchonia have buttressed (de Zeeuw, manuscript), and Planchonia (de
trunks (e.g. most species of Couratari). The con- Zeeuw, manuscript). Information from these
siderable variation in bark characteristics is studies is summarized below.
discussed in Roth ( 1969) and Prance and Mori Foetidia possesses uniformly encapsulated crys-
(1979). tals in strands in the axial parenchyma and silica
bodies in the ray cells. Ray crystals, however,
Wooo ANATOMY. The wood anatomy of New are not present. De Zeeuw's studies of Careya,
World Lecythidaceae has been studied by the late Chydenanthus, and Planchonia support a close
Carl de Zeeuw (1987, 1990) who summarized its relationship among these genera. He reports
structure (1990) as follows. "Pores diffuse, moder- septations in the fibers of all three genera and
ately small to moderately large, numerous to very crystals in the axial parenchyma of Careya for the
few, elliptical in outline, solitary or in radial mul- first time. De Zeeuw also found septate fibers in
tiples of 2 to 3 with a small component of long Barringtonia samoensis and suggests that their
radial strings or clusters. Vessel elements medium presence supports a close relationship between
length, with ends at a large angle to cell axis; per- Barringtonia and other genera of Plancho-
forations simple in all genera, with the addition of nioideae. He reports the presence of prismatic
some reticulate or scalariform plates in Grias; crystals in the axial parenchyma and rays of Plan-
intervessel pitting alternate (except in Grias and chonia for the first time. Although the two species
Gustavia which have a mixture of alternate and of Petersianthus are geographically isolated from
opposite to irregular pitting), the pits usually one another, de Zeeuw found them to be similar in
medium size (except very small in Gustavia); their general wood anatomy but different enough
ray-vessel pitting in part similar to intervessel to be distinguished by their wood alone. Peter-
pitting and in part half-bordered to simple linear- sianthus macrocarpus emits a strong, disagreeable
irregular pits of larger size; tyloses present or odour from the green wood, and silica is virtually
absent, usually thin walled. Rays entirely hetero- absent in both the rays and the axial parenchyma.
cellular Type II or III in Grias and Gustavia, mixed In contrast, the wood of P. quadrialatus lacks a
heterocellular and homocellular in other genera, strong odour but possesses rough, rounded silica
mostly less than one millimeter in height, 2- to 3- bodies in the rays and axial parenchyma.
seriate, the tallest in Grias and Gustavia, the widest
up to 6-seriate in Grias and Gustavia; crystals FLOWER STRUCTURE. Flowers are rather large and
present in Gustavia and in some samples of Grias; have an extremely variable morphology. They are
silica sparse to abundant in all but Bertholletia, actinomorphic in the Planchonioideae, Foetid-
Couroupita, Grias, and Gustavia; dark gum often ioideae and basal members of Lecythidoideae,
present. Axial parenchyma apotracheal banded in but highly zygomorphic in the more advanced
all genera except Grias and Gustavia where it is Lecythidoideae. The calyx is usually imbricate in
reticulate to diffuse-in-aggregates, and with scanty bud and 4-6-lobed (2-lobed in Bertholletia) but is
paratracheal in addition in all genera, mostly in valvate in Foetidia and unlobed in Barringtonia
1- to 2-seriate bands but uniseriate in Grias and section Barringtonia. Petals vary from 3 to 18 and
Gustavia and up to 6-seriate in part of Lecythis; are free. There are numerous stamens in most
strands average 4-7 cells long; strands of crystal genera, up to 1200 in Gustavia and as few as 10 in
Lecythidaceae 223
Cariniana. In all actinomorphic androecia, except

Foetidia, the stamens are united at their base.
Zygomorphy is caused by variations of the androe-
cium which, in some genera of Lecythidoideae,
forms a strap-like structure to one side which
arches over the summit of the ovary to form a
hood over a ring of fertile stamens at its base
(Fig. 74). The hood appendages may bear anthers
(Lecythis, Corythophora) or may be sterile and
produce nectar at their base. In zygomorphic
androecia of the Planchonioideae, the inner whorl
of stamens is often reduced and staminodial. An
annular nectar-secreting disk occurs in the Plan-
chonioideae and the Foetidioideae but not in
the Lecythidoideae. The ovaries are inferior or
semi-inferior. Short styles are characteristic of
most Lecythidoideae except Bertholletia and a few
species of Lecythis, whereas the styles are long and
filamentous in Planchonioideae and Foetidioideae
and usually equal the filaments in length. There
are few to numerous ovules attached in various
positions to the axile placenta.
EMBRYOLOGY. The embryology has been stud-

ied in detail by Tsou {1994a). The ovules are
anatropous, bitegmic, tenuinucellate with multi-
layered outer and inner integuments, the inner
integument forms the micropyle, and one or more
vascular bundles lead to the outer integument.
Anther wall formation is of the basic type and the
tapetum is glandular.
POLLEN MORPHOLOGY. The pollen of Lecythi-

daceae has been studied by Muller {1972, 1979)
and by Tsou {1994a, 1994b). Two distinct types
of pollen occur in the family, syntricolpate in Fig. 74. Lecythidaceae. Androecial structures in Lecythi-
Planchonioideae and tricolpate or tricolporate in doideae, showing increasing zygomorphy in connection with
Lecythidoideae and Foetidioideae (Fig. 75). The formation of a strap-like hood over the ovary and the ring of
fertile stamens. A Gustavia hexapetala, stamens often >1000,
latter type also occurs in the related Napoleon- offering pollen reward. B Grias cauliflora, androecium a sym-
aeaceae. The Planchonia pollen type is character- metric stamen ring; stamens all fertile, offering pollen reward;
ized by grains which are usually larger than 40 J..l.m, pollination probably by beetles. C Allantoma lineata, androe-
possess a thick ectexine and syntricolpate colpi, cium fused into an actinomorphic tube; stamens all fertile;
and generally lack clearly defined endoapertures. pollination probably by beetles. D Cariniana pauciramosa,
antheriferous tube slightly zygomorphic; stamens all fertile.
Specialized structures in the marginal zones of the E Lecythis corrugata, the androecium zygomorphic, the hood
ectoaperture and in the polar area are found in without anthers but nectariferous. F Couroupita guianensis,
some species. The Lecythis type is less specialized, fertile stamens around gynoecium with monad pollen, sterile
smaller (the polar axis usually less than 35 ~m), stamens on hood with tetrad fodder pollen. G Corythophora
alta, the closed antheriferous hood with sterile fodder pollen.
tricolporoidate or tricolporate, and the exine is
H Lecythis pisonis, hood flat, its proximate appendages anther-
generally simpler, composed of a thin endexine, a iferous with sterile fodder pollen. I Eschwei/era /ongipes,
layer of more or less distinct columellae, and a androecium zygomorphic, the hood closed with a triple coil,
rather thin tectum which is smooth, finely reticu- offering nectar. J Couratari stellata, androecium zygomorphic,
late, foveolate or scabrate-verrucate. hood with double coil inwards and an external flap outwards,
offering nectar from closed hood. (Modified from Mori and
Dimorphic pollen occurs in Couroupita and Prance 1987)
some species of Lecythis (Jacques 1965; Mori et al.
1980; Mori and Boeke 1987). In species with
KARYOLOGY. The most detailed study of chromo-

some number is that of Kowal et al. (1977). The
Lecythidoideae has a basic number of x = 17 and
the Planchonioideae has x = 13. Tetraploid counts
I Basic tricolpate prototype occurring of n = 34 have been made for Gustavia superba and
j'"' "' ' "'""""'

Lecythidioideae,
in Foetidioideae,
n = 26 for Petersianthus macrocarpus (Mangenot
and Mangenot 1962) and Barringtonia racemosa
(Morawetz 1986). The genera separated from
Lecythidaceae have different numbers -Napoleon-
aea n = 16 and Asteranthos n = 21 (Kowal1989).
JI Syntricolpate Barringtonia
calyptrata type
POLLINATION. Details of the pollination of New
World Lecythidaceae can be found in Prance
(1976), Mori and Kallunki (1976), Mori et al.
j (1978), Prance and Mori (1979), Mori and Boeke

(1987), Nelson et al. (1987), Mori and Prance
(1990b), and Knudsen and Mori (1996). Observed
pollinators are principally bees, but bats and
beetles have also been observed visiting the
ill Syntricolpate type with marginal flowers.
j'""''"'"'"fu
ridges and grooves, found in Careya Species of Gustavia, all with actinomorphic
flowers, non-differentiated pollen, and open
flowers, offer pollen as a reward to their bee polli-
nators (Mori and Boeke 1987). Species of another
New World, actinomorphic-flowered genus, Grias,
possess floral scents with fatty acid derivatives
ISZ Syntricolpate Barringtonia typical of beetle-pollinated flowers (Knudsen and
Mori 1996). The discovery of beetles in the flowers
calyptrocalyx type with marginal
ridges, polar cushions and of an Ecuadorian species of Grias suggests that
marginal grooves this genus is indeed beetle-pollinated (J. Knudsen,
personal communication to S.A. Mori 1995).
Fig. 75. Lecythidaceae. Main pollen types of Lecythidaceae;
pollen grains drawn in polar view. (Muller 1973)
Nothing is known about the pollination of Allan-
toma but, based on the similarity of the floral
structure of this genus with that of Grias, beetle
pollination is also suspected.
dimorphic pollen, the stamina! ring pollen is Pollination of zygomorphic-flowered species of
fertile and the hood pollen does not germinate. Lecythidaceae is more complicated. Species with
In Couroupita guianensis the ring pollen is in this floral type vary in the type of pollinator
monads and the sterile hood pollen remains in reward, the degree of openness of the androecial
tetrads. In Lecythis pisonis the hood pollen turns hood, and in the complexity of the androecial
black 24 hours after an thesis while that of the ring hood (Mori 1989). Some species possess fodder
remains yellow. In species of Corythophora, the pollen on an open hood (e.g. Lecythis zabucajo),
sterile pollen is yellow whereas the fertile pollen is others have fodder pollen on a closed hood (e.g.
white. In this genus, the sterile pollen may be sit- Corythophora rimosa), others seem to offer a com-
uated in the stamina! ring (C. amapaensis) or it bination of fodder pollen and nectar on a closed
may be located on the hood (C. rimosa). In the few hood (e.g. Lecythis corrugata; Mori and Boeke
species that have been studied, the sterile or 1987), and the most advanced species florally offer
fodder pollen serves as the pollinator reward. For only nectar from a closed hood (e.g. species of
example, in C. amapaensis, euglossine bees enter Eschweilera and Couratari) as rewards to the floral
the flower, and extract yellow, fodder pollen which visitors. The floral visitors are mostly bees, but
they subsequently place in their corbiculae. At the bats have also been reported to visit at least two
same time, fertile, white pollen is deposited on the species (Mori et al. 1978; Mori and Prance 1990b).
head and back of the bee from where it is trans- A large number of different types of bees have
ferred to the stigma of subsequent flowers visited been reported to visit the flowers of zygomorphic-
(Mori and Boeke 1987). flowered Lecythidaceae. Detailed studies of indi-
Lecythidaceae 225
vidual species of Lecythidaceae have not shown a riverine species and are wind-dispersed. The seeds
one-to-one relationship between species of plant of Lecythis sections Pisonis and Lecythis have a
and species of bee. The most important trait of the swollen funicular aril (see Fig. 77G) and remain in
bee pollinator of zygomorphic-flowered Lecythi- the fruit after dehiscence of the operculum and are
daceae with closed hoods is that the pollinator be bat-dispersed. The large, globose, woodypyxidium
large enough to force open the hood, and possess of Bertholletia is secondarily indehiscent and has
a tongue long enough to reach the nectar. For only a small operculum that falls inwards (see Fig.
example, in the Brazil nut (Bertholletia excelsa), 76E). This species drops the pyxidium to the
species of Bombus, Centris, Epicharis, Eulaema, ground and is dispersed by agoutis. Lecythis lurida
and Xylocopa have been observed entering the is also secondarily indehiscent. The seeds of Allan-
flowers (Mori and Prance 1990a; Mori 1992). Some toma lineata are water-dispersed. The seeds of
species with more complicated androecial hoods, Bertholletia excels a, Allantoma lineata and Abdul-
e.g. Eschweilera pedicellata, may be exclusively majidia spp. all have an extremely hard, woody
visited by euglossine bees because they are the testa.
only New World bees with tongues long enough to Three embryo types occur in the seeds of
easily reach the nectar. It is noteworthy that the Lecythidaceae:
flowers of E. pedicellata are pollinated by trap-
1) macropodia! or undifferentiated in Grias,
lining euglossine bees and that the large and
Abdulmajidia, Allantoma, Chydenanthus, Cory-
showy flowers do not have any scent (Knudsen and
thophora, Eschweilera, Lecythis, Barringtonia
Mori 1996).
and Careya;
Two species, Lecythis poiteaui and L. barnebyi,
2) Gustavia type with fleshy, plano-convex
are known to be pollinated by bats. Both of these
cotyledons;
species produce flowers on racemes that project
3) Couroupita type with leaf-like cotyledons and a
from the ends of the branches, have very large
long, well-differentiated radicle in Couroupita,
flowers with more than 1,000 stamens, emit a
Cariniana, Foetidia, Planchonia and Peter-
musty scent from the flowers, and produce
sianthus.
copious, relatively dilute nectar (Mori et al. 1978;
Mori and Prance 1990a). Many species of the The germination of all species observed so far is
actinomorphic-flowered, Old World Plancho- phanerocotylar.
nioideae, especially species of Barringtonia and
Abdulmajidia, are also known to be bat-pollinated. PHYTOCHEMISTRY. Some species are accumula-
tors of selenium. The seeds of two species of
FRUIT AND SEED. The fruits of Lecythidaceae Lecythis, L. ollaria and L. minor are toxic because
are extremely variable. Those of Foetidia and of the presence of the selenium-containing analog
Planchonioideae are indehiscent drupes with 1 to of the sulphur amino acid cystathionine: HOOC-
many seeds. Petersianthus is remarkable for the CH (NH 2)-CH Se-CH2CH (NH 2) COOH (Kerdel-
large wings on the exterior of the fruit and Bar- Vargas 1966). The bark of Planchonia careya
ringtonia for their rectangular 4-ridged or 4- (F. Muell.) Kunth contains saponins and is used by
winged fruit. The fruits of several species of natives of New Guinea to stun fish.
Barringtonia are buoyant and are dispersed by
ocean currents which accounts for their wide- AFFINITIES. The family has traditionally been
spread distribution, e.g. B. racemosa and B. acu- placed in the Myrtales. Thorne (1968) suggested
tangula. The varied structure of the fruit of a Thealean relationship, and Cronquist (1981)
Lecythidoideae is described and illustrated in and Takhtajan (1987) both established the order
Prance and Mori (1978, 1979) and Mori and Prance Lecythidales somewhere between the Theales and
(1990b). The basal genera Gustavia, Grias and Malvales. A Myrtalean relationship is no longer
Couroupita have indehiscent fruits with the seeds acceptable on the basis of differences in wood
embedded in pulp. These are dispersed by a anatomy, pollen and floral morphology such as the
variety of forest-floor mammals such as agouti, basal staminal ring. Tsou (1994a) was the first to
arboreal mammals such as monkeys and, for river- suggest a relationship with Scytopetalaceae, which
ine species, by fish. The remaining genera of seems highly likely. Asteranthos, which was for-
Lecythidoideae have a great variety of dehiscent merly in the Lecythidaceae, is better placed in the
fruits. Cariniana has unilaterally winged seeds Scytopetalaceae. The Lecythidaceae, Napoleon-
and Couratari has wings that surround the seed. aeaceae and Scytopetalaceae all seem to be closely
Both genera are tall, emergent trees, or savanna or related. This is supported by DNA sequence analy-
ses (Morton et al. 1997); however, if Asteranthos is in highly threatened areas such as western Colom-
removed from Lecythidaceae, it is also necessary bia and Ecuador, eastern Brazil, and Sumatra and
to remove Napoleonaea to obtain a monophyletic Java, and are therefore potentially threatened with
group of Lecythidoideae, Planchonioideae and extinction. The 15 Madagascan species of Foetidia
Foetidioideae. The multigene sequence analysis by are in particular danger.
Anderberg et al. {2002) places these families sister
to Sapotaceae within the broadly defined Ericales INFRAFAMILIAL CLASSIFICATION
sensu APG {1998). Flowers actinomorphic or zygomorphic; calyx
usually imbricate in bud; petals present; styles
PALAEOBOTANY. There are few, reliable fossil usually short or of medium length; stamens united
records of Lecythidaceae. Lecythidoanthus kugleri at base, sometimes prolonged into a hood or tube;
described by Berry ( 1924a, 1924b) from the pollen tricolpate or tricolporate. Fruit often dehis-
Miocene of Trinidad almost certainly represents a cent, less frequently indehiscent. Cortical bundles
lecythidaceous flower. Milanez {1935) described in with normal orientation; secondary xylem with
detail a Cretaceous wood from Piauf, Brazil which crystal chains.
he named Lecythioxylon brasiliense. Huertas
Subfam. Lecythidoideae (genera 1-10)
{1969) described a fossil fruit of an Eschweilera
Flowers actinomorphic; calyx imbricate in bud;
from Cundinamarca, Colombia which he named
petals present; styles long and linear; stamens
Lecythidopyxion girardotanum. Barringtonia-
united at base; pollen syntricolpate. Fruit indehis-
like fossils have frequently been described, for
cent. Cortical bundles with reversed orientation
example, Barringtonioxylon deccanense from the
(xylem outside; phloem inside); secondary xylem
upper Cretaceous or Early Eocene of Muhurzari,
without crystal chains.
India and B. eopterocarpum (Prakash and Dayal
1965). Subfam. Planchonioideae (genera 11-16)
Fossil pollen that is similar to some extant Bar- Flowers actinomorphic; calyx valvate in bud;
ringtonia and Chydenanthus species is known petals absent; styles long and linear; stamens free
from the lower Eocene of Borneo and India; a to base; pollen syntricolpate. Fruit indehiscent.
slightly different type, referable to recent Cortical bundles with reversed orientation; sec-
Abdulmajidia, Barringtonia and Planchonia, first ondary xylem without crystal chains.
appeared in the lower Eocene of Cameroon (see
Subfam. Foetidioideae (genus 17)
Tsou 1994b for references).
KEY TO THE GENERA
ECONOMIC IMPORTANCE. The most well-known
economic product from the family is the Brazil nut 1. Petals present; calyx usually imbricate in bud; stamens
from the seeds of Bertholletia excelsa (see Mori united at base or into tube or hood 2
- Petals absent; calyx valvate in bud; stamens free to base
and Prance 1990a; Mori 1992). Various other
17. Foetidia
species of Lecythis also have edible seeds but are 2. Floral nectary disc absent; styles usually short; flowers
only used locally. Two species of Grias have actinomorphic or zygomorphic; fruit dehiscent or inde-
avocado-like fruit, the pulp of which is relished in hiscent; secondary xylem with crystal chains 3
Amazonian Peru, and the yellowish pulp around - Floral nectary disc present; styles long and linear; flowers
actinomorphic; fruit indehiscent; secondary xylem
the seeds of Gustavia speciosa subsp. speciosa is without crystal chains 14
eaten in Colombia. The fruit and seeds of Plan- 3. Androecium actinomorphic 4
chonia careya are eaten in New Guinea. The wood - Androecium zygomorphic 7
of many species of Lecythidoideae is used locally 4. Flowers 2.5-20cm diam. at anthesis; stamens united only
for general construction, but the quantity of silica at base; ovules inserted on upper 1/2 of septum; fruits
fleshy, globose, to ovoid indehiscent 5
in some species restricts their use. Cariniana pyri- - Flowers less than 2.5 em diam. at anthesis; stamens united
formis has been overexploited for its timber which into a tube with some anthers inserted on interior; ovules
has been exported from Colombia as Colombian inserted on lower 1/2 of septum; fruits woody, campanu-
mahogany or albarco. The wood of Planchonia late or cylindrical, dehiscent 6
valida is much used in house construction in 5. Petals 6-8{-18); stamens 500-1210, the anthers linear, 2-5
mm long, dehiscing by apical pores; placentae expanded,
Indonesia. The seeds and bark of various species the ovules 7-93 per locule, horizontal or slightly descend-
of Barringtonia are used locally as a fish poison. ing; fruits usually with 2 or more seeds; embryo with
plano-convex, fleshy cotyledons 1. Gustavia
CoNSERVATION. Many species of Lecythidaceae - Petals 4; stamens 85-210, the anthers globose, less than
1 mm long, dehiscing by longitudinal slits; placentae not
are of extremely restricted distribution and grow
Lecythidaceae 227
expanded, the ovules 2-4, pendulous; fruits 1-seeded; 15. Ovules 4 per loculus, inserted at apex of axis, pendulous;
embryo undifferentiated 2. Grias fruit one-seeded 12. Barringtonia
6. Buds oblong; calyx rim-like or with 5 inconspicuous, - Ovules 8-20 per loculus and usually inserted all along axis
broadly triangular lobes at anthesis; petals 5, the apices or towards base; or ovules 1-2 per loculus, inserted at base
pointed; ovary usually 4-locular; seeds not winged; embryo of axis and ascending; fruit usually 2-many-seeded 16
undifferentiated 3. Allantoma 16. Pedicel articulate; ovules 1-2 per loculus, inserted at base
- Buds globose; calyx with 6 triangular lobes at anthesis; of axis and ascending; fruit 1-2-seeded; carpels 2
petals 6, the apices rounded; ovary usually 3-locular; seeds 13. Chydenanthus
unilaterally winged; embryo with 2 foliaceous cotyledons - Pedicel not articulate; ovules 8-20 per loculus, inserted all
4. Cariniana along axis, at top or middle of axis; fruit usually 5-many-
7. All or at least three-fourths of hood appendages bearing seeded, rarely 2-4-seeded; carpels 3 or 4 17
anthers 8 17. Petals 3; ovules inserted on upper part of axis; outer
- Usually with all of hood appendages sterile, or rarely with integument of seed thin; carpels 3 14. Abdulmajidia
less than half of hood appendages bearing anthers 10 - Petals 4; ovules inserted along or on mid part of axis; outer
8. Ovary 6-locular; ovules 30-115 per locule, on bilamellar integument of seed arilloid; carpels 4 18
placentae throughout length of locule; fruit globose, inde- 18. Embryo undivided, undifferentiated 15. Careya
hiscent; seed as long as broad; embryo with 2 foliaceous, - Embryo divided, with two foliaceous plicate cotyledons
highly convoluted cotyledons 5. Couroupita 16. Planchonia
- Ovary 2-3( -5)-locular; ovules 5-8 per locule, the placen-
tae not bilamellar, attached towards base of locule; fruit
GENERA OF LECYTHIDACEAE
campanulate or cylindric, dehiscent; seeds longer than
broad; embryo undifferentiated or with 2 foliaceous Users wanting to make species determinations are
cotyledons, these are highly convoluted 9 referred to the Flora Neotropica monograph 21
9. Androecium elongated on one side into a strap-like struc- (1, 2), in which all Neotropical genera have
ture which curves over summit of ovary; ovary 2( -5)- been revised by the authors. The Pflanzenreich
locular; seeds not winged; embryo undifferentiated
6. Corythophora
treatments of Knuth (1939a, 1939b) are largely
- Androecium elongated on one side but not forming a outdated.
strap-like structure over summit of ovary; ovary 3-locular;
seeds with unilateral wings; embryo with foliaceous
cotyledons 4. Cariniana 1. Gustavia L.
10. Buds enclosed by calyx except for horizontal slit at apex;
Gustavia L., Pl. Surinam. 12: 17-18 (1775).
calyx with 2 lobes at anthesis; style greater than 10mm
long; fruit functionally indehiscent, with small, inwardly
falling operculum only; seeds with thick, honey testa, Small to large trees, leaves often clustered at end
remaining inside fruit at maturity 7. Bertholletia of branches. Inflorescences suprafoliar, axillary or
- Buds not enclosed by calyx; calyx with 6 lobes at anthesis; cauline, solitary or racemose. Flowers actinomor-
style usually less than 10 mm long; fruit usually dehiscent,
with freely falling operculum; seeds without thick, honey phic. Calyx nearly entire to 4-6-lobed. Petals 6,
testa, usually falling from fruit at maturity 11 8, 12 or 18. Androecium a symmetrical ring of
11. Androecial hood coiled inwards, with outwardly extended 500-1210 stamens, the filaments fused at base, all
flap at apex of coil; fruit cylindric or campanulate; ovary fertile. Ovary 4-6(-10)-locular, 7-93 ovules per
3-locular; seeds with wing around circumference; embryo loculus. Fruit berry-like, indehiscent, with seeds
with 2 foliaceous cotyledons 8. Couratari
- Androecial hood fiat or if coiled inward without outwardly embedded in pulp; seeds with yellow, expanded,
extended flap at apex of coil; fruit usually globose; contorted funicles or without well-developed
ovary not 3-locular; seeds without wings; embryo funicles. Embryo with large, fleshy, plano-convex
undifferentiated 12 cotyledons and minute hypocotyl and plumule. 2n
12. Androecial hood usually forming complete coil inwards,
with blunt tipped appendages at apex of coil, these differ-
= 34, 68. 40 species, from Costa Rica to Amazonia
entiated from more abundant, echinate hood appendages; and coastal NE Brazil.
ovary usually 2-, infrequently 4-locular; seeds with lateral
arils or, less frequently, aril completely surrounding testa
9. Eschweilera 2. Grias L.
- Androecial hood fiat or expanded at apex but not forming
Grias L., Syst. Nat. ed. 10: 1075 (1759).
complete coil inwards, the hood appendages not differen-
tiated; ovary usually 4-, less frequently 2-locular; seeds
usually with basal aril, less frequently without aril 13 Trees to 30m tall, leaves clustered at end of
13. Ovary 2-locular, the style not differentiated from summit branches. Inflorescences on trunk or branches,
of ovary, the summit umbonate; fruit campanulate racemes or fascicles, rarely in axils of lower leaves.
6. Corythophora
- Ovary 4-locular, the style usually differentiated from Flowers actinomorphic. Calyx 4-lobed. Petals 4.
summit of ovary, the summit usually truncate; fruit usually Androecium a symmetrical ring of 85-210
globose 10. Lecythis stamens, the filaments fused at base, all fertile,
14. Fruit broadly winged; anthers introrse 11. Petersianthus curved inwards. Ovary 4-locular, 2-4 pendent
- Fruit not winged; anthers extrorse 15 ovules per loculus. Fruit a drupe, indehiscent, with
1 seed embedded in a fleshy pulp; seeds fusiform, ramiflorous racemes or panicles with continuous
lacking cotyledons or developed funicle. 2n = 34. growth at apex. Flowers zygomorphic. Calyx 6-
Six species, Central America, Jamaica, NW South lobed. Petals 6. Androecium with strap-like hood
America. arching over summit of ovary, with anthers
bearing sterile pollen in hood and fertile pollen
in basal ring, with open gap between ring and
3. Allantoma Miers
hood stamens, stamens numerous. Ovary 6-
Allantoma Miers, Trans. Linn. Soc. Lond. 30: 170, 294 (1874), locular with numerous ovules attached to septum.
pro parte, androphoro excl. Fruit round, large and indehiscent, exocarp
woody, numerous seeds embedded in a fleshy
Small to large trees. Leaves not clustered at end of pulp; seeds with funicle 10-15 mm long, cotyle-
branches. Inflorescences terminal or subterminal dons foliaceous. 2n = 34. 3 species, Neotropical,
racemes or once-branched panicles. Flowers actin- Nicaragua to Colombia, Venezuela, Amazonia and
omorphic. Calyx 5-6-lobed. Petals 5-6. Androe- the Guianas.
cium a slightly asymmetrical urceolate tube with
apex divided in 8-10, inwardly reflexed lacinae
bearing anthers and additional stamens inserted 6. Corythophora R. Knuth
sparsely over interior; stamens ca. 30, all fertile. Corythophora R. Knuth in Engl. Pflanzenr. IV. 219a: 50-51
Ovary (3)4-5 locular, with ca. 20 ovules inserted (1939).
all along septum of each loculus. Fruit an elongate,
cylindrical, circumscissile capsule, with tack- Large trees. Leaves not clustered at end of
shaped operculum, dehiscent from base; seeds branches. Inflorescences terminal or subterminal
numerous, narrowly linear longate, with hard, racemes or panicles. Flowers zygomorphic. Calyx
woody testa, and caducous flattened stipe-like 6-lobed. Petals 6. Androecium with strap-like
funicle, not embedded in pulp. One species, hood arching over summit of ovary, the hood
Neotropical, Amazonian Venezuela and Brazil appendages with or without anthers, 40-230
along upper Orinoco, Rio Negro and lower stamens in staminal ring, hood closed over stami-
Amazon rivers. na! ring. Ovary 2-5-locular, with 4-26 ovules in
each loculus, ovules attached to base or lower part
of septum. Fruit a woody, campanulate or cylin-
4. Cariniana Casar.
drical circumscissile capsule; seeds elongate, not
Cariniana Casaretto., Nov. Stirp. Bras. Dec. 4: 34-37 (1842). embedded in pulp, with basal aril and macropo-
dia! embryo. 2n = 34. Four species, Neotropical, E
Small to large trees to 55 m tall. Leaves not clus- Amazonian Brazil and Guianas.
tered at end of branches. Inflorescences terminal
or axillary racemes or panicles. Flowers nearly
7. Bertholletia Humb. & Bonpl. Fig. 76
actinomorphic to zygomorphic. Calyx 5-6-lobed.
Petals 5-6. Androecium obliquely hood-shaped, Bertholletia Humb. & Bonpl., Pl. Aequinoct. 1: 122-127, t. 36
almost symmetrical to markedly asymmetric, with (1807).
stamens inserted in a complete circle all over inte-
rior or at apex only; stamens 10-150, all fertile. Large trees to 55 m tall. Leaves not clustered at
Ovary 3-locular with ovules inserted all along ends of branches. Inflorescences terminal and axil-
septum. Fruit a cylindrical or campanulate cir- lary spikes or panicules of spikes. Flowers zygo-
cumscissile capsule, with tack-shaped operculum; morphic. Calyx 2-lobed. Petals 6.Androecium with
seeds numerous, unilaterally winged, not embed- strap-like hood arching over summit of ovary
ded in pulp. 15 species, Neotropical, from north- and tightly pressed against staminal ring; hood
ern Colombia and Venezuela to Central Brazil, appendages all sterile, ring stamens 80-135. Ovary
predominantly Amazonian. (3)4{ -6)-locular, with 16-25 ovules inserted at
base of septum of each loculus. Fruit a large,
globose, woody circumscissile capsule with small,
5. Couroupita Aubl. internally detaching operculum, thus secondarily
Couroupita Aubl., Hist. Pl. Guiane 2: 708-711, t. 282 (1775). indehiscent; seeds 10-25, triangular in cross
section, testa hard and woody, not embedded in
Medium- to large-sized trees. Leaves clustered at pulp, the embryo undifferentiated, cotyledons
end of branches. Inflorescences cauliflorous or represented by two small scales only. 2n = 34.
Lecythidaceae 229
Fig. 77. Lecythidaceae. Lecythis pisonis. A Flowering branch.

B Flower. C Androecium, medial section. D Ovary, vertical
Fig. 76. Lecythidaceae. Berthol/etia excelsa. A Flowering section. E Ovary, transverse section. F Pyxidium with detached
branch. B Androecium, medial section. C Calyx and pistil. D operculum. G Seed with fleshy aril and cord-like funicle at
Ovary, vertical section. E Pyxidium. F Seed. G Seedling. Drawn base. Drawn by B.Angell. (Mori and Prance 1990b)
by B. Angell. (Mori and Prance 1990b)
One species, B. excelsa Humb. & Bonpl., confined 9. Eschweilera Mart. ex A.P. DC.
to Amazonia.
Eschweilera Martius ex A.P. DC., Prodr. 3: 393 (1828).
8. Couratari Aubl. Small to large trees. Leaves not clustered at end of

branches. Inflorescences terminal, axillary, rami-
Couratari Aubl., Hist. Pl. Guiane 2: 723, t. 290 (1775).
florous or cauliflorous racemes, spikes or panicles.
Flowers zygomorphic. Calyx 4- or 6-lobed. Petals 4
Medium to large trees. Leaves not clustered at ends
or 6. Androecium with strap-like hood arching
of branches. Inflorescences terminal or axillary
over summit of ovary, the appendages sterile,
racemes or panicles. Flowers zygomorphic. Calyx
coiled inwards; ring stamens 82-500. Ovary
6-lobed. Petals 6. Androecium with strap-like
usually 2-locular, rarely 4-locular, the ovules
hood arching over summit of ovary, the hood
usually basal. Fruit small to medium-sized, cir-
apically coiled inwards, the apical part abruptly
cumscissile capsules always dehiscent; seeds 1- 35,
folded back over hood to form outwardly folded
not embedded in pulp, often with a lateral
flap; hood appendages all sterile; ring stamens
appressed aril, rarely without aril. 2n = 34. About
10-75. Ovary 3-locular, with numerous ovules
85 species, Neotropical, Mexico, Central America,
inserted along septum or at base of loculus. Fruit
Colombia, Venezuela, Amazonia to E and C Brazil.
an elongate, cylindrical or campanulate circum-
scissile capsule; operculum tack-shaped, seeds
numerous, not embedded in pulp, with symmetri- 10. Lecythis Loefl. Fig. 77
cal wings, cotyledons foliaceous. 20 species,
Lecythis Loefl., Iter Hispan.: 189 (1758).
Neotropical from Costa Rica to E Brazil.
Small to very large trees. Leaves not clustered at

end of branches. Inflorescences terminal or axil-
lary racemes, spikes or once-branched panicles,
rarely on young stems below leaves. Flowers zygo-
morphic. Calyx 6-lobed, the lobes often with
mucilage ducts. Petals 6. Androecium with strap-
like hood arching over summit of ovary, the hood
appendages sterile or with anthers; ring stamens
70-1000. Ovary {3)4{-6)-locular, ovules inserted
on lower part of septum. Fruits small to very large,
woody circumscissile capsules, sometimes inde-
hiscent and falling to the ground with seeds inside,
generally campanulate or urceolate, the base
usually remaining on tree after dehiscence; seeds,
often with a basal swollen aril projecting beyond
micropylar end, rarely with lateral aril or exaril-
late. 2n = 34. 26 species. Neotropical, Central
America to Sao Paulo, Brazil.
11. Petersianthus Merrill

Petersianthus Merrill, Philipp J. Sci. C. Bot. 11: 200 (1916).
Combretodendron A. Chev.
Large trees. Leaves not clustered at ends of

branches. Inflorescences terminal panicles.
Flowers actinomorphic. Calyx 4-lobed. Petals 4,
less than 10 mm long. Disc well developed, lining Fig. 78. Lecythidaceae. Barringtonia calyptrata. A Habit. 8
area above receptacle.' Androecium a symmetrical Flower buds, beneath with circumscissile calyx. C Flower in
ring of numerous stamens, fused at base to 4 mm, vertical section, note nectary disk and ramification of androe-
all fertile. Ovary 4-locular, 3-8 ovules per loculus, cium. D Fruits with persistent style. Drawn by R. van Creve!.
(Payens 1967)
placentation central axile. Fruits dry, indehiscent,
fibrous, fusiform, broadly 3-4-winged. Seeds
4-5; embryo with 2 plicate cotyledons and a embryo undifferentiated. 2n = 26. 41 species,
long radicle. 2 species, tropical W Africa and E Africa, Madagascar and tropical Asia.
Philippines.
13. Chydenanthus Miers

12. Barringtonia Forst. Fig. 78
Chydenanthus Miers, Trans. Linn. Soc. II, 1: 111 (1875).
Barringtonia Forst., Char. Gen.: 75, t. 38 (1776); Payens, Blumea
15: 157-263 (1967), monogr.
Medium to large trees. Leaves not clustered at
ends of branches. Inflorescences terminal pani-
Small to large trees. Leaves clustered at ends of
cles. Flowers actinomorphic. Calyx cupuliform,
branches. Inflorescences axillary or less frequently
with 4 small lobes. Petals 4, exceeding 10 mm long.
terminal or cauline racemes or spikes, usually pen-
dulous. Flowers actinomorphic. Calyx distinctly 4- Disc a w ell-developed ring. Androecium a sym-
metrical ring of numerous stamens in 3-4 whorls,
lobed or completely fused or with only an apical
the innermost of which is staminodial, distinctly
pore in early development. Petals {3)4{5), exceed-
fused at base. Ovary cylindrical, 2-locular, 1- 2
ing 10mm long. Disc a well-developed ring.
erect ovules per loculus, placentation basal, axile.
Androecium a symmetrical ring of numerous
stamens in 3-8 whorls, fused for 1-10mm, inner Fruits long, elliptic, unridged 1-seeded drupes;
whorl staminodial, second inner whorl often sta- embryo undifferentiated. 2n = 26. 2 species, Java
and Sumatra.
minodial. Ovary {2- )4-locular, 4 p endulous ovules
per loculus, placentation axile. Fruits 1-seeded
drupes, often rectangular, 4-ridged or 4-winged;
Lecythidaceae 231
14. Abdulmajidia Whitmore 17. Foetidia Commerson

Abdulmajidia Whitmore, Kew Bull. 29: 207-211 (1974). Foetidia Commerson ex Lam. Encyl. 2:457 (1786).
Trees. Leaves clustered at ends of branches. Inflo- Small to large trees. Leaves clustered at ends
rescences terminal, axillary or cauline racemes or of branches. Inflorescences terminal or axillary
spikes. Flowers actinomorphic. Calyx 3-4-lobed, racemes or flowers solitary. Flowers actinomor-
coloured deep red. Petals 3, ca. 28 mm long. Disc a phic. Calyx 3-5-lobed, valvate, lobes thick, hard,
well-developed ring. Androecium a symmetrical purple on both surfaces. Petals absent. Disc a
ring of numerous stamens in 7-8 whorls, the slightly elevated area on surface of ovary adjacent
innermost of which is staminodial, distinctly to androecium, purple coloured. Stamens numer-
fused at base. Ovary 3-4-locular, 8-20 ovules per ous, free to base, all fertile. Ovary 3-4-locular,
loculus, inserted on upper portion of placenta. 10-15 ovules per loculus, inserted in two rows on
Fruits 2-5-seeded drupes, rounded not ridged; central placenta. Fruits turbinate, 1-4-locular,
seeds with hard, woody testa. 2 species, Malay few-seeded drupes; embryo slightly curved, with
Peninsula. long radicle. 17 species, Madagascar, Mauritius
and E Africa.
15. Careya Roxburgh
Careya Roxburgh, Hort. Bengal.: 52 (1814). Selected Bibliography
Shrubs to large trees. Leaves clustered at ends Anderberg, A.A. 2002. See general references.
of branches. Inflorescences terminal or axillary APG (Angiosperm Phylogeny Group) 1998. See general
references.
spikes. Flowers actinomorphic. Calyx 4-lobed. Berry, E.W. 1924a. A fossil flower from the miocene of
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ring. Androecium a symmetrical ring of numerous Berry, E. W. 1924b. The Tertiary Flora of the Island of Trinidad.
stamens in 5-8 whorls, the innermost of which is B.W.I. Johns Hopkins Studies in Geology 6: 120-123.
staminodial, outermost also staminodial, dis- Cronquist, A. 1981. See general references.
Huertas, G.G. 1969. Un nuevo genero y especie fosiles de las
tinctly fused at base. Ovary 4(5)-locular, 20-40 lecitidaceas. Caldasia 10(48): 365-369.
ovules per loculus inserted in two or four rows on Jacques, F. 1965. Morphologie de pollen et des ovules de
placenta; placentation central axile. Fruits drupes, Couroupita guianensis Aubl. (Lecythidacees). Pollen Spores
4-locular, with several seeds in each loculus, 7: 175-180.
crowned by 4 persistent sepals; seeds with undif- Kartawinata, E.K.1965. The genus Planchonia Blume (Lecythi-
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and the putatatively related Lecythidaceae. Brittonia 41:
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1-many-seeded drupes; embryo with two folia- chromosomiques dans une collection d'especes tropicales.
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Mori, S.A., Prance, G.T. 1990a. Taxonomy, ecology and eco- daceae from Malaysia. Kew Bull. 29: 207-211.
nomic botany of the Brazil nut (Bertholletia excelsa Humb. Zeeuw, C.H. de. 1987. Wood anatomy. In: Mori, S.A. et al. The
& Bonpl.: Lecythidaceae). Adv. Econ. Bot. 8: 130-150. Lecythidaceae of a lowland neotropical forest: La Fumee
Mori, S.A., Prance, G.T. 1990b. Lecythidaceae - part II. The Mountain, French Guiana, chap. IX. Mem. New York Bot.
zygomorphic-flowered New World genera. Fl. Neotrop. Gard. 44: 100-112.
Monogr. 21(2): 1-376. Zeeuw, C.H. de. 1990. Secondary xylem of Neotropical Lecythi-
Mori, S.A., Prance, G.T., Bolten, A.B. 1978. Additional notes on daceae. In: Mori, S.A., Prance, G.T. (eds.) Lecythidaceae -
the floral biology of neotropical Lecythidaceae. Brittonia 30: part II. The zygomorphic-flowered New World genera, chap.
113-130. II. Fl. Neotrop. Monogr. 21 (II): 4-59.
Mori, S.A., Orchard, J.E., Prance, G.T. 1980. Intrafloral pollen Zeeuw, C.H. de. Manuscript. Wood anatomy of the Old World
differentiation in the New World Lecythidaceae subfamily Lecythidaceae. Part I - Foetidia Commerson ex Lamarck,
Lecythidoideae. Science 209: 400-403. Foetidioideae.
Mori, S.A., Black, D., de Zeeuw, C. 1987a. Habit and bark. In: Zeeuw, C.H. de. Manuscript. Wood anatomy of the Old World
Mori, S.A. et al. The Lecythidaceae of a lowland Neotropical Lecythidaceae. Part II - Crateranthus E.G. Baker and
forest: La Fumee Mountain, French Guiana, chap. VIII. Mem. Napoleonaea P. Beauv., Napoleonaeoideae.
New York Bot. Gard. 44: 86-99. Zeeuw, C.H. de. Manuscript. Wood anatomy of the Old World
Mori, S.A. et al. 1987b. The Lecythidaceae of a lowland Lecythidaceae. Part III. Careya Roxb. and Chydenanthus
neotropical forest. La Fumee Mountain, French Guiana. (Bl.) Miers, Planchonioideae.
Mem. New York Bot. Gard. 44: 1-190. Zeeuw, C.H. de. Manuscript. Wood anatomy of the Old
Moritz, A. 1984. Estudos biol6gicos da florayao e da frutifi- World Lecythidaceae. Part IV - Petersianthus Merrill,
cayao da Castanha-do-brasil (Bertholletia excelsa H.B.K.). Planchonioideae.
Belem: EMBRAPA-CPATU, Documentos 29: 1-82. Zeeuw, C.H. de. Manuscript. Wood anatomy of the Old World
Morton, C.M., Mori, S.A., Prance, G. T., Karol, K.G., Chase, M.W. Lecythidaceae. Part V - Planchonia King, Planchonioideae.
1997. Phylogenetic relationships of Lecythidaceae: a cladis- Zeeuw, C.H. de. Manuscript. Wood anatomy of the Old World
tic analysis using rbcL sequence and morphological data. Lecythidaceae. Part VI - Barringtonia J.R. & G. Forster,
Am. J. Bot. 84: 530-540. Planchonioideae.
Muller, J. 1972. Pollen morphological evidence for subdivision
and affinities of Lecythidaceae. Blumea 20: 351-355.
Lepidobotryaceae 233
Lepidobotryaceae
K. KUBITZKI
Lepidobotryaceae J. Leonard in Bull. Jard. Bot. Etat 20: 38 1993). The wood of Lepidobotrys is similar but has
(1950), nom. cons. no vestured pits.
Germination of Ruptiliocarpon is epigeal
Dioecious trees with very sparse indumentum of (Hammel and Zamora 1993).
unicellular hairs. Leaves unifoliolate, rachis with
a stipel and disarticulation at base of pulvinate FLORAL MoRPHOLOGY. The nectary disk of Lepi-
petiolule; stipules and stipels fugaceous. Inflo- dobotryaceae has attracted considerable attention;
rescences racemes or spikes, sometimes com- it is shallowly bowl-shaped in Lepidobotrys and
pound; flowers unisexual; sepals and petals 5, tubular in Ruptiliocarpon. The disk is found in
free, imbricate (calyx with quincuncial preflo- male and female flowers but in the latter is usually
ration); stamens 5 + 5, the antesepalous longer smaller. Its nectar-secreting function, obvious
than the antepetalous; filaments fused at base into from the rich development of glandular tissue, has
a nectariferous disk or tube; anthers tetrasporan- been observed in the greenhouse (Link 1991). In
giate, dehiscing lengthwise; gynoecium 2-3- Lepidobotrys the stamens are inserted with long
carpellate, ovary syncarpous, with apical-axile filaments at the margin of the disk which pro-
placentation; ovules 2 per locule, collateral, trudes between the filament bases (Fig. 79D).
anatropous, with placental obturator above the The disk is comprised of glandular tissue which
micropyle; stylodia 2 or 3, nearly free or very extends deeply into the floral axis but is never
short. Fruit a septicidal capsule, exocarp and vascularized itself but closely associated with the
endocarp separating; seeds black, usually 1 per sepal and carpel strands; it is covered by an epi-
fruit, with orange aril attached to apex of locule; dermis on which nectarial stomata are sparsely
embryo oily, endosperm 0. dispersed (Link 1991). In Ruptiliocarpon the
A family comprising two monotypic genera; nectary disk is tubular and the stamens are
West Africa, C and Andean South America south inserted on its rim. The antesepalous stamens have
to Peru. short free filaments, whereas the anthers of the
antepetalous stamens are sessile on the margin of
VEGETATIVE STRUCTURES. Both genera agree in the tube. In the male flowers the adaxial side of the
the characteristic leaf structure. The petiole of the stamina! tube has profusely branched vascular
unifoliolate leaf is pulvinate at its base (Fig. 79H). tissue and densely stained cells (To be and Hammel
The petiolulus is pulvinate for its entire length, 1993). The disk of Lepidobotrys is interpreted
articulated at the juncture with the petiole and is by Link (1991) as receptacular, whereas Tobe and
provided with an adaxial stipel. In Lepidobotrys Hammel (1993) contend that the tubular structure
this stipel has been found to be 2-nerved and in Rutiliocarpon is the product of fusion of the
is probably the fusion product of two stipels stamen filaments.
(Leonard 1950).
Leaf anatomy of Lepidobotrys is noteworthy for EMBRYOLOGY. Ovules of Ruptiliocarpon are ana-
paracytic and laterocytic stomata, vertical bundle tropous, bitegmic, and crassinucellar. The inner
sheath extensions of some of the veins, and cristar- and outer integument are multiplicative; both take
que cells (van Welzen and Baas 1984). In the wood part in the formation of the micropyle. The ovules
of Ruptiliocarpon growth rings are discernable; and young seeds are pachychalazal. The embryo
the vessels are solitary or grouped and have simple sac is 8-celled. An obturator is formed from
perforations with oblique plates; intervascular pits funicular tissue near the micropyle (Tobe and
are vestured; fibres are thin-walled, non-septate; Hammel1993).
rays are uniseriate homogeneous or nearly so, with
mostly procumbent cells; parenchyma is scattered, POLLEN MORPHOLOGY. The pollen of Lepi-
in bands, paratracheal, and terminal (Mennega dobotrys is tricolporoidate, subprolate,
234 K. Kubitzki
Fig. 79. Lepidobotryaceae. Lepidobotrys staudtii. A Inflores- Erythroxylaceae, Meliaceae, and several other
cence. B Flower bud. C Inflorescence. D Flower, vertical families (see Hallier 1923; Leonard 1950; Hamel
section. E Stamens. F Pistil. G Inflorescence, vertical section. H
Base of leaf and petiole showing joint. (Hutchinson and Dalziel and Zamora 1993). Leonard (1950), who proposed
1927) family rank for Lepidobotrys, described various
characters distinguishing it from Oxalidaceae:
crassitegillate-intectate, with intraluminal col- - Lepidobotryaceae: dioecious; stipules and
umellae and excessivley wide ectoapertures; the stipels +; disk +; ovary 3-2-merous; ovules
nexine has a granular inner surface with which it collateral; capsule septicidal.
is linked together with the intine (Erdtman 1952; - Oxalidaceae: hermaphrodite or androdioecious;
Oltmann 1971; Huynh 1969). The exine of stipules and stipels 0; disk 0; ovary 5-merous;
Rupotiliocarpon is described as verrucate to fos- ovules superposed.
sulate/foveolate (obviously from SEM analysis)
(Hammel and Zamora 1993). Both Oltmann (1971) In a molecular analysis (rbcL), Savolainen, Fay
and Huynh (1969) stressed the palynological sim- et al. (2000) found strong support for a Celastrales
ilarity between Lepidobotrys, Sarcotheca and clade, in which Lepidobotryaceae are basal to
Dapania. Parnassiaceae and Celastraceae. Since in this
analysis Celastrales appear in relatively: -close
FRUIT AND SEED. The capsule of Lepidobotrys proximity to Oxalidaceae, the matter may require
dehisces with 2 or 3 valves, while that of Ruptilio- further attention.
carpon ruptures irregularly. Both integuments The relationship between Lepidobotrys and
take part in the formation of the seed coat but Ruptiliocarpon, doubted by Takhtajan (1997), is
mechanical strength is provided by the fibrous so close that, to my mind, the generic rank of
exotegmen (only present in Ruptioliocarpon, Tobe Ruptiliocarpon is only weakly justified.
and Hammel1993).
KEY TO THE GENERA
AFFINITIES. Lepidobotryum and Ruptiliocarpon 1. Flowers pedicellate; ovary 3-merous 1. Lepidobotrys
have been compared with Linaceae, Oxalidaceae, - Flowers sessile; ovary 2-merous 2. Ruptiliocarpon
Lepidobotryaceae 235
1. Lepidobotrys Engl. Fig. 79 Selected Bibliography

Lepidobotrys Engl., Bot. Jahrb. Syst. 32: 108 (1902); Leonard,
Bull. Jard. Bot. Etat 20: 33 (1950). Erdtman, G. 1952. See general references.
Hallier, H. 1923. Lepidobotrys Engl.: Die Oxalidaceen und die
Geraniaceen. Beih. Bot. Centralbl. 39, II: 163.
Flowers pedicellate, male in many-flowered stro- Hammel, B.E., Zamora, N.A. 1993. Ruptiliocarpon (Lepi-
biliform racemes, female in few-flowered, short dobotryaceae): a new arborescent genus and tropical
fasciculate racemes, unisexual with rudimentary American link to Africa, with a reconsideration of the
family. Novon 3: 408-417.
organs of opposite sex; stamens with versatile Hutchinson, J., Dalziel, J.M. 1927. Flora of west tropical Africa,
anthers on long filaments inserted at margin of vol. 1. London: Crown Agents.
fleshy disk surrounding the ovary; ovary 3-locular; Huynh, K.-L. 1969. Etude du pollen des Oxalidaceae I. Mor-
stylodia 3. Exocarp leathery, endocarp pergamen- phologie generale - palynotaxonomie des Oxalis ameri-
taceous. A single species, L. staudtii Engl., Africa, cains. Bot. Jahrb. Syst. 89: 271-303.
Leonard, J. 1950. Lepidobotrys Engl., type d'une famille nou-
Guinea-Congo region. velle des Spermatophytes: les Lepidobotryaceae. Bull. Jard.
Bot. Nat. Belg. 20: 31-40.
Link, D.A. 1991. The floral nectaries of Geraniales. III. Lepi-
2. Ruptiliocarpon Hammel & N. Zamora dobotryaceae J. Leonard. Bull. Jard. Bot. Nat!. Beige 61:
Ruptiliocarpon Hammel & N. Zamora, Novon 3: 408 (1993). 347-354.
Mennega, A.M.W. 1993. Comparative wood anatomy of
Ruptiliocarpon caracolito (Lepidobotryaceae). Novon 3:
Flowers sessile, in leaf-opposed compound spikes, 418-422.
cryptically unisexual (with rudimentary ovules or Oltmann, 0. 1971. Pollenmorphologisch-systematische Unter-
anthers without pollen); stamens with basifixed suchungen innerhalb der Geraniales. Diss. bot. 11, 163 +XI
anthers and filaments fused into a nectariferous pp., 30 t. Lehre: Cramer.
Savolainen, V., Fay, M.F. et al. 2000. See general references.
tube; stamens inserted on margin of tube, the Takhtajan, A. 1997. See general references.
antesepalous on short filaments, the antepetalous Tobe, H., Hammel, B. 1993. Floral morphology, embryology,
with sessile anthers; connectives produced to form and seed anatomy of Ruptiliocarpon caracolito (Lepi-
a small pubescent appendage; ovary 2-locular; sty- dobotryaceae). Novon 3: 423-428.
lodia very short. Exocarp coriaceous to woody, Welzen, P. van, Baas, P. 1984. A leaf anatomical contribution to
the classification of the Linaceae complex. Blumea 29:
irregularly rupturing and exposing 2 horny endo- 453-479.
carps, one nearly completely surrounding the
seed, the other usually empty and smaller. Only
one species, R. caracolito Hammel & N. Zamora, in
humid lowland forests of Costa Rica, Colombia,
Peru, and Suriname.
236 B. Wallniifer
Lissocarpaceae
B. wALLNOFER
Lissocarpaceae Gilg in Engler, Syllabus, ed. 9 & 10: 324 (1924), For further details see Schadel and Dickison
nom. cons. (1979); compare also the description of the leaf
anatomy of L. tetramera by Busch (1913).
Small, glabrous trees without latex. Leaves alter- In the wood the vessels are solitary and in radial
nate, simple, exstipulate, finely and pinnately multiples of 2-9 cells; perforation plates are simple
veined, with entire margins; extrafloral nectaries or less frequently scalariform; pits among vessels
often present on abaxial leaf surface. Inflores- and ray cells or parenchyma are small (mostly
cences axillary, racemose. Flowers with prophylls, 4-8 Jlm in diameter); axial parenchyma is pre-
articulated at base, sessile or shortly pedicellate, dominantly apotracheal; imperforate tracheary
actinomorphic, 4-merous, unisexual (plants prob- elements are of the libriform fiber type; rays are
ably dioecious); calyx campanulate with 4 imbri- heterocellular, 1-2 cells wide, predominantly more
cate, retuse lobes, persistent but not enlarging in than 1 mm high; fibers are without spiral thicken-
fruit; corolla sympetalous, isomerous with calyx, ings (Ng 1971, 1991; Dickison and Phend 1985).
with lobes contorted sinistrorsely; tube promi- The wood of Ebenaceae is very similar except that
nent, bearing distally a corona of 8 lobes which, the rays are mostly over 1 mm high.
however, is missing in some species; male flowers
with 8 stamens in one whorl, the filaments shortly INFLORESCENCE AND FLOWER STRUCTURE. The
connate and attached to the corolla tube below the inflorescence is of racemose nature (for further
middle; anthers linear, erect, basifixed, with an details, see Walln6fer, in press). Erroneously
apiculate-prolonged connective, 4-sporangiate, flowers were often considered to be hermaphro-
dehiscing by longitudinal slits; locules and ovules ditic, but Ng (1971, 1991) pointed out that they are
not developed; female flowers: staminodes resem- unisexual and the plants probably dioecious.
bling stamens, but anthers collapsed and without Gynoecium structure is quite similar to Ebenaceae
pollen; gynoecium syncarpous, 4-carpellate; ovary except that the ovary is inferior and false septa are
inferior, 4-locular; style terminal; stigma shallowly not developed (Ng 1971, 1991). Each of the four
4-lobed; ovules 2 per locule, anatropous; placenta- locules bears a pair of anatropous ovules at the
tion apical. Fruit an ovoid berry with a persistent apex of the ovary. The four locules all open into a
calyx; seeds 1-2 by abortion, pendulous; hilum rel- common stylar passage. As in Ebenaceae, the
atively small, apical; testa smooth, thin, coria- radicle of the embryo is superior.
ceous; endosperm horny, abundant, smooth;
embryo straight, with two small foliaceous cotyle- EMBRYOLOGY AND SEED. The ovules are pendu-
dons and a strongly developed radicle. lous, oblong, anatropous, with the raphe descend-
A monogeneric family with 8 species from trop- ing on the outer side, but otherwise unknown. The
ical South America. vascular system of the seed consists of one bundle
descending from the placenta to the chalaza which
VEGETATIVE ANATOMY. Extrafloral nectaries then sends 6-12 branches back up to the apex of
occur on abaxial leaf surfaces scattered along the the locule. These branches are visible as promi-
midvein, but are lacking on some leaves (Busch nently raised, longitudinal ridges on the seed
1913; Gentry 1996). The stomata are anomocytic surface (Ng 1971; White 1981).
and are restricted to the abaxial leaf surface. Dif-
fuse vesiculose sclereids are present in the petiole POLLEN MORPHOLOGY. The pollen is triporate
and mesophyll of the leaves. The vasculature of the and suboblate to oblate-spheroidal or somewhat
petioles consists of an arc with outwardly curving flattened; the pores are tenuimarginate and about
ends. The marginal ultimate venation is fimbriate. 7 )liD wide. The sexine is thinner than the nexine
Lissocarpaceae 237
that Lissocarpa is the closest relative of Ebenaceae.

This was strongly endorsed by the molecular
studies of Berry et al. (2001) and Anderberg et al.
(2002).
DISTRIBUTION AND HABITATS. Lissocarpa OCCUrS

in Bolivia, Peru, Colombia, Venezuela, northern
Brazil, and Guyana (White 1981; Berry 1999, 2001;
Walln6fer, in press). Four species are characteris-
tic elements of the specialized vegetation on white
sand and along blackwater rivers, four are found
in montane forests.
One genus:
Lissocarpa Benth. Fig. 80

Lissocarpa Benth. in Benth. & Hook. f., Gen. Pl. 2: 671 (1876);
Berry, Brittonia 51 : 214-216 (1999), synop.
Description as for the family.
Anderberg, A.A.2002. See general references.

Bentham, G. 1876. Styraceae. In: Bentham, G., Hooker, J.D.
Genera plantarum 2: 666-671. London: Reeve & Co.
Berry, P.E. 1999. A synopsis of the family Lissocarpaceae.
Brittonia 51: 214-216.
Berry, P.E. 2001. Lissocarpaceae. In: Berry, P.E., Yatskievych, K.,
Holst, B.K. (eds.) Flora of the Venezuelan Guayana. 6: 19-20.
St. Louis: Missouri Botanical Garden Press.
Fig. 80. Lissocarpaceae. Lissocarpa benthamii. A Flowering Berry, P.E., Savolainen, V., Sytsma, K.J., Hall, J.C., Chase, M.W.
branch. B Flower with prophylls. C Flower with calyx and 2001. Lissocarpa is sister to Diospyros (Ebenaceae). Kew
ovary in vertical section. D Corolla laid open. E Stamen. F Bull. 56: 725-729.
Ovary in transverse section. G Fruit. H Apex of fruit with Busch, P. 1913. Anatomisch-systematische Untersuchung der
persistent calyx. I Seed. J Same in vertical section. (Oliver Gattung Diospyros. Crefeld: Wilhelm Greven, 95 pp.
1895) Dickison, W.C., Phend, K.D. 1985. Wood anatomy of the
Styracaceae: evolutionary and ecological considerations.
I.A.W.A. Bull., N.S., 6: 3-22.
Franceschi, D. de 1993. Phylogenie des Ebenales: analyse de
and columellate (Erdtman 1952). According to l'ordre et origine biogeographique des especes indiennes.
Pub!. Dept. Ecol. Institut Fran<;ais de Pondicherry 33: 1-153
Ng (1971) and White (1981), the surface shows a (+ annexe A-E, + 61 planches).
prominent reticulate sculpturing. Later workers, Gentry, A.H. 1996. A field guide to the families and genera
however, found the tectum fossulate to somewhat of woody plants of northwest South America. Chicago:
rugulate (Morton and Dickison 1992) or granu- University of Chicago Press.
lated and undulate. Size ranges from 46 x 53 to Gilg, E. 1924. Lissocarpaceae. In: Engler, A. Syllabus der
Pflanzenfamilien. Ed. 9 & 10: 324.
65 X 70~-Lm (Ng 1971). Morton, C.M., Dickison, W.C. 1992. Comparative pollen mor-
phology of the Styracaceae. Grana 31 : 1-15.
AFFINITIES. When Bentham (1876) described Ng, F.S.P. 1971. A taxonomic study of the Ebenaceae with
Lissocarpa he placed it in Styracaceae. Oliver special reference to Malesia. Doctoral Thesis, University of
(1895) considered that it better belonged in Ebe- Oxford, 221 pp.
Ng, F.S.P. 1991. The relationships of the Sapotaceae within the
naceae. Perkins (1907) excluded it from Styra- Ebenales. In: Pennington, T.D. The genera of Sapotaceae,
caceae, and Gilg (1924) placed it in a family of its chap. l. Kew: Royal Botanic Gardens & New York: New York
own. Ng (1971, 1991) and Franceschi (1993) found Botanical Garden, pp. 1-13.
238 B. Wallnofer
Oliver, D. 1895. Lissocarpa benthamii Giirke. Hooker's Icon. Pl. Wallnofer, B., in press. A revision of Lissocarpa Benth (Lisso-
25: pl. 2413. carpaceae). Ann. Naturhist. Mus. Wien, B, 105.
Perkins, J. 1907. Styracaceae. In: Engler, A. (ed.) Das Pflanzen- White, F. 1981. Lissocarpaceae. In: Maguire, B. et al. The botany
reich IV 241: 1-lll. Leipzig: Engelmann. of the Guayana Highland - part XI. Mem. New York Bot.
Schadel, W.E., Dickison, W.C. 1979. Leaf anatomy and venation Gard. 32: 329-330.
patterns of the Styracaceae. J. Arnold Arbor. 60: 8-37.
Loasaceae 239
Loasaceae
M. WEIGEND
1-4 microsporangia; connective usually undiffer-

Loasaceae Juss., Ann. Mus. Natl. Hist. Nat. 5: 19-27 (1804),
"Loaseae". entiated; staminodia highly diversified, free and
filiform or petaloid or united to form elaborate
Annual or perennial, erect, decumbent, or winding staminodial complexes (nectar scales); nectaries
herbs, rarely rosette plants, vines, woody Hanas, antepetalous glands, or ring- or cup-shaped
subshrubs, shrubs or small trees, from 5 em to disc, or absent; ovary completely inferior to 4/5
10m tall; stem with white pith, rarelysolid, terete, superior; style filiform, included or exserted;
rectangular or grooved, often with lenticels and/or stigma punctiform or with 2-5 papillose, parallel
tuberculate; rhizome, xylopodium or runners or divergent lobes; ovules anatropous, either
sometimes present; roots fibrous, primary root single, pendent, or numerous on simple or divided
dominant or evanescent, occasionally primary parietal placentae, in some Petalonyx with ob~ura
and/or secondary roots tuberous, or only adventi- tor, in Gronovioideae and Petalonych01deae
tious roots present in mature plant; indumentum possibly crassinucellar, in Mentzelioideae and
typically with scabrid/glochidiate trichomes and Loasoideae unitegmic and tenuinucellar with well-
urticant setae, sometimes with uniseriate and/or developed chalaza! and micropylar haustoria.
glandular trichomes. Leaves opposite or some Fruit a capsule or cypsela, typically dehiscentwith
alternate, petiolate or sessile, exstipulate or pseu- 3-5 apical valves, septicidal or rarely septifragous,
dostipulate; lamina linear, ovate, or circular, rarely with 3-5 apical valves and 1 longitudinal
membranaceous to coriaceous, 2-50 em long, base suture, or only with 3-5 longitudinal sutures,
cuneate, cordate or peltate, usually divided, rarely rarely indehiscent; seeds 1-many, globose, ovoidal
pinnatisect, trifoliate, bipinnatifid, or palmate; or angular, sometimes winged; testa usually pale
margin serrate or mucronate, rarely entire, with or dark brown to black, reticulate with or without
hydathode teeth. Inflorescences often terminal central papillae, sometimes with fenestrate anti-
thyrsoids, sometimes reduced to dichasia, mono- clinal walls, or striate; embryo straight, embedded
chasia or monads, rarely racemes; bracts present in copious and oily endosperm. Seedling with 2
or rarely. absent, mostly green. Flowers perfect, ovate to obcordate, densely trichomatose, apically
proterandric, usually actinomorphic, chasmoga- emarginate cotyledons, midvein ending in hydath-
mous, rarely cleistogamous; perianth heteroch- ode tooth.
lamydeous, (4)5( -8)-merous; calyx tube conical Comprising 20 genera in four subfamilies with
to globose, densely covered with trichomes; calyx ca. 330 species, mostly American, outliers in Africa
lobes usually persistent and accrescent, rarely and on the Marquesas Islands (Polynesia).
caducous; aestivation apert; corolla choripetalous,
sympetalous or pseudosympetalous, aestivat~on VEGETATIVE MoRPHOLOGY. The vast majority of
apert, contort or imbricate; petals erect, spreadmg taxa are annual to perennial herbs and this
or reflexed, linear, spatulate, ovate or circular, appears to be the plesiomorphic condition for
planar or deeply boat-shaped, margin entire, most groups. Subshrubs (frequently with ligneous
irregularly serrate, or laciniate, sometimes with underground structures - Cevallia, some Peta-
filiform apical appendages or longitudinallamel- lonyx, Mentzelia) and shrubs (e.g. in Me~tzel~a,
liform flaps, often with short claw and one lateral Aosa, Presliophytum, Nasa, Loasa, Ktssenza)
tooth on each side, membranaceous to carnose, evolved several times independently in desert
green, white, yellow, orange or red; androecium habitats from herbaceous ancestors. Gronovia and
haplostemonous, obdiplostemonous or polyste- Fuertesia are the only winding genera in
monous (150 or more), all stamens fertile or some Loasaceae with alternate leaves. Fuertesia is the
staminodial; initiation centripetal; filaments only woody liana in the family. Winding herbs
filiform, inserted basally or epipetalous, rarely with opposite leaves are restricted to one mono-
very short, or petaloid; anthers basifixed, with phyletic assemblage of primarily South Andean
240 M. Weigend
genera: Loasa, Caiophora, Blumenbachia, Scyphan- trichomes are called scabrid if the processes are
thus. Irrespective of habit, the basic structure of shorter than the diameter of the shaft, and
the plants follows one pattern: at least the first two glochidiate if they are longer and typically hook-
foliage leaves are always opposite (usually there shaped (Fig. 81). There are numerous derivations
are additional opposite leaf pairs). These are from these two basic types (e.g. Dostert and
mostly followed by alternate leaves and the shoot Weigend 1999). Stinging hairs or setae are also
nearly always terminates in a distal dichasium or unicellular, but they have a smooth surface, a
an inflorescence with opposite leaves (and para- slightly bulbous base, a sharp tip, and they are
cladia) at least in its terminal element(s). Typically filled with a clear to brownish fluid. A more or less
all internodes are elongated, but there are rosulate violent skin rash develops upon contact with these
taxa with shortened basal internodes (e.g. in Loasa stinging hairs. The chemical identity of the irri-
and Caiophora). tants is unknown. Flexible (non-silicified) uniseri-
Underground structures are diverse. Nasa has an
evanescent primary root which is quickly replaced
by adventitious roots from the decumbent basal
portion of the stem. In some species of Nasa (e.g.
N. cymbopetala) this basal, decumbent region
turns into a thick, horizontal rhizome. Caiophora,
Blumenbachia, Loasa and Scyphanthus have a
strong primary root which develops either into a
more or less thickened tap-root (C. pentlandii, B.
prietea) or into a spindle-shaped root tuber (L.
asterias); sometimes the primary root is fibrous
and only the secondary roots are tuberous (C. con-
torta), sometimes both are tuberous (C. pteros-
perma, B. dissecta). Additionally, runners with
cataphylls are widespread but not universally
present in Loasa (L. gayana, L. nana) and Caio-
phora (C. nivalis, C. contorta). A long-lived {>7
years) xylopodium is found in Schismocarpus and
Xylopodia, with annual and perennial shoots,
respectively, arising from them.
Foliage in Loasaceae is usually evergreen, but
some taxa are deciduous during the dry season
(Xylopodia, some Mentzelia) or during winter
(south temperate taxa, many species of Loasa,
Blumenbachia). Some of these deciduous taxa
produce narrowly spaced, spirally inserted leaves
in a rapid flush at the beginning of the moist
season (e.g. some Nasa and Aosa).
VEGETATIVE ANATOMY. The stem is typically

covered with numerous trichomes and terete,
rarely quadrangular (Blumenbachia sect. Blumen-
bachia), or deeply grooved (some species of the
Nasa triphylla group). The stem is usually filled
with a white pith and may be externally covered
with dark green to black lenticels (many Nasa)
or cork layers (some Petalonyx). Frequently the
bark starts exfoliating in the second year (e.g.
Petalonyx, Presliophytum, many Mentzelia).
The numerous different trichome types can be
technically divided into three basic types. Scabrid-
glochidiate trichomes are unicellular, silicified or Fig. 81. Loasaceae. Klaprothia fasciculata, glochidiate tri-
calcified with one to numerous processes. These chome from fruit surface, SEM (xSOO). Photo M. Weigend
Loasaceae 241
ate hairs represent a third trichome type. They are phora. Many species in Loasa and some species in
two to over 20 cells long and usually gland-tipped. Blumenbachia have lateral inflorescences which
Setae (stinging hairs) and scabrid-glochidiate probably evolved from a terminal thyrsoid via
trichomes are widespread in Loasaceae, but un- early proliferation. Other derivations concern the
iseriate trichomes are largely restricted to the number and size of bracts: Aosa lacks bracts; in
genus Nasa. Subspecies, species, species groups Nasa only one instead of two bracts is present
and sometimes genera can be defined by trichome on the monochasial branches. Mentzelia sect.
cover: slender glochidiate trichomes with only two Mentzelia has terminal thyrsoids with dichasial
apical hooks are found in Gronovia and Fuertesia, branches and long internodes. All other sections
glochidiate hairs with porrect instead of defiexed in Mentzelia have strongly basitonic inflores-
branches in Petalonyx. Long (>10 cells), brown or cences with shortened distal internodes and often
white uniseriate trichomes are typical of the Nasa single terminal flowers (distal dichasium reduced
grandiflora group, and T-shaped (medifixed) tri- to terminal flower); the primary flower is often
chomes are restricted to the Nasa triphylla group; preceded by 1-12 sterile bracts. Petalonyx is
symmetrically T-shaped trichomes occur only in aberrant with racemose inflorescences with one
Nasa humboldtiana. recaulescent bract and two prophylls per flower,
The brittle wood of Loasaceae is always soft and departing strongly from the thyrsoid/cymose pat-
sometimes slightly succulent. The data here follow terns of the rest of the family.
Carlquist (1984, 1987) unless otherwise indicated.
Growth rings are often visible, e.g. in Xylopodia FLOWER STRUCTURE. Gronovioideae and Peta-
(pers. obs.) and some Eucnide, Mentzelia and lonychoideae have petals with a single principal
Petalonyx, but sometimes absent (Plakothira vein, a cup-shaped nectary, a single pendent
frutescens). Vessels have simple (most) or scalari- ovule and only five stamens. In Gronovioideae the
form (rare: Nasa picta) perforation plates. Lateral calyx lobes are usually larger than the petals and
wall pitting consists typically of circular to oval persist after anthesis, petals are free and the
(6 x 6-8 J..lm) pits. Fibre tracheids are present; they anthers have introrse dehiscence and are densely
have bordered pits or the borders are vestigial. Tall trichomatose. In Petalonychoideae the calyx lobes
and wide, homogeneous or heterogeneous vascu- are smaller than the petals, the petals are free or
lar rays (multi- and uniseriate) are usually present, united, the anthers have lateral dehiscence and are
rarely missing (Petalonyx). The ray cells have thin glabrous, 2-3 of the stamens can be staminodial
but lignified walls. Axial parenchyma is diffuse and filiform (Davis and Thompson 1967). Mentze-
and abundant (most), or absent (Presliophytum), lioideae and Loasoideae share the presence of
or vasicentric (Plakothira frutescens). Sieve- petals with numerous veins from the base, an
elements are of the S-type. annular nectary or distinct antesepalous nectar
Cystoliths (calcium carbonate, silica) and crystal glands, and numerous ovules and stamens.
druses/raphides (calcium oxalate) are frequently Mentzelioideae have free, fiat staminodia, if any,
present, e.g. in the wood parenchyma, in the leaf petals are planar and membranaceous. Loasoideae
lamina or the ovary roof. have antesepalous staminodial complexes of 3-12
staminodia in an inner and an outer tier, the outer
INFLORESCENCE STRUCTURE. Urban (1892, 1910) ones are usually united into a nectar scale. The
provided the first detailed analyses of some taxa, typical staminodial complex in Loasoideae con-
and a comprehensive survey was carried out by sists of three outer, united staminodia forming a
Weigend (1997). Most Loasaceae have acrotonic, nectar scale and two more or less enclosed, free,
terminal thyrsoids with a distal dichasium, or inner staminodia. The nectar scales are extremely
structures derived from it (see Figs. 84D, 85A). Re- diverse morphologically and can be decorated
and concaulescence is widespread in Loasaceae with dorsal calli, arches or filaments, apical wings,
inflorescences, and only relatively few taxa (some and colour patterns. Petals are usually shortly
Mentzelia and Loasa, all Blumenbachia, Scyphan- clawed and deeply boat-shaped (cymbiform),
thus and Caiophora) lack metatopy. Reductions of white or yellow, refiexed or spreading, and the
dichasia to monochasia have occurred repeatedly. immature stamens are refiexed into the petals.
In Caiophora and Scyphanthus the inflorescences Stamens move autonomously into an upright posi-
are winding monochasia derived from the distal tion when mature, but this process can also be
dichasium. By a shortening of the internodes, triggered by manipulation of the nectar scale in
rosulate species with apparently axillary flowers many species (Wittmann and Schlindwein 1995;
have evolved from these monochasial taxa in Caio- Schlindwein and Wittmann 1997). More or less
242 M. Weigend
campanulate corollas with red or orange, carnose, KARYOLOGY. Chromosome counts are available
unclawed petals and erect stamens have evolved in for the majority of genera and some of these are
hummingbird-pollinated groups. The staminodial strongly supported by karyology. Gronovioideae
complexes are here robust and provide a mechan- have 2n = 26 (Cevallia) or 2n = 74 (Gronovia).
ical guide to the copiously produced nectar. Schis- Petalonychoideae (Petalonyx) have 2n = 44, 46
mocarpus and some species of Mentzelia (M. sect. (Davis and Thompson 1967). Base numbers of
Mentzelia) show distinct heteranthery. these two groups are unknown and all of the
Ovary position is completely inferior to largely genera may represent palaeopolyploids. Mentze-
superior, both conditions are sometimes found in lioideae seem to have a base number of x = 7. From
very closely related taxa. The parietal placentae in Mentzelia the numbers 2n = 18, 20, 22, 28, 36, 54,
Loasoideae provide good characters for generic 72 have been reported (Hill 1976), the apparent
delimitation: Most taxa have placentae which are tetraploid 2n = 28 being the most frequent one.
either flat or globose in transverse section but in Numbers over n = 14 are exclusively from weedy,
Caiophora they are Y-shaped with ovules sitting annual Mentzelia sect. Trachyphytum (Hill1976).
only on the morphologically adaxial side of the Eucnide is apparently strictly hexaploid with 2n =
placenta, and in Presliophytum and Huidobria they 42 (Thompson and Ernst 1967). Loasoideae seem
are cross-shaped in section and have ovules all to have the base number x = 6. Loasa malesher-
over their surface. bioides has the lowest chromosome number so far
known from the family, 2n = 12 (Grau 1988), and
EMBRYOLOGY. The only detailed studies on it is possibly the only diploid, all other taxa being
embryology available are from Loasa and Blumen- polyploids. The tetraploid chromosome number
bachia (Garcia 1962): the ovules are unitegmic 2n = 24 is widely found (Blumenbachia, Aosa,
and tenuinucellate, with very distinctive micro- Presliophytum, Kissenia, Klaprothia, Xylopodia;
pylar and chalaza! haustoria and cellular endo- Coleman and Smith 1969; Poston and Thompson
sperm. This type is widespread in Loasoideae 1977; pers. obs.). Higher ploidy levels are encoun-
(Loasa, Nasa, Blumenbachia, Presliophytum, tered in Klaprothia (Poston and Thompson 1977)
Caiophora). Mentzelia has basically similar ovules and Plakothira (pers. obs., octaploid with 2n = 48),
(Vijayaraghavan and Prabhakar 1984). The embry- Nasa (octaploid with 2n =56, N. dyeri, pers. obs.),
ology of Schismocarpus and Eucnide is still Loasa (2n = 36, L. triloba, Grau 1988) and Huido-
unknown. Gronovia and Petalonyx have surficially bria (hexaploid with 2n = 36, Grau 1997). Most
very distinctive ovules which are probably crassin- species of Loasa have 2n = 24,26 (Grau 1988) and
ucellar and certainly lack well-developed micropy- most of the genus Nasa 2n = 28 (pers. obs.). In all
lar and chalaza! haustoria. Petalonyx thurberi (and groups of Loasaceae, polyploidization and dys-
maybe other Petalonyx) ovules have an obturator. ploid changes thus seem to have been fairly
More data and a critical anatomical study of the common events.
embryology of Loasaceae are evidently required.
PoLLINATION. Loasaceae are animal- or rarely
POLLEN MORPHOLOGY. Pollen morphology is self-pollinated. The reward in polystemonous
comparatively uniform in Loasaceae. The pollen Loasaceae is usually pollen and nectar, but some
grains are usually spherical to prolate and tricol- taxa (e.g. Mentzelia) lack nectaries and offer
pate or tricolporate. The exine can be punctate and pollen as the only reward (Thompson and Ernst
spinose, echinate or rugulose (Gronovioideae) or 1967). A floral odour is usually not perceptible but
striate with thin lirae (Petalonyx, Poston and has been reported from the nectariferous flowers
Nowicke 1993). In Mentzelia, Schismocarpus and of Mentzelia decapetala (Keeler 1981) and Nasa
Eucnide the pollen is striate to striate-reticulate ferruginea (pers. obs.). Facultative autogamy is
with exclusively longitudinal striations. The pollen widespread in annuals and has been reported
in Loasoideae is usually reticulate, but sometimes from Mentzelia, Eucnide (Thompson and Ernst
equatorially striate (Loasa ser. Floribundae and 1967), Petalonyx (Davis and Thompson 1967),
Deserticolae). The colpus margin may be differen- Gronovia, Klaprothia, Nasa, Blumenbachia and
tiated or undifferentiated. Kissenia has syncolpate Loasa. Cleistogamy is known from Loasa triloba
pollen grains with microreticulate exine. The rain- (Gilg 1925) and has now also been observed in
forest genus Chichicaste is the only taxon with tri- Nasa chenopodiifolia. In Eucnide hawkmoths and
colporoidate pollen with conspicuous stoppers, a bees have been reported as pollinators, and polli-
feature that has evolved convergently in many nation by colibris has been suggested for some
plant species from perhumid habitats. taxa (Thompson and Ernst 1967). Klaprothia is
Loasaceae 243
only visited by Empid flies, whereas Presliophytum papillae - one of the very few morphological
is visited by a very wide range of pollinators characters separating these two genera (Hill1976;
including Lepidoptera and Hymenoptera (Euglos- Hufford 1988).
sine and Colletid bees, Xylocopa, Apis, Bombus ), all Corner (1976) interpreted the seeds as neotenic
of which are likely to effect pollination. Many taxa enlarged ovules with an exotesta often character-
of Loasoideae share pendent flowers with spread- istic of the genera. For example, in Loasa he found
ing petals and strongly contrasting nectar scales the exotesta as a palisade of cells with strongly
and thigmonasty (stamen movement triggered thickened outer walls, and in Eucnide as tabular
by manipulation of the nectar scale in Caiophora, polygonal cells with thick cuticle.
Loasa, Blumenbachia, Aosa, Nasa, Xylopodia) . Seeds are usually barochoric, i.e. they have no
These are primarily pollinated by specialized Col- obvious means of dispersal. In Caiophora the
letid bees (Schlindwein and Wittmann 1997). Col- seeds remain trapped in hanging fruits with lon-
letids are the only regular visitors of these flowers. gitudinal slits which only release them during dry
Many high Andean members of Nasa and Caio- and windy conditions. Anomochory is the dis-
phora have large, orange corollas with large nectar persal mechanism of Caiophora, Scyphanthus and
scales and copious nectar production. These are many Loasa (seeds very light with a deeply pitted
visited and pollinated by humming birds and surface) and Presliophytum (seeds very small).
bumble bees. Caiophora coronata has bowl-shaped Epizoochory is found in Blumenbachia, Klaprothia
flowers close to the ground and is pollinated by mentzelioides and Mentzelia aspera: fruits are
small rodents (Cocucci and Sersic 1997). tardily deshiscent and densely covered with hook-
shaped trichomes. Developing fruits of some
FRUIT, SEED AND DISPERSAL. Loasaceae have Eucnide are negatively phototropic and grow into
capsular fruits, with the only exception of rock crevices where they dehisce and shed their
Gronovioideae and Petalonychoideae which have seeds (Thompson and Ernst 1967). The seeds of
cypselas. The capsules are typically many-seeded Blumenbachia sect. Gripidea have two air-filled
and dehiscent and the seeds have a dark brown to wings which considerably slow down the speed
nearly black testa. of the falling seed in the air but they also lend
Most Loasoideae have a reticulate testa; a group buoyancy to the seed when it falls into water and
of closely allied genera from southern South may thus represent an adaptation to hydrochory
America (Caiophora, Loasa, Scyphanthus) shares or/and anemochory. The seeds of Loasa ser.
very high, fenestrate anticlinal walls (Fig. 82). Nasa Macrospermae are large, round and heavy. They
has low, apically thickened anticlinal walls. The roll down between the rock debris (scree slope
seeds of Eucnide are striate whereas the seeds of habitat), thus reaching the moister lower layers.
Mentzelia are reticulate or polyhedral with central There they germinate and have enough stored
assimilates to produce a long hypocotyl that
carries them back to the light. Nasa lacks obvious
dispersal mechanisms, apart from a pronounced
elongation of the pedicel (in taxa with erect, cylin-
drical capsules) so that the seeds are sprinkled out
by strong wind in dry weather. Other species
(many Nasa ser. Grandiflorae) have horizontal
capsules opening with apical valves and a longitu-
dinal slit, thus forming an open platform on which
the seeds are exposed to rain and wind. Anemo-
chory of the entire fruit is found in Gronovioideae
(accrescent calyx) and Petalonyx (fruit remains
attached to bract and prophylls).
PHYTOCHEMISTRY. The phytochemistry of Loas-

aceae is poorly known, with the only exception
of iridoids, on which extensive data are available
(Hegnauer 1989; Mi.iller et al. 1999; Weigend
et al. 2000; Rodriguez et al. 2002). Iridoids and
Fig. 82. Loasaceae. Caiophora madrequisa, deeply pitted seed secoiridoids are nearly universally present in
with thin anticlinal cell walls, SEM (x40). Photo M. Weigend Loasaceae and only very few taxa have no or very
244 M. Weigend
low concentrations of these compounds. Some four lineages, representing the four subfamilies
monomeric iridoids are widespread throughout Gronovioideae, Petalonychoideae, Mentzelioideae
the group, e.g. sweroside, 8-epi-kingiside and and Loasoideae (Weigend 1997), but indicating a
loganin, loganic acid. Others are restricted to basal position of Loasoideae rather than Petalony-
certain groups: tricoloroside and similar hetero- choideae. This is congruent with morphological
oligomeric compounds are found only in some data: Loasoideae show the closest similarity to the
taxa of Loasa, 10-hydro:xyoleoside dimethylester sister group of Loasaceae (Hydrangeaceae). Both
is widespread in Caiophora and has not been groups share predominantly opposite phyllotaxy,
reported from any other taxon in the family. reticulate testa, parallel stigmatic lobes, pluriovu-
Loasaceae contain relatively small amounts of phe- late ovaries, polystemonous androecia, petals
nolic compounds such as quercetin and caffeic with numerous veins from the base, and annular
acid (Hegnauer 1966); cyanogenic compounds and nectaries. Loasoideae contain approximately 200
alkaloids have not been reported from the family. species in 13 genera. Mentzelioideae are recog-
The chemical composition of the seeds has been nized as a natural group of genera (comprising
investigated for only very few taxa (three species of Mentzelia, Schismocarpus and Eucnide) which is
Mentzelia) and these contained ca. 20% storage difficult to define morphologically, apart from the
protein and 34-43% fat (oleic, linolic and palmitic alternate phyllotaxy and lack of specialized sta-
acids, Hegnauer 1966). Many other interesting phy- minodial complexes. A basal position of Eucnide
tochemical questions have not yet been addressed, in Loasaceae has recently been proposed (Hufford
e.g. the identity of the irritant substances in the 1998), but morphological support for this is
stinging hairs or the type of floral pigments. weak and a more detailed molecular analysis
will be required to sort out the exact relationships
DISTRIBUTION AND HABITATS. Loasaceae are between the subfamilies of Loasaceae. Both
largely restricted to the Americas, ranging form Mentzelioideae and Loasoideae have retained
the northern temperate zone to the southern tem- various plesiomorphic characters such as pluri-
perate zone. They are absent from northern tem- ovulate ovaries, polystemonous androecia, petals
perate forests, but are found in the Mediterranean with numerous veins from the base, dilated fila-
regions of both hemispheres, in open vegetation ments and annular nectaries. However, these ple-
such as prairie, pampa, semi-desert, desert, lorna, siomorphic characters can not be used to define a
and in forest vegetation such as rain-green forests, core family Loasaceae, from which the haploste-
rainforests, cloud forests, and southern temperate monous subfamilies Gronovioideae and Petalony-
forests. The family extends from sea level up to the choideae are excluded, or united in a different
subparamo (ca. 3500m) and puna (ca. 4800m) of subfamily (Takhtajan 1997). Mentzelioideae and
the Andes. The highest concentration of taxa and Loasoideae share only plesiomorphic characters,
the most narrowly endemic taxa are found in the and the haplostemonous subfamilies seem to have
largely cloud forest-dwelling genus Nasa between arisen from a common ancestor with Mentze-
elevations of 1500 and 3500m in the northern and lioideae (Xiang 1999; Moody et al. 2001). Apart
central Andes. Peru is the most important centre from haplostemonous androecia, they share uni-
of diversity for the family, with at least 80 species ovulate ovaries, petals with a single vein from the
in seven genera (mostly subfamily Loasoideae), base, and cup-shaped nectaries but, in anther and
two genera and most species being endemics. fruit morphology, perianth structure, and pollen
Chile and Mexico, each with approximately 40 surface, they are very distinct from each other.
species and seven genera, are secondary centres of Within subfamily Loasoideae only two tribes
diversity. Loasoideae are mainly South American are accepted here - tetramerous Klaprothieae and
and extend north into southern Mexico. This sub- penta- to octamerous Loaseae. Kissenieae Urb. &
family has outliers in Africa (Kissenia) and Poly- Gilg {1900)) can not reasonably be treated as a
nesia (Plakothira). Mentzelioideae, Gronovioideae group distinct from Loaseae, and are here reduced
and Petalonychoideae have their centre of diver- under that tribe. Klaprothieae (6 species) are very
sity in Mexico and the south-western U.S.A., and clearly defined by a variety of characters, such as
only the former two reach South America with one the lack of urticant setae, strictly opposite phyl-
genus each (Mentzelia and Gronovia respectively). lotaxy, and longitudinal lamellae on the petals.
Loaseae can not be similarly defined by derived
SuBDIVISION. The relationships within the family characters. They comprise ca. 196 species and are
are now well understood and the molecular ecologically and morphologically the most diverse
data presented by Xiang et al. {1998) recognize group within the family. Loaseae can be informally
Loasaceae 245
divided into "Lower Loaseae" with erect flowers examination of their morphology reveals that they
with white (rarely yellowish or greenish white) also display the closest morphological similarity
petals and pale (white, greenish or yellowish to Loasaceae: characteristic "Loasaceae" trichomes
white) nectar scales and without thigmonasty (Behnke and Barthlott 1983) are found in ]amesia
(eight species in the genera Huidobria, Presliophy- (scabrid) and Fendlera (glochidiate). In Loasaceae
tum, Chichicaste, Kissenia). These "Lower Loaseae" and Fendlera they cover even petals, style and
include all taxa without specialized pollination by anthers. Filaments (esp. the first, antesepalous one)
Colletid bees or ornithophily. They are a grade, are dilated Uamesia) to forked (Fendlera): this is a
not a clade. The "Higher Loaseae" typically have character which is widespread in, and highly char-
pendent flowers, brightly coloured petals, vari- acteristic of Loasaceae, and it gave rise to both the
ously elaborated and coloured nectar scales and simple staminodes of Mentzelioideae and the
show thigmonasty (ca. 188 species in the genera staminodial complexes of Loasoideae (Weigend
Aosa, Blumenbachia, Caiophora, Loasa, Scyphan- 1997). The seeds of ]amesia and Fendlera and all
thus, Nasa) and are pollinated by specialized Col- primitive members of Loasoideae have a straight
letid bees or hummingbirds. The "Higher Loaseae" embryo surrounded by fleshy endosperm (Takhta-
are probably monophyletic and fall into three, jan 1997) and a dark brown, reticulate testa with
probably natural groups: the eastern Brazilian/ slightly elevated anticlinal walls. The basic pat-
Caribbean genus Aosa (6 spp.), the large North/ terns of centripetal stamen initiation are also
Central Andean genus Nasa (predominantly identical between Loasaceae and Hydrangeaceae
tropical montane in mesic habitats), and the (Payer 1853; Leins and Winhard 1973; Hufford
South Andean group comprising Blumenbachia, 1998). Other characters uniting at least some
Caiophora, Loasa and Scyphanthus (high Andean, Hydrangeaceae with Loasaceae are fruit morphol-
and south temperate, in Mediterranean and desert ogy (semi-superior, septicidal capsules opening
habitats). with apical valves in both ]amesia/Fendlera and
Within subfamily Mentzelioideae two taxa most Loasaceae), petal morphology (clawed,
were recognized by Gilg (1925): Eucnideae with boat-shaped petals), style and stigma morphology
Eucnide, and Mentzelieae with Mentzelia and (style short, parallel stigmatic branches in
Schismocarpus. The relationships between these Fendlera and Loasaceae), stem filled with white
tribes are still unclear and no subdivision of sub- pith, bark exfoliating, flower typically 4-5-merous,
family Mentzelioideae is proposed here. aestivation contort or imbricate, androecium
mostly polystemonous in both groups, ovary
AFFINITIES. Until very recently the affinities of unilocular, typically of 3-5 carpels, with intruding,
Loasaceae were highly controversial. Traditionally parietal placentae, embryology (unitegmic, ten-
the family had been placed near Turneraceae and uinucellate ovules, cellular endosperm formation,
Passifloraceae (Cronquist 1981; Weigend 1997) endosperm haustoria; Takhtajan 1980), pollen typ-
or even united with Turneraceae in one family ically prolate to spherical, tricolpate to tricolpo-
(Humboldt, Bonpland and Kunth 1823). The rate, inflorescence morphology (thyrsoids with a
superficial similarity between the Turneraceae distal dichasium in Fendlera and Fendlerella and
and some Loasaceae is considerable, but refers most Loasaceae), presence of hydathode teeth on
only to characters which are quite variable within the leaves (e.g. in Philadelphus, ]amesia and in all
Loasaceae themselves (Gilg 1925). Phytochemistry Loasaceae), phytochemistry (very similar iridoid
and embryology indicated that the affinities of spectra; Takhtajan 1980; Hegnauer 1989), and the
Loasaceae lie elsewhere, and Takhtajan (1980) was occurrence of raphides or oxalate druses. A close
the first to propose a placement close to Corniflo- relationship between Cornales and Loasaceae is
rae as a separate superorder Loasiflorae. Recent also congruent with data from wood anatomy
molecular studies confirmed a relationship to Cor- (Carlquist 1992).
nales (Hempel et al. 1995, Xiang et al. 1998, Xiang On the basis of the phylogenetic hypothesis
1999), and the Loasaceae are now known to be adopted here, subfamily Loasoideae appear to
sister to the Hydrangeaceae (s.l., incl. Philadel- have retained a proportionately larger number of
phus, ]amesia, Fendlera, Fendlerella, Deutzia plesiomorphic character states than the other
and allies). A taxonomically isolated position of three subfamilies. A basal position of Eucnide
Loasaceae (order or superorder), as repeatedly (in Mentzelioideae; Hufford 1990) is currently
proposed, is not justified. The genera ]amesia and supported by neither published molecular data
Fendlera are the most basal branch of the (Xiang 1999) nor morphology. Gronovioideae and
Hydrangeaceae clade in Xiang (1999), and a close Petalonychoideae seem to represent the most
246 M. Weigend
derived clade in the family (haplostemonous - Flowers in cymoids or thyrsoids; calyx lobes persistent;
flowers, one-seeded, inferior cypsela, exclusively mature fruit free of bracts and prophylls. Gronovioideae
3
alternate phyllotaxy, testa reduced, petals one- 3. Erect subshrub with narrow, sinuate-lobate leaves; calyx
veined) and are clearly sister groups (Weigend lobes and petals linear, isomorphic; connective protracted
1997; Moody et al. 2001). The family Loasaceae as into long appendage 19. Cevallia
such is strongly supported as monophyletic by its - Liana or climber with cordate to subpalmately lobed
general morphological coherence (Weigend 1997) leaves; calyx lobes spatulate; petals lanceolate; connective
not differentiated 4
and by molecular markers (Hempel et al. 1995; 4. Leaves cordate, margin entire; inflorescence branches
Xiang 1999), but it can not be defined by autapo- dichasial; petals laciniate 18. Fuertesia
morphic characters, since the "typical" trichomes - Leaves subpalmately lobed; inflorescence branches mono-
are now known to be present also in hydrangea- chasial; petals entire 17. Gronovia
ceous genera such as Fendlera. Characters pre- 5. Petals fiat and membranaceous; staminodia, if present, not
in antesepalous groups. Mentzelioideae 6
sent in Loasaceae and altogether absent from - Petals deeply boat-shaped or carnose if fiat; staminodia
Hydrangeaceae are the stinging hairs and pro- always present, in antesepalous groups. Loasoideae 8
nounced recaulescence and concaulescence in 6. Flowers obdiplostemonous with 5 large and 5 small
the inflorescence, but there are some Loasaceae anthers; xylopodium present 16. Schismocarpus
- Flowers polystemonous, if stamens few then all the same
(e.g. Mentzelia sect. Mentzelia) which lack both. size; xylopodium absent 7
7. Testa tuberculate; stinging hairs absent; petals always free
EcoNOMIC UsEs. No Loasaceae are currently cul- 14. Mentzelia
tivated for economic purposes, but many taxa - Testa striate; stinging hairs usually present; petals free or
are collected from the wild and play some role in united IS. Eucnide
8. Flowers tetramerous; petals with longitudinal lamellae;
folk medicine: species of Caiophora and Nasa are urticant hairs always absent. Klaprothieae 9
used to treat a variety of disorders (e.g. allergies, - Flowers penta- to octamerous; petals without longitudinal
bronchial diseases, and liver complaints) in lamellae; urticant hairs often present. Loaseae 11
Andean South America. Species of Mentzelia are 9. Lamina with two lobes on each side, margin serrate; petals
used against stomach complaints in South green, narrowly obovate; erect shrub from underground
xylopodium 13. Xylopodia
America and exported to Europe for that purpose. - Lamina entire, margin serrate; petals usually white, widely
The seeds of Mentzelia used to be an important ovate if pale green; erect annual or perennial herbs or
food source for Native Americans in western shrubs without xylopodium 10
North America, and seeds and other parts of the 10. Capsule straight, dehiscent, with >20 seeds; erect perennial
plants were used in medicinal preparations herbs or shrubs 12. Plakothira
- Capsule straight or twisted, if twisted seeds <15; erect
(Moerman 1998). annual or decumbent perennial herbs 11. Klaprothia
11. Flowers erect; petals white, cream or greenish white (never
CONSPECTUS OF LOASACEAE with serrate margin); nectar scales pale yellow, white or
cream coloured, fiat, dorsal threads filiform if present
I. Subfamily Loasoideae Urb. & Gilg (1900).
12
1. Tribe Loaseae Urb. & Gilg (1900).
- Flowers usually pendent, very rarely erect (petals always
Genera 1-10.
with deeply serrate margin if flowers erect); petals and/or
2. Tribe Klaprothieae Urb. & Gilg (1900). nectar scales brightly coloured, scales usually red and
Genera 11-13.
yellow or green and with various morphological elabora-
II. Subfamily Mentzelioideae (Rchb.) Urb. & Gilg (1900). tions such as dorsal calli, wings, or a double arch, dorsal
Genera 14-16. threads often dilated if present 15
12. Nectar scales of more than 3 staminodia, 4-5 free stamin-
III. Subfamily Gronovioideae (Rchb.) Link (1833). odia present; plant without urticant setae 3. Huidobria
Genera 17-19. - Nectar scales of 3 staminodia, 2 free staminodia; plant with
IV. Subfamily Petalonychoideae Weigend ( 1997) or without urticant setae 13
Genus 20. 13. Calyx lobes much larger than petals; nectar scales >Sx as
long as wide; desert shrub without urticant setae
2. Kissenia
KEY TO THE GENERA
- Calyx lobes smaller than petals, nectar scales as long as
1. Flowers haplostemonous; petals with 1 principal vein, wide or up to 2x as long as wide; plants with urticant
membranaceous; ovary with 1 pendent ovule; seed without setae 14
dark testa 2 14. Erect desert shrubs with leaves up to 5 em in diam.; petals
- Flowers polystemonous or obdiplostemonous; petals with with long claw, narrowly ovate; nectar scales with dorsal
3-5 principal veins; ovary with 1-many ovules on parietal threads much exceeding the scale neck in length
placentae; testa dark 5 4. Presliophytum
2. Flowers in racemes; calyx lobes caducous; bracts - Tall rainforest herb (to 4m) with leaves up to 40cm long;
and prophylls remaining attached to mature fruit. petals with short claw, subcircular to widely ovate; nectar
Petalonychoideae 20. Petalonyx scales without threads 1. Chichicaste
Loasaceae 247
15. Inflorescence ebracteose; testa tuberculate 5. Aosa 3-7 apical valves and/or 1-7longitudinal sutures,
- Inflorescence bracteose or frondose; testa deeply pitted, very rarely indehiscent, straight or narrowly
irregularly fibrous, reticulate or rugulate, never
tuberculate 16 twisted; seeds usually numerous; testa mostly
16. Every flower on the branches of the inflorescence with one reticulate, pale brown to black.
bract only (primary flower usually with two); nectar scales
with conspicuous dorsal sacs and apical wings, often with
dorsal callus 10. Nasa 1. TRIBE LOASEAE Urb. & Gilg (1900).
- Every flower of the inflorescence with two bracts; nectar
scales rectangular or with double arch and/or flags but
never with conspicuous dorsal sacs or apical wings or Leaves alternate or opposite; stinging hairs mostly
dorsal calli 17 present. Flowers 5( -8)-merous, mostly pendent;
17. Fruits always twisted anticlockwise; seeds winged or with petals lacking longitudinal lamellae, yellow,
fibrous testa or angular, but never deeply pitted with fen-
estrate anticlinal walls nor rugulose; nectar scales rectan-
orange, red, white, very rarely green; staminodial
gular, equalling the free staminodia in size complexes always present, outer staminodes
6. Blumenbachia united to form a nectar scale. Capsule usually with
- Fruits straight or twisted, if twisted fruits twisted clock- 10 prominent veins on the outside.
wise and anticlockwise alternating in the inflorescence;
testa deeply pitted with fenestrate anticlinal walls, very
rarely with irregularly rugulose testa; nectar scales with 1. Chichicaste Weigend
double arch and often with flags, or nectar scales reduced
in size (<<than free staminodia) 18 Chichicaste Weigend, Nasa and the Conquest of South
18. Fruits completely inferior, narrowly cylindrical, straight, America: 215 (1997).
more than lOx as long as wide 9. Scyphanthus
- Fruits partially superior to completely inferior, cylindrical, Erect, sparsely branched, short-lived herb up to
ovoidal, clavate or globose, never more than 7X as long as
wide, often twisted 19 4 m tall, with stinging hairs. Leaves opposite below,
19. Fruits straight, opening with apical valves 7. Loasa alternate above; lamina ovate with shallowly lobed
- Fruits straight or twisted, opening with longitudinal slits and serrate margin, up to 40 em long. Inflores-
(very rarely: rosette herb with oblong, cream-coloure~ cences terminal thyrsoids with monochasial or
petals, and flowers singly on ebracteose peduncles If rarely dichasial paracladia; each flower with 2
capsule opening with apical valves) 8. Caiophora
small prophylls; flower erect; petals cream white to
green; nectar scale white, apex forming 3-4 trian-
gular lobes. Capsule 1/3 superior, subglobose,
I. SUBFAMILY LOASOIDEAE Urb. & Gilg (1900).
opening with 5 apical valves; testa reticulate. Only
one species, Chichicaste grandis (Standi.) Weigend
Leaves opposite below and alternate above or
from the lowland rainforests of NW Colombia to
opposite throughout; stinging hairs sometimes
Costa Rica.
present. Inflorescences thyrsoids, often with
numerous paracladia and strong metatopies, or
dichasia, rarely reduced to few-flowered monocha- 2. Kissenia Endl.
sia; flowers pendent or erect, 4-8-merous; petals
Kissenia End!., Gen. Pl. suppl.: 76 (1842), (sphalm. Fissenia
free, mostly clawed, either deeply boat-shaped or Endl.).
carnose (never flat and membranaceous), or some-
times with longitudinallamelliform flaps, margin Erect, densely branched shrubs without stinging
entire or serrate, with >3 principal veins from hairs. Dominant, carnose tap-root present. Leaves
base; polystemonous, fertile stamens usually in opposite below, alternate above; lamina ovate
antepetalous position, filaments filiform; anthers to reniform with shallowly lobed and crenate
latrorse; staminodia in antesepalous groups of margin. Inflorescences terminal dichasia or thyr-
3-11, free or united with outer staminodia united soids with 1-3 paracladia; each flower with 2
to form a nectar scale (typically outer 3 united small, entire prophylls; flowers erect, numerous;
and inner 2 free), scales variously decorated petals cream coloured, much shorter than calyx
with calli, filiform appendages and/or saccate lobes; nectar scale oblong, yellowish; ovary largely
thickenings, sometimes winged apically, white, inferior, placentae reduced with few ovules.
red, yellow, usually with colour patterns; free sta- Capsule inferior, indehiscent, crowned with con-
minodia usually S-shaped, papillose; parietal pla- spicuously accrescent calyx lobes; seeds 1-2, irreg-
centae simple or divided, usually with numerous ularly ovoidal; testa poorly developed. 2n = 24. Two
ovules. Capsule terete, often with 10 prominent species from arid SW and NE Africa and the
veins on the outside, calyx persistent, opening with Arabian Peninsula.
248 M. Weigend
3. Huidobria Gay green and brown, or red and yellow, with 3 dorsal
Huidobria Gay, Fl. Chil. 2: 440 (1847); Grau, Sendtnera 4: 77-93
filaments, scale apex forming 3 lobules, or entire
( 1997), rev. and recurved. Capsule inferior to 3/4 superior,
s~bglobos~ to clavate, sometimes curved, opening
Erect, densely branched annual herbs or shrubs w1th 5 ap1cal valves; testa reticulate, epidermis
without stinging hairs. Dominant, carnose tap- cells longer than wide (210 x 60Jlm), at their cha-
root present. Leaves opposite below, alternate laza! pole with crescent-shaped, rugulose bulge.
above; lamina linear with entire margin or ovate 2n = 24. Seven spp. from E Brazil (6 spp.) and
and with shallowly lobed and crenate margin. Hispaniola (1 sp.).
Inflorescences complex asymmetrical dichasia
(fl~wers apparently irregularly alternating with 6. Blumenbachia Schrad.
foliage leaves); each flower with 2 frondose pro-
phylls; flowers erect; petals white to cream; nectar Blumenbachia Schrad., Goett. Gel. Anz. 3/171: 1706 (1805);
Weigend, Sendtnera 4: 202-220 (1997), rev.
scale formed from 5-7 outer staminodia, white,
with 5-7 long dorsal appendages, inner 3-5 sta-
minodia free, white; placentae deeply divided into Scandent or ascending, annual or perennial herbs,
3 lamellae. Capsule largely inferior, subglobose, with stinging hairs. Root system fibrous, rarely
opening with 4-5 apical valves; testa striate or primary root developing into root tuber or thick-
e~ed tap-root. Leaves opposite; lamina usually
reticulate. 2n = 36. Two spp., deserticolous shrubs
~1dely ovate, subpalmately lobed, sometimes bip-
from Chile.
l~natl~ect. Inflorescences terminal thrysoids with
d1ehas1al or monochasial branches (sect. Angu-
4. Presliophytum (Urb. & Gilg) Weigend latae), or dichasia (sect. Gripidea) or borne singly
Presliophytum (Urb. & Gilg) Weigend, Nasa and the Conquest
(sect. Blumenbachia ); each flowers with 2 frondose
of South America: 215 (1997). or bracteose prophylls; petals white; nectar scale
with 3 long, dorsal, filiform appendages, white, red
Erect, densely branched shrubs with stinging and yellow; placentae simple. Capsule inferior,
hairs. Dominant, carnose tap-root present. Leaves cylindrical to globose, twisted anticlockwise only,
opposite below, often alternate above, lamina septicidal or septifragous, apex coherent; seeds
ovate to reniform with shallowly lobed and crenate angular; testa polyhedral with irregular lateral
margin. Inflorescences complex, asymmetrical wings (sect. Angulatae), or ovoidal and testa
dichasia; each flower with 2 frondose prophylls fibrous with narrow epidermis cells (sect. Blumen-
(flowers apparently irregularly alternating with bachia), or widely reticulate, with 1-2 terminal
foliage leaves); erect; petals white; nectar scale win~s (sect. Gripidea). 2n = 24, 26. Twelve spp.,
white, with 3 long dorsal appendages; placentae Brazil, Uruguay, Paraguay, Argentina, Chile.
deeply divided into 3 lamellae. Capsule inferior,
subglobose, opening with 4-5 apical valves; testa 7. Loasa Adans.
foveate-reticulate. 2n = 24. Three species from the
desert of western Peru (below 2000 m), on rocky Loasa Adans., Fam. Pl. 2: 501 (1763); Urban & Gilg, Nova Acta
Acad. Caes. Leop.-Carol. German. Nat. Cur. 76 (1900), rev.;
slopes. Loasa longiseta Phil. from N Chile proba- Sleumer, Bot. Jahrb. Syst. 76: 411-462 (1955), rev.
bly also belongs into this genus, differing form the
other members only in fruit and seed morphology. Rosulate, erect or scandent annual or perennial
rarely small shrubs, stinging hairs present.
he~bs,
5. Aosa Weigend Pnmary root dominant, thin, or forming a tap-
root or thickened to a root tuber, sometimes rhi-
Aosa Weigend, Nasa and the Conquest of South America: 214 zomatose. Leaves
(1997).
opposite throughout or opposite
below and alternate above; lamina most often
ovate, pinnatifid or bipinnatisect, sometimes
Annual or perennial herbs or shrubs, with sting-
widely ovate or subcircular and subpalmately
ing hairs. Leaves opposite below, often spirally
inserted above; lamina oblong, ovate to suborbi- lobed or ternate. Inflorescences terminal thyrsoids
cular, lobed or entire, margin crenate or serrate. or dichasia, rarely flowers single; each flower with
2 frondose prophylls; flowers pendent, or erect in
Inflorescence complex terminal thyrsoids with
some rosulate species; petals white, yellow or red;
monochasial or rarely dichasial paracladia; pro-
nectar scale with (rarely without) 3 long, often
phylls absent; petals cream to green; nectar scale
Loasaceae 249
flag-shaped dorsal appendages and decorated with with 3 flag-shaped dorsal appendages and double
a double arch on back, usually white, red and arch, white, red and yellow. Capsule inferior, nar-
yellow. Capsule cylindrical to subglobose, opening rowly cylindrical, opening with longitudinal slits;
with 3-5 apical valves; seeds ovoidal; testa deeply seeds angular; testa deeply pitted with fenestrate
pitted, anticlinal walls fenestrate, rarely seeds anticlinal walls. One or two spp., Mediterranean
large, globose and testa rugose. 2n = 12, 24, 26, 36. scrub lands in Chile, very close to Caiophora and
Ca. 36 species, mostly Chile and adjacent to Loasa ser. Pinnatae.
Argentina, one species ranging into coastal Peru.
Loasa is very closely related to Caiophora,
10. Nasa Weigend Fig. 83
Scyphanthus and Blumenbachia and is likely to be
paraphyletic. On the other hand, Loasa malesher- Nasa Weigend, Nasa and the Conquest of South America: 214
bioides Phil. from Chile and Argentina is aberrant (1997); Dostert and Weigend, Harvard Papers Bot. 4:
in this generic group and should probably be seg- 439-468 (1999), rev.; Weigend, Arnaldoa 5: 159-170 (1998),
rev.; Weigend, Nordic J. Bot. 20: 15-31 (1999), rev.; Weigend,
regated from Loasa. Flora of Ecuador 64: 1-90 (2000), rev.
8. Caiophora K. Presl Herbs, subshrubs or shrubs, 5-400 em, with sting-

ing hairs. Mature plant only with adventitious
Caiophora K. Presl., Reliq. Haenk. 2:41, tab. 42 (1836) (sphalm. from the decumbent basal stem portion (primary
Cajophora); Weigend, Sendtnera 4: 221-242 (1997), rev.
root evanescent). Leaves opposite or alternate;
lamina ovate to orbicular, lobulate, palmate
Rosulate, erect or scandent perennial herbs with
or pinnatifid to bipinnate, sometimes peltate.
stinging hairs. Primary root dominant, sometimes
Inflorescences terminal thyrsoids, dichasia or
thickened, or primary and/or secondary roots
monochasia, each flower with one recaulescent
tuberous, often rhizomatose. Leaves opposite;
bract; petals white, yellow, orange or red, often
lamina mostly ovate, pinnatifid or bipinnatisect,
bicoloured; nectar scales with 0-3 calli on back
sometimes ternate. Inflorescences terminal sym-
and/or apical wings and/or 2 dorsal nectar sacs;
metrical or highly asymmetrical thyrsoids or
staminodia L-shaped, sometimes with basal
dichasia, rarely flowers borne singly from basal
appendages; placentae simple. Capsule cylindrical
rosette; each flower with 2 frondose prophylls;
to globose, usually opening with 3-5 apical valves;
flowers mostly pendent, 5-8-merous; petals green,
seeds ovoidal, globose or slightly angular; testa
white, yellow or red; nectar scale usually with 3
shallowly reticulate with wide anticlinal walls.
long, often flag-shaped dorsal appendages and
2n = 28, 56. Ca. 100 spp., most in Colombia,
decorated with a double arch on back (occasion-
Ecuador and Peru, few in Chile, Bolivia, Venezuela
ally dorsal appendages lost, arches poorly differ-
and C America (to S Mexico). Most diverse in
entiated), usually white, red, yellow, or green; free
cloud forests, some species weeds in cultivated
staminodia sometimes with basal appendages;
lands, a few found in semi-deserts, rain-green
placentae Y-shaped, ovules only the morphologi-
coastal forests, rainforests, puna, subparamo or
cally adaxial side. Capsule mostly inferior,
paramo.
subglobose to narrowly cylindrical, frequently
antidromously twisted, usually opening with 3-5
longitudinal slits, rarely opening with 3 apical
2. TRIBE KLAPROTHIEAE Urb. & Gilg (1900).
valves; seeds angular; testa deeply pitted with fen-
estrate anticlinal walls. 2n = 14, 16. Ca. 56 spp.,
Leaves opposite; stinging hairs always absent.
Peru, Bolivia, Argentina, Chile, one species each in
Flowers 4-merous, mostly erect; petals each
Uruguay, Brazil and Ecuador.
with 2 longitudinal lamellae, white or green; sta-
minodial complex with outer staminodia free or
9. Scyphanthus D. Don rarely united to form a nectar scale; placentae
simple. Capsule without prominent veins on the
Scyphanthus D. Don in Sweet, Brit. Fl. Gard. I, 3: tab.238 (1828).
outside.
Scandent annual herbs, without stinging hairs.
Leaves opposite; lamina widely ovate, bipinnati- 11. Klaprothia Kunth. Fig. 81
sect. Root system fibrous. Inflorescences terminal,
Klaprothia Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp. 6:
highly asymmetrical dichasia; each flower with 2 96 (1823); Poston & Nowicke, Syst. Bot. 15: 671-677 (1990),
frondose prophylls; flowers yellow; nectar scale rev.
250 M. Weigend
Capsule inferior, ovoidal and straight to slightly

twisted and tardily dehiscent with apical valves, or
clavate and twisted and opening with longitudinal
slits; seeds narrowly ovoidal; testa reticulate with
aequatorial striae. 2n = 24, 48. Two widespread and
K weedy species from S Mexico to Brazil and Bolivia,
also Caribbean, Galapagos.
12. Plakothira Florence

Plakothira Florence, Adansonia II, 7: 240 (1985); Florence,
Allertonia 7: 238-253 (1997).
Shrub or perennial herb with fibrous roots.

Lamina undivided. Inflorescences terminal dicha-
sia or complex thyrsoids; each flower with 2
minute prophylls; flowers erect; petals green or
white; staminodial groups antesepalous with 7-9
staminodia each, 4-6 outer ones fused to form a
green and yellow nectar scale or free to base, often
papillose-hairy. Capsule semisuperior, subglo-
bose, straight; seeds narrowly ovoidal; testa retic-
ulate with aequatorial striae. 2n = 48. Three spp.,
\
Marquesas Is. (Polynesia), probably congeneric
with Klaprothia (Plakothira is morphologically
and karyologically much closer to K. mentzelioides
than K. mentzelioides is to K. fasciculata) .
'J
J
13. Xylopodia Weigend Fig. 84C-I
Xylopodia Weigend, Nasa and the Conquest of South America:
215 (1997).
F G H
Shrub with erect branches from horizontal
Fig. 83. A-Q Loasaceae-Loasoideae. A-1 Nasa argemonoides.
A Habit. B Mature Fruit. C Seed. DDetail of testa epidermis. E xylopodium. Lamina with 2-3 lobes on each side,
Floral diagram, staminodial complexes black, U-shaped struc- margin serrate. Inflorescences terminal dichasia;
ture: nectar scale of three united staminodes. F-G Nectar scale. each flower with 2 minute prophylls; flowers
H Free staminode. I Uniseriate trichomes (stem). J-M Nasa pendent; petals green; staminodial groups with
rudis subsp. australis. 1- K Nectar scale. L Free staminode.
M Flower. N Nasa humboldtiana, T-shaped, scabrid trichome
7-9 staminodia, outer 4-6 staminodes fused to
(stem). 0 Nasa triphylla, glochidiate trichome. P Nasa aequa- form a green and yellow scale, densely papillose-
toriana, glochidiate trichome. (Drawn by M. Weigend) hairy, inner ones free, greenish yellow. Capsule
semisuperior, subglobose, straight, opening with 4
apical valves; seeds narrowly ovoidal; testa reticu-
late. 2n = 24. Only one species, X. klaprothioides
Weigend, rocky slopes in Cajamarca, Peru.
Erect or ascending annual or perennial herbs.

Lamina undivided. Inflorescences terminal, dicha- II. SUBFAMILY MENTZELIOIDEAE (Rchb.) Urb. &
sia (K. mentzelioides Kunth) or complex thyrsoids Gilg (1900).
(K. fasciculata (K. Presl) Poston); each flower with
2 minute prophylls; flowers erect; petals white; sta- Leaves usually opposite below and alternate above,
minodial groups with 2- 6 staminodia each, all free with scabrid-glochidiate trichomes, sometimes
or (K. mentzelioides) 3-4 outer ones only slightly with stinging hairs. Inflorescences terminal thyr-
connate basally, very densely papillose-hairy, apex soids, often without metatopy; flowers erect,
often irregularly dub-shaped and lobed, yellow. merous, sessile or pedicellate; prophylls frondose
Loasaceae 251
or bracteose; petals free or rarely united, planar

or shallowly boat-shaped, never clawed, margin
entire, membranaceous, with >3 principal veins
from base; polystemonous, rarely obdiplostemo-
nous; filaments filiform or sometimes widened or
apically forked; anthers latrorse; staminodia, if
present, petaloid and free to base; parietal placen- D
tae simple, with numerous ovules. Capsule terete,
without prominent veins on the outside, calyx per-
sistent, opening with 3-5 apical valves, straight;
seeds ovoidal or angular or laterally compressed;
testa mostly polyhedral or striate, grey or brown.
14. Mentzelia L. Fig. 84A, B

Mentzelia L., Sp. Pl. 1: 516 (1753); Darlington, Ann. Missouri
Bot. Gard. 21: 103-226 (1934), rev.; Hill, Brittonia 28: 86- ll2
(1976), seed morph.
Bartonia Pursh ex Sims (1812).
(t)G
Acrolasia K. Pres! (1835).
Trachyphytum Nutt. ex Torrey & Gray (1840).
Bicuspidaria Rydb. (1903).
Annual or perennial herbs, subshrubs, shrubs or

small trees with scabrid-glochidiate trichomes,
without stinging hairs. Dominant and persistent
tap-root present. Leaves opposite below and
alternate above, rarely opposite throughout
(M. arborescens Urb. & Gilg); lamina ovate to
H
triangular-ovate, occasionally lyrate, usually
lobed, margin serrate or mucronate. Inflorescences
complex, terminal, basitonic thyrsoids; each flower
with 2 frondose prophylls; petals white, yellow, or
orange; stamens numerous (rarely 10); filaments
equal to very unequal in size, sometimes outer
filaments dilated or apically forked; sometimes
with petaloid staminodes (M. decapetala (Pursh)
Urb. & Gilg). Stigma punctiform. Capsule inferior,
sessile or pedicellate, opening with 3-7 apical
valves; seeds angular or laterally compressed; testa Fig. 84. A, B Loasaceae-Mentzelioideae, Mentzelia scabra. A
reticulate or polyhedral, sometimes with central Inflorescence branch. B Inner and outer stamen showing
distinct heteranthery. C-1 Loasaceae-Loasoideae, Xylopodia
papillae, sometimes narrowly winged. 2n = 9, 10, klaprothioides. C Habit. D Infructescence. E Flower. F Floral
11, 14, 18, 27, 36. Ca. 80 spp., from Argentina to diagram, staminodial complexes black. G Petal with refiexed
Canada incl. Caribbean and Galapagos Is., most in anthers held by longitudinal lamellae. H Nectar scale. I Free
SW USA and Mexico. The majority of species in staminode. Drawn by M. Weigend
desert and semi-desert habitats, some penetrate
into grasslands and rain-green forests.
serrate or mucronate. Inflorescences terminal

15. Eucnide Zucc.
thyrsoids (sometimes with long, terminal mono-
Eucnide Zucc., Del. Hort. Monac. 1844/4:28 (1844); Thompson chasia) or simple dichasia; each flower with 2 fron-
& Ernst, J. Arnold Arbor. 48: 56-88 (1967), rev.; Thompson dose or bracteose prophylls; petals narrowly ovate
& Powell, Phytologia 49: 16- 32 (1981), rev.
or ovate, sometimes post- or congenitally united
(sympetalous), yellowish white, yellow, or orange;
Annual or perennial herbs; stinging hairs usually filaments free to base or stamens epipetalous, all
present. Leaves petiolate; lamina widely ovate to equal, staminodia absent; stigma with 3-5 diver-
subcircular, margin shallowly lobed and deeply
252 M. Weigend
gent lobes. Capsule inferior, cylindrical to conical, Inflorescences terminal thyrsoids, typically with
pedicellate, pedicel sometimes strongly elongating distal dichasium and one additional, monochasial
after anthesis (chasmocarpous), opening with 3-5 paraclade; calyx lobes free or united nearly to
apical valves; seeds ovoidal; testa striate. 2n = 42. apex, yellow or yellowish green; petals entire,
Thirteen spp., Guatemala to SW USA, most species included or long and exserted from the calyx,
in Mexico. thinly membranaceous, pale yellow; stamens
without differentiated connective. n = 37. Two spp.,
fast-growing annual herbs in rain-green forests,
16. Schismocarpus Blake
NW Peru to Mexico.
Schismocarpus Blake, Contrib. Gray Herb. II, 53: 61 (1918).
18. Fuertesia Urb. Fig. 85A-L

Subshrub without stinging hairs. Annual shoots
arising from an underground xylopodium. Leaves Fuertesia Urb., Ber. Deutsch. Bot. Ges. 28: 515 (1910).
alternate, petiolate; lamina widely ovate to subcir-
cular, entire, margin serrate. Inflorescences termi- Liana with splinter hairs and glochidiate hairs
nal thyrsoids, typically with one distal dichasium with 2 hooks at the tip. Lamina cordate, coria-
and one additional monochasial paraclade; each ceous, margin entire. Inflorescences terminal
flower with 2 bracteose prophylls; petals narrowly thyrsoids, typically with distal dichasium and 1-3
ovate or ovate, membranaceous, planar, apex additional dichasial paracladia; calyx lobes free,
acute, green, white; stamens 10, 5 shorter, 5 longer, yellowish green; petals laciniate, included in
obdiplostemonous; staminodia absent; stigma the calyx, thinly membranaceous, pale yellow;
with 5 divergent lobes. Capsule semisuperior, stamens without differentiated connective. One
ovoidal, pedicellate, opening with 5 apical valves; species, F. dominguensis Urb. from Hispaniola.
seeds ovoidal; testa striate. Only one species, S.
pachypus Blake, from Oaxaca, Mexico.
19. Cevallia Lagasc.
Cevallia Lagasc., Gen. Sp. Pl.: 35 (1805); Davis & Thompson,
III. SUBFAMILY GRONOVIOIDEAE (Rchb.) Link Madroflo 19: 1-18 (1967).
{1833).
Subshrub with glochidiate hairs with spreading
Leaves (upper) always alternate (first pair in (not reflexed) branches, without stinging hairs.
seedling opposite), always with glochidiate tri- Lamina narrowly ovate, margin sinuate. Inflores-
chomes, sometimes with stinging hairs. Inflo- cences terminal thyrsoids, typically with distal
rescences terminal thyrsoids with a terminal dichasium and 1 additional, mono- or dichasial
dichasium and 1-2 additional paracladia, each paraclade, strongly condensed; calyx lobes and
flower with 2 filiform prophylls; flowers erect, 5- petals identical, free to base, whitish, membrana-
merous; petals free, spatulate, planar, not clawed, ceous; stamens with connective long protracted
margin entire or laciniate, thinly membranaceous, into club-shaped appendage. n = 13 (Davis and
with one principal vein from base; haplostemo- Thompson 1967). One species, C. sinuata Lagasc.,
nous, staminodia absent; filaments filiform; in the deserts of SW USA and N Mexico.
anthers introrse; ovary inferior, with 1 pendent
ovule. Fruit a cypsela, pentagonous in cross
section, alate on ribs, calyx persistent and accres- IV. SUBFAMILY PETALONYCHOIDEAE Weigend
cent, fruit free from bracts and prophylls; testa {1997).
white to beige.
Leaves alternate with glochidiate hairs with short,
spreading (not reflexed) branches, without sting-
17. Gronovia L. ing hairs. Inflorescences terminal racemes; each
Gronovia L., Sp. Pl. 1: 202 (1753). flower with 2 prophylls, in the axil of 1 ovate
pherophyll; flowers erect, 5-merous; petals free to
Scandent, annual herbs; stinging hairs and base or postgenitally fused, often clawed, with
glochidiate hairs with 2 hooks at the tip always one principal vein from base; haplostemonous,
present. Lamina reniform, deeply sinuate-lobate stamens antesepalous, sometimes displaced out-
with 3-4lobes on each side, lobes ovate-lanceolate, side the pseudosympetalous corolla or filaments
long acuminate, membranaceous, margins entire. locked between the petal margins; filaments
Loasaceae 253
Behnke, H.-D., Barthlott, W. 1983. New evidence from the ultra-

structural and micromorphological fields in angiosperm
classification. Nordic J. Bot. 3: 43-66.
Carlquist, S. 1984. Wood anatomy of Loasaceae with relation
to systematics, habit and ecology. Aliso 10: 583-602.
Carlquist, S. 1987. Wood anatomy of Plakothira (Loasaceae).
Aliso 11: 563-569.
Carlquist, S. 1992. Wood anatomy of sympetalous dicotyledon
families: a summary, with comments on systematical rela-
tionships and evolution of woody habit. Ann. Missouri Bot.
Gard. 79:303- 332.
Cocucci, A.A., Sersic, A.N. 1997. Evidence of rodent pollination
in Cajophora coronata (Loasaceae). Plant Syst. Evol. 211:
113-128.
Coleman, J.R., Smith, L.B. 1969. Chromosome numbers of
some Brazilian Angiosperms. Rhodora 71: 548-551.
M Darlington, J. 1934. A monograph of Mentze/ia. Ann. Missouri
Bot. Gard. 21: 103-226.
Davis, W.S., Thompson, H.J. 1967. A revision of Petalonyx
~ ®1
(Loasaceae) with a consideration of affinities in subfamily
Gronovioideae. Madroiio 19: 1-18.
Dostert, N., Weigend, M.1999. A synopsis of the Nasa triphy lla
0 p Q complex (Loasaceae), including some new species and sub-
H
species. Harvard Pap. Bot. 4: 439-467.
Garcia, V. 1962. Embryological Studies in the Loasaceae: devel-
Fig. 85. A-L Loasaceae-Gronovioideae, Fuertesia domingen- opment of endosperm in Blumenbachia hieronymi Urb.
sis. A Habit. BFruit. C Floral diagram. D Calyx lobe. E Petal. F Phytomorphology 12: 307-312.
Stamen. G Style and stigma in cup-shaped nectary. H Stinging Gilg, W. 1925. Loasaceae. In: Engler, A., Prantl, K. Die
hair (seta). I Hook-shaped, glochidiate hairs. J Modified nattirlichen Pftanzenfamilien ed. 2., 21. Leipzig: W.
glochidiate trichomes. K Splinter hair. L-Q Loasaceae- Engelmann, pp. 522-543.
Petalonychoideae, Petalonyx L P. parryi, flower. M- Q P. lin- Grau, J. 1988. Chromosomenzahlen chilenischer Loasaceae.
earis. M Fruit, enclosed in bract and bracetoles. N Fruit, bracts Mitt. Bot. Staatssamml. Mtinchen 27: 7-14.
and bracteoles removed. 0 Sepal. P Petal. Q Stamen. (Original; Grau, J. 1997. Huidobria, eine isolierte Gattung der Loasaceae
L Davis and Thompson 1967) aus Chile. Sendtnera 4: 77- 93.
Hempel, A.L., Reeves, P.A., Olmstead, R.G., Jansen, R.K. 1995.
Implications of rbcL sequence data for higher order rela-
tionships of the Loasaceae and the anomalous aquatic plant
Hydrostachys (Hydrostachyaceae). Plant Syst. Evol. 194:
25-37.
Hill, R.J. 1976. Taxonomic and phylogenetic significance of
seed coat microsculpturing in Mentzelia (Loasaceae) in
Wyoming and adjacent western States. Brittonia 28: 86-112.
filiform; anthers laterorse; filiform staminodia Hufford, L. 1988. Seed morphology of Eucnide and other
sometimes present; ovary with 1 pendent ovule. Loasaceae. Syst. Bot. 13: 154-167.
Cypsela remaining attached to bract and pro- Hufford, L. 1990. Androecial ontogeny and the problem of
monophyly of Loasaceae. Can. J. Bot. 68: 402-419.
phylls, irregularly pentagonous, not alate on ribs; Hufford, L. 1998. Early development of androecia in polyste-
calyx caducous; testa white to beige. 2n = 46. monous Hydrangeaceae. Am. J. Bot. 85: 1057- 1067.
Humboldt, F.H.A. v., Bonpland, A.J., Kunth, C.S. 1815-1823
Only one genus: [1825] Nova genera et species plantarum. Paris.
Keeler, K.H. 1981. The nectaries of Mentzelia nuda: from pol-
linator attraction to seed protection. Am. J. Bot. 68: 295-299.
20. Petalonyx A. Gray Fig. 85M-R Leins, P., Winhard, W. 1973. Entwicklungsgeschichtliche
Studien an Loasaceen-Bltiten. Osterr. Bot. Zeitschr. 121/122:
Petalonyx A. Gray, Pl. Nov. Thurb.: 319 {1854); Davis & 145- 165.
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Timber Press.
Moody, M.L., Hufford, L., Douglas E. Soltis, D.E., Soltis P.S.
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254 M. Weigend
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tions in Loasaceae subfamily Loasoideae. Syst. Bot. 2: 28-35. Flora de Colombia, vol. 22. Sta. Fe de Bogota: Instituto de
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Schlindwein, C., Wittmann, D. 1997. Microforaging routes of (Loasales). Am. J. Bot. 87: 1202-1210.
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Takhtajan, A.L. 1980. Outline of the classification of flowering Xiang, Q.- Y. 1999. Systematic affinities of Grubbiaceae and
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Takhtajan, A.L. 1997. See general references. sequences. Harvard Pap. Bot. 4: 527-542.
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Urban, I. 1892. Die Bliitenstande der Loasaceen. Ber. Deutsch. and rbcL sequences. Am. J. Bot. 85: 285-297.
Bot. Ges. 10: 220-225.
Maesaceae 255
Maesaceae
B. STAHL and A.A. ANDERBERG
Maesaceae (A. DC.) Anderberg, Stahl & Kallersjo, Taxon 49: primulalean families in having a mixture of uni-
185 (2000). and multiseriate rays (Janssonius 1920).
Small to medium-sized trees, shrubs or sometimes INFLORESCENCE AND FLORAL STRUCTURE. Inflo-
lianas with schizogenous cavities; indumentum of rescences are axillary, usually appearing along
non-glandular, uniseriate trichomes and scales. a major part of the young shoots. Most species
Leaves simple, petiolate, exstipulate, distinctly have compound racemes (Fig. 86), but species with
alternate (generally not clustered), with prismatic simple racemes are not uncommon. The flowers
crystal druses; margins entire or variously serrate are perigynous. The calyx lobes are usually well
or crenate. Inflorescences axillary racemes or com- demarcated, being broadly obtuse-ovate to trian-
pound racemes; pedicels subtended by a bract and gular in shape; the corolla is campanulate or some-
bearing prophylls just below the calyx. Flowers times urceolate with a well-developed tube which,
(6)5-4-merous, bisexual or functionally female, if depending on species, varies from being some-
unisexual then plants polygamous or functionally what shorter to more than twice the length of the
dioecious; calyx lobes mostly broadly ovate, per- lobes. There is a single whorl of antepetalous
sistent; corolla white, yellow or pinkish, campanu- stamens, which are usually completely included in
late or sometimes urceolate, lobes broadly ovate to the corolla. The filaments are basally fused to the
subrotund, quincuncial in bud; stamens homo- corolla tube. The anthers are basifixed, shortly
merous, antepetalous, included or slightly exserted; oblong-ovoid and obtuse or retuse at apex; they
filaments inserted on the corolla tube; anthers open introrsely by means of longitudinal slits.
almost square, or somewhat shorter than wide, The ovary is conical and tapers into a short but
apically rounded or retuse, dorsifixed, introrsely well-demarcated style; the stigma is capitate or
dehiscent by longitudinal slits; ovary semi-inferior; truncate, or sometimes 3-5-lobed. There is no
style well demarcated; stigma truncate or capitate, nectariferous disk. The ovules are spirally inserted
entire or 2-5-lobed; placentation free central; in one or 2-8 series on the central placenta.
ovules many, arranged in 1-5 whorls, not or only Floral development in the family was studied by
slightly immersed into the placenta. Fruit berry- Caris et al. (2000).
like with a somewhat woody meso carp, globular or
ovoid, indehiscent. Seeds numerous, small, angular, EMBRYOLOGY. According to Johri et al. (1992), the
dark brown; seed coat reticulate, two-layered, anthers of Maesa have a fibrous endothecium and
with rhomboid crystals; endosperm abundant with a secretory tapetum and, like in other primulalean
evenly thickened cell walls; embryo with short families, the ovules are bitegmic, tenuinucellate
hypocotyl and short, narrow cotyledons. x = 10. and have a Polygonum-type embryo sac and
A monogeneric family with about 150 species, nuclear endosperm formation. An integumentary
distributed in tropical and subtropical regions of tapetum is present and the embryogeny is of the
the Old World. onagrad type.
VEGETATIVE STRUCTURES. Most species are POLLEN MORPHOLOGY. Pollen of Philippine

shrubs, but some are small or medium-sized trees, Maesa cumingii is 3-colporoidate, ± spheroidal
scandent shrubs or lianas. (11 x 9J.lm) and subreticulate (Erdtman 1952),
Schizogeneous secretory cavities are common and similar to pollen of most other members of
and probably occur in the leaves of all species; they primulalean families.
are often visible as brownish-orange lines on
vegetative and reproductive parts (Smith 1973). KARYOLOGY. A basic chromosome number of x =
Calcium oxalate druses are found in the meso- 10 has so far been recorded in about five species
phyll. The wood in Maesaceae is unique among of Maesa.
256 B. Stahl and A.A. Anderberg
Fig. 86. Maesaceae. A- C Maesa lanceolata. D-F M. japonica. brown to black; the testa is two-layered and con-
A Habit, flowering branch. B, C, E, F Flowers (E in vertical tains rhomboid crystals; endosperm is abundant
section showing placentation and ovules). D Fruit. (Anderberg
and Stahl1995) and has smooth cells walls. The embryo is short
with cotyledons of about the same length and
width as the hypocotyl.
REPRODUCTIVE SYSTEMS AND POLLINATION.
Morphologically, the flowers of all species appear PHYTOCHEMISTRY. A special kind of quinone,
to be hermaphrodite. However, Mez (1902) and maesaquinone, occurs in fruits of Maesa lanceo-
Sleumer (1987) described Maesa as dioecious, but lata and M. japonica, and other quinones are
there is little support of this in other treatments of present in vegetative parts of the former species
the genus. Although admitting that it is difficult to (Hegnauer 1969). Saponins have been reported
ascertain whether the anthers are functional or from several species of the family, both in vegeta-
not, Sleumer ( 1987) found the anthers of female tive parts and fruits. In addition, ubiquitous
flowers reduced in size, thus suggesting cryptic or flavonols have been isolated from leaves of M.
functional dioecy. Flowers within a given individ- chisa, and alkaloids have been detected in leaves of
ual are sometimes unisexual and the plants func- M. ramentacea and M. perlarius. The endosperm
tionally polygamous. Flower morphology suggests contains amyloid and fatty oil, although the
pollination by insects, but anemophily may be former appears to be sparse in the family (Seegeler
expected in some species. 1983).
FRUIT AND SEED. Being many-seeded, the fruits SUBDIVISION AND RELATIONSHIPS WITHIN THE
are best described as berries, although the meso- FAMILY. The single genus, Maesa, has been
carp is often thin and the endocarp rather hard divided into two subgenera (Mez 1902). Thus, a
and brittle. The seeds are small, angular, and dark few African species with ovules arranged in a
Maesaceae 257
single series on the placenta are referred to subg. Maesa Forssk. Fig. 86
Monotaxis Mez and the remaining species (also
Maesa Forssk., Fl. Aegypt. Arab. 66 (1775); Mez, Pfianzenreich
some African), with ovules in 2 or several series, IV 236: 15-54 (1902); Ridley, Fl. Malaya Peninsula 2:227-229
are placed in subg. Maesa. (1923), reg. rev.; Walker, Philippine J. Sci. 73: 12-48 (1940),
reg. rev.; Smith, J. Arnold Arb. 54: 3-36 (1973), Fijian spp.;
AFFINITIES. Maesaceae have mostly been treated Fosberg & Sachet, Phytologia 44: 362-369 (1979), Pacific
spp.; Taton, Fl. d'Afrique Centrale (no vol.) 3-12 (1980), reg.
as a subfamily of Myrsinaceae, both agreeing in rev.; Halliday, Fl. Trop. E. Afr. (no vol.) 2-5 (1984), reg. rev.;
woody habit, sympetalous, 5-merous flowers, free Sleumer, Blumea 32: 39-65 (1987), New Guinean spp.
central placentation, and schizogenous secretory
cavities. However, the differences in reproductive Description as for the family. About 150 spp.,
structures and wood anatomy were noted by throughout the Old World tropics and subtropics;
earlier workers (de Candolle 1841; Janssonius two subgenera: subg. Monotaxis Mez with ovules
1920). Recent phylogenetic studies, including in a single row, and subg. Maesa in 2 or more
both morphological and DNA sequence data series.
(Anderberg and Stahl1995; Anderberg et al. 1998;
Kallersjo et al. 2000; Savolainen, Fay et al. 2000),
strongly support the position of Maesaceae dis- Selected Bibliography
tinct from Myrsinaceae and Primulaceae. Accord-
ing to Kallersjo et al. (2000), Maesaceae are sister Ambasta, S.P. (ed.) 1986. The useful plants oflndia. New Delhi:
to Theophrastaceae-Primulaceae-Myrsinaceae. Publication and Information Directorate.
Anderberg, A.A., Stahl,B.1995. Phylogenetic interrelationships
DISTRIBUTION AND HABITATS. The family is in the order Primulales, with special emphasis on the family
circumscriptions. Can. J. Bot. 73: 1699-1730.
distributed throughout the Old World tropics, Anderberg, A.A., Stahl, B., Kallersjo, M. 1998. Phylogenetic
extending into subtropical regions in Asia. The relationships in the Primulales inferred from rbcL sequence
largest concentration of species is found in SE data. Plant Syst. Evol. 211: 93-102.
Asia, but many species are also present in India, Anderberg, A.A., Stahl, B., Kallersjo, M. 2000. Maesaceae, a new
China, and the Pacific. About six species occur in primuloid family in the order Ericales s.l. Taxon 49: 183-187.
Candolle, A. de 1841. Troisieme memoire sur la famille de
Africa, one of which has seemingly been intro- Myrsinacees. Ann. Sci. Nat. Bot. Bioi. Veg. 16: 129-176.
duced in Madagascar (Perrier de la Bathie 1953), Caris, P., Ronse Decraene, L.P., Smets, E., Clinckemaillie, D.
and three species occur in tropical Australia 2000. Floral development of three Maesa species, with
(Queensland). In China,Maesa grows in evergreen special emphasis on the position of the genus within
broad-leaved or mixed forests at 600-2700m Primulales. Ann. Bot. II, 86: 87-97.
altitude, often in damp areas such as riverbanks. Hegnauer, R. 1969. See general references.
Towards the Equator, the family continues as Janssonius, H.H. 1920. Mikrographie des Holzes der auf Java
primary lowland or mid-altitude element with vorkommenden Baumarten 4. Leiden: Brill.
few species reaching subalpine elevations. Several Johri, B.M. et a!. 1992. See general references.
Kallersjo, M. et a!. 2000. See general references.
species occur in disturbed and secondary forests.
Mez, C. 1902. Myrsinaceae. In: Engler, A. (ed.) Das Pfianzenre-
ich IV, 236. Leipzig: W. Engelmann.
ECONOMIC IMPORTANCE. In Ethiopia, the oily Perrier de la Bathie, H. 1953. Myrsinacees. In: Humbert, H.
seeds of Maesa lanceolata are used for frying, and (ed.) Flore de Madagascar et des Comores. Paris: Mus. Nat!.
crushed fruits are used to treat infections of intes- Hist. Nat.
Savolainen, V., Fay, F. et a!. 2000. See general references.
tinal parasites (Seegeler 1983). The fruits of some Seegeler, C. J.P. 1983. Oil plants in Ethiopia, their taxonomy and
Indian species are sometimes used as a fish- agricultural significance. Wageningen: Pudoc.
poison, and the leaf of one species, M. indicia Sleumer, H. 1987. A revision of the genus Maesa Forssk.
Wall., is a curry ingredient (Ambasta 1986). The (Myrsinaceae) in New Guinea, the Moluccas, and the
wood is sometimes used for construction works. Solomon Islands. Blumea 32: 39-65.
Smith, A.C. 1973. Studies of Pacific island plants, XXV. The
Extracts of leaves, twigs and bark have insecticidal Myrsinaceae of the Fijian region. J. Arnold Arbor. 54: 1-41,
effects (Hegnauer 1969). 228-292.
One genus:
258 S. Dressler
Marcgraviaceae
S. DRESSLER
Marcgraviaceae Choisy in DC., Prodr. 1: 565 {1824), nom. cons. inflorescences. Both forms are connected by inter-
mediate leaf-forms.
Terrestrial, hemiepiphytic or epiphytic lianas or A noteworthy feature of the family is the pres-
shrubs, rarely small trees. Hypophyllous glands, ence of solid or poriform "hypophyllous glands",
raphide cells and variously shaped sclereids which occur often in specific patterns on the
frequently present. Leaves simple, alternate, ex- abaxial face of the leaf. In Marcgravia, they can
stipulate, glabrous, margins entire or (some intergrade with "marginal glands". They have been
Marcgravia) minutely crenate. Inflorescences ter- regarded as domatia, hydathodes, extrafloral nec-
minal, racemose, sometimes resembling umbels or taries, or resin-secreting organs, which certainly is
spikes, erect or pendulous; bracts transformed not appropriate. A nectar-secreting activity has
into variously shaped nectaries; pedicels mostly been observed in early developmental stages of the
with two sepaloid prophylls. Flowers hermaphro- leaf. There is still no certainty about their ecolog-
ditic, actinomorphic, hypogynous; sepals (4)5, ical significance. They may attract ants as protec-
unequal, free or nearly so, imbricate, persistent; tion against predators while the leaf tissue is still
petals 3-5, imbricate, free or connate; stamens soft (Dressler 1997).
3-many; filaments free or basally connate, uni- to Leaf venation is relatively uniform, pinnate-
biseriate; anthers basifixed or nearly so, tetraspo- brochidodromous with ascending secondary
rangiate, introrse, longitudinally dehiscent. Ovary arches. Leaf venation on juvenile branches in
superior, completely or incompletely 2-20-locular; Marcgravia displays an earlier ontogenetic stage.
ovules few to numerous, anatropous. Fruit subglo-
bose, apiculate with persistent style and stigma, VEGETATIVE ANATOMY. The leaves of Marcgravi-
capsular, loculicidally and septifragously dehis- aceae are dorsiventral and hypostomatic. Stomata
cent from the base or berry-like and irregularly (at least of Marcgravia) belong to the staurocytic
dehiscent, pulpy inside. Seeds exotestal-theoid, type (Fryns-Claessens and van Cotthem 1973). In
hemispherical to reniform, few to numerous with the adult leaves of Marcgravia a hypodermis and
a shiny reticulate testa; endosperm scanty or a thick cuticle are usually present, and the palisade
lacking; embryo straight. parenchyma includes one to three layers. Leaves
A family comprising 7 genera and about 130 of the juvenile branches lack a hypodermis, the
species, distributed in tropical Central and South upper epidermis is often papillate and stomata
America including the West Indies. occur also on the upper side. The other genera lack
a hypodermis as well as marginal glands, and the
VEGETATIVE MORPHOLOGY. The developing palisade parenchyma usually includes only one
leaves convolutely enclose the shoot-tip, which is layer.
rather characteristic of the family and similar to Different kinds of idioblasts occur in the family.
some Theaceae. All Marcgraviaceae develop an Very characteristic is the presence of various types
apical mucro that is often caducous at the fully of sclereids in the leaves (de Roon 1967), nectaries,
expanded leaf. perianth, pistil and pericarp as well as in the
In Marcgravia, the leaves are distichously cortex and medulla of the shoots. Rap hide cells are
arranged and often provided with a drip-tip, found in parenchymatous tissues. Styloids and
whereas phyllotaxis of the other genera is spiral. oil-containing or mucilaginous cells occur rarely
Marcgravia has dimorphic branches: sterile, (de Roon 1975).
creeping or root-climbing branches that are An unilacunar node with a single trace in the
angled and bear two rows of small, juvenile, petiole closed to a siphonostele is common
cordate leaves; and fertile and pendulous branches throughout the family (Schofield 1968).
that are rootless and terete. The latter bear the The main features of the wood are as follows:
typical adult leaves and may develop terminal vessels mostly solitary or in radial multiples of
Marcgraviaceae 259
Ruyschia orantea Schwartzia Sarcopera

Souroubea
en
B
Marcgraviastrum Marcgravia Marcgravia
2-4, medium-sized (Marcgravia) to rather long Fig. 87. Marcgraviaceae. Inflorescence structure with inser-
(Norantea s.l., Souroubea), perforations simple, tion of the nectarial bracts and the degree of fusion of their
stalk with the pedicel. A Fertile flower. B Nectarial bract. C
rarely scalariform (e.g. in No ran tea s.l., Nectary with abortive flower. (after de Roon 1975)
Souroubea); fibres septate, c. 800 f.!m long in
Marcgravia, 1200-1800f..lm in Norantea s.l. and
Souroubea; parenchyma scanty paratracheal to inflorescence that consist of a bract fused with
vasicentric; rays homogeneous, uni- to multiseri- its petiole (Fig. 87). These nectaries are generally
ate, cells mostly upright (de Roon 1975). Norantea developed on all flowers of the inflorescence in
s.l. has been reported being a caulinar myrmeco- all Marcgraviaceae except Marcgravia. In this
phyte with a non-disintegrating pith (Jolivet genus the apical (central) flowers are sterile and
1996). provided with a nectary that is fused with the
The sieve-element plastids belong to the Ss-type abortive pedicel, whereas the fertile flowers lack
(Behnke 1991). The epicuticular wax platelets are nectaries.
unspecific (Ditsch and Barthlott 1994). An indu- The nectaries are foliaceous to gibbose in
mentum of multicellular hairs is rarely present on Ruyschia; cup- or spur-shaped and often auricu-
young shoots or the inflorescence. late in Souroubea; cup-, sac,- or pitcher-shaped
in Norantea, Sarcopera, Schwartzia, and Marc-
INFLORESCENCE STRUCTURE. All inflorescences graviastrum; and tubular-clavate to pitcher- or
are racemose. In Sarcopera they are spike-like, boat-shaped in Marcgravia. Often they are
whereas in Marcgraviastrum and Marcgravia conspicuously coloured and, from a pair of glands,
the inflorescence axis is extremely contracted, secrete a sweetish liquid as an attractant for polli-
which leads to umbellate inflorescences. A char- nators. Inflorescence type and insertion of nec-
acteristic, autapomorphic trait of the family is taries are highly characteristic for each genus and
the presence of extrafloral nectaries in the facilitate identification.
260 S. Dressler
In Sarcopera, the apical flowers often have fully

developed nectaries, while the nectaries become
reduced towards the base of the inflorescence
and the lowermost flowers lack the nectaries.
This feature as well as the contraction of the axis
(Marcgraviastrum) are interpreted as advanced
states. A combination of both is found in
Marcgravia.
FLORAL STRUCTURE AND ANATOMY. The flowers

are pentamerous (except Marcgravia). The
aestivation is imbricate and the sepals, if five, are
quincuncially arranged. The petals are free or
somewhat fused and reflexed at anthesis. In Marc-
gravia, the corolla is tetramerous and completely
fused into a calyptra that falls off at anthesis.
The stamens are arranged in one to two whorls;
their number is (3- )5 in Ruyschia and Souroubea,
and ranges from (5-)7-100 (e.g. in Marcgravias-
trum and Marcgravia). The anthers are tetraspo-
rangiate, dorsifixed and open introrsely by
longitudinal slits. The filaments arefiliform to flat-
tened and free, rarely adnate to the base of the
petals. Pentandrous flowers always have antesepa-
lous stamens.
The ovaries are superior, (in)completely 2-15-
locular and have axile placentation with the pla-
centae proliferating into the locules. In Marcgravia
and Marcgraviastrum ovules are numerous,
whereas the other genera have only 5-20. The style
is distinct but in Souroubea, Sarcopera, and Marc-
gravia the stigma is mostly (sub)sessile.
Sclereids are common in the pericarp and all
floral tissues except the androecium. For further
anatomical details see Juel (1887), Weber (1956),
and de Roon (1975).
EMBRYOLOGY. Only Marcgravia spp. have been

studied. The anther tapetum is secretory and con- Fig. 88. Marcgraviaceae. Pollen in equatorial view, SEM
sists of uninucleate cells. Ovules are anatropous, (x4000). A Marcgravia sintenisi, sexine tectate-perforate.
bitegmic and tenuinucellar, with the micropyle B Marcgravia umbellata, sexine coarsely reticulate. Photo
S. Dressler
formed by the inner integument. The nucellus is
highly reduced. The chalazal megaspore develops
into a Polygonum-type embryo sac. Endosperm
development is cellular and the cell closest to the from the rest of the family in having an imperfo-
micropyle develops into a non-aggressive hausto- rate or perforate tectum whereas the other genera
rium. For further details see Davis ( 1966) and may also have reticulate pollen. Another palyno-
Johri et al. {1992). logical trait of Souroubea and Ruyschia is the
comparatively thin (<2 1-tm) exine. Punt (1971)
POLLEN MORPHOLOGY. The pollen grains are postulates evolutionary trends: from tectate to
mostly spheroidal (suboblate to subprolate), 3(4)- coarsely reticulate and tectate-perforate sexine;
colporate and relatively small (12- 31 J.lm long) from long and slit-like exocolpi to shorter and
(Fig. 88). The colpi are usually long and narrow, broader ones; and from small, distinct endoaper-
only some Marcgraviastrum and Marcgravia spp. tures to large, lalongate endoapertures with paral-
have shorter colpi. Souroubea and Ruyschia differ lel to divergent or diffuse lateral ends.
Marcgraviaceae 261
KARYOLOGY. No data on chromosome numbers Fig. 89. Marcgraviaceae. Marcgravia serrae with flower bat
are published, but preliminary studies revealed Hylonycteris underwoodii approaching for nectar bracts
central in the inflorescence. (Tschapka and von Helversen
small chromosomes (0.7-2.5Jlm) with semi- 1999)
reticulate interphase nuclei and proximal-anterior
chromosome condensing in prophase to pro-
metaphase. For Marcgravia evenia ssp. calcicola mium, erect inflorescences, many light stamens
2n = 36 has been determined (A.M. Benko- and nocturnal anthesis facilitate sternotribic pol-
Iseppon, pers. comm.). lination by phyllostomid bats. Opossum pollina-
tion was observed in Marcgravia nepenthoides
POLLINATION. The pollination system m (Tschapka and von Helversen 1999).
Ruyschia is unknown; fly pollination was sug- Although formerly reported as being generally
gested (Delpino 1869; Vogel1993). In some species protandrous, as observed in some Marcgravia
of Souroubea the flowers emit a very strong, sweet spp. (Tschapka and von Helversen 1999) and
smell and are psychophilous (Gentry in Putz and Souroubea guianensis (Machado and Lopes 2000),
Mooney 1991). Sphingophily has also been sug- stigma and anthers mature at about the same time
gested (Vogel1993). A detailed study of S. guianen- in other taxa (e.g. Schwartzia brasiliensis, cf.
sis (Machado and Lopes 2000) revealed Sazima et al. 1993). Despite their elaborate inflo-
"non-legitimate" visits of a hummingbird and the rescences, many Marcgraviaceae are probably
occurrence of "tapetal oil". Norantea, Sarcopera, autogamous and even cleistogamous (Bailey
and Schwartzia brasiliensis show large, lively 1922), since self-pollination takes place in the bud.
coloured nectaries and bird pollination has been
observed in a few species (Sazima et al. 1993). FRUIT, SEED AND DISPERSAL. All Marcgraviaceae
Species of Marcgraviastrum and Marcgravia, have depressed globose fruits with a persistent
genera that for long have been regarded as excel- calyx. The pericarp is more or less coriaceous
lent examples of bird-pollination, have proved to rather than woody, and uqripe fruits could be
be rather chiropterophilous (Fig. 89; see also Vogel described as baccate. The pericarp is reddish-
1958; Sazima and Sazima 1980; Tschapka and von green to purple or brownish and eventually
Helversen 1999; Zusi and Hamas 2001; Dressler dehisces irregularly or loculicidally and sep-
and Tschapka 2002). These genera may have inde- tifragously from the base and the apex, exposing a
pendently shifted from entomophily to vertebrate pulpy, purple placentar tissue with numerous
pollination. Ornithophily seems well developed in small seeds. Thus, the fruits are capsular.
the Galeatae group of Marcgravia subgen. Marc- The seeds are flattened, hemispherical to reni-
gravia (syn. Plagiothalamium), in which red form, and have a shiny, reddish, reticulate testa.
nectaries, which subtend the pseudo-umbel, en- They are irregularly distributed in the pulp, where
sure nototribic pollination by hummingbirds. In often also some black aborted seeds are found.
certain spp. of Marcgravia subgen. Orthothala- Endosperm is starchy but scanty or lacking. The
262 S. Dressler
embryo is straight, relatively large and provided analyses place Tetrameristaceae and Pellicieraceae
with a short radicle and long cotyledons. sister to Marcgraviaceae and all these close to Bal-
At maturity, the pericarp and the pulpy placen- saminaceae, somewhat distant from Theaceae and
tae are often reddish. When the fruit dehisces, Ternstroemiaceae in the Ericales (Savolainen et al.
the seeds, pulp, and pericarp attract birds and 2000; Anderberg et al. 2002).
mammals. Toucans, aracaries and others are
reported to consume the fruits (Davis and Yost DISTRIBUTION AND HABITATS. The family is
1983 ). Vegetative propagation by sterile shoots also confined to the Neotropics, where it extends from
plays a role. S Mexico to N Bolivia including the West Indian
islands with higher altitudes. Members of the
PHYTOCHEMISTRY. Melchior (1924) reported the family are relatively rare and prefer primary
occurrence of inulin crystals from the leaves of vegetation ranging from humid tropical lowland
two Marcgravia species, but Weber (1955) was forests to montane rain and cloud forests.
unable to detect this substance in a different Many species prefer higher altitudes and the
Marcgravia species and Norantea guianensis. At- genus Ruyschia is restricted to them. Sometimes
tempts to trace inulin or fructan with fructanase altitudinal vicariance between species is ob-
in several taxa were unsuccessful (Albrecht and served.
Dressler, unpubl. data).
Unspecified terpenes, tannins, saponins, alka- PALAEOBOTANY. Single pollen grains of Marc-
loids, and phenolics including leucopelargonidin gravia and of Norantea s.l. were recorded from
and myricetin have been reported from the family the Lower Oligocene (c. 45 million years ago) of
(Hegnauer 1969; Saleh and Towers 1974; Merz San Sebastian, Puerto Rico (Graham and Jarzen
1991). 1969). Lacking a detailed description, however,
this record was not confirmed by Muller (1981).
RELATIONSHIPS WITHIN THE FAMILY. Norantea Fossilized wood was described as Ruyschioxylon
s.l., which shows a high diversity in inflorescence sumatrense from the Tertiary of Sumatra [sic!] by
structure and pollen morphology, was recognised Hofmann (1884).
as a heterogeneous assemblage by de Roon (1975)
and Bedell (1985). It was separated into four ECONOMIC IMPORTANCE. Apart from occasional
genera by de Roon and Dressler (1997). horticultural use in the tropics, the family has no
The unspecialised inflorescence morphology economic significance. Miscellaneous applications
and pollination system of Ruyschia and Souroubea in folk medicine have been reported, which include
may be plesiomorphic within the family. In con- use against wounds, headaches, sleeplessness,
trast, characters such as heterophylly, contraction rheumatism, amoebiasis, centipede stings, haem-
of the inflorescence axis, differentiation of sterile orrhaging during menstruation, diarrhoe and
and fertile flowers, increase in number of stamens syphilis (Descourtilz 1827; Davis and Yost 1983;
and ovary locules, oligomerisation of the corolla, Schultes and Raffauf 1990; Putz and Mooney 1991;
pollination system and specialised wood anatomy Merz 1991; herbarium labels). Some local tribes in
characterise Marcgravia as a highly advanced Peru and Ecuador eat the fruits of Marcgravia
group. ("purum hijos" =false figs) or drink the sap; Angely
( 1977) reports that Schwartzia adamantia is
AFFINITIES. Many traits of Marcgraviaceae such believed to indicate the presence of diamonds.
as external morphology, polymorphic sclereids,
pollen morphology and epicuticular waxes are KEY TO THE GENERA
also found in Theaceae/Ternstroemiaceae. Seed
1. Inflorescence umbellate or subumbellate 2
structure has its next equivalent in tribe Adinan- - Inflorescence spicate or racemose 3
dreae of Ternstroemiaceae. The possession of 2. Inflorescence completely fertile (all flowers developed);
raphide bundles is shared with Tetrameristaceae sepals and petals 5; petals free or variously connate; leaves
and Pellicieraceae, both long regarded as close re- spiral 6. Marcgraviastrum
- Inflorescence partly sterile (central flowers aborted with
latives of Theaceae. In both families each locule
only the bracteal nectaries developed); sepals and petals 4;
contains only a solitary ovule and terminal inflo- petals calyptrately connate; leaves distichous
rescences are lacking. Pelliciera has solitary sub- 7. Marcgravia
sessile axillary flowers with two large, coloured 3. Inflorescence spicate; nectaries inserted next to the sessile
prophylls. Tetrameristaceae have axillary pedun- flowers 5. Sarcopera
culate umbelliform racemes. Recent molecular - pedicel Inflorescence racemose; nectaries variously inserted on the
4
Marcgraviaceae 263
4. Stamens 3-5; nectariferous bracts inserted at the base of

the calyx 5
- Stamens (5-) 12 or more; nectariferous bracts inserted at
various distances from the calyx, but never at its base 6
5. Ovary 2-locular; bracts gibbose or somewhat leaf-like,
solid or almost so 1. Ruyschia
- Ovary 3-5-locular; bracts spur-like, tubular, hollow, often
auriculate 2. Souroubea
6. Inflorescence an elongated raceme, {20-)35-65 em long;
pedicels short, 2-5{-7)mm long; nectariferous bracts
inserted above the middle of the pedicel, never attached at
the base 3. Norantea
- Inflorescence a short raceme, 4-25{ -35) em long; pedicels
elongate, (20- )30-70 mm long; nectariferous bracts
inserted below the middle of the pedicel, rarely at the
base 4. Schwartzia
1. Ruyschia Jacq.
Ruyschia Jacq., Enum. Syst. Pl. 2: 17 {1760); de Roon, Contrib.
tow. Monogr. Marcgraviaceae: 141-150 {1975), rev.
Caracasia Szyszyl. {1893).
Climbing shrubs or lianas. Leaves shortly petio-

late. Inflorescences dense multiflorous racemes
with 20-30(-50) flowers. Flowers small, shortly
pedicellate. Nectaries small, sessile, inserted at the
base of flowers or on upper half of pedicel, gibbose
to semi- or subglobose, mostly solid. Petals free or A .
basally slightly united. Stamens 3 or 5. Filaments±
fused with the base of petals. Ovary 2-locular, Fig. 90. Souroubea vallicola. A Habit. B Flower with nectary. C
ovules few to up to 20, stigma capitate. Seven Flower at anthesis. DPetals with stamens. E Ovary with stigma.
spp., mostly from higher altitudes of Central (de Roon 1970)
America (3), northern Andes (3), and Lesser
Antilles (1).
Lianas or sprawling shrubs, often epiphytic. Leaves
2. Souroubea Aubl. Fig. 90 petiolate. Inflorescences long, dense racemes with
100-300 flowers. Flowers shortly pedicellate. Nec-
Souroubea Aubl., Hist. Pl. Guiane 1: 244 {1775); de Roon,
Contrib. tow. Monogr. Marcgraviaceae: 151-187 {1975), rev.
taries on the upper half of the pedicel, stipitate,
saccate. Sepals 5. Petals 5, free to slightly basally
Climbing shrubs or lianas, often epiphytic. Leaves connate. Stamens 20-35; filaments basally adnate
shortly petiolate. Inflorescences lax or dense to corolla. Ovary 5-locular with 10-20 ovules per
racemes with 15-60(-100) flowers. Nectaries on locule. Two spp., N South America and S Brazil,
the upper part of the pedicel, mostly below calyx, Bolivia.
sessile or sometimes stipitate, hollow, mostly spur-
like and auriculate. Flowers 5-merous (rarely 3- to 4. Schwartzia Veil.
6-merous). Petals free or connate up to 2/3 of their
Schwartzia Vel!., Fl. Flumin.: 221 (1829); Bedell, Generic rev.
length. Filaments free or basally adnate to corolla. Marcgraviac. I. Norantea complex: 106-215 (1985), rev.
Ovary (3-)5-locular, often pentagonal, stigma with Norantea subgen. Cochliophyllum Delpino (1869).
(3- )5 radiating lobes. Seeds rather few. Nineteen
spp., Mexico to Bolivia, not in the West Indies. Sprawling shrubs, occasionally small trees. Leaves
sessile to petiolate. Inflorescences short racemes
3. Norantea Aubl. with 8-60 flowers (only S. brasiliensis with long
racemes of 60- 300 flowers) . Pedicels elongate.
Norantea Aubl., Hist. Pl. Guiane 1: 554 {1775); Bedell, Generic Nectaries adnate to the lower third of the pedicel,
rev. Marcgraviac. I. Norantea complex: 69-105 (1985), rev.
Norantea subgen. Sacciophyllum Delpino {1869).
mostly stipitate, cup-, sac- or boat-shaped. Sepals
5. Petals 5, free to variously connate. Stamens
264 S. Dressler
(5-)12-27 or 50-80 in one or several whorls. Fila- Climbing shrubs or vines with dimorphic
ments free or basally connate or adnate to corolla. branches: sterile, juvenile branches creeping,
Ovary (in)completely 3-5-locular. Fourteen spp., appressed to the substrate, and attached by roots;
from Costa Rica along the Andes to Bolivia, fertile ones free, mostly pendulous and provided
Caribbean, E Brazil. with wart-like lenticels. Leaves distichous, those of
sterile branches small, thin, sessile, mostly asym-
metrically cordate; those of fertile ones larger,
5. Sarcopera Bedell
thicker, often with a drip-tip. Inflorescences
Sarcopera Bedell in de Roon & S. Dressler, Bot. Jahrb. Syst. 119: umbelliform, the apical (central) flowers abortive
328 (1997); Bedell, Generic rev. Marcgraviac. I. Norantea with only the bracts (nectaries) well developed, the
complex: 226-295 (1985), rev. fertile flowers without bracts, long-pedicellate.
Norantea subgen. Pseudostachyum Delpino (1869).
Nectaries tubular-saccate or boat-shaped. Sepals 4,
decussate. Petals 4, connate into a deciduous cap.
Sprawling shrubs and lianas, rarely small trees,
Stamens 6-many, free. Ovary 3-20-locular with
often epiphytic. Leaves sessile to petiolate, occa-
numerous ovules per locule. Stigma capitate to
sionally asymmetrical. Inflorescences spicate
umbonate. About 60 spp. in Central and South
with (35-) 100-450 flowers. Pedicels absent (rarely
America and the West Indies.
short). Flowers brilliantly coloured. Nectaries peti-
Two subgenera (sections according to Gilg &
olate, inserted to the inflorescence axis at flower
Werdermann) are recognised: subgen. Marcgravia
base (at least flowers of the upper portion of inflo-
(=subgen. Plagiothalamium Wittm.) with flowers
rescence), cup- or sac-shaped. Sepals 5. Petals 5,
deflexed on the pedicels and stalked leaves, and
free or slightly basally connate. Stamens (6-)8-25;
subgen. Orthothalamium Delpino with flowers
filaments free or variously connate or adnate to
erect on the pedicels and mostly (sub)sessile
corolla. Ovary completely 2-, 3- (5-)locular with
leaves.
4-8(-12) ovules per locule; style lacking. About
10 spp., from Honduras through the Andean
Cordillera to N Bolivia, Guayana Highlands.
6. Marcgraviastrum (Wittm. ex Szyszyl.) Anderberg, A.A. et al. 2002. See general references.
de Roon & S. Dressler Angely, J. 1977. Flora descritiva do Parana, ed. 2, 10 vols. Sao
Paulo: Centro de Pesquisas Basicas e Instituto Paranaense
Marcgraviastrum (Wittm. ex Szyszyl.) de Roon & S. Dressler, de Botfmica.
Bot. Jahrb. Syst. 119: 332 (1997); Bedell, Generic rev. Marc- Bailey, I.W. 1922. The pollination of Marcgravia: a classical
graviac. I. Norantea complex: 296-420 (1985), rev. case of ornithophily? Am. J. Bot. 9: 370-384.
Norantea subgen. Byrsophyllum Delpino (1869). Bedell, H. G. 1985. A generic revision of Marcgraviaceae I. The
Norantea sub sect. Marcgraviastrum Wittm. ex Szyszyl. ( 1893 ). Norantea complex. Ph.D. Dissertation. College Park, MD:
University of Maryland.
Behnke, H.-D. 1991. See general references.
Sprawling shrubs or lianas, often epiphytic. Leaves Davis, G.L. 1966. See general references.
sessile to petiolate. Inflorescences umbelliformly Davis, W.E., Yost, J.A. 1983. Ethnobotany of the Waorani of
contracted racemes with (2-)5-14( -22) flowers. Eastern Ecuador. Bot. Mus. Leafi. 29: 159-211.
Flowers erect on pedicels, subtended by sessile, Delpino, F. 1869. Rivista monografica della famiglia delle
rarely stalked, pendulous, saccate to tubular Marcgraviaceae. Nuovo Giorn. Bot. Ital. 1: 257-290.
Descourtilz, M.E. 1827. Flore medicale des Antilles. 8 vols.
nectaries attached to the lower part of the pedicel. Paris: Pichard.
Sepals 5. Petals 5, free to variously connate. Ditsch, F., Barthlott, W. 1994. Mikromorphologie der Epicuti-
Stamens 12-many; filaments mostly free with cularwachse und die Systematik der Dilleniales, Lecythi-
the outer whorl basally adnate to corolla. Ovary dales, Malvales und Theales. Trop. Subtrop. Pflanzenwelt 88.
(in)completely 5-9-locular with numerous ovules Dressler, S. 1997. 321. Marcgravia umbellata. [Curtis's] Bot.
Mag., ser. 6, 14: 130-136.
per locule. Fifteen spp.; Nicaragua along Andean Dressler, S., Tschapka, M. 2002. Bird versus bat pollination in
Cordillera to Peru and Surinam, 2 spp. E Brazilian the genus Marcgravia and the description of a new species.
Shield. [Curtis's] Bot. Mag. VI, 19: 104-114.
Fryns-Claessens, E., van Cotthem, W. 1973. A new classification
of the ontogenetic types of stomata. Bot. Rev. (Lancaster) 39:
7. Marcgravia L. 71-138.
Gilg, E., Werdermann, E. 1925. Marcgraviaceae. In: Engler &
Marcgravia L., Sp. Pl. 1: 503 (1753); de Roon, Ann. Missouri Prantl, Die natiirlichen Pfianzenfamilien, ed. 2, vol. 21.
Bot. Gard. 57: 39-50 (1970), reg. rev. Leipzig: W. Engelmann, pp. 94-106.
Marcgraviaceae 265
Graham, A., Jarzen, D.M. 1969. Studies in neotropical paleo- Roon, A. C. de 1970. Flora of Panama, VI. 121. Marcgraviaceae.
botany. 1. The Oligocene communities of Puerto Rico. Ann. Ann. Missouri Bot. Gard. 57: 29-50.
Missouri Bot. Gard. 56: 308-357. Roon, A. C. de 1975. Contributions towards a monograph of the
Hegnauer, R. 1969. See general references. Marcgraviaceae. Ph.D. Thesis. Utrecht.
Hofmann, H.L. 1884. Untersuchungen tiber fossile Kiesel- Roon,A.C. de, Dressler, S.l997. New taxa of Norantea Aubl. s.l.
holzer. Diss. Univ. Leipzig. Z. Naturwiss. 57: 156-195. (Marcgraviaceae) from Central America and adjacent South
Howard, R.A. 1970. The ecology of an elfin forest in Puerto America. Bot. Jahrb. Syst. 119: 327-335.
Rico. X. Notes on two species of Marcgravia. J. Arnold Arbor. Saleh, N.A.M., Towers, G.H.N. 1974. Flavonol glycosides of
51: 41-55. Norantea guianensis flowers. Phytochemistry 13: 2012.
Johri, B.M. et al. 1992. See general references. Savolainen, V. et al. 2000. See general references.
]olivet, P. 1996. Ants and plants. An example of coevolution. Sazima, M., Sazima, I. 1980. Bat visits to Marcgravia
Leiden: Backhuys Publishers. myriostigma Tr. et Pl. (Marcgraviaceae) in southeastern
Juel, H.O. 1887. Beitrage zur Anatomie der Marcgraviaceae. Brazil. Flora 169: 84-88.
Kongl. Svenska Vetenskapsacad. Handl. 12, Afd. 3, Nr. 5, Sazima, 1., Buzato, S., Sazima, M. 1993. The bizarre inflores-
Bihang: 1-28. cence of Norantea brasiliensis (Marcgraviaceae): visits of
Machado, I.C., Lopes, A.V. 2000. Souroubea guianensis Aubl.: hovering and perching birds. Bot. Acta 106: 507-513.
quest for its legitimate pollinator and the first record of Schofield, E.K. 1968. Petiole anatomy of the Guttiferae and
tapetal oil in the Marcgraviaceae. Ann. Bot. II, 85: 705-711. related families. Mem. New York Bot. Gard. 18: 1-55.
Melchior, H. 1924. trber das Vorkommen von Inulin in den Schultes, R.E., Raffauf, R.F. 1990. The healing forest. Portland:
Blattern der Marcgraviaceen. Ber. Deutsch. Bot. Ges. 42: Dioscorides Press.
198-204. Tschapka, M., von Helversen, 0. 1999. Pollinators of syntopic
Merz, K.S. 1991. Untersuchungen an tropischen epiphytischen Marcgravia species in Costa Rican lowland rainforest: bats
Lebensgemeinschaften: Biomasse, Wasserretention, Natur- and opossums. Plant Biol. 1: 382-388.
stoffe. Dissertation. Heidelberg: Universitat Heidelberg. Vogel, S. 1958. Fledermausblumen in Siidamerika. Oesterr. Bot.
Muller, J. 1981. Fossil pollen records of extant angiosperms. z. 104: 491-530.
Bot. Rev. (Lancaster) 47: 1-142. Vogel, S. 1993. Floral adaptive radiation in the Marcgraviaceae
Nandi, O.I., Chase, M.W., Endress, P.K. 1998. A combined (abstr.). Sprengel Symposium Berlin-Spandau 1993: 21.
cladistic analysis of angiosperms using rbcL and non- Weber, H. 1955. Haben die Marcgraviaceen "Inulinblatter"?
molecular data sets. Ann. Missouri Bot. Gard. 85: 137-212. Ber. Deutsch. Bot. Ges. 68: 408-412.
Punt, W. 1971. Pollen morphology of the genera Norantea, Weber, H. 1956. trber die Bliitenstande und die Hochblatter
Souroubea, and Marcgravia (Marcgraviaceae). Pollen Spores von Norantea Aubl. (Marcgraviaceae). Beitr. Biol. Pflanzen
13: 199-232. 32: 313-329.
Putz, F.E., Mooney, H.A. (eds.) 1991. The biology of vines. Zusi, R.L., Hamas, M.J. 2001. Bats and birds as potential polli-
Cambridge: Cambridge University Press. nators of three species of Marcgravia Hanas on Dominica.
Roon,A.C. de 1967. Foliar sclereids in the Marcgraviaceae.Acta Caribbean J. Sci. 37: 274-278.
Bot. Neerl. 15: 585-628.
Myrsinaceae
B. STAHL and A.A. ANDERBERG
Myrsinaceae R. Br., Prodr.: 399 (1810). fruits sometimes ruminate; embryo straight to
Aegicerataceae Blume (1833). curved, hypocotyl short or long, cotyledons short
Coridaceae J. Agardh, Theoria Syst. Pl.: 332 (1858).
and narrow.
An almost cosmopolitan family of 49 genera and
Perennial or sometimes annual herbs, subshrubs, about 1500 species.
shrubs, trees or lianas; secretory cavities with a red
or brownish content often present in vegetative CHARACTERS OCCURRING IN RELATIVELY FEW
and reproductive parts; indumentum of multicel- GENERA AND SPECIES. Anthers transversely
lular, branched or unbranched trichomes, peltate septate in Aegiceras and a few species
of Ardisia.
and capitate trichomes common. Leaves simple, Flowers apetalous in Glaux, choripetalous
in
exstipulate, alternate, sometimes opposite or Embelia and Heberdenia; zygomorphic in Caris;
whorled, if alternate then sometimes in a basal 3-merous in Cybianthus subg. Triadaphara and
rosette or condensed into pseudowhorls; margin Pelletiera; 7-merous in Trientalis. Fruits vivipa-
entire or variously serrate. Inflorescences of termi- rous in Aegiceras; operculate in Anagallis.
Leaves
nal or axillary racemes, if axillary then sometimes with subepidermal fibre strands in Cybianthus
reduced into fascicles or solitary flowers. Flowers subg. Triadaphara. Most species of
Embelia a~d
5-merous, sometimes 3- (Cybianthus subg. Tri- Grenacheria scandent. Tuberous hypocotyl m
adaphara, Pelletiera), 4- (e.g. Cybianthus), or 7-9- Cyclamen. Grammadenia is mostly
epiphytic.
merous ( Trientalis ), hermaphrodite or unisexual, if
unisexual then plants mostly dioecious; calyx VEGETATIVE MORPHOLOGY. Most temperate
herbaceous, rotate to campanulate, rarely hypocra- genera are perennial herbs, although some are
teriform; corolla rotate, campanulate, urceolate or annuals (Asteralinan, Pelletiera, some Anagallis)
rarely absent (Glaux ), if rotate then often divided or subshrubs (Caris and some Lysimachia),
to near base or with free lobes; stamens in one whereas the tropical genera are mostly
shrubs or
whorl, antepetalous, filaments partly to entirely small to medium-sized trees. Among the tropical
fused with the corolla, sometimes united to form a groups, however, some are herbaceous
represent.a-
temporary or permanent tube; anthers tetraspo- tives, such as Labisia and Ardisia subg. Bladhza.
rangiate, dithecal, introrse, opening by longitudi- Some tropical Myrsinaceae are pachycaul (e.g.
nal slits or apical pore-like slits, rarely transversely Badula, Discacalyx). In Cyclamen, the hypocotyl
septate, free or rarely fused, if free then often con- develops into a large tuber.
nivent and forming a protruding cone; gynoecium The leaves are mostly alternate, and in some
syncarpous, 5-carpellate; ovary superior, unilocu- groups they are condensed into pseudowhorls at
lar, with a free central placental column; style short shoot apices. Opposite leaves characterise Aster-
to long, sometimes exserted; stigma punctate, alinan and Pelletiera as well as several species of
truncate to capitate, or rarely discoid; ovules few to Anagallis and Lysimachia; some species of Lysi-
many, in one to several series and often immersed machia have whorled leaves. In most taxa the
into the placenta, anatropous and bitegmic, or leaves are short-petiolate, an exception being
campylotropous and unitegmic (Cyclamen), ten- Grammadenia with sessile leaves and Ardisiandra
uinucellate. Fruits fleshy, often berry-like drupes and Cyclamen with mostly long-petiolate leaves. In
(mainly tropical groups) or capsules opening with the latter two genera the leaf blade is distinct from
valves, a circular lid, or by disintegration (mainly the petiole and has a palmate venation. Otherwise,
temperate groups), rarely viviparous (Aegiceras). most members of the family have oblanceolate to
Seeds of capsular fruits mostly angular, papillose, oblong leaf blades. The leaf margins vary from
reticulate or rugose, those of drupaceous fruits entire to serrulate or serrate. In Amblyanthus,
subglobose, with or without immersed hilum area; Amblyanthapsis and Ardisia subg. Crispardisia,
endosperm abundant, in seeds from drupaceous the leaf margins are crenulate, a feature related to
Myrsinaceae 267
bacterial symbiosis (Lersten and Horner 1976). Maesaceae does not occur in the Myrsinaceae. In
The leaves of many species of Cyclamen have car- lignified Hawaiian species of Lysimachia the wood
tilaginous outgrowths marginally, but these are lacks rays, but has septate fibres with small simple
not homologous with the leaf-teeth of other pits (Carlquist 1992).
genera. The young, developing leaves are typically
folded inwards (involute or conduplicate), except INFLORESCENCE STRUCTURE. The inflorescence
in Ardisiandra which has revolute vernation. in the Myrsinaceae is basically racemose, being
developed as a terminal or axillary raceme, spike,
VEGETATIVE ANATOMY. Conspicuous secretory compound raceme, pseudo-umbel or fascicle (Mez
cavities, appearing as reddish brown or black dots 1902). In addition, several herbaceous genera and
and streaks, are present in vegetative and repro- species have solitary flowers in the leaf axils. In
ductive parts of virtually all tropical Myrsinaceae, contrast to Maesaceae, prophylls are lacking.
but occur also in Caris and several species of Lysi-
machia. Extraxylary foliar bundles of unbranched FLOWER STRUCTURE. The flowers are hypogy-
fibres like those of Theophrastaceae are lacking nous and obhaplostemonous. The corolla is
in the Myrsinaceae, but sclereids arranged in a usually open and rotate or campanulate (see Fig.
similar way adjacent to the leaf epidermis occur in 91). A few genera deviate with respect to corolla
Cybianthus sub g. Triadaphara (Grosse 1908). Stone shape: Caris with a two-lipped, zygomorphic
cells occur in the mesophyll of many tropical corolla, and Emblemantha, Sadiria and Synardisia
members of the family, being particularly with urceolate corollas. Generally the corolla is
common in Tapeinasperma. sympetalous, but in Embelia and Heberdenia the
Multicellular trichomes occur in great variety, lobes are distinct and in many other genera they
both glandular and non-glandular (Grosse 1908). are united at the very base only. Asteralinan and
Immersed trichomes with a short, one- or few- Pelletiera have inconspicuous petals which are
celled stalk and multicellular head are common in much shorter than the calyx. In Glaux, the corolla
tropical groups, a feature shared among related is lacking and the synsepalous calyx is pink and
families only by Theophrastaceae. Likely homolo- showy. Usually, corolla aestivation is either imbri-
gous to these immersed trichomes are the scales cate, sometimes quinquncial, or twisted. However,
present in several tropical genera. In Cybianthus Lysimachia subg. Seleucia, Parathesis, and many
subg. Canamarpha, these scales often cover major species of Cybianthus subg. Canamarpha have
parts of the plant giving it a brownish colour. valvate corollas. Labisia has a conduplicate corolla
Small but protrusive, capitate trichomes are also aestivation with the margins of each lobe clasping
present on the inner surface of the corolla in many one of the anthers. Generally the corolla lobes are
woody Myrsinaceae, being particularly abundant entire but emarginate lobes occur in flowers of
on the corollas of Embelia, Heberdenia, Laheria, Amblyanthus, Discacalyx, Manaparus, Oncaste-
Pleiameris and many Cybianthus, in which the mum, and Wallenia. In most genera, the corolla
petal lobes often are more or less papillose. Artic- lobes are spreading or erect but in Aegiceras,
ulated hairs are found in Ardisiandra and temper- Cyclamen, Geissanthus and many Ardisia the
ate genera except Cyclamen, and non-articulated, lobes are reflexed at anthesis, and in Parathesis
unbranched or irregularly branched trichomes in they are recurved.
many other genera. Parathesis is characterised by The stamens, which develop from the same pri-
stellate trichomes, which occur on both young mordium as the petals (Sattler 1962), are placed in
shoots and floral parts. Short-lived, secretory front of these. In many genera the stamens are
trichomes in the buds of some Ardisia subg. Cris- more or less free and connate only at the very base
pardisia evidently function as colleters and are near the receptacle, whereas in others the fila-
important for the bacterial symbiosis in this group ments are partly or entirely fused. In addition, the
(Lersten 1977). filaments or filament ring are often fused with the
According to Metcalfe and Chalk (1950), the corolla for a longer or shorter distance. Several
wood is characterised by rather small vessels genera have more or less exserted stamens but this
arranged in radial multiples of 3 or 4 cells and in depends also on how much the petals are reflexed.
irregular groups; perforation plates are simple or In some cases this feature is combined with the
scalariform. The parenchyma is paratracheal and occurrence of versatile anthers, e.g. in Aegiceras
usually sparse. The rays are typically multiseriate and Wallenia. In Cyclamen, Lysimachia, Ardisia
and although some uniseriate rays may occur, the and several other genera, the filaments are shorter
mixture of uni-and multiseriate rays present in than the anthers, which are connivent to form a
Fig. 91. Myrsinaceae. A Embelia philippinensis, flower. B Wal- narrowly (Cyclamen, Hymenandra and some
lenia laurifolia, flower. C Heberdenia bahamensis, flower. D species of Ardisia) or triangularly sagitatte.
Parathesis cubana, flower. E Myrsine africana, flower. F
Aegiceras corniculatum, flower. GOncostemum sp., flower. H, I
Staminode structures present in related families
Ardisia crenata. H Inflorescence. I Fruit. J, K Lysimachia are lacking in Myrsinaceae. The alleged stamin-
nemorum. J Habit. K Flower. L, M Ardisiandra wettsteinii. L odes of Lysimachia subg. Seleucia (= Stieronema)
Fruit. M Habit. N, 0 Cyclamen hederifolium, unopened and lack vascular strands and may be protuberances
dehisced fruit. P, Q Anagallis arvensis, same. Drawn by rather than staminodes.
Pollyanna von Knorring
The ovary is unilocular and evidently syncar-
pous, although it is initiated from a ring-shaped
primordium. The ovules vary from few to numer-
prominent cone. In Amblyanthus, Conandrium ous and are arranged in one or several series on
and Hymenandra, the anthers are laterally connate the placenta; they are immersed into the placental
to form a permanent cone surrounding the ovary. tissue, except in Ardisiandra. The stigma is punc-
In Oncostemum, both the filaments and anthers tate, truncate or capitate, rarely discoid. In the
are fused to form a ring-like, tubular structure. short -styled genera Myrsine and Rapanea, the
The anthers are introrse, have a fibrous endothe- stigma is often irregularly shaped, e.g. fimbriate
cium and usually open with longitudinal slits. In or ligulate.
some groups (e.g. Grammadenia, Monoporus, and
species of Lysimachia) the anthers open by apical EMBRYOLOGY. Although data from many, partic-
pores. Aegiceras and some Asiatic Ardisia (Stone ularly tropical groups are lacking, the embryology
1993) have transversely septate, locellate anthers. of the Myrsinaceae seems to agree largely with
The anthers vary in shape from short, globose or that of other primulalean families (Davis 1966;
almost square (e.g. Asterolinon, Cybianthus) to Johri et al. 1992). The anther tapetum is secretory
Myrsinaceae 269
with uninucleate cells. Pollen is binucleate when members seem to have dry, nectarless flowers with
shed. The ovules are anatropous, tenuinucellate only pollen offered as a reward. A structure similar
and usually bitegmic with the micropyle formed to a nectar disc is present in Ardisiandra but does
by both integuments. The inner integument forms not seem to produce any nectar. A unique feature
an endothelium, except in Caris, Cyclamen, is the oil-producing glandular hairs on the petals
Aegiceras, Ardisia, and Myrsine. The archespore is and stamen filaments of certain Lysimachia
multicellular and the embryo sac is of the Poly- species. These species do not produce nectar, and
gonum type with ephemeral antipodes, but lacks Vogel ( 1986) showed that they are pollinated by a
endosperm haustoria. The seeds are albuminous specific genus of oil-collecting bees, Macropis.
with nuclear endosperm formation, excepting Some genera have flowers with a protruding
Aegiceras which has viviparous fruits and exalbu- anther cone and these are likely to be buzz-
minous seeds. Embryogeny follows the caryophyl- pollinated by pollen-gathering bees, as was shown
lad type except in Trientalis, which seems to have by Pascarella (1997a) for Ardisia escallonioides.
an onagrad type of embryogeny. The embryo is Autogamy has been reported for several species
short, or long and curved, the latter condition of Ardisia (Pascarella 1997b), and at least Cycla-
being found in taxa with drupaceous fruits; the men shows an increased tendency for selfing if not
cotyledons are generally short and narrow. The visited by pollinators. Anemophily has been
embryo is dicotyledonous except in Cyclamen, in reported in Rapanea (Otegui and Cocucci 1999).
which the development of one of the cotyledons is
suppressed. Polyembryony has been recorded in FRUIT AND SEED. With the present circumscrip-
species of Ardisia, Parathesis, and Rapanea (Braun tion, both single-seeded drupes and many- to few-
1859; Anderberg and Stahl1995). seeded capsules are present in the Myrsinaceae.
Aegiceras and Cyclamen differ from other Drupaceous fruits characterise all woody tropical
genera by having unitegmic ovules. However, and subtropical groups except Aegiceras, the fruits
during development the inner integument is of which are viviparous. When ripe, the drupes are
partly resorbed, at least in Cyclamen, indicating red to black and provided with a variously thick
that this feature is a reduction from the prevailingand juicy mesocarp and a woody but usually
condition in the family. brittle endocarp. They are often somewhat com-
pressed and usually less than 1 em wide. Some-
POLLEN MORPHOLOGY. The pollen grains of what larger fruits occur in Elingamita and
Myrsinaceae are oblate-spheroidal to almost Tapeinosperma, and in the latter genus the meso-
spherical, and 10-45 f.lm in diameter (Erdtman carp is more or less fibrous. In Fittingia, which also
1952). Most taxa have 3-colporate pollen but has comparatively large fruits, the soft mesocarp
4-colporate grains have been reported from forms a distinct wing in herbarium specimens.
Hawaiian Lysimachia (subg. Lysimachiopsis), More or less globose capsules characterise the
Myrsine, and Rapanea, and pantocolporate grains remaining, chiefly herbaceous genera. In most
are known from Lysimachia subg. Idiophyton species the capsules open with apical valves or
(Nowicke and Skvarla 1977; Stahl1996; Hope 2000). crumble irregularly, but in Anagallis the capsules
open with a circular lid. The valvate capsule in
KARYOLOGY. Particularly the herbaceous Trientalis has a papery wall that falls off early,
members of the Myrsinaceae show a great vari- leaving the seeds on the placenta. A similar capsule
ability in chromosome number, but numbers is found also in certain species of Lysimachia.
based on x = 9, 10, 11, and 12 are common. At least The seeds of all Myrsinaceae except Aegiceras
in Lysimachia and Trientalis polyploidy seems to have copious endosperm. Seeds in taxa with dru-
be frequent. In woody, tropical members of the paceous fruits are more or less globose and have
family 2n = 46 is by far most commonly reported an immersed hilum area forming a concavity at
chromosome number. However, 2n = 96 reported the base of the seed; the testa is thin and devoid of
in some species of Ardisia (Faure 1968) suggests crystals and the endosperm is often ruminate with
some variability in the woody groups as well. irregularly thickened cells walls (Anderberg and
Stahl 1995). In genera with capsular fruits the
POLLINATION AND REPRODUCTIVE SYSTEMS. seeds are many, except in Asterolinon and Pel-
Most Myrsinaceae appear to be pollinated by letiera which have few seeds; the seeds are nor-
insects, notably bees and flies. However, although mally angular with smooth, reticulate or papillose
species of Anagallis and Elingamita have nectar- surfaces, and the hilum is more or less flush with
producing trichomes on floral parts, most the surface; the testa is rather thick, usually dis-
tinctly two-layered and generally provided with The present circumscription of Myrsinaceae is
rhomboid crystals. The seeds of Asterolinon based on phylogenetic research by Anderberg and
linum-stellatum and Pelletiera deviate in being Stahl (1995),Anderberg et al. (1998), Kallersjo et al.
coarsely rugose, and by having a distinctly im- (2000), and Anderberg et al. (2001) and differs in
mersed, concave hilum area. Seeds of Trientalis two important aspects from most previous classi-
have a white outer reticulum covering the seeds. fications. Firstly, the genus Maesa, which often has
Some Lysimachia, e.g. L. vulgaris, have an outer been treated as a subfamily of Myrsinaceae, had to
porous, almost spongy tissue covering the seed be transferred to a family of its own (Anderberg et
surface, and in Neotropical Lysimachia subg. al. 2000), and secondly, several genera previously
Theopyxis the seeds are winged (Stahl 1990). The treated as members of Primulaceae have been
endosperm is composed of cells with smooth and included in Myrsinaceae as circumscribed here.
evenly thickened walls except in Cyclamen, which It should also be noted that Coris and Aegiceras,
has irregularly thickened endosperm cell walls two genera that in some earlier classifications
provided with distinct narrow constrictions. were treated in separate families, Coridaceae
and Aegicerataceae respectively, are now included
DISPERSAL. Most groups with drupaceous fruits in Myrsinaceae. In the classification of Primu-
are probably dispersed by birds and other fruit- laceae by Pax and Knuth (1905), the genera now
eating animals. In groups with capsular fruits transferred to Myrsinaceae belonged to four dif-
the seeds are released ballistically through the ferent tribes: Androsaceae (Ardisiandra, Stimpso-
opening of the capsule. However, in most Cycla- nia), Lysimachieae (Anagallis, Asterolinon, Glaux,
men the fruit is pulled to the ground by coiling of Lysimachia, Pelletiera, Trientalis), Cyclamineae
the pedicel, and upon drying the capsule opens (Cyclamen), and Corideae (Coris ). In the analysis of
near the ground to expose the seeds, the coat of Kallersjo et al. (2000), based on a combination of
which contains edible substances that attract morphological and cpDNA sequence data, Coris is
dispersing insects. placed as sister to the rest of Myrsinaceae, with
Ardisiandra being placed as the sister group to a
REPRODUCTIVE BIOLOGY. Most Myrsinaceae more inclusive group formed by three clades: (1)
have perfect, bisexual flowers, but many of the the genera of the erstwhile Lysimachieae, (2) Cycla-
tropical and subtropical woody genera are dioe- men, and (3) the genera of Myrsinaceae s. str.
cious. In addition, due to incomplete separation of (except Maesa but including Aegiceras ). The analy-
sexes or incomplete reductions of reproductive ses by Kallersjo et al. (2000) suggest that both
organs, many woody genera are often referred to Anagallis and Glaux are nested within Lysimachia
as polygamous or cryptically dioecious. Vegetative as presently circumscribed. Among woody tropical
propagation is known in many species of Anagal- Myrsinaceae, generic alignments are also rather
lis, Ardisia subg. Bladhia and Lysimachia, which unclear, although many genera with few ovules in
are prostrate and root at the nodes. Propagation by a single series on the placenta tend to appear in the
means of root suckers has been reported in same general clade (Stahl1996).
Myrsine (Burrows 1999), and may also occur in
other shrubby taxa. Trientalis propagates with DISTRIBUTION AND HABITATS. Myrsinaceae have
rhizomes ending in a bulb-like winter bud, and a worldwide distribution. The herbaceous genera
a similar mode is known from Lysimachia are distributed mainly in (cool) temperate regions,
thyrsiflora and Glaux. whereas the woody genera are mainly tropical.
However, several herbaceous or suffrutescent rep-
PHYTOCHEMISTRY. Similarly to the other primu- resentatives occur in tropical or subtropical areas,
lalean families, most Myrsinaceae produce and many woody genera show high species diver-
saponins. Calcium oxalate in the form of druses sity in tropical-montane habitats. Coris and Cycla-
and simple crystals are present in the leaves of men are chiefly distributed in the Mediterranean
most tropical woody genera. The seeds store oil region and North Africa; Glaux is confined to
and amylose, but no starch (Hegnauer 1969). grassy seashores in the North Temperate zone. The
largest genera, Ardisia and Rapanea, are pantrop-
SUBDIVISION AND RELATIONSHIPS WITHIN THE ical, occurring in lowland as well as in montane
FAMILY. Myrsinaceae form the largest family of forests. Some genera have more restricted distri-
the order Primulales of earlier classifications (e.g. butions, like Heberdenia and Pleiomeris, endemic
Cronquist 1988). It is sister to Primulaceae, the two to Macaronesia, Sadiria (Assam-Bhutan), Solonia
being the sister group of Theophrastaceae. (Cuba), and Vagaea (Hispaniola). Among the
Myrsinaceae 271
genera with regionally restricted distributions - Style shorter than or slightly longer than the ovary, not
should be mentioned Ardisiandra (East Africa), slender; stigma usually truncate, capitate, or discoid 33
18. Filaments (or filament tube) longer than the anthers 19
Wallenia (West Indies), and Oncostemum (Mada- - Filaments shorter or as long as the anthers 23
gascar). The most widely distributed species is 19. Anthers versatile 20
found in Anagallis, viz. A. arvensis and A. - Anthers not versatile 22
foemina, two weedy, cosmopolitan species with a 20. Fruits viviparous, mangrove plants 32. Aegiceras
European/Mediterranean origin, A. pumila, with - Fruits not viviparous 21
21. Filaments fused into a tube, free towards apex
a pantropical (submontane) distribution, and 22. Solonia
A. minima which is cosmopolitan. - Filaments not fused 23. Geissanthus
22. Flowers bisexual; ovules in two or several series
FossiL RECORD. Pollen of the Myrsine type 26. Gentlea
- Flowers unisexual; ovules 1-seriate 27. Stylogyne
appears in the Oligocene of Australia and New 23. Corolla tube about as long as the lobes, or longer 24
Zealand (Mildenhall1980; Martin 1994). - Corolla tube much shorter than the lobes 26
24. Ovules in two or more series; style very slender
KEY TO THE GENERA (Mesoamerica) 19. Synardisia
- Ovules 1-seriate; style comparatively thick (Asia) 25
1. Herbs or subshrubs, rarely shrubs; fruit dry, capsular; 25. Subshrubs; leaf blades decurrent 24. Emblemantha
seeds usually angular 2 - Shrubs or small trees; leaf blade not decurrent
- Shrubs, trees or Hanas, sometimes subshrubs; fruit± fleshy, 25. Sadiria
drupaceous (or rarely viviparous); seeds subglobose 11 26. Corolla lobes valvate in bud, stellate- or dendroid-
2. Flowers with corolla and calyx 3 pubescent 31. Parathesis
- Flowers with calyx only 8. Glaux - Corolla lobes imbricate or contorted in bud, not stellate-
3. Flowers regular, actinomorphic 4 or dendroid-pubescent 27
- Flowers two-lipped, distinctly zygomorphic 1. Coris 27. Corolla twisted to the left 30. Antistrophe
4. Corolla-lobes conspicuously reflexed 10. Cyclamen - Corolla twisted to the right 28
- Corolla -lobes spreading or erect, never reflexed 5 28. Flowers 4-merous; ovules 1-seriate 21. Tetrardisia
5. Capsule opening with a circular lid 9. Anagallis - Flowers typically 5-merous; ovules in one or more series
- Capsule opening with valves or irregularly 6 29
6. Corolla much smaller than calyx 7 29. Ovules 1-seriate, erect at the base of the placenta 30
- Corolla equal to, or larger than calyx 8 - Ovules in two or more series, if 1-seriate then with ovules
7. Petals and capsule five-merous 6. Asterolinon inserted around the middle of the placenta 31
- Petals and capsule three-merous 7. Pelletiera 30. Inflorescences long-pedunculate, ovules 10-15
8. Flowers 7-9-merous; flowers white; leaves in a terminal 28. Ctenardisia
pseudowhorl; seeds with white reticulum, persistent - Inflorescences without long peduncles; ovules 6-8
5. Trientalis 29. Yunckeria
- Flowers usually 5-merous; flowers white, pink or yellow; 31. Anthers connate, forming an anther tube
leaf arrangement variable; seeds not with loose white 20. Hymenandra
reticulum 9 - Anthers often coherent at the beginning of anthesis but not
9. Leaves coarsely dentate 10 connate 32
- Leaf margins entire 4. Lysimachia 32. Ovules 1-seriate; inflorescences of axillary, bracteate clus-
10. Flowers in axillary fascicles 3. Ardisiandra ters (African) 18. Afrardisia
- Flowers solitary in the shoot apex 2. Stimpsonia - Ovules mostly in two or more series; inflorescences axil-
11. Lianas or scandent shrubs 12 lary or terminal, usually appearing as panicles
- Plants not scandent 13 17. Ardisia
12. Corolla lobes free to base 11. Embelia 33. Free parts of filaments as long as or longer than the
- Corolla lobes united to form a distinct tube anthers, at least in male flowers 34
12. Grenacheria - Free parts of filaments shorter than the anthers, or anthers
13. Flowers arranged in fascicles 14 sessile or fused 39
- Flowers arranged in panicles, racemes or umbel-like 34. Leaf margins crenate; petals emarginate 35
inflorescences 17 - Leaf margins entire or serrate to serrulate; petals emar-
14. Ovules many, in 2-4 series on the placenta (Macaronesia) ginate or not 36
15 35. Anthers connate, forming a cone 33. Amblyanthus
- Ovules few, in a single series on the placenta 16 - Anthers coherent but not connate 34. Amblyanthopsis
15. Corolla lobes free to base 13. Heberdenia 36. Corolla cup-shaped, shorter than the calyx
- Corolla lobes united to form a distinct tube 35. Elingamita
14. Pleiomeris - Corolla with well-developed lobes, longer than the calyx
16. Style well demarcated; stigma discoid or discoid-fimbriate 37
15.Myrsine 37. Corolla usually ± papillose within, tube shorter than the
- Style very short and inconspicuous; stigma variously lobes 38
shaped (ligulate, capitate), but not discoid 16. Rapanea - Corolla not papillose within (or near base only), tube as
17. Style two to several times longer than the ovary, often long as lobes or longer 36. Wallenia
slender; stigma mostly punctate (without obvious stig- 38. Flowers borne in panicles on short lateral shoots
matic surface) 18 subtended by numerous bracts (Malesia) 37. Loheria
- Inflorescences variable but not as above (Neotropics) ovate, with articulated hairs. Flowers solitary in
38. Cybianthus p.p. the leaf-axils along the stem; calyx campanulate;
39. Ovules 2- or 3-seriate, anthers connate 49. Conandrium
- Ovules 1- or rarely 2-seriate, and if 2-seriate, then anthers corolla white, hypocrateriform with a short tube,
not connate 40 hairy; corolla-lobes spreading, emarginate; sta-
40. Leaves sessile; epihytic shrubs 39. Grammadenia mens adnate to the tube; stigma capitate. Capsule
- Leaves petiolate; terrestrial plants 41 globose, opening with apical valves; seeds many,
41. Plants suffrutescent or herbaceous; petals conduplicate
and largely concealing the anthers
reticulate. One sp., S. chamaedryoides Wright, Asia.
45. Labisia
- Plants woody; petals not conduplicate 42
42. Leaves sclerophyllous, linear (Hispaniola) 40. Vegaea
- Leaves not linear 43
3. Ardisiandra Hook. f. Fig. 91L, M
43. Filaments and anthers united into a distinct ring or tube Ardisiandra Hook. f., J. Proc. Linn. Soc. Bot. 7: 205 (1864);
41. Oncostemum Rosvik, Arb. Univ. Bergen Ser. Mat.-Nat. 7: 1-15 (1969).
- Stamens not united into a tube 44
44. Flowers bisexual 45
- Flowers unisexual 47 Perennial herbs. Stem prostrate, with copious
45. Anthers sessile or subsessile 42.Badula articulated hairs. Leaves alternate, petiolate,
- Stamens with well-developed filaments 46 coarsely dentate, revolute when young. Flowers
46. Anthers dorsifixed, style longer than the ovary 5-merous, solitary or in few-flowered fascicles in
43. Tapeinosperma
- Anthers broadly basifixed, style shorter than the ovary
the leaf-axils; calyx campanulate, lobes cordate;
44. Discocalyx corolla campanulate, white or yellow; corolla-lobes
47. Corolla papillose on inside 38. Cybianthus p.p. erect, entire, imbricate in bud; stamens connivent,
- Corolla not papillose on inside 48 filaments basally connate into a ring, anthers sagit-
48. Flowers 4-merous, borne on short, densely bracteate, axil- tate forming a cone; style protruding from the
lary inflorescences 46. Systellantha
- Flowers 4- or 5-merous, arranged in ± lax panicles or anther cone; stigma subcapitate. Capsule sub-
racemes 49 globose, often thin-walled; seeds many, papillose.
49. Anthers opening by one or two apical pores 2n = 34. Three spp., E African mountains.
47. Monoporus
- Anthers opening by longitudinal slits 50
50. Fruit with spongy or fleshy exocarp (fruits "winged" in 4. Lysimachia L. Fig. 91J, K
herbarium specimens); endocarp longitudinally ridged
48. Fittingia Lysimachia L., Sp. Pl.: 146 (1753); Handel-Mazzetti, Not. Roy.
- Fruit with thin exocarp; endocarp smooth Bot. Gard. Edinburgh 16: 51-122 (1928), rev.; Ray, Illinois
44. Discocalyx Bot. Monogr. 24: 1-160 (1956), rev. New World spp.; Huynh,
Candollea 25: 267-296 (1970), pollen.
Naumburgia Moench (1802).
GENERA OF MYRSINACEAE
Perennial, biennial or annual herbs, rarely shrubs.

1. Coris L.
Leaves entire, alternate, opposite, or whorled,
Goris L., Sp. Pl.: 177 (1753). sometimes in terminal pseudowhorls. Flowers 5-
merous, in terminal racemes, thyrses, or solitary in
Perennial herbs, woody at the base. Leaves small, the leaf-axils; calyx rotate to campanulate; corolla
alternate, linear, flat or revolute. Flowers zygomor- deeply lobed, rotate to campanulate, white, yellow,
phic, 5-merous, in a terminal spike-like raceme; red, pink, or purple, twisted or valvate in bud;
calyx campanulate, spinose, with conspicuous stamens spreading or connivent and forming a
dark resin dots; corolla pink or white, two-lipped cone; stigma truncate or subcapitate. Capsule
with a short tube; corolla-lobes bifid, imbricate in globose, opening with valves or disintegrating
bud; stamens adnate to the tube; stigma capitate. irregularly; seeds reticulate or papillose, some-
Capsule ovoid, opening with apical valves; seeds times winged, or with outer spongy tissue. 2n = 16,
few, papillose. 2n = 18. Two spp., Mediterranean 20, 24, 28, 30, 32, 34, 36, 40, 42, 56, 60, 68, 72, 84,
region, Somalia. 92, 96, 98, 100, 102, 108, 112. About 150 spp.,
worldwide.
2. Stimpsonia C. Wright ex A. Gray
5. Trientalis L.
Stimpsonia C. Wright ex A. Gray, Mem. Amer. Acad. Arts II, 6:
401 (1858); Rosvik, Arb. Univ. Bergen Ser. Mat.-Nat. 7: 1-15 Trientalis L., Sp. Pl.: 344 (1753).
(1969); Anderberg eta!., Bot. Jahrb. Syst.123: 369-376 (2001).
Perennial herbs. Leaves entire, elliptic, in a termi-
Annual or biennial herb. Leaves in a basal rosette nal pseudowhorl. Flowers generally 7-merous,
and scattered along the stem, dentate, elliptic to
Myrsinaceae 273
solitary in the leaf-axils; calyx rotate; corolla solitary in the leaf-axils or in loose terminal
rotate, deeply lobed, white, twisted in bud; stamens racemes; calyx rotate, deeply lobed; corolla rotate
spreading; stigma truncate. Capsule globose, or campanulate, deeply lobed, blue, red, pink or
valves caducous; seeds with white outer reticulum. white, twisted in bud; stamens free, filaments often
2n = 88, 90, 96, 100, 130, 160. Two spp., North hairy; stigma truncate or subcapitate. Capsule
temperate regions. globose, operculate; seeds papillose. 2n = 20, 22,
40, 66, 75, 80. About 20 spp., Africa, S America,
Europe.
6. Asterolinon Hoffmannsegg & Link
Asterolinon Hoffmannsegg & Link, Fl. Portug. 1: 332
(1813-1820). 10. Cyclamen L. Fig. 91N, 0
Cyclamen L., Sp. Pl.: 145 (1753); Grey-Wilson, The genus
Annual herbs. Leaves opposite, entire, ovate or Cyclamen (1988); Anderberg, Kew Bull. 49: 455-467
lanceolate. Flowers inconspicuous, 5-merous, soli- (1994); Anderberg eta!., Pl. Syst. Evol. 220: 147-160 (2000).
tary in the leaf-axils; calyx rotate, deeply lobed;
corolla rotate, deeply lobed, white; corolla-lobes Perennial herbs with tuberous hypocotyl. Leaves
rotundate, obtuse, twisted in bud; stamens spread- basal, petiolate, cordate, glabrous, rather fleshy,
ing; stigma subcapitate. Capsule globose, disinte- often with pale green or white patterns. Flowers
grating irregularly, or with caducous valves; seeds hypogynous, 5-merous, nodding, axillary, solitary;
few and rugose, or many and papillose. 2n = 40. calyx campanulate; corolla campanulate, white to
Two spp., Europe and N & NE Africa. pink often with purple blotches or stripes; corolla-
lobes entire and distinctly reflexed, twisted in bud;
peduncles coiling at anthesis; stamens connivent,
7. Pelletiera A. St.-Hil. forming a cone; style often exserted; stigma trun-
Pelletiera A. St.-Hi!., Mem. Mus. Hist. Nat. 9: 365 (1822). cate. Capsule globose, opening with apical valves.
Seeds almost smooth. 2n = 20, 22, 30, 34, 48, 68, 72,
Annual herbs. Leaves opposite, entire, lanceolate. 84, 96. 19 spp., S & C Europe, Turkey, Lebanon,
Flowers inconspicuous, solitary in the leaf-axils; N Africa, Somalia.
calyx 5-merous, rotate, deeply lobed; corolla 3-
merous, rotate, deeply lobed, white; corolla-lobes 11. Embelia Burm. f. Fig. 91A
lanceolate, acute; stamens spreading; stigma sub-
capitate. Capsule globose, disintegrating irregu- Embelia Burm. f., Fl. Ind. 62, tab. 23 (1786), nom. cons.; Mez in
larly; seeds few, rugose. Two spp., South America Pflanzenreich IV. 236:295-332 (1902); Taton, Fl.Afr. Central,
Myrsinaceae, pp. 30-49 (1980), reg. rev.; Walker, Philippine
and Macaronesia. J. Sci. 73: 155-184 (1940), rev. E Asian spp.; Perrier de Ia
Bathie, H., Fl. Madagascar 161: 124-137 (1953), reg. rev.;
Smith, J. Arnold Arb. 54: 274-278 (1973), rev. Fijian spp.;
8. Glaux L. Halliday, Fl. Trop. Afr. (no vol.) 11-16 (1984), reg. rev.;
Sleumer, Blumea 32: 385-394 (1987), rev. New Guinean spp.;
Glaux L., Sp. Pl.: 207 (1753).
Stone in Ng, Tree Fl. Malaya 4: 278 (1989), reg. rev.; Chen &
Pipoly, Fl. China 15: 30-34 (1996), reg. rev.
Perennial herb. Leaves small, fleshy, opposite,
broadly triangular, entire. Flowers apetalous, 5- Lianas, sometimes shrubs or small trees. Leaves
merous, solitary in the leaf-axils; calyx campanu- petiolate or rarely subsessile, sometimes distic-
late, pink; corolla absent; stamens spreading; hous, margins crenate or entire. Inflorescences
stigma truncate. Capsule globose, opening with terminal or axillary panicles, racemes or fascicles.
apical valves; seeds few, reticulate. 2n = 30. One Flowers 4-5-merous, hi- or unisexual and then the
sp. (G. maritima L.), North temperate region. plants dioecious; corolla greenish or white, rarely
reddish, rotate, the inside densely papillose, lobes
9. Anagallis L. Fig. 91P, Q free or united at base only, narrowly ovate to
elliptic, imbricate in bud, usually spreading and
Anagallis L., Sp. Pl.: 148 (1753); Taylor, Kew Bull. 3: 321-350 somewhat recurved at anthesis; stamens usually
(1955), African spp.
Centunculus L. (1753 ).
exserted, filaments distally adnate to and decur-
rent on lower part of corolla lobes; anthers dorsi-
Annual or perennial herbs. Leaves ovate to linear, or basifixed, broadly ovoid or oblong, sometimes
opposite or alternate, entire. Flowers 5-merous, recurved at anthesis, opening by longitudinal
slits; ovary subglobose to ovoid, style short or
somewhat longer then the ovary, stigma discoid; fused to corolla tube; anthers basifixed, broadly
ovules few, 1-seriate. Fruit subglobose, sometimes sagittate, opening through longitudinal slits; ovary
depressed; endosperm ruminate or sometimes ovoid, tapering into a rather slender style equal or
smooth. About 130 spp., Old World tropics. slightly longer than the ovary, stigma punctate;
ovules 15-20, 2- or 3-seriate. Fruit globose;
endosperm ruminate. One sp., P. canariensis
12. Grenacheria Mez
(Willd.) A. DC., Macaronesia.
Grenacheria Mez in Pflanzenreich IV. 236: 292-294 (1902);
Sleumer, Blumea 32: 394-396 (1987), rev. New Guinean spp.
15. Myrsine L. Fig. 91E
Scandent shrubs or lianas. Leaves short-petiolate, Myrsine L., Sp. Pl. 196 (1753); Mez in Pflanzenreich IV. 236:
margins entire. Inflorescences axillary racemes 338-342 (1902); Taton, Fl. Afr. Centr., Myrsinac. 49-52
forming synflorescences at stem apices. Flowers (1980), reg. rev.; Halliday, Fl. Trop. E. Afr., Myrsinac. 6-8
(1984), reg. rev.; Larsen & Hu, Fl. Thailand 6(2): 163-164
5-merous, unisexual, the plants dioecious; corolla
( 1996), reg. rev.
whitish, rotate or short-campanulate, densely
papillose on inside, lobes ovate, imbricate in bud;
Shrubs or small trees. Leaves petiolate, margins
stamens included, filaments partly fused with and
serrate or entire. Inflorescences axillary, few-
decurrent on the corolla tube; anthers dorsifixed,
flowered fascicles. Flowers 4-5-merous, unisexual
ovoid, opening by longitudinal slits; ovary ovoid,
or (morphologically) bisexual, plants chiefly dioe-
tapering into a style of ± equal length, stigma
cious; corolla rotate, greenish, white or pink, petals
discoid; ovules few, 1-seriate. Fruit globose;
ovate, imbricate in bud; stamens in male flowers
endosperm smooth. About six spp., Malesia.
exserted, filaments united for most of their length,
forming a distinct collar more or less adnate to the
13. Heberdenia Banks ex A. DC. Fig. 91C corolla tube; anthers dorsifixed, sagittate, opening
through longitudinal slits; ovary in female flowers
Heberdenia Banks ex A. DC., Ann. Sci. Nat. II, 16: 79 (1841); de
Wit, Bull. Jard. Bot. Bruxelles 27: 233-242 (1957); Stahl, Bot. globose-ovoid, style short but well demarcated,
J. Linn. Soc. 122: 315-333 (1996). stigma discoid, lobed or fimbriate; ovules few,
1-seriate. Fruit globose, endosperm ruminate.
Small trees. Leaves short-petiolate, margins entire. 2n = 46. About four spp., Azores, Africa, tropical
Inflorescences axillary fascicles, appearing just and subtropical Asia.
below the leaves. Flowers 5-merous, bisexual;
corolla pale green, rotate, the inner surface papil- 16. Rapanea Aubl.
lose, lobes narrowly oblong, free to base, imbricate
in bud, recurved at anthesis; stamens exposed at Rapanea Hist. Pl. Guiane 1: 121 (1775); Mez in Pflanzenreich
anthesis; filaments long, at base adnate to and IV. 236: 342-396 (1902); Lundell, Fl. Guatemala, Fieldiana
Bot. 24,8: 190-192 (1966), reg. rev.; Smith, J. Arnold Arb. 54:
decurrent on corolla lobes; anthers dorsifixed, ver- 278-292 (1971), Fijian spp.; Pipoly, Sida 17: 115-162 (1996),
satile, narrowly sagittate, recurved at anthesis, Philippine spp.; Ricketson & Pipoly, Sida 17: 579-589 ( 1997),
opening by longitudinal slits; ovary broadly ovoid, Mesoamerican spp.; Ricketson & Pipoly, Sida 18: 1095-1144
style rather long, stigma punctate; ovules many, (1999), Venezuelan spp.
Suttonia A. Rich. (1832).
2-4-seriate. Fruit subglobose; endosperm rumi-
nate. One sp., H. bahamensis (Gaertn.) Sprague,
Macaronesia. Shrubs or trees. Leaves short-petiolate, margins
entire, rarely serrate. Inflorescences axillary fasci-
cles, sometimes on short bracteate shoots. Flowers
14. Pleiomeris A. DC. 4-5(6)-merous, unisexual or bisexual, if unisexual
Pleiomeris A. DC., Ann. Sci. Nat. II, 16: 79 (1841); Mez in
then plants dioecious or polygamous; corolla
Pflanzenreich IV. 236:337-338 (1902); de Wit, Bull. Jard. Bot. rotate, rarely campanulate, greenish, lobes nar-
Bruxelles 27: 233-242 (1957). rowly ovate to ovate, usually granular on margins
and inner surface, valvate in bud; stamens
Small trees or shrubs. Leaves short-petiolate, enclosed, or exposed, filaments inconspicuous,
margins entire. Inflorescences axillary, few- largely fused to the corolla; anthers basifixed,
flowered fascicles. Flowers 5-merous, bisexual; sagittate to oblong, opening by longitudinal slits;
corolla greenish, papillose on inside, rotate with ovary globose to ovoid, style short and inconspic-
well-developed tube, lobes narrowly ovate, subval- uous, stigma variable and often irregular, subglo-
vate in bud; stamens enclosed, filaments largely bose, conical, or ligulate; ovules few, 1-seriate.
Myrsinaceae 275
Fruit subglobose; endosperm smooth or ruminate. Fruit globose; endosperm smooth. 16 spp., tropi-
2n = 46, 48. About 300 spp., pantropical. Often cal Africa. Often included in Ardisia.
included in Myrsine.
19. Synardisia (Mez) Lundell
17. Ardisia Sw. Fig. 91H, I
Synardisia (Mez) Lundell, Wrightia 3: 88-90 (1963); Lundell,
Ardisia Sw., Nov. Gen. Sp. Pl. 3, 48 (1788), nom. cons.; Mez in Fl. Guatemala, Fieldiana Bot. 24,8: 195-197 (1966).
Pflanzenreich IV. 236: 57-154 (1902); Walker, Philippine J. Ardisia subg. Synardisia Mez (1902).
Sci. 73: 48-155 (1940), E Asian spp.; Lundell, Fl. Guatemala,
Fieldiana Bot. 24, 8: 136-156 (1966), reg. rev.; Stearn, Bull. Shrubs or trees. Leaves short-petiolate, margins
Brit. Mus. Nat. Hist., Bot. 4: 156-165 (1969), Jamaican spp.; entire. Inflorescences terminal panicles. Flowers 5-
Lundell, Fl. Panama, Ann. Missouri Bot. Gard. 58: 313-352
(1971), reg. rev.; Stone in Ng, Tree Fl. Malaya 4: 268-278
merous, bisexual; corolla pink, campanulate to
(1989), reg. rev.; Sleumer, Blumea 33: 115-140 (1988), New urceolate, lobes contorted in bud, broadly ovate;
Guinean spp.; Yang & Dwyer, Taiwania 34: 192-297 (1989), stamens enclosed, filaments short; anthers shortly
rev. subg. Bladhia; Larsen & Hu, Fl. Thailand 6 (2): 82-151 sagittate, shortly produced at apex, opening
(1996), reg. rev.; Chen & Pipoly, Fl. China 15: 10-30 (1996), through longitudinal slits; ovary subglobose, style
reg. rev.; Pipoly & Ricketson, Sida 18: 433-472 (1998), rev.
subg. Graphardisia. long and slender, stigma punctate; ovules 8
Icacorea Adans. (1775). or more, 2- or 3-seriate. Fruit depressed-globose.
Bladhia Thunb. (1781). One sp., S. venosa (Mez) Lundell, Mexico to
Auriculardisia Lundell (1981). Honduras.
Ibarraea Lundell ( 1981).
Parardisia Nayar & Giri (1986).
20. Hymenandra A. DC.
Shrubs or treelets, sometimes subshrubs or herbs.
Hymenandra (A. DC.) A. DC. ex Spach, Hist. Veg. Phan. 9: 374
Leaves petiolate, margins entire, crenate or serrate. (1840); Stone, Gard. Bull. Singapore 43: 1-17 (1991).
Inflorescences terminal or axillary panicles or Ardisia sect. Hymenandra A. DC., Trans. Linn. Soc. London 27:
racemes, often with flowers arranged in corymbs 126 (1834); Ann. Sci. Nat. II, 2: 297 (1834).
or umbels. Flowers 5(4)-merous, bisexual; corolla
rotate, pink or white, petals contorted in bud, often Subshrubs, sometimes unbranched. Leaves petio-
recurved at anthesis, ovate-lanceolate; stamens late, sometimes shortly so, margins entire or
enclosed to exposed, filaments short; anthers sometimes crenate. Inflorescences axillary, or
narrowly sagittate, opening by longitudinal slits terminal on reduced lateral branches, often
or subapical pores; ovary ovoid, style long and pedunculate and subtended by foliaceous bracts,
slender, stigma punctate; ovules few to numerous, paniculate but often umbelliform. Flowers 5-
one- to several-seriate. Fruit subglobose, endo- merous, bisexual; corolla pink, purple or white,
sperm smooth or ruminate. 2n = 46, 48, 96. About rotate, lobes ovate to narrowly ovate, contorted or
250 spp. pantropical, mainly tropical Americas and imbricate in bud; stamens included, filaments very
Asia, a few in temperate Japan. short; anthers basifixed, narrowly sagittate, later-
ally connate along thecal margins, opening by lon-
gitudinal slits; ovary ovoid, style long and slender,
18. Afrardisia Mez
stigma punctate; ovules 5-12, 1- or 2-seriate. Fruit
Afrardisia Mez in Pflanzenreich IV. 236: 183-187 (1902); de subglobose; endosperm smooth. Eight spp.,
Wit, Blumea Suppl. IV, 242-262 (1958); Taton, Fl. Afr. Centr., SE Asia (mainly Borneo). Pipoly and Ricketson
Myrsinac. 12-29 (1980), reg. rev.; Halliday, Fl. Trop. E Afr.,
Myrsinac. 16-19 (1984), reg. rev.
(1999) argued for the inclusion of several, chiefly
Mesoamerican species of Ardisia in Hymenandra.
Shrubs, trees or subshrubs. Leaves short-petiolate,
margins serrate or entire. Inflorescences axillary 21. Tetrardisia Mez
clusters, the peduncles short, bracteate. Flowers 5-
Tetrardisia Mez in Pflanzenreich IV. 236: 189 (1902); Stone,
merous, bisexual, or female and the plants then Malayan Nat. J. 46:1-11 (1992).
gynodioecious; corolla rotate pink, red or white,
lobes contorted in bud, spreading at anthesis, Shrubs or small trees. Leaves petiolate, margins
ovate-lanceolate; stamens enclosed, filaments crenate, serrulate or subentire. Inflorescences
short; anthers narrowly sagittate, opening by axillary or terminal panicles. Flowers usually 4-
longitudinal slits; ovary subglobose, style long merous, bisexual; corolla purplish or pink, rotate,
and slender, stigma punctate; ovules 6-8, !-seriate. lobes lanceolate, imbricate in bud; stamens
enclosed, filaments very short; anthers narrowly gular; stamens enclosed, filaments short; anthers
sagittate, opening by longitudinal slits; ovary sub- basifixed, sagittate, apiculate; ovary subglobose,
globose, style long and slender, stigma punctate; style rather stout, somewhat exserted, stigma
ovules few, 1-seriate. Fruits globose, endosperm punctate; ovules few, 1-seriate. Fruits unknown.
smooth or ruminate. Three of four spp., Java, One sp. (E. urnulata B. C. Stone), Sumatra.
Borneo, Malaya Peninsula, Thailand. Treated as a
subgenus of Ardisia by Larsen and Hu (1995).
25. Sadiria Mez
Sadiria Mez in Pflanzenreich IV. 236: 181-183 (1902).
22. Solonia Urb.
Solonia Urb., Fedde Repert.18: 22- 23 {1922); Leon & Alain, Fl. Shrubs or trees. Leaves petiolate, margins crenate
Cuba 4: 113 {1957). or entire. Inflorescences axillary panicles, usually
appearing as few-flowered fascicles or umbels.
Shrubs. Leaves short-petiolate, margins dentate. Flowers 5-merous, bisexual; corolla campanulate
Inflorescences terminal panicles. Flowers 5- or urceolate, lobes ovate, contorted in bud;
merous, bisexual; corolla rotate, lobes ovate to stamens included, filaments very short, adnate to
broadly ovate, contorted in bud, reflexed at anthe- lower part of corolla; anthers basifixed, narrowly
sis; stamens exposed at anthesis, filaments united cordate, sometimes apically produced, opening by
into a tube adnate to the corolla; anthers dorsi- longitudinal slits; ovary subglobose, style long and
fixed, versatile, sagittate, opening through longitu- thick, stigma punctate; ovules few, 1-seriate. Fruit
dinal slits; ovary broadly ovoid, style thick, globose, endosperm ruminate. Four spp., Assam,
somewhat longer than the ovary, stigma punctate; Bhutan.
ovules 12-15, 2- or 3-seriate. Fruits globose. One
sp., S. reflexa Urb., Cuba.
26. Gentlea Lundell
23. Geissanthus Hook. f. Gentlea Lundell, Wrightia 3: 100 {1964); Lundell, Fl. Guatemala,
Fieldiana Bot. 24, 8: 156-160 {1966), reg. rev.; Ricketson &
Geissanthus Hook. f. in Benth. & Hook. f., Gen. 2: 642 (1876); Pipoly, Sida 17: 697-707 ( 1997).
Mez in Pflanzenreich IV. 236:232-241 (1902); Pipoly, Novon Ardisia subg. Walleniopsis Mez {1902).
3: 463-474 {1993), Colombian spp.
Trees or shrubs. Leaves petiolate, margins entire or entire or serrulate. Inflorescences terminal pani-
crenulate. Inflorescences terminal panicles. cles or umbels. Flowers mostly 5-merous, bisexual;
Flowers basically 5-merous, unisexual, the plants corolla greenish white or pink, campanulate or
dioecious or polygamous; calyx split open by rotate, lobes occasionally free to base, imbricate in
(2)3-6( -8) unequal lobes; corolla white, campan- bud; stamens exserted, filaments long; anthers
ulate, lobes imbricate in bud, ovate-oblong, dorsifixed, shortly obtuse-cordate, opening by
recurved at anthesis; stamens exserted, filaments longitudinal slits; ovary subglobose or ovoid,
long; anthers dorsifixed, versatile, sagittate, style long and slender, stigma punctate; ovules
opening by longitudinal slits; ovary subglobose, few to many, in 2 or more series. Fruit subglobose;
style long and slender, stigma truncate-capitate; endosperm smooth or ruminate. Nine spp.,
ovules 3-5, 1-seriate. Fruit globose; endosperm Mexico to N and NW South America.
smooth. About 30 spp., Andes, Venezuela to
Bolivia.
27. Stylogyne A. DC.
Stylogyne A. DC., Ann. Sci. Nat. Bot. II, 16: 78 (1841); Mez in
24. Emblemantha B. C. Stone Pflanzenreich IV. 236: 263-279 (1902); Lundell, Fl. Panama,
Ann. Missouri Bot. Gard. 58: 308-312 (1971), reg. rev.
Emblemantha B. C. Stone, Proc. Acad. Sci. Philadelphia 140:
275-280 {1988).
Low subshrubs. Leaves petiolate with decurrent entire or sometimes crenulate. Inflorescences
leaf blade, margins crenate, lower surface greyish, terminal or axillary panicles, often reduced to
veins red-purple above. Inflorescences axillary umbels or fascicles. Flowers 5(4)-merous, mostly
panicles with short lateral branches. Flowers 5- unisexual and the plants dioecious; corolla white,
merous, bisexual; corolla urceolate-tubular and green or pinkish, rotate, lobes oblong, contorted in
almost completely closed, lobes very short-trian- bud; stamens included, filaments long and slender;
Myrsinaceae 277
anthers dorsifixed or basifixed, narrowly sagittate, Four or five spp., Assam, Malaya.
opening by longitudinal slits; ovary ovoid, style
long and slender, stigma truncate; ovules few, 1- or 31. Parathesis (A. DC.) Hook. f. Fig. 91D
2-seriate. Fruit globose; endosperm smooth.
About 60 spp., Neotropics. Parathesis (A. DC.) Hook. f. in Benth. & Hook. f., Gen. 2: 645
(1876); Mez in Pflanzenreich IV. 236: 173-181 (1902);
Lundell, Contrib. Texas Res. Found., Bot. Stud. 5: 1-206
28. Ctenardisia Ducke (1966); Lundell, Fl. Guatemala, Fieldiana Bot. 24,8: 160-189
(1966), reg. rev.; Lundell, Fl. Panama, Ann. Missouri Bot.
Ctenardisia Ducke, Arch. Jard. Bot. Rio de Janeiro 5: 179-180 Gard. 58: 292-304 (1971), reg. rev.
(1930); Lundell, Wrightia 7: 42-44 (1982).
Trees or shrubs, often with stellate or dendroid
Shrubs, monocaulous. Leaves psuedoverticillate at hairs on young shoots and floral parts. Leaves
stem apices, short-petiolate, margins serrate or petiolate, margins entire or rarely crenulate. Inflo-
entire. Inflorescences terminal panicles, long- recences terminal or axillary panicles. Flowers 5-
pedunculate. Flowers 5-merous, bisexual; corolla merous, bisexual; corolla pink or white, rotate with
greenish, rotate, lobes lanceolate to oblong, short tube, lobes narrowly ovate to lanceolate,
reflexed at antheis, contorted in bud; stamens recurved at anthesis, valvate in bud; stamens
enclosed, filaments rather short; anthers sagittate, exposed due to recurving petals, filaments short,
dorsifixed, opening by longitudinal slits; ovary fused to corolla tube; anthers narrowly saggitate,
ovoid, style long and slender, stigma punctate; basifixed, opening through longitudinal slits,
ovules 10-14, 1-seriate at base of placenta. Fruit sometimes shortly produced at apex; ovary sub-
globose. Two spp., Brazil. globose to ovoid, style long and slender, stigma
punctate; ovules few, 1-seriate. Fruit globose;
29. Yunckeria Lundell endosperm ruminate. About 75 spp., Mesoamer-
ica, Greater Antilles, Andes.
Yunckeria Lundell, Wrightia 3: 111-114 (1964); Lundell, Fl.
Guatemala, Fieldiana Bot. 24,8: 197-200 (1966).
32. Aegiceras Gaertn. Fig. 91F
Shrubs or small trees. Leaves long-petiolate, Aegiceras Gaertn., Fruct. 1: 216 (1788); Mez in Pflanzenreich
margins entire. Inflorescences terminal panicles. IV. 236: 55-57 (1902); Walker, Philippine J. Sci. 73: 47-48
Flowers 5(4)-merous, bisexual; corolla rotate, (1940), E Asian spp.; Stone in Ng, Tree Fl. Malaya 4: 266
lobes narrowly oblong, imbricate in bud; stamens (1989), reg. rev.
included but filaments rather short; anthers
narrowly sagittate, dorsifixed, opening by apical Shrubs or small trees. Leaves petiolate, coriaceous,
pores; ovary ovoid, style long and slender, stigma margins entire. Inflorescences terminal or axillary
punctate; ovules 6-8, 1-seriate at base of placenta. umbels or panicles. Flowers 5-merous, bisexual;
Fruit globose. Three spp., Mexico to Nicargua. corolla campanulate, white, petals contorted in
Often included in Ctenardisia. bud, reflexed at anthesis; stamens exserted, fila-
ments distally united into a ring adnate to the
corolla base; anthers sagittate, dorsifixed, versatile,
30. Antistrophe A. DC. transversely septate, opening by longitudinal slits;
Antistrophe A. DC., Ann. Sc. Nat. II, 16: 79 (1841); Mez in ovary ovoid, gradually tapering into a thick style
Pflanzenreich IV. 236: 150-151, 187-189 (1902); Grosse, Bot. of more than twice the length of the ovary, stigma
Jahrb. Beibl. 96: 22 (1908), anatomy; Stone in Ng (ed.), Tree punctate; ovules numerous, pluriseriate. Fruit
Fl. Malaya 4: 268 (1989), reg. rev.
viviparous, curved and elongate. 2n = 46.
Mangrove genus of two spp., SE Asia, Pacific, NE
Shrubs or subshrubs. Leaves petiolate, margins Australia.
entire or serrate. Inflorescences axillary, few-
flowered panicles, often reduced to umbels or fas-
cicles. Flowers 4-5-merous, bisexual; corolla 33. Amblyanthus A. DC.
rotate, petals contorted to the left, lanceolate; Amblyanthus A. DC., Ann. Sc. Nat. II, 16: 79 (1841); Mez in
stamens enclosed, filaments very short; anthers Pflanzenreich IV. 236: 208-210 (1902).
narrowly sagittate with long apices, dorsifixed,
opening by longitudinal slits; ovary subglobose, Trees or shrubs. Leaves petiolate, margins
style long and slender, stigma punctate; ovules glandular-crenate. Inflorescences axillary panicles
2-4, 1-seriate. Fruit globose, endosperm ruminate.
but appearing as long-pedunculate corymbs. plants dioecious; corolla greenish or cream, cam-
Flowers 5-merous, bisexual; corolla short- panulate (tube shorter in female flowers), lobes
camapanulate to rotate, petals contorted in bud, broadly ovate, sometimes emarginate, imbricate in
very broadly ovate, emarginate; stamens included, bud; stamens in male flowers exserted, filaments
filaments free; anthers fused, ovoid, dorsifixed, inserted at base of the corolla; anthers sagittate,
opening through longitudinal slits; ovary ovoid, usually recurved at anthesis, dorsifixed, versatile,
tapering into a style of about the same length, opening by longitudinal slits; ovary in female
stigma capitate; ovules 2-4, 1-seriate. Fruit flowers ovoid, tapering into a style of about equal
globose; endosperm ruminate. Three spp., Bengal, length or shorter, stigma truncate; ovules 3-5, 1-
NE India. seriate. Fruit globose, endosperm smooth. About
20 spp. West Indies.
34. Amblyanthopsis Mez
37. Loheria Merr.
Amblyanthopsis Mez in Pfianzenreich IV. 236: 210-211 (1902).
Loheria Merr., Philippine J. Sci. Bot. 5: 373 (1910); Sleumer,
Shrubs. Leaves petiolate, margins glandular- Blumea 33: 100-102 (1988), New Guinean spp.; Stone,
crenate. Inflorescences axillary, few-flowered Micronesica 24: 65-80 (1991).
panicles appearing as corymbs or umbels. Flowers
5-merous, bisexual; corolla shortly campanulate to Small trees, sparsely branched or unbranched.
rotate, petals contorted in bud, broadly ovate, Leaves short-petiolate or subsessile, often large,
emarginate; stamens included, filaments rather clustered at branch tips, margins entire through-
long; anthers ovoid, dorsifixed, opening by longi- out or distally crenate. Inflorescences lateral, lax
tudinal slits; ovary subglobose, style thick and panicles (subg. Loheria) or arranged in globular
long, stigma capitate; ovules c. 3, 1-seriate. Fruit clusters on pendant peduncles (subg. Longicorna
globose; endosperm smooth. Two spp., NE India, B. C. Stone). Flowers 4-5-merous, unisexual, the
Bhutan. plants dioecious; corolla red, pink, white or green-
ish, rotate (subg. Loheria) or ± tubular (subg.
Longicorna), lobes oblong to elliptic, ±papillose
35. Elingamita Baylis on inside, sometimes reflexed at anthesis, imbri-
Elingamita Baylis, Rec. Auckland Inst. Mus. 4: 99-102 (1951); cate in bud; stamens enclosed or exposed due to
Heenan, New Zeal. J. Bot. 38: 569-574 (2000). recurving petals, filaments adnate to base of
corolla tube; anthers obtusely sagittate, basifixed,
Trees. Leaves short-petiolate, margins entire. Inflo- opening through longitudinal slits; ovary ovoid
rescences terminal panicles. Flowers 4-6-merous, with a rather stout style of equal length, stigma
unisexual, the plants dioecious; corolla yellowish, capitate to discoid; ovules 3-5, 1-seriate. Fruit sub-
tubular, shorter than the calyx, obscurely lobed; globose; endosperm ruminate. Six spp., Malesia
stamens exserted, filaments adnate to corolla at (Philippines, New Guinea).
base only; anthers oblong, dorsifixed, opening by
longitudinal slits; ovary ovoid tapering into a short 38. Cybianthus Mart.
style, stigma punctate and somewhat excavate;
ovules 2-4, 1-seriate. Fruit subglobose and rather Cybianthus Mart., Nov. Gen. Sp. 3: 87 (1829), nom. cons.; Mez
large, exocarp thick; endosperm smooth. 2n = 46. in Pfianzenreich IV. 236: 215-229,249-263,283-292 (1902);
Agostini, Acta Bioi. Venez.lO: 129-185 (1980); D'Arcy,Ann.
One sp., E. johnsonii Baylis, New Zealand. Missouri Bot. Gard. 60: 442-448 (1973), rev. subg. Tri-
adophora; Pipoly, Brittonia 35: 61-80 (1983), rev. subg.
Laxiflorus; Pipoly, Ann. Missouri Bot. Gard. 79: 908-957
36. Wallenia Sw. Fig. 91B (1992), reg. rev., subg. Conomorpha; Pipoly, Sida 18: 1-160
(1998), rev. Ecuadorean and Peruvian spp.
Wallenia Sw., Nov. Gen. Sp. Pl. 2:31 (1788); Mez in Pfl.anzenre-
Conomorpha A. DC. (1834).
ich IV. 236: 241-249 (1902); Stearn, Bull. Br. Mus. Nat. Hist.,
Weigeltia A. DC. (1834).
Bot. 4(4): 165-174 (1969), Jamaican spp.
Comomyrsine Hook. f. in Benth. & Hook. (1876).
Corelliana D'Arcy (1973).
Trees or shrubs. Leaves petiolate or rarely sessile,
margins entire, sometimes clustered at branch Shrubs or small trees, vegetative and floral parts
tips. Inflorescences terminal panicles (subg. Wal- sometimes brownish-lepidote. Leaves petiolate,
lenia) or axillary racemes (subg. Homowallenia sometimes pseudoverticillate, margins entire or
Mez). Flowers 4-5-merous, unisexual and the sometimes serrate. Inflorescences axillary panicles
Myrsinaceae 279
or racemes. Flowers 4-5-merous (3-merous in 41. Oncostemum A. Juss. Fig. 91G

subg. Triadophora (Mez) Agostini), mostly uni-
Oncostemum A. Juss., Mem. Mus. Paris 19: 133 (1830); Mez in
sexual and the plants dioecious; corolla green, Pfianzenreich IV. 236: 189-208 (1902); Perrier de Ia Bathie,
greenish-brown or yellowish, rotate or short- Fl. Madagascar 161: 16-120 (1953).
campanulate, the inner surface often granular,
lobes imbricate in bud, ovate, sometimes reflexed Shrubs or small trees. Leaves petiolate, margins
at anthesis; stamens enclosed or exposed because entire or rarely crenulate. Inflorescences axillary
of the reflexed petals; filaments short to long; panicles or racemes, through reductions often
anthers subglobose to broadly oblong, dorsifixed, appearing as umbels. Flowers (4)5-merous, bisex-
opening by oblique to round apical pores or lon- ual; corolla white, purple or greenish, rotate or
gitudinal slits; ovary subglobose, style short, short-campanulate, lobes oblong to ovate, con-
stigma capitate or truncate; ovules 2-5, uniseriate. torted in bud; stamens enclosed, united into a tube
Fruit globose, endosperm smooth. About 150 spp., with upper part free from the corolla; anthers ±
Neotropics. connate, obtuse-triangular, opening by longitudi-
nal slits; ovary ovoid, tapering into a style of equal
39. Grammadenia Benth. length, stigma discoid, rarely 5-lobed; ovules 3-5,
1-seriate. Fruit globose, endosperm ruminate.
Grammadenia Benth., Pl. Hartw. 218 (1846); Mez in About 90 spp., Madagascar. Distinction from
Pfianzenreich IV. 236: 228-232 (1902); Pipoly, Mem. New Badula unclear (Coode 1976).
York Bot. Gard. 43: 1-76 (1987).
Shrubs or small trees, often epiphytic. Leaves 42. Badula Juss.

sessile, basally auriculate, margins entire or den-
Badula Juss., Gen. Pl. 420 (1789); Perrier de Ia Bathie, Fl.
ticulate. Inflorescences axillary racemes. Flowers Madagascar 161: 120-124 (1953); Coode, Fl. Mascareignes
mostly 5-merous, uni- or bisexual, the plants dioe- 115: 6-25 (1981).
cious, monoecious or hermaphroditic; corolla
greenish white, rotate, lobes oblong to ovate, Trees or shrubs, often unbranched or sparsely
imbricate in bud; stamens included; anthers sessile branched. Leaves petiolate, condensed at branch
or subsessile (filaments forming a sheath adnate to tips, margins entire. Inflorescences axillary pani-
the corolla tube), almost square, opening by sub- cles. Flowers 5-merous, bisexual; corolla rotate,
apical slits; ovary subglobse, usually depressed, white or reddish, petals ovate to subrotund, imbri-
style short, stigma truncate; ovules few, 1- or 2- cate in bud; stamens enclosed, filaments very
seriate. Fruit ellipsoid or subglobose, endosperm short; anthers shortly sagittate, opening by longi-
ruminate. Seven spp., C America, Lesser Antilles, tudinal slits; ovary ovoid, style of about the same
tropical Andes. Sometimes treated as a subgenus length as the ovary, stigma discoid; ovules 3-5,
of Cybianthus s.l. 1-seriate. Fruit subglobose, endosperm smooth.
About 17 spp., Mascarenes, Madagascar.
40. Vegaea Urb.
Vegaea Urb., Symb. Antill. 7: 535-537 (1913). 43. Tapeinosperma Hook. f.
Tapeinosperma Hook. f. in Benth. & Hook., f., Gen. Pl. 2: 647
Trees. Leaves clustered at branch tips, thick- (1876); Mez in Pfianzenreich IV. 236: 162-171 (1902); Smith,
coriaceous, sessile, linear and spine-tipped, mar- J. Arnold Arb. 54: 228-263 (1973), Fijian spp.; Sleumer,
Blumea 33: 102-107 (1988), New Guinean spp.
gins entire. Inflorescences axillary, few-flowered
racemes. Flowers 5(6)-merous, bisexual; corolla
campanulate, white, waxy and thick, lobes broadly Trees or shrubs, often unbranched. Leaves often
ovate, imbricate in bud; stamens included, fila- large and clustered at branch tips, petiolate or
ments short; anthers ovoid-oblong, dorsifixed, sessile, margins entire. Inflorescences axillary or
opening by longitudinal slits; ovary ovoid, style terminal panicles. Flowers mostly 5-merous, bisex-
thick, somewhat longer than the ovary, curved ual; corolla red, rotate or campanulate, lobes ovate,
below apex, stigma truncate; ovules few, 1-seriate. contorted in bud; stamens included, filaments
Fruits not known. One sp., V. pungens Urb., connate at base and adnate to the corolla tube, dis-
Hispaniola. tally free; anthers oblong-ovoid, dorsifixed near
base, opening by longitudinal slits starting as sub-
apical pores; ovary globose or ovoid, style rather
thick, longer than the ovary, stigma truncate, 46. Systellantha B. C. Stone
capitate or somewhat discoid; ovules (2- )6-12,
Systellantha B. C. Stone, Malayan Nat. J. 46: 13-24 (1992).
1-seriate. Fruits large, variable in shape, often
depressed-globose, fleshy when fresh; endosperm
Shrubs or treelets with compressed and somewhat
smooth or ruminate. About 50 spp., Borneo, New
Guinea, New Hebrides, Fiji, New Caledonia, trop. sinuous branches. Leaves petiolate, margins entire
to crenulate. Inflorescences short axillary racemes.
Australia.
Flowers 4-merous, unisexual with flowers of both
sexes occurring on the same inflorescence; corolla
44. Discocalyx (A. DC.) Mez pink, rotate, lobes ovate, imbricate in bud; stamens
Discocalyx (A. DC.) Mez in Pflanzenreich IV. 236: 211-214
included, filaments short, fused at base of corolla;
(1902); Smith, J. Arnold Arb. 54: 236-273 (1973), Fijian spp.; anthers dorsifixed near base, narrowly sagittate,
Sleumer, Blumea 33: 83-93 (1988), New Guinean spp. opening by longitudinal slits starting as subapical
pores; ovary subglobose, style thick, longer than
Shrubs or trees, sparsely branched or unbranched. the ovary, stigma truncate; ovules 1-3, 1-seriate.
Leaves clustered at branch tips, petiolate, margins Fruit globose, seed not known. Two spp., Borneo.
entire. Inflorescences axillary racemes or panicles.
Flowers 4-5-merous, unisexual and plants dioe- 47. Monoporus A. DC.
cious, or bisexual; corolla urceolate, campanulate
or rotate, white or sometimes yellowish, green or Monoporus A. DC., Ann. Sci. Nat. II, 16:78 (1841); Perrier de Ia
red, lobes imbricate in bud, ovate or oblong, some- Bathie, Fl. Madagascar 161: 6-14 (1953).
times emarginate; stamens enclosed; filaments
very short; anthers appearing sessile, basifixed, Trees or shrubs. Leaves petiolate, often condensed
broadly sagittate or ovoid, truncate at apex, at branch tips, margins entire. Inflorescences axil-
opening by longitudinal slits; ovary ovoid or sub- lary panicles. Flowers 5-merous, unisexual, the
globose, style very short, stigma discoid, some- plants dioecious; corolla pink to orange-red,
times with erose margins; ovules 2-5, 1-seriate. rotate, the tube very short, the lobes oblong or
Fruit subglobose; endosperm smooth. About 50 ovate, spreading at anthesis, imbricate in bud,
spp., SE Asia (especially the Philippines), the emarginate; stamens included, filaments very
Pacific. short; anthers sagittate, opening apically by one or
two irregular pores; ovary subglobose to cylindri-
cal, style very short or absent, stigma punctate;
45. Labisia Lindl. ovules many, 1-seriate in several parallel rows.
Labisia Lind!., Bot. Reg. 31, tab. 48 (1845), nom. cons.; Mez in Fruit globose; endosperm ruminate. Eight spp.,
Pflanzenreich IV. 236: 171-172 (1902); Stone, Malayan Nat. Madagascar.
J. 42: 43-51 (1988), synopsis; Stone in Ng, Tree Fl. Malaya 4:
281 (1989), reg. rev.
48. Fittingia Mez
Perennial herbs or subshrubs, usually un- Fittingia Mez, Bot. Arch. 1: 105-106 (1922); Sleumer, Blumea
branched. Leaves long- to short-petiolate, blade 33: 94-100 (1988); Stone, Sida 16: 267-270 (1994).
base decurrent, the blade with numerous, densely
set secondary veins, margins entire or crenulate- Small trees or shrubs, mostly unbranched. Leaves
serrulate. Inflorescences axillary panicles with clustered at branch tips, petiolate, margins entire.
reduced lateral branches. Flowers 5-merous, bisex- Inflorescences axillary racemes. Flowers 4-5-
ual; corolla white or pink, rotate, lobes broadly merous, unisexual, the plants dioecious; corolla
ovate with incurved margins largely concealing pale red white or greenish, campanulate to
the anthers, valvate in bud; stamens included; urceolate, lobes imbricate in bud, ovate to oblong;
anthers sessile, sagittate, apically produced, stamens included, filaments short; anthers
opening by longitudinal slits; ovary globose, style obtusely sagittate, basifixed, opening by longitudi-
of about the same length as the ovary, stigma nal slits; ovary subglobose, style short, stigma cap-
punctate; ovules few, 1-seriate. Fruit globose; itate to discoid; ovules unknown. Fruit subglobose
endosperm smooth. n = 23-25. Six spp., W Malesia, with a thick and soft exocarp (flattened in her-
New Guinea. barium specimens); endosperm smooth. Six spp.,
New Guinea.
Myrsinaceae 281
49. Conandrium (K. Schum.) Mez Hegnauer, R. 1969. See general references.
Hope, G. 2000. Australian national university pollen database.
Conandrium Mez in Pflanzenreich IV. 236: 156-157 (1902); http/www.geo.arizona.edu/palynologfy/sem/anu.html
Sleumer, Blumea 33: 109-113 (1988). Johri, B.M. et a!. 1992. See general references.
Kallersji:i, M. et a!. 2000. See general references.
Shrubs or small trees. Leaves petiolate, sometimes Larsen, K., Hu, C.M. 1995. Reduction of Tetradisia to Ardisia.
Nord. J. Bot. 15: 161-162
shortly so, margins entire or vaguely crenate Lersten, N.R. 1977. Trichome forms in Ardisia (Myrsinaceae)
towards apex. Inflorescences axillary panicles or in relation to the bacterial leaf nodule symbiosis. Bot. J.
racemes with long slender pedicels. Flowers 5- Linn. Soc. 75: 229-244.
merous, bisexual; corolla rotate, red to pink, lobes Lersten, N.R., Horner, H.T. 1976. Bacterial leaf nodule
contorted in bud, broadly ovate; stamens enclosed, symbiosis in angiosperms with emphasis on Rubiaceae and
Myrsinaceae. Bot. Rev. 42: 145-214.
filaments free, short; anthers broadly sagittate, Martin, H.A. 1994. Australian Tertiary phytogeography:
laterally connate over the ovary, opening by evidence from palynology. In: Hill, R.S. (ed.) History of
longitudinal slits; ovary ovoid to globose, style the Australian vegetation: Cretaceous to Recent. Cambridge:
short, stigma punctate; ovules many, 2- or 3- Cambridge University Press, pp. 104-142.
Metcalfe, C.R., Chalk, L. 1950. See general references.
seriate. Fruit globose. Two spp., New Guinea, Mez, C. 1902. Myrsinaceae. In: Engler, A. (ed.) Das Pflanzenre-
Moluccas, Bismarck Archipelago. ich 9, IV 236: 1-437.
Mildenhall, D.C. 1980. New Zealand Late Cretaceous and
Cenozoic plant biogeography: a contribution. Palaeogeogr
Selected Bibliography Palaeoclimatol Palaeoecol31: 197-233.
Nowicke, J.W., Skvarla, J.J. 1977. Pollen morphology and the
relationships of the Plumbaginaceae, Polygonaceae, and
Anderberg, A.A., Stahl, B.1995. Phylogenetic interrelationships Primulaceae to the order Centrospermae. Smithsonian
in the order Primulales, with special emphasis on the family Contrib. Bot. 37: 1-64.
circumscriptions. Can. J. Bot. 73: 1699-1730. Otegui, M., Cocucci, A. 1999. Flower morphology and biology
Anderberg, A.A., Stahl, B., Kallersji:i, M. 1998. Phylogenetic of Myrsine laetevirens, structural and evolutionary implica-
relationships in the Primulales inferred from rbcL sequence tions of anemophily in Myrsinaceae. Nord. J. Bot. 19: 71-85.
data. Pl. Syst. Evol. 211: 93-102. Pascarella, J.B. 1997a. Pollination ecology of Ardisia escallo-
Anderberg, A.A., Stahl, B., Kallersji:i, M. 2000. Maesaceae, a new nioides (Myrsinaceae). Castanea 62: 1-7.
primuloid family in the order Ericales s.l. Taxon 49: 183-187. Pascarella, J.B. 1997b. Breeding systems of Ardisia Sw. (Myrsi-
Anderberg, A.A., Peng, C.-I., Trift, I., Kallersji:i, M. 2001. The naceae). Brittonia 49: 45-53.
Stimpsonia problem; evidence from DNA sequences of Pax, F., Knuth, R. 1905. Primulaceae. In: Engler, A. (ed.) Das
plastid genes atpB, ndhF and rbcL. Bot. Jahrb. Syst. 123: Pflanzenreich 22, IV 237: 1-386.
369-376. Pipoly, J.J., Ricketson, J.M. 1999. Discovery of the Indo-
Braun, A. 1859. Uber Polyembryonie und Keimung von Caele- Malesian genus Hymenandra (Myrsinaceae) in the Neo-
bogyne. Ein Nachtrag zu der Abhandlung iiber Partheno.- tropics, and its boreotropical implications. Sida 18: 701-746.
genesis bei Pflanzen. Abh. K. Acad. Wiss. Berlin, 1859: Sattler, R. 1962. Zur friihen Infloreszenz- und Bliitenentwick-
109-263. lung der Primulales sensu Jato mit Beriicksichtigung der
Burrows, J. 1999. Myrsine. Plant Life (S. Afr.) 20: 25-26. Stamen-Petalum-Entwicklung. Bot. Jahrb. Syst. 81: 358-396.
Carlquist, S. 1992. Wood anatomy of sympetalous dicotyledon Stahl, B.1990. Primulaceae. In: Harling, G.,Andersson, L. (eds.)
families: a summary with comments on systematic rela- Fl. Ecuador 39: 23-35.
tionships and evolution of the woody habit. Ann. Missouri Stahl, B. 1996. The relationships of Heberdenia bahamensis
Bot. Gard. 79: 303-332. and H. penduliflora (Myrsinaceae). Bot. J. Linn. Soc. 122:
Co ode, M.J.E. 1976. Notes on Pittosporaceae and Myrsinaceae 314-333.
of the Mascarenes. Kew Bull. 31: 221-225. Stone, B.C. 1993. New and noteworthy Malesian Myrsinaceae,
Cronquist, A. 1988. See general references. VI. Scherantha, a new subgenus of Ardisia. Pacific Sci. 47:
Davis, G.L. 1966. See general references. 276-294.
Erdtman, G. 1952. See general references. Vogel, S. 1986. Olblumen und i:ilsammelnde Bienen. Zweite
Faure, P. 1968. Contribution a !'etude caryo-taxonomique des Folge, Lysimachia und Macropis. Akad. Wiss. Lt. Mainz, Abh.
Myrsinacees et des Theophrastacees. Mem. Mus. Hist. Nat. Math.-Naturwiss. Kl., Trop. Subtrop. Pflanzenwelt 54: 1-168.
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Grosse, A. 1908. Anatomisch-systematische Untersuchungen
der Myrsinaceen. Bot. Jahrb., Beibl. 95: 1-46.
282 G.T. Prance
Napoleonaeaceae
G.T. PRANCE
line of vessels at growth boundary, moderately

Napoleonaeaceae A. Rich. in Bory, Diet. Class. Hist. Nat. 11: 432
(1827). large, mean diam. 76 J.lm in Crateranthus, 88 J.lm in
Napoleonaea, over 50% solitary, the rest in groups
Trees, shrubs or suffrutices; young axis with cor- of 2-6; perforations mostly simple, in part multi-
tical vascular bundles. Leaves alternate, simple, ple to branched scalariform; intervessel pits
entire or serrate, pinnately nerved; stipules absent. alternate to subopposite, closely spaced, elliptic to
Inflorescences usually of solitary, axillary flowers, angular, 4-7 J.lm; ray-vessel pits linear, elongate,
less frequently in cauline fascicles or axillary pan- opposite, tyloses often present in heartwood. Rays
icles; 4-7 bracts subtending each flower, often with heterocellular with 2 classes in Crateranthus, 60%
paired glands on abaxial surface. Flowers large, 3-7 -seriate with uniseriate tails, the rest uniseriate
actinomorphic, hermaphrodite; sepals (2)3 and or slightly biseriate; rays at broad end up to 14
imbricate or 5-6 and valvate, often with paired cells wide in Napoleonaea; crystals present in
glands towards apex; corolla sympetalous, plicate, Napoleonaea only. Axial parenchyma apotracheal
with 30-35 ribs, with a wavy margin, probably in uniseriate tangential lines or reticulate in areas
representing an outer ring of stamina! tissue or of closely spaced, wide rays, paratracheal in
pseudo-corolla; androecium either 8-10 whorls of limited amounts where lines of parenchyma touch
numerous, free stamens fused to base of pseudo- tangential faces of vessels, strands of moderate
corolla, or of 3 concentric whorls, fused at base, the length, crystal strands common in Crateranthus
outer and middle antherless, the inner one bearing and small prismatic crystals only in Napoleonaea,
10 anthers, and 10 staminodes; anthers 2- or 4- silica absent. Fibres moderately long in Napoleon-
locular, longitudinally dehiscent; ovary inferior or aea with mean length 2295 J.lm, and long in Crat-
semi-inferior, 3-5-locular, with 2-numerous anat- eranthus with mean length 2779 J.lm, 16.3-21 J.lm in
ropous ovules in each loculus, the placentation diameter, thick walled.
apical-axile with 15-18 pendulous ovules or - The wood of the two genera is quite similar indi-
central-axile with 2-4 ovules per locule; style cating a close relationship. The partly scalariform
slender with a pointed stigma or short with a perforation and scalariform to linear ray-vessel
pentagonal or hexagonal stigma; nectary disk well pits set Napoleonaeaceae apart from Lecythi-
developed, annular. Fruit indehiscent with 1 to daceae but not from Asteranthos (Scytopeta-
several exalbuminous seeds; embryo with 2 plano- laceae), which shares those characters.
convex cotyledons, and a short radicle and large
plumule. FLOWER STRUCTURE. The large, usually solitary
Two genera and 10 species in tropical West hermaphrodite flowers are unique among
Africa. angiosperms because of the structure of the
corolla and androecium. Both genera have a
VEGETATIVE MORPHOLOGY. Most species are showy, many-ribbed, united corolla with a dentate,
shrubs or small trees, at least one species reaches wavy margin. Masters (1869) and others have con-
30m in height. sidered this structure to be of staminodial origin
The wood anatomy of Napoleonaeaceae was which seems likely from the ribs. The numerous,
described by the late Carl de Zeeuw in an un- long free stamens of Crateranthus (Fig. 92) are
published paper from which this description is fused to the base of the corolla and are arranged
condensed (for reference see Bibliography of in 8-10 whorls. In Napoleonaea the androecium
Lecythidaceae, this volume). The wood is yellow- consists of 3 concentric whorls. The outer whorl
brown without differentiation between sapwood consists of 60-70, narrow strap-like appendages
and heartwood, and possesses a fine texture with which are free or slightly connate at their base; the
the vessels indistinct to the naked eye. Vessels middle whorl is of 30-40, ribbon-like appendages
diffuse, uniformly distributed but rarely with a which are wider and longer than the outer ones;
Napoleonaeaceae 283
they are also fused laterally up to 2/3 of their Planchonioideae (Lecythidaceae) x = 13, and from
length. These two whorls are delicately folded and Asteranthos (Scytopetalaceae), x = 21.
hooked together. The inner whorl consists of± 20
stamens which are fertile and are similar mor- FRUIT AND SEED. The fruit is a 1- to several-
phologically to the middle whorl. These stamens seeded drupe. In Crateranthus the calyx is persist-
curve 180° inwards and their apex is inserted into ent at the proximal end and the style at the distal
the narrow space below the larger broad stigma, end. In Napoleonaea, the calyx scars or small per-
so that the anthers are situated between the sistent calyx are at the distal end of the fruit, there
annular nectary disk and the short column of the are 1 or 2 seeds, and the embryo consists of 2
style. They are grouped into five groups of four, plano-convex cotyledons and has a short radicle
each group coinciding with one side of the pen- and a large plumule.
tagonal stigma. The outer two stamens of each
group develop extremely large, bisporangiate AFFINITIES. An Ebenalean affinity for Napoleon-
anthers and the inner two bear only vestigial aea was suggested by Lindley ( 1830) and Miers
anthers. The anthers remain hidden inside the (1874). There is no doubt that Lecythidaceae,
flowers even at anthesis. The anthers are latrorse Napoleonaeaceae and Scytopetalaceae remain a
in Crateranthus and extrorse in Napoleonaea. The closely related group of families and that their
calyx of the two genera is quite different: Crater- affinities are with Sapotaceae, Styracaceae and
anthus has a 3-lobed imbricate calyx which Ebenaceae. Napoleonaea has frequently been
persists in fruit, and Napoleonaea possesses a placed in a family of its own (e.g. de Candolle 183 7;
5-6-lobed valvate calyx. In Napoleonaea there are Airy Shaw 1973), and this was also suggested by
two large glands towards the apex of the outer Tsou (1994: 101), but usually Napoleonaeaceae
surface. The ovary of Napoleonaea is inferior, that have been regarded as a subfamily of Lecythi-
of Crateranthus is semi-inferior and there is a well- daceae. They are most closely related to the genus
developed, annular nectary disk in both genera. Asteranthos, which has a similar pseudo-corolla
For further details of flower structure and its and has recently been related or transferred to
development see Cruger (1860) and Tsou (1994), Scytopetalaceae (Tsou 1994; Morton et al. 1997). If
and the detailed study of the flower of Napoleon- Asteranthos is placed in Scytopetalaceae, it is then
aea by Masters (1869). necessary to recognise the Napoleonaeaceae as a
distinct family to maintain a monophyletic family
EMBRYOLOGY. The embryology of the family was Lecythidaceae (see Tsou 1994; Morton et al. 1997).
studied by Tsou (1994). The anthers have a glan-
dular tapetum and the endothecial wall thicken- DISTRIBUTION AND HABITATS. The family is con-
ings are rod-like. The ovules are bitegmic and fined to W Tropical Africa where most species
anatropous or either anatropous or campy- occur in the understorey of lowland, tropical moist
lotropous in Napoleonaea. The nucellus is tenuin- forests, especially near to the coast and along
ucellate and embryo sac formation is of the rivers. Napoleonaea gossweilera, the southernmost
Polygonum type. The micropyle is formed from species, occurs in savannah and open forests in
the inner integument. Crateranthus differs from Zambia.
Napoleonaea in the long-sagittate anthers and a
completely closed endothecium that extends to the EcoNOMIC USES. No general uses of species of the
filament, the extremely long micropyle formed by family are recorded. The bark of Napoleonaea
very slender cells of the inner integument, and vogelii and N. imperialis is used locally as a cough
non-branching ovular vasculature. medicine. The pulp around the seeds of some
species of Napoleonaea is eaten. The hard wood of
POLLEN MORPHOLOGY. The pollen is 3-colporate N. imperialis is used locally to beat mud floors and
with a granulate colpus membrane, and is more to make clogs.
similar to that of Lecythidoideae than to Plancho-
nioideae (Lecythidaceae) (Muller 1972; Tsou KEY TO THE GENERA
1994).
1. Sepals S-6, valvate; androecium of 3 whorls, the outermost
and middle of sterile, strap-like appendages, the inner
KARYOLOGY. The basic chromosome number of bearing 10 large fertile, 2-locular anthers and 10 small
two species of Napoleonaea has been reported as sterile ones; style short, with an expanded pentagonal or
x = 16 (Mangenot and Mangenot 1957, 1962), hexagonal stigma; ovules 2-4 per loculus, central-axile
which differs from that of Lecythidoideae x = 17, 1. Napoleonaea
284 G.T. Prance
flattened pentagonal or hexagonal stigma. Ovary

5-locular, inferior. Fruit with 1-2 seeds per locule.
x = 16. Eight spp. in W Tropical Africa.
2. Crateranthus Baker Fig. 92

Crateranthus Baker in Rendle, Cat. Talbot's Nig. Pl.: 35 (1913);
Knuth in Pflanzenreich IV. 219: 65-67 (1939).
Small trees, stem of young branches winged in 2

species. Flowers solitary and axillary. Calyx (2)3-
lobed, imbricate. Corolla plicate, ca. 30-ribbed,
fused. Stamens inserted on base of corolla, the fil-
aments free, in 8-10 whorls; anthers introrse. Style
filamentous, the stigma pointed. Ovary 3-4-
locular, semi-inferior. Fruit with persistent calyx at
proximal end. Three spp. in W Tropical Africa.
Airy Shaw, H.K. 1973. Adictionary of the flowering plants and
ferns of the late J.C. Willis, 8th edn. Cambridge University
Press.
Candolle, A.P. de 1837. Prodr. systematis naturalis, Vol. 7. Paris:
Treuttel & Wiirtz, pp. 550-551.
Criiger, H. 1860. Westindische Fragmente: 11. Die Entwicklung
der Blume von Napoleona imperialis Beauv. Bot. Z. 18:
361-367.
Knuth, R. 1939. Barringtoniaceae. In: Engler, A. (ed.) Pflanzen-
Fig. 92. Napoleonaeaceae. Crateranthus talbotii. A Flowering reich IV: 219. Leipzig: W. Engelmann.
branch. B Flower bud. C Flower, vertical section. D Stamen. E Liben, L. 1971. Revision du genre africain Napoleonaea P.
Young fruit. (Knuth 1939) Beauv. (Lecythidaceae). Bull. Jard. Bot. Nat!. Belg. 41:
363- 382.
Lignier, 0. 1890. Recherches sur l'anatomie des organes vege-
- Sepals (2)3, imbricate; androecium of 8-10 whorls of tatifs des Lecythidees, des Napoleonees et des Barring-
numerous free stamens inserted on base of corolla; style toniees. Bull. Sci. France Belgique 21: 291-420.
filamentous, with a pointed stigma; ovules 15-18 per Lindley, J. 1830. Introduction to a natural system of botany, 44,
loculus, apical-axile and pendulous 2. Crateranthus Lecythideae, 177, Belvisiaceae. London: Longman, Rees,
Orme, Brown & Green.
Mangenot, S., Mangenot, G. 1957. Nombres chromosomiques
1. Napoleonaea P. Beauv. nouveaux chez diverses Dicotyledones et Monocotyledones
d' Afrique occidentale. Bull. Jard. Bot. Etat 27: 639-654.
Napoleonaea P. Beauv. ex Fr. Fischer, Mem. Soc. Nat. Mosc. 1: Mangenot, S., Mangenot, G. 1962. Enquete sur les nombres
92 (1806); Fl. Oware 2, 29: 7 (1807); Liben, Bull. Jard. Bot. chromosomiques dans une collection d'especes tropicales.
Nat!. Belg. 41: 363-382 (1971), rev. Rev. Cytol. Bioi. Veg. 25: 411-447.
Masters, M.T.1869. On the structure of the flower in the genus
Trees or shrubs. Flowers usually solitary and axil- Napoleona, etc. J. Linn. Soc. 10: 492-504.
lary, less frequently in cauline fascicles or axillary Miers, J. 1874. On Napoleona, Omphalocarpum, and Asteran-
thos. Trans. Linn. Soc. II, 1: 17- 19.
panicles. Calyx 5(6)-lobed, valvate. Corolla plicate, Morton, C.M., Mori, S.A., Prance, G.T., Karol, K.G., Chase, M.W.
35-ribbed, fused. Androecium of 3 whorls, the 1997. Phylogenetic relationships of Lecythidaceae: a cladis-
outer two of free or fused, strap-like sterile tic analysis using rbcL sequence and morphological data.
appendages, the inner of 20 strap-like filaments, Am. J. Bot. 84: 530-540.
10 with large fertile, extrorse anthers, 10 with Muller, J. 1972. Pollen morphological evidence for subdivision
and affinities of Lecythidaceae. Blumea 20: 351-355.
small sterile anthers. Stamens and staminodes in Tsou, C.-H. 1994. The embryology, reproductive morphology
5 groups of 4, folded inwards beneath stigma and systematics of Lecythidaceae. Mem. New York Bot.
between style and disk. Style short, with large, Gard. 71: 1- 110.
Oxalidaceae 285
Oxalidaceae
A.A. Cocucci
Oxalidaceae R. Br. in Tuckey, Narr. Exp. Zaire: 433 (1818), nom. MORPHOLOGY. Oxalidaceae are commonly
cons. perennial herbs, sometimes cushion-forming,
sometimes succulent, or shrubs, trees (Averrhoa,
Perennial, rarely annual herbs, sometimes succu- Sarcotheca and some Oxalis species) or Hanas
lent, often with underground storaging bulbs, (Dapania and a few species of Oxalis). The herba-
tubers or rape-like roots, or shrubs, small trees, or ceous species normally form rosettes; some South
sometimes vines; nodes trilacunar. Leaves alter- African species of Oxalis may have an evident
nate, often clustered, digitate or pinnate, rarely aerial stem. Bulbs, which are more frequent among
unifoliolate, the terminal leaflet lacking in Biophy- the species of Oxalis of the Cape and in the South
tum, leaflets articulate; petiole usually well- American section ]onoxalis, are formed by fleshy
developed, sometimes expanded into blade-like leaf bases that may or may not be protected by a
phyllodium, or woody and persistent, at the base tunica of leathery, scale-like leaves. These bulbs
and insertion of the blade articulated; stipules may contribute to propagation by forming new
present or not. Inflorescences thyrso-paniculate, bulbs. The bulbs may form at the end of sinking
with monochasia or dichasia in spike-like, umbel- internodes (Fig. 95A, p. 289); exceptionally bulbs
late or capitate arrangement, or racemes. Flowers arise on aerial shoots.
regular, perfect, usually heterotri(di)stylous, or The leaves are borne alternately and are often
plants androcioecious (Dapania), rarely cleistoga- clustered. Rarely they are sessile, as in Oxalis
mous and apetalous; calyx lobes 5, quincuncially sect. Sessilifoliolatae, and usually have well-
overlapping, the three external ones often much developed petioles. These are sometimes
larger, all persistent in fruit; petals 5, mostly con- expanded into blade-like phyllodia (species of
tartly overlapping, free or often postgenitally Oxalis sect. Heterophyllum such as 0. rusciformis),
united at the base above the free claws, caducous, or woody and persistent (Oxalis sect. Thamnoxys ).
white, red, violet to purple or yellow, never blue; At or somewhat above the leaf base and at the
stamens obdiplostemonous, antepetalous stamens insertion of the leaf blade, there are well-
shorter than antesepalous or sterile (Averrhoa p. developed articulations that are engaged in leaf
p.); filaments united at the base in an annulus; movements.
anthers introrse with small connective protrusion; The leaf blade is usually digitate or pinnate; the
nectaries outside the base of antepetalous stamens terminal leaflet is lacking in Biophytum, Dapania,
(actually at the level of a short androgynophore); Sarcotheca, and a few species of Oxalis have simple
ovary superior; carpels 5, antepetalous, united to leaf blades articulated from the petiole, which sug-
form a 5-locular ovary; placentas axile or rarely gests that the blade is actually unifoliolate by
parietal (Oxalis aberrans); stylodia always dis- reduction from a compound leaf. In Oxalis the leaf
tinct, stigmata capitate or punctiform; ovules 2 per blade is usually trifoliolate (Oxalis sect. Tham-
locule, rarely more, anatropous, bitegmic, crassin- noxys), but species with two (Oxalis sect. Pteropo-
ucellate or tenuinucellate, with endothelium, the dae), four (0. deppei and 0. tetraphylla) or 5-12
micropyle directed upwards, slit-like (Averrhoa leaflets ( Oxalis sect. Multifoliolatae, Palmatifoli-
and Biophytum) or funnel-shaped (Oxalis) with atae and Ionoxalis) exist as well.
chalazal appendages. Fruit a loculicidal, some- The digitate leaf blades are considered to be the
times fleshy capsule, or a 5-ribbed berry. Seeds result of peltation (Troll 1937-1943). This is par-
with straight embryo and fleshy endosperm, aril- ticularly evident in four-segmented leaf blades,
late in Dapania. where one leaflet is produced from the transverse
A family comprising five genera and about 880 zone (Querzone) and is opposite to the terminal
species, widespread in tropical and temperate leaflet. Leaf blades with many leaflet pairs are con-
zones. sidered to be derived from more simple forms
(Knuth 1930).
286 A.A. Cocucci
Leaf sensitivity is common in the family. It was daceae are one of the four families where trimor-
early reported for Biophytum, the "living herb". phic heterostyly is known to occur (the others
Leaves are sensitive in response to light (sleeping being Connaraceae, Lythraceae and Pontederi-
position) or touch (evident in Averrhoa, Biophy- aceae). Oxalis is mostly tristylous. Tristyly is also
tum, less so in Oxalis) by moving petioles and leaf found in Biophytum. Loss of tristyly to secondary
blades. Leaves are also sensitive in Sarcotheca. In distyly has taken place in some species of Oxalis
Oxalis each leaflet folds lengthwise along the (Orndurff 1972). Distyly is present in Dapania
middle nerve and sinks to lie against the petiole. and Sarcotheca (Weeler 1992; Richards 1997). Sar-
In some Oxalis and in Biophytum even the cotyle- cotheca celebica, although morphologically disty-
dons are sensitive. In Oxalis acetosella the sleeping lous, is functionally dioecious, with a short-styled,
position is attained 1-3 min after darkening while female-sterile morph, and a long-styled, male-
return to the horizontal position takes place sterile morph (Lack and Kevan 1987). Style
within 1/2-1 h after the leaves are exposed the light heteromorphism is linked with an outcrossing
again. Spontaneous leaf blade oscillations have breeding mode, although autogamy and even
been reported for Oxalis and Averrhoa (Knuth cleistogamy (Oxalis acetosella) are also present.
1930). Flowers are short-lived and open only for a few
hours on sunny days. Bee pollination prevails.
VEGETATIVE ANATOMY. Bladder-like trichomes Some Oxalis of South Africa (0. tubiflora or 0.
occur on the leaf underside (Knuth 1930) and annae) and South America (0. macrostylis) have
uniseriate, slightly moniliform trichomes are comparatively long, salverform corollas and are
found on the stamen filaments (Matthews and probably butterfly pollinated (Vogel 1954). Bees
Endress 2002). Lysigenous secretory spaces are are the most important pollinators in Sarcotheca
commonly situated at the leaf or sepal margins in (Lack and Kevan 1987).
some African Oxalis. Calcium oxalate crystals may
be found in the parenchyma cells of every plant DISPERSAL. Seeds of Oxalis and Biophytum are
organ and in the xylem bundles of the bulb scales. ballistic: the elastic self-everting endotesta ejects
Each cell bears normally one large cubic or pris- the seed explosively (Overbeck 1923). The outer,
matic crystal. Crystal-bearing cells may be elastic skin of the testa splits off from the inner
arranged in cell chains beneath the epidermis. portion of the seed by rupturing of the cell walls
Sieve elements contain a unique variant of P-type of the crystalliferous endotestal cell layer. The
plastids (Behnke 1982). inner tangential walls, the thickened bases of the
radial walls and the crystals remain attached to
FLOWER STRUCTURE. Floral morphology of Oxal- the sclerotic exotegmic layer of the ejected seed.
idaceae has been studied by Matthews and Endress The berries of Averrhoa and Sarcotheca are
(2002) within a broader comparative context. probably zoochorous.
FRUITS, SEEDS AND SEEDLINGS. The fruits are POLLEN MORPHOLOGY. Pollen grains are
more (Dapania) or less (Oxalis) fleshy capsules, described as tricolpate or tricolporoidate; some
berries (Averrhoa, Sarcotheca), or fleshy capsules Oxalis are 3-colporate and exceptionally panto-
eventually becoming schizocarpic (Biophytum). colp( or )ate or 4-colp( or )ate; the exine is finely
The seed coat has a crystalliferous endotesta and reticulate (Fig. 93). The shape is oblate to spher-
a well-developed exotegmen; the endosperm is oidal (Erdtman 1952; Huynh 1969).
oily (Bouman 1974; Corner 1976; Boesewinkel
1985). EMBRYOLOGY. Pollen is usually shed in the binu-
In some species ( 0. rubella) the primary shoot cleate state; that of Biophytum has been reported
is initially enclosed in a sheathing cotyledonary to be trinucleate. The ovules are crassinucellate in
tube. The growing petiole of the first leaf pushes Averrhoa, and tenuinucellate in Oxalis and Bio-
and presses the epicotyl down into the deeper phytum. The ovules have a slit-like (Averrhoa and
parts of the cotyledonary tube. Biophytum) or funnel-shaped ( Oxalis) micropyle
and have chalaza! appendages; in Oxalis the nucel-
BREEDING SYSTEMS AND POLLINATION BIOLOGY. lus together with the inner integument is elevated
At least in Oxalis, the stigma is of the dry type and and separated from the outer integument by a
formed by multicellular and multiseriate papillae stalk (Matthews and Endress 2002). Megagameto-
(Heslop-Harrison and Shivanna 1977). Oxali- phyte development is of the Polygonum type
Oxalidaceae 287
with their margins above the claw, multicellular

hairs on the petals, sieve-tube plastid type and
absence of oxalate druses. In addition, the absence
of ellagic acid and the presence of the rare benzo-
quinone rapanone are shared by both families
(Hegnauer 1969, 1990). Pentamery and obdiplo-
stemony are also shared but probably as ple-
siomophic conditions within the Rosid clade
(Matthews and Endress 2002). Geographical dis-
tribution patterns suggest the origin of the family
in the southern hemisphere, prior to the separa-
tion of South America and Africa (Raven and
Axelrod 1974).
Within the family, Dapania, Sarcotheca and
Averrhoa are closely related (Veldkamp 1967). The
Malagasy species of Dapania is intermediate to
Sarcotheca. This group has retained several possi-
ble plesiomorphic character states: a higher chro-
Fig. 93. Oxalidaceae. Oxalis lasiopetala, pollen in polar view, mosome base number, multifoliolate leaf blades,
REM, bar = 10 Jlm. (Palynological Laboratory Stockholm) and woody habit.
UsEs. Oxalis tuberosa (oca) tubers, a crop from

(Davis 1966) or Allium type (Herr and Dowd Andean civilizations, are locally important as an
1968). Endosperm development is nuclear. alternative to potatoes. Its wild relatives come
from Bolivia and Peru (Emswhiller 2002;
KARYOLOGY. The numerous chromosome counts Emswhiller and Doyle 2002). Fruits of Averrhoa
available for Oxalis reveal virtually every gameto- carambola (starfruit or bilimbi) are edible and
phytic chromosome number from 5 to 42, with also used as a bleaching agent.
predominance of n = 7 (Fedorov 1969). Biophytum
has n = 7-16 with a mode at n = 9; Averrhoa has KEY TO THE GENERA
n = 11 and 12. This pattern has been interpreted as
1. Fruit a dry capsule 2
being due to aneuploid reduction from ancient - Fruit fleshy, sometimes dehiscent 3
ployloids with n = 12 (Lewis 1979) or, according 2. Capsule valves not much spreading, ± united with the
to De Azkue (2000), from primitive diploids with central columella 5. Oxalis
n = 6. - Capsule valves, when spreading out, completely detaching
from the central columella 4. Biophytum
3. Leaves plurifoliolate (5 or more leaflets); ovules and seeds
AFFINITIES. Oxalidaceae have traditionally been 3- 7 per locule 3. Averrhoa
related to Geraniaceae (Engler 1931; Cronquist - Leaves unifoliolate or trifoliolate; ovules and seeds 0-2 per
1981; Thorne 2000), with which they share pen- locule 4
tamerous flowers, obdiplostemony, a syncarpous 4. Shrubs or trees; inflorescences thyrso-paniculate; fruit
indehiscent; seeds exarillate 1. Sarcotheca
gynoecium and stamina} nectaries. However, - Lianas (rarely shrubs); inflorescences racemes; fruit dehis-
studies of plastid and nuclear gene sequences cent into a 5-rayed star; seeds arillate 2. Dapania
place Oxalidaceae close to Connaraceae and
families such as Cephalotaceae, Cunoniaceae and
Elaeocarpaceae (Chase et al. 1993; Price and 1. Sarcotheca Blume Fig. 94
Palmer 1993; Savolainen, Fay et al. 2000; Soltis et Sarcotheca Blume, Mus. Bot. Ludg. Bat. 1: 241 (1850);
al. 2000). The affinity between Oxalidaceae and Veldkamp, Blumea 15: 527-543 (1967), rev.
Connaraceae had already been recognized in the Roucheria Miq. (1859).
pre-molecular era. Recent morphological studies Connaropsis Planch. ex Hook. f. (1860).
(Mathews and Endress 2002) strongly supported
this affinity by a set of morphological and anatom- Shrubs or trees. Leaves unifoliolate or trifoliolate,
ical synapomorphies such as imbricate petal aes- when trifoliolate, lateral leaflets early caducous.
tivation, trimorphic heterostyly, postgenital union Inflorescences thyrso-paniculate, axillary or
of the petals which appear to be hooked together pseudoterminal; flowers small, heterodistylous,
288 A.A. Cocucci
Trees. Leaves alternate, pinnate, plurijugate,

terminal leaflet lacking. Inflorescences thyrso-
paniculate. Flowers small, downy, purple streaked;
stamens all fertile or 5 staminodial; ovules 3-7 per
locule. Fruit baccate. Seeds sometimes arillate. Two
species, A. carambola L. and A. bilimbi L., proba-
bly of Indomalaysian origin and spread by man
throughout the tropics.
4. Biophytum DC.
Biophytum DC., Prodr. 1: 689 (1824).
Herbs up to 1 m, stem sometimes woody. Leaves

sometimes rosulate, pinnate, plurijugate, terminal
leaflet lacking. Inflorescences umbelliform or ±
capituliform. Flowers yellow, heterotristylous (at
least in part); stamens united at the base; ovules
many per locule. Fruit a loculicidal capsule. About
50 species widespread in the tropical zones of
America, Africa and Asia, reaching the Himalayas
and Mexico in the North, and the Zambezi River
Fig. 94. Oxalidaceae. Sarcotheca glauca. A H abit. B Flower.
C Fruit. D Androecium and gynoecium of short-styled flower. and Madagascar in the South; particularly species-
E Same, gynoecium. F, G Same of long-styled flower. Drawn by rich in Madagascar; growing in dry and wet open
R. van Creve!. (Veldkamp 1967) places.
5. Oxalis L. Fig. 95
red or white; stamens 10; ovules 2 per locule. Fruit Oxalis L.,Sp. Pl.: 433 (1753); Lourteig,Bradea 7(1): 1-99 (1994),
a globose berry with 5 episeptal furrows. Eleven subg. Thamnoxys, 7(2): 1- 629 (2000), subg. Oxalis and
Trifidus.
species or probably less, each of local range
in Thailand, Malay Peninsula, Borneo and
Philippines. Annual and perennial herbs, sometimes succulent,
including rhizome- or bulb-bearing geophytes,
chamaephytes, shrubs or vines ( 0. rhombifolia
2. Dapania Korth. Jacq. and 0. scandens H.B.K. from South America)
Dapania Korth., Nederl. Kruidk. Arch. 3: 381 (1854); Veldkamp, and an aquatic of the Western Cape ( 0. natans
Blumea 15: 523-527 (1967), rev. Eckl. & Zeyh.) with a long internode holding
a cluster of floating leaves. Leaf blades with
Woody lianas, rarely erect shrubs. Leaves unifolio- ( 1-)3-many leaflets. Flowers solitary, thyrsic or
late, stipulate. Inflorescences axillary racemes. subumbellate, heterotristylous, distylous, or
Flowers red to white, heterodistylous or androdi- homostylous. Fruit capsular. Seeds with self-evert-
oecious, stamens all fertile or 5 staminodial; ing endostesta. About 500 species mainly in tropi-
ovules in hermaphrodite flowers 1-2 per locule. cal and subtropical zones, extending into arctic
Fruit fleshy, loculicidal to the base with patent (Lappland) and subantartic (Falkland Islands)
valves, 1-6-seeded. Seeds arillate, aril bright to zones, particularly species-rich in South Africa
whitish yellow. Three species, two of which and Andean South America. 0. magellanica Forst.
in Malaya, Sumatra and Borneo, and one in is distributed in S South America, New Zealand,
Madagascar. Tasmania and Australia. Some species are widely
naturalized weeds. The genus is divided into four
subgenera [subg. Oxalis, 412 spp., subg. Monoxalis
3. Averrhoa L. (Small) Lourteig, 2 spp., subg. Thamnoxys (Endl.)
Averrhoa L., Sp. Pl.: 428 (1 753); Lourteig & Cerceau-Larrival, Reiche, 71 spp., subg. Trifidus Lourteig, 2 spp.] and
Phytologia 56: 381-412 (1984), rev. 34 sections.
Carambola Adans. (1763).
Oxalidaceae 289
Selected Bibliography Fig. 95. Oxalidaceae. A Oxalis contracta, floriferous plant with
subterranean tubers. B-H Oxalis leptocaulos. B Habit. C Leaf.
D Flower. E Androecium and gynoecium. F Pistil. G Capsule.
Behnke, H.-D. 1982. Sieve-element plastids of Connaraceae H Seed. (Knuth 1930)
and Oxalidaceae. A contribution to the knowledge of P-type
plastids in dicotyledons and their significance. Bot. Jahrb.
Syst. 103: 1-8.
Boesewinkel, F.D. 1985. Development of ovule and seed-coat in
Averrhoa (Oxalidaceae) with notes on some related genera. Erdtman, G. 1952. See general references.
Acta Bot. Neerl. 34: 413- 424. Federov, A. 1969. See general references.
Bouman, F. 1974. Developmental studies of the ovule, integu- Hartl, D. 1957. Die Pseudosympetalie von Correa speciosa
ments, and seeds in some angiosperms. Doctoral Thesis, (Rutaceae) und Oxalis tubiflora (Oxalidaceae). Abh. Akad.
University of Amsterdam. Naarden: LOS. Wiss. Lit. Mainz, Abh. Math. Naturwiss. Kl. 2: 53-63.
Chase, M.W.et al. 1993. See general references. Hegnauer, R. 1969, 1990. See general references.
Corner, E.J.H. 1976. See general references. Herr, J.M., Dowd, M.L. 1968. Development of the ovule and
Cronquist, A. 1981. See general references. megagametophyte in Oxalis corniculata L. Phytomorphol-
Davis, G.L. 1966. See general references. ogy 18: 43- 53.
De Azkue, D. 2000. Chromosome diversity of South American Heslop-Harrison, Y., Shivanna, K.R.1977. The receptive surface
Oxalis (Oxalidaceae) Bot. J. Linn. Soc. 132: 143-152. of the Angiosperm stigma. Ann. Bot. 41: 1233-1258.
Emswhiller, E. 2002. Biogeography of the Oxalis tuberosa Huynh, K.-1.1969. Etude du pollen des Oxalidacees. Bot. Jahrb.
alliance. Bot. Rev. 68: 128- 152. Syst. 89: 272- 303.
Emswhiller, E., Doyle, J.J. 2002. Origin of domestication and Knuth, R. 1930. Oxalidaceae. In: Engler, A. (ed.) Das Pflanzen-
polyploidy in oca (Oxalis tuberosa: Oxalidaceae). 2. Chloro- reich IV. 95. Weinheim: Engelmann, pp. 1-481.
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Unwin.
Parnassiaceae 291
Parnassiaceae
M.P. SIMMONS
Parnassiaceae Gray, Nat. arr. Brit. pl. 2: 623, 670 (1821), nom. leaf of Parnassia appears sessile. However, the
cons. single leaf trace divides from the stem vasculature
Lepuropetalaceae (Engl.) Nakai (1943). well below the leaf, and runs parallel to the stem
vasculature before entering the lamina. This
Perennial rosulate herbs or diminutive winter indicates that the leaf is pseudo-sessile, with the
annuals, glabrous or young petiole bases puberu- petiole fused to the stem. Other cauline leaves, if
lent, with conspicuous tannin sacs in epidermis. present, are truly sessile (Watari 1939). Leaf vena-
Leaves alternate or subopposite, spathulate, ovate, tion is campylodromous in Parnassia, acrodro-
reniform, or orbicular, entire, exstipulate, all peti- mous in Lepuropetalon. Leaf crystals are absent.
olate or cauline leaf(s) appearing sessile. Flowers Stomata are anomocytic, on the abaxial leaf
solitary on scapes or on lateral shoots, hermaph- surface in Parnassia, on both leaf surfaces in
rodite, 5-merous, weakly zygomorphic; sepals Lepuropetalon (Gornall and Al-Shammary 1998).
(4)5( -7), persistent; petals (0, 4)5( -7), minute or Fimbriate appendages on the young leaves of Par-
showy; stamens 5, antesepalous, anthers lon~it~ nassia secrete mucilage (Solereder 1908; Metcalfe
dinally dehiscent, introrse or extrorse and mdl- and Chalk 1950).
vidually dehiscent above stigmas; staminodes
antepetalous, glandular, dilated distally; ovary FLORAL MORPHOLOGY. The flowers of Parnassia
superior to half-inferior, 3-4(5)-carpellate, placen- are weakly zygomorphic (Martens 1936; Hultgard
tation axile or parietal; ovules horizontal, numer- 1987). The calyx is quincuncial, with both right-
ous, bitegmic or unitegmic; style terminal, short and left-handed spirals equally frequent. The
or obsolete; style branches or stylodia distinct; immature stamens are introrse. Individually, at the
stigmas commissural. Fruit a membranous rate of about one stamen per day, the stamens
capsule, loculicidally dehiscent a~ apex. Se~ds elongate and bend inwards, dehiscing extrorsely
numerous, cylindrical or oblong, mmute, blackish upwards, directly over the stigmas. After dehis-
or testa transparent and embryo opaque, endo- cence, the stamens bend outwards, lying between
sperm a single cell layer or 0. . the petals. The stamens move in order of th~ir age
A family of two genera and about 71 species, (the stamen opposite the largest sepal first) m one
widely distributed in the Northern Hemisphere, of two zigzag orders (Gris 1868; Martens 1936;
most diverse in China and the Himalayas, and in Hultgard 1987). Nectar is secreted from the pad of
South America. tissue bearing the staminodial rays and not from
the glistening globules at the apices of the rays
VEGETATIVE STRUCTURES. Rhizomes are present (Fig. 96B, C). A central stalk is evident on eac.h
only in Parnassia. The surface of the rhizome is staminode, such that the number of stalks IS
composed of a 3-layered exoderm with suberised generally uneven. In addition to visually attracting
cells. The cortex and solid pith of the rhizome insects (Daumann 1935), the globules may serve to
consist of starch-filled parenchyma cells. Adventi- transfer the released pollen onto the insects
tious roots originate from the pericambium. The (Bennett 1871). The gynoecium of Parnassia is
pericycle of Parnassia is 6-layered and fibrous, in syncarpous at the base, but paracarpous and
Lepuropetalon 1-2-layered and sclerenchymatous. unilocular above, with T-shaped placentae. In con-
Three collateral vascular bundles surround a solid trast, the placentation of Lepuropetalon is strictly
pith in Lepuropetalon, a hollow pith in Parnassia parietal, without T-shaped placentae.
(Korta 1972; Gornall and Al-Shammary 1998).
Secretory cells with tanniniferous contents occur FLORAL ANATOMY. In Parnassia, the vascular
in the epidermis of Lepuropetalon (Fig. 96F) and traces of the nectaries separate from the petal
Parnassia (Metcalfe and Chalk 1950). The lower- traces above the point at which the stamen traces
most or only, depending on the species, cauline separate from the sepal traces. Therefore, the nee-
292 M.P. Simmons
taries have been interpreted as an inner androe- hybrid polyploids (P. palustris; Hultgard 1987). P.
cial whorl of staminodes (Arber 1913; Eames 1961; palustris includes diploid, triploid, tetraploid, pen-
Sharma 1968). The same pattern appears to apply taploid, and hexaploid individuals (2n = 17, 18, 27,
to Lepuropetalon (Murbeck 1918). The globiferous 36, 32-37,43-45 and 54; Funamoto et al. 1998 and
filaments of the staminodes of Parnassia are each references cited therein). The reports of 2n = 20 for
supplied by separate vascular traces. Based on this P. palustris are probably erroneous (Gastony and
vasculature pattern, the individual stamens Soltis 1977). The hexaploids of P. palustris have
and staminodes have been interpreted as derived been considered to be the product of triploid
from stamen-fascicles that have become connate hybrids that have undergone chromosome dou-
(Drude 1875; Arber 1913; Eames 1961; Bensel and bling (Spongberg 1972). The only differences in
Palser 1975; but see Klopfer 1972). gross morphology between diploid and tetraploid
populations of P. palustris are the generally larger
EMBRYOLOGY. The embryology of Parnassia has pollen grains and seeds in tetraploids (Hultgard
been examined by Pace (1912), Saxena (1964), and 1987).
Sharma (1968). Anthers are tetrasporangiate (bis-
porangiate at anthesis in Lepuropetalon), have a POLLINATION. The pollinators of P. palustris are
fibrous endothecium, and a secretory tapetum. non-social wasps (Pompiloidea, Apoidea, and
Tapetal cells are one- or two-nucleate. Microspore Vespoidea), flies (Syrphidae, Diptera), crane flies
tetrads are tetrahedral. Mature pollen is bi-nucle- (Tipulidae), bottle flies (Calliphoridae), mos-
ate, with equally-sized vegetative and generative quitoes (Culicidae), butterflies and moths
nuclei. (Lepidoptera), and ants (Myrmica; Sprengel 1793;
Ovules are anatropous, unitegmic in Lep- Daumann 1935; Spongberg 1972; Hultgard 1987;
uropetalon (Murbeck 1918), bitegmic in Parnassia, Proctor et al. 1996). Insects are visually attracted
tenuinucellate, and have Polygonum type of by the globules of the staminodes and the promi-
embryo sac formation. The megaspore mother cell nent, dark petal venation on the white petals that
generally forms a linear tetrad of megaspores, act as nectar guides to the nectaries at the
although T-shaped tetrads also occur. The second, staminode bases (Sprengel 1793; Daumann 1935;
third, or fourth megaspores may develop, and Spongberg 1972). Once in the vicinity of the
two megaspores may develop simultaneously. flowers, the scent from the nectaries induces flies
Endosperm formation is nuclear. The endosperm to land on the flowers (Daumann 1935).
is consumed by the embryo until only a single cell
layer remains, or is entirely absent in mature seeds REPRODUCTIVE SYSTEMS. The reproductive
(Arber 1913). The endosperm is composed of a system of Lepuropetalon is essentially unknown.
single cell layer in Lepuropetalon (Murbeck 1918). Based on the winter-annual habit and the
Embryos are straight, with short cotyledons. dehiscence of the anthers directly over the
stigmas, Lepuropetalon is probably self-pollinated
POLLEN MORPHOLOGY. Pollen grains are shed as (Spongberg 1972). In contrast, the reproductive
monads (Pace 1912). The grains are prolate to system of Parnassia palustris has been extensively
spheroidal and tricolporate; tetracolporate and studied. P. palustris has been reported to be pro-
syncolpate grains occur in Parnassia. The sexine tandrous, requiring insect pollination (Sprengel
is as thick or slightly thicker than the nexine, 1793; Gris 1868; Bennett 1871; Hultgard 1987).
with reticulate sculpturing that is less-developed However, protandry has been shown to be imper-
towards the poles and colpi (Erdtman 1952; fect as the styles are receptive when the anthers
Hideux and Ferguson 1976; Hultgard 1987). dehisce individually above them, and self-
fertilization does occur (Martens 1936; Hultgard
KARYOLOGY. Chromosome numbers have been 1987). Based on seed set and germination, Martens
extensively studied in Parnassia. Parnassia has ( 1936) concluded that autogamy is superior to
three base chromosome numbers: x = 7, 8, and 9 outcrossing in P. palustris, and that the stamen
(Funamoto et al. 1998). P. kotzebuei (2n = 18, 36), dehiscence above the styles favors autogamy.
P. laxmannii (2n = 18, 36), P. oreophyla (2n = 18, In contrast, based on seed set, Hultgard (1987)
36) include both diploid and tetraploid individu- concluded that outcrossing is superior to
als (Funamoto et al. 1996). The tetraploids autogamy. Hultgard suggested that the species
have been considered to be autopolyploids (P. primarily reproduces by outcrossing, and inter-
oreophyla; Funamoto et al. 1996; P. palustris; preted the method of stamen dehiscence to favor
Wentworth and Gornall 1996) or intraspecific outcrossing.
Parnassiaceae 293
FRUIT AND SEED. The fruit is a loculicidal capsule closely related or transitional to Droseraceae
with a persistent calyx, which is borne erect and (based on characters of the gynoecium; Drude
opens at the apex, with numerous tiny seeds. 1875; Pace 1912; Arber 1913) and/or Hypericaceae
From about 100 to 2000 pale- to dark-brown seeds (based on characters of the androecium or
develop in capsules of Parnassia palustris flavonoids; Lindley 1846; Drude 1875; Arber 1913;
(Hultgard 1987). Jay 1971), or even Nymphaeaceae (based primarily
The testa of P. nubicola is 5-6 cells thick. The on vegetative characters; Hallier 1901). Many
outer epidermal cells enlarge and accumulate workers (Drude 1875; Murbeck 1918; Sharma
tannin-like deposits (Sharma 1968). In P. palustris, 1968; Klopfer 1972; Hultgard 1987) have supported
these cells are tetragonal or pentagonal, isodia- recognition of Parnassiaceae as a distinct family
metric to elongated, and have raised borders. An (or order, Takhtajan 1997).
air cavity develops between the testa and the In several broad-scale molecular phylogenetic
embryo. The testa is transparent, but the embryo analyses, Parnassiaceae have been resolved as the
is not (Hultgard 1987). Seed germination of Par- sister group of Celastraceae (e.g., Chase et al.1993;
nassia is epigeal (Spongberg 1972). The testa of Savolainen, Chase et al. 2000; Soltis et al. 2000). In
Lepuropetalon is 2 cells thick, membranous, and a well-sampled phylogenetic analysis of Celas-
the cells have raised borders. There are small inter- traceae using 26S nrDNA and a simultaneous
cellular spaces between the two layers of cells analysis of morphological and molecular charac-
forming the testa. An air cavity does not develop ters (Simmons et al. 2001), Parnassiaceae were
between the testa and the embryo (Murbeck resolved, in a weakly-supported clade, as part
1918). of an early-derived lineage within Celastraceae.
However, because of their distinctive morphology,
DISPERSAL. The capsules of Parnassia are borne and the fact that the clade from Simmons et al.
erect, so that the very light seeds, which are mainly (2001) was weakly-supported and based only on
wind-dispersed (Ridley 1930), only fall out by characters from 26S nrDNA, Parnassiaceae are
wind blowing on the capsule (Sprengel1793). The retained as a separate family.
seeds are buoyant and float for several weeks,
allowing dispersal by water (Praeger 1913). The air DISTRIBUTION AND HABITATS. The center of
cavity between the testa and the embryo facilitates diversity of Parnassia is in China and the
dispersal by wind and water (Hultgard 1987). Himalayas, to which most (49) species are endemic
(Gu and Hultgard 2001). Parnassia palustris is the
PHYTOCHEMISTRY. Flavonoids of Lepuropeta[on most widespread species of the genus, with popu-
and/or Parnassia have been investigated by Jay lations in North America, Europe, and Asia, and
(1971) and Bohm et al. (1986). The flavonoid south to Morocco (Korta 1972). In the New World,
profiles support the close relationship of the two P. townsendii is the southernmost species and is
genera and their separation from Saxifragaceae. endemic to the Sierra Madre Occidental of Mexico.
Methanolic extracts of P. nubicola were shown to P. townsendii may represent relictual populations
have antiproliferative activity against the growth of the P. fimbriata complex, the other members of
of human keratinocytes (KC and Muller 1997). which (P. fimbriata and P. intermedia) have dis-
persed northward (Bye and Soltis 1979). Parnassia
AFFINITIES. The close relationship of the two grows in temperate to arctic regions, preferentially
genera is supported by floral and vegetative char- in open, moist habitats from shore meadows
acters (Murbeck 1918; Spongberg 1972; Gornall to mountainsides. Lepturopetalon has a disjunct
and Al-Shammary 1998). Several workers have distribution in North and South America, where
treated Lepuropetalon and Parnassia as members it grows in seasonally-moist rock outcrops and
of Saxifragaceae (Hooker and Thomson 1858; sandy soils, including disturbed areas. It is proba-
Engler 1930; Cronquist 1981). However, this puta- bly more widespread than currently known (Ward
tive relationship has been recognized as problem- and Gholson 1987).
atic based on various characters from morphology
(Drude 1875; Hallier 1901; Murbeck 1918), stamen ECONOMIC IMPORTANCE. Several species of
vasculature (Eames 1961), embryology (Sharma Parnassia are cultivated in rock gardens and
1968), flavonoids (Jay 1971), floral development bog gardens (Huxley et al. 1992).
(Klopfer 1972), floral anatomy (Bensel and Palser
1975), and pollen structure (Hideux and Ferguson CoNSERVATION. Parnassia caroliniana and P.
1976). Parnassia has also been suggested to be cirrata are rare (Walter and Gillett 1998).
294 M.P. Simmons
Fig. 96. Parnassiaceae. A-B Parnassia palustris. A Habit. B above stigmas; ovary 3-4(5)-carpellate. Seeds
Flower. C Parnassia alpicola, flower. D-F Lepuropetalon oblong, testa transparent, embryo opaque,
spathulatum. D Habit. E Flowering plant. F Flower with sepals
showing the pattern of epidermal tannin sacs. (Takhtajan endosperm minimal or 0. 2n = 14, 17,18,27, 32, 36,
1981) 32-37, 43-45 and 54. About 70 spp., northern
hemisphere-holarctic, south to northern Mexico,
Morocco, and India, various open, moist habitats
including rock outcrops, bogs, and wet meadows,
KEY TO THE GENERA
grasslands, forests, and disturbed areas.
1. Diminutive winter annuals forming hemispherical clumps, Drude (1875) recognized four sections based on
without rhizomes; leaves spathulate; flowers inconspicu- characteristics of the staminodes, ovary position,
ous, petals smaller than sepals. Northern and Southern and carpel number. Engler (1930) followed
Hemispheres of the New World 2. Lepuropetalon
- Perennials forming rosettes, usually with rhizomes or root- Drude's classification, and added a fifth section.
stocks; leaves ovate, reniform, or orbicular; flowers con- Building on Drude and Engler's classifications, Ku
spicuous, petals larger than sepals. Northern Hemisphere (1987, 1995) recognized nine sections and nine
of the Old and New World 1. Parnassia series for the Chinese species, based on character-
istics of the staminodes, petal margins, and cauline
GENERA OF p ARNASSIACEAE leaf number.
1. Parnassia L. Fig. 96A-C

2. Lepuropetalon Elliott Fig. 96D-F
Parnassia L., Sp. Pl.: 273 (1753); Drude, Linnaea 39: 239-324
(1875), rev.; T.C. Ku, Bull Bot. Res., Harbin 7: 1-59 (1987), Lepuropetalon Elliott, Sketch Bot. S. Carolina 1: 370 (1817);
reg. rev.; Gu and Hultgard, Parnassia, pp. 358-379, in: Fl. Spongberg, J. Arnold Arbor. 53: 409-498 (1972), part. rev.
China vol. 8 (2001).
Diminutive winter annuals, caulescent or acaules-
Perennials, forming rosettes, usually with rhi- cent, forming hemispherical clumps, glabrous.
zomes or rootstocks, glabrous or young petiole Leaves alternate or subopposite, spathulate.
bases puberulent. Leaves alternate, ovate, reni- Flowers minute; petals (0, 4)5, minute; staminodes
form, or orbicular; basal leaves petiolate, cauline unornamented; anthers introrse; ovary 3-carpel-
leaf(s) appear sessile. Flowers borne on scape; late. Seeds cylindrical, blackish, endosperm
sepals (4)5(-7); petals (4)5(-7), showy; stamin- minimal. 2n = 46. Only one sp., L. spathulatum
odes glandular, ornamented with many terminal Elliott, with a disjunct distribution in southeastern
globular stalks or 2- 5-lobed; immature anthers U.S.A., the Gulf Coast of Mexico, Uruguay, central
introrse, mature extrorse, individually dehiscent Chile.
Parnassiaceae 295
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Hallier, H. 1901. Ober die Verwandtschaftsverhaltnisse der
Angiosperm phylogeny inferred from a combined data set
Tubifloren und Ebenalen den polyphyletischen Ursprung
of 18 S rDNA, rbcL, and atpB sequences. Bot. J. Linn. Soc.
der Sympetalen und Apetalen und die ~nordnung ~er 133: 381-461.
Angiospermen iiberhaupt. Abh. Naturw1ss. Naturw1ss.
Spongberg, S.A. 1972. The genera of Saxifragaceae in the
Verein Hamburg 16: 1-112.
southeastern United States. J. Arnold Arbor. 53: 409-
Hideux, M.J., Ferguson, I.K. 1976. The stereostructure of the 498.
exine and its evolutionary significance in Saxifragaceae
Sprengel, C.K. 1793. Das entdeckte Geheimnis d.er N~tur .im
sensu lato. In: Ferguson, I.K., Muller, J. (eds.) The evolu-
Bau und in der Befruchtung der Blumen. Berhn: Fnednch
tionary significance of the exine. London: Academic Press,
Vieweg dem aeltern.
pp. 327-378.
Takhtajan, A. (ed.) 1981. See general references.
Hooker, J.D., Thompson, T. 1858. Prrecursores ad floram
Takhtajan, A. 1997. See general references.
Indicam. J. Proc. Linn. Soc. 2: 54-103.
296 M.P. Simmons
Walter, K.S., Gillett, H.J. 1998. 1997 IUCN red list of threatened Watari, S. 1939. Anatomical studies on the leaves of some Sax-
plants. Gland, Switzerland: IUCN- The World Conservation ifragaceous plants, with special reference to the vascular
Union. system. J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 5: 195-316.
Ward, D.B., Gholson, A.K. 1987. The hidden abundance of Wentworth, J.E., Gornall, R.J. 1996. Cytogenetic evidence
Lepuropetalum spathulatum (Saxifragaceae) and its first for autopolyploidy in Parnassia palustris. New Phytol. 134:
reported occurrence in Florida. Castanea 52: 59-67. 641-648.
Pellicieraceae 297
Pellicieraceae
K. KUBITZKI
giotropic axes are produced that bear terminal leaf

Pellicieraceae (Triana & Planchon) L. Beauvis. ex Bullock in
Taxon 8: 182 (1959). clusters.
Apart from subterranean roots, aboveground
Usually small (more rarely up to 18m high) man- aerial roots are formed on the stem base in numer-
grove trees with fluted trunk bases. Leaves spirally ous orthostichies (Fig. 97). These roots do not
arranged, asymmetric, involute in bud, glabrous, immediately penetrate the surface of the stem base
sessile, exstipulate, the wider half of the blade ini- but grow downwards inside the bark. Thus, the
tially bearing a series of prominent but ephemeral cortical tissue is elevated but the surface remains
(salt?) glands rolled innermost in bud but after unbroken, except for numerous prominent
expansion becoming entire; leaf base narrowed to lenticels, so that the vertical series of adventitious
the insertion with a pair of extrafloral nectaries roots below a covering layer of bark forms the
and decurrent. Flowers solitary in the 1-3leaf axils fluted buttresses (Tomlinson 1986). These but-
below the resting terminal bud, pentamerous, tresses form a conical mantle around the stem
sessile, enclosed by 2 prophylls before anthesis, which at soil level may be up to 1.5 m wide and
large (-12 em wide) at anthesis; calyx lobes short, may extend on the stems up to the high-tide mark
free, quincuncially arranged, with numerous small (Fuchs 1970).
glands internally at the base; petals free, lanceate, Anatomically, Pelliciera is characterized by the
about 6 em long, ephemeral and falling with the presence of raphides in idioblasts of the parenchy-
sepals and stamens; stamens 5, free, lying within matous tissue; vessel segments with oblique, sim-
the alternate groves of the ovary; anthers narrow, ple perforations; imperforate tracheary elements
about 2.5 em long, sagittate, distally pointed, with inconspicuously bordered pits; elongate,
extrorsely dehiscing by longitudinal slits; gynoe- sinuous fibres in the mesophyll; encyclocytic
cium 2-carpellate, syncarpous; ovary superior, stomata; branched, sclerenchymatous idioblasts
tapering into a narrow style with a bifid stigma; and elongated thick-walled fibres in the cortex and
ovary cylindrical, woody, about 6cm long with 10 pith; and petioles with an almost cylindrical vas-
longitudinal grooves; locules 2, each containing a cular system (Beauvisage 1920; Metcalfe and Chalk
single anatropous ovule pendulous from the inner 1950; Baretta-Kuipers 1976).
angle, only one of them developing, or one locule
sterile. Fruit indehiscent, napiform, 8-12 em long EMBRYOLOGY AND CHROMOSOMES. Unknown.
and about as wide, somewhat flattened, the style
persisting; fruit wall ridged, leathery externally POLLEN MORPHOLOGY. Pollen grains are 3-colpo-
but spongy within; seed solitary, exalbuminous; rate, spheroidal, crassi-exinous, with lalongate ora
seed coat at maturity of ribbon-like fragments; (Erdtman 1952). Ornamentation of the grains vary
embryo cordate, with elongate hypocotyl pointing both in Recent and fossil material from scabrate to
into the stylar beak; plumule reddish, long, slender distinctly reticulate, and a size range of 40-90 J..lm
and hooked, enclosed by 2 fleshy cotyledons. is found likewise in modern and fossil pollen
Monotypic, Pelliciera rhizophorae Triana & (Graham 1977).
Planchon a mangrove of Central and northern
South America. PHYTOCHEMISTRY. Tests for aluminium accumu-
lation were negative, in contrast to the rich
MORPHOLOGY AND ANATOMY. The evergreen occurrence of aluminium in Theaceae and
trees have a well-developed, erect trunk and a Ternstroemiaceae (Chenery 1948).
narrow, elongate crown. They develop terminal
"naked" buds (not protected by bud scales) AFFINITIES. Traditionally included in Tern-
in association with flowering. Branching is dis- stroemiaceae, Pelliciera was removed from this
continuous (Attim's model) and tiers of pla- family on anatomical grounds by Beauvisage
298 K. Kubitzki
Fig. 97. Pellicieraceae. Fluted trunk bases of mangrove species Oligocene, Pelliciera was a common member of
Pelliciera rhizophorae, Piedra Blancas N.P., Costa Rica. Cour- coastal communities throughout the Caribbean
tesy W. Huber, A. Weissenhofer, Vienna
region and from southern Mexico to Panama and
Guyana, and also was present in Nigeria but, by the
end of the Tertiary, it was reduced to its present
range (Fuchs 1970; Graham 1977}. Its increasing
(1920}. It differs from Ternstroemiaceae and range restriction is attributed, i.a., to changes in
Theaceae in possessing raphides, in having wood climate, sea level, soil salinity (Jimenez 1984} and
with pore multiples and simple vessel perfora- competition by Rhizophora. With Rhizophora,
tions, and in lacking aluminium accumulation, Pelliciera occupies the intertidal zone, but with a
features shared with Tetrameristaceae and much more restricted ecological range, typically
Marcgraviaceae. Indeed, recent molecular studies in sheltered sites such as estuarine banks or
employing both plastid and nuclear genes protected beaches (Tomlinson 1986}.
(Savolainen, Chase et al. 2000; Savolainen, Fay The fallen fruits release the seeds which are
et al. 2000; Soltis et al. 2000; Anderberg et al. 2002} dispersed in the water and germinate immediately
provide strong support for a clade consisting after becoming stranded. The poor protection of
of Pelliciera, Tetramerista, Marcgraviaceae, and the seed by the fruit wall is of short duration and
Impatiens, whereas Ternstroemiaceae and precludes dispersal of the fruits by ocean currents.
Theaceae occupy more distant positions within Seeds in water swell after a few hours and sink
the broadly construed Ericales. Pelliciera is closest (Collins et al. 1977}.
to Tetramerista, and the two share, among other
features, pits on the upper surface of the petals. Only one monotypic genus:
DISTRIBUTION, HABITATS AND SEED DISPERSAL.

Pelliciera is distributed on the Pacific coast of Pelliciera Planchon & Triana Fig. 98
Central and northern South America from Costa Pelliciera Planchon & Triana in Benth. & Hook., Gen. Pl. 1: 186
Rica to northernmost Ecuador, and more sparsely (1862); Kobuski, J. Arnold Arbor. 32: 256- 262 (1951), rev.
on the Atlantic coast of Colombia. The microfossil
record shows that, during the Eocene and Description as for family.
Pellicieraceae 299
Beauvisage, L. 1920. Contribution a !'etude anatomique de Ia

famille des Ternstroemiacees. Tours.
Chenery, E.M. 1948. Aluminium in the plant world. Pt. I.
General survey in dicotyledons. Kew Bull. 1948: 173-183.
Collins, J.P., Berkelhamer, R.C., Mesler, M. 1977. Notes on the
natural history of the mangrove Pelliciera rhizophorae Tr. &
Pl. (Theaceae). Brenesia 10/11: 17-29.
Fuchs, H.P. 1970. Ecological and palynological notes on
Pelliciera rhizophorae. Acta Bot. Neerl. 19: 884-894.
Graham, A. 1977. New records of Pellicieria (Theaceae/
Pellicieraceae) in the Tertiary of the Caribbean. Biotropica
9: 48- 52.
Jimenez, J.A. 1984. A hypothesis to explain the reduced distri-
bution of the mangrove Pelliciera rhizophorae. Tr. & Pl.
Biotropica 16: 304-308.
Kobuski, C.E. 1951. Studies in the Theaceae XXII. The genus
Pelliciera. J. Arnold Arbor. 32: 256-262.
Metcalfe, C.E., Chalk, L. 1950. See general references.
Robyns, A. 1967. Flora of Panama, Pt. IV. Fam. 122. Theaceae.
Ann. Missouri Bot. Gard. 54:41- 56.
Savolainen, V., Chase, M.W. et al. 2000. See general references.
Soltis, D. E. et al. 2000. See general references.
Szyszylowicz, I. von 1893. Theaceae. In: Engler, K., Prantl,
A K. Die nattirlichen Pflanzenfamilien III, 6. Leipzig: W.
Engelmann, pp. 175-192.
Tomlinson, P.B. 1986. The botany of mangroves. Cambridge:
Fig. 98. Pellicieraceae. Pelliciera rhizophorae. A Flowering
Cambridge University Press.
branch. B Androecium and gynoecium. C Fruit. Drawn by
H.J. Cuddy. (Robyns 1967)

Baretta-Kuipers, T. 1976. Comparative wood anatomy of
Bonnetiaceae, Theaceae and Guttiferae. Leiden Bot. Ser.
3: 76-101.
300 D.H.Wilken
Polemoniaceae
D.H. WILKEN
Polemoniaceae Juss., Gen. Pl.: 136 (1789), nom. cons. carps living from 2-10 years as vegetative rosettes
are known in Aliciella and Ipomopsis. Vines char-
Annual or perennial herbs, sometimes vines or acterize Cobaea, while Acanthogilia and Cantua
woody shrubs, rarely small trees; indumentum of are exclusively shrubs. Suffrutescent perennials or
multicellular and uniseriate trichomes, eglandular subshrubs also occur in several genera with a
or with terminal unicellular to multicellular majority of annuals, including Aliciella, Collomia,
glands, rarely plants completely glabrous. Leaves Eriastrum, Gilia, Ipomopsis, and Linanthus.
alternate to opposite, pinnately or less often Loeselia, Phlox and Polemonium are primarily
palmately veined, entire to deeply divided, some- perennial but each includes annual species.
times compound, petiolate or sessile, exstipulate. Leaf arrangement is usually alternate, but oppo-
Inflorescences dichasial, in racemose to paniculate site in Phlox and Linanthus. Many annuals develop
or capitate clusters, rarely flowers solitary. Flowers basal rosettes but in some genera (Collomia,
hermaphroditic, actinomorphic, sometimes zygo- Eriastrum, Gilia, Navarretia) the basal leaves are
morphic, hypogynous; sepals (4)5( 6), connate, weakly developed or senesce during bolting and
rarely free, persistent, the tube herbaceous along efflorescence. The leaves of Gymnosteris are
the midribs, hyaline between the midribs, some- reduced to a single whorl of foliaceous bracts sub-
times herbaceous throughout; petals (4)5(6), the tending the terminal flower cluster. Leaves of Pole-
corolla rotate to salverform, funnelform or bilabi- moniaceae lack stipules and can be petiolate or
ate, the lobes mostly convolute in bud; stamens sessile. Leaf blades are pinnately veined, varying
(3-)5(6), alternate with the petals, the filaments from entire to compound. Simple leaf blades with
equal to unequal in length and attached to the entire to toothed or lobed margins predominate
tube, sometimes at differing levels; anthers in Cantua, Collomia, Gymnosteris, Loeselia, and
basifixed to dorsifixed, dithecal, tetrasporangiate, Phlox. Although other genera may have entire
dehiscing by longitudinal slits; ovary inserted on a leaves, the blades of at least the basal and
nectariferous disk, syncarpous, (2)3-locular, pla- lower cauline leaves are deeply lobed or divided.
centae axile; style with (2)3 stigmatic branches; Palmately divided leaves occur in some species of
ovules 1-many per locule, in 2 rows, (hemi)anat- Collomia, Ipomopsis and Linanthus. Cobaea and
ropous, tenuinucellate, with one thick integument. Polemonium have compound leaves, with the ter-
Fruit a dehiscent dry capsule, loculicidal or minal leaflet in Cobaea modified into a branched
rarely septicidal (Acanthogilia, Cobaea), some- tendril.
times explosively dehiscent (Collomia, Phlox),
rarely indehiscent; seeds smooth to angled, some- VEGETATIVE ANATOMY. Primary stems are
times winged, the epidermal cells with spiral characterized by a well-developed, often collen-
thickenings and included mucilage, the cells burst- chymatous cortex and a well-defined endodermis
ing when wetted and forming a mucilaginous coat; (Metcalfe and Chalk 1950). Nodes are unilacunar.
mature embryos straight; endosperm oily. Phloem elements are arranged either as a contin-
The family consists of 18 genera and about 380 uous cylinder or as separate strands associated
species, distributed primarily in North America, with sclerenchyma, but the xylem is in a continu-
extending into Central and South America, with a ous ring. Woody taxa are generally characterized
few species in Eurasia. by nonstoried wood, relatively few vessels per
group, vessel-elements with simple perforation
VEGETATIVE MORPHOLOGY. Most members of plates and alternate circular bordered pits on their
the Polemoniaceae are annuals or herbaceous lateral walls, axial vasicentric scanty parenchyma,
perennials. Herbaceous perennials survive dor- and both multiseriate and uniseriate rays with
mancy either as rhizomes, basal rosettes, or buds mostly erect cells (Carlquist et al. 1984). Mean
at the tips of vertical, underground axes. Mono- vessel diameters range from over 60 f..Lm in Cobaea
Polemoniaceae 301
to less than 30 jlm in Leptodactylon and Phlox. aceae was given by Weberling {1998). Most Pole-
Mean vessel-element lengths range from 116- moniaceae have terminal, determinate inflores-
250 jlm in Leptodactylon to over 400 j.!Iil in Acan- cences. Many- to few-flowered inflorescences are
thogilia and Cantua buxifolia. Tracheids are the most common but solitary flowers occur in some
primary imperforate tracheary elements in such species. All inflorescences found in Polemoniaceae
temperate genera as Eriastrum, Leptodactylon, and can be related to determinate panicles, which
Phlox, whereas tropical genera (Cantua, Cobaea, are of wide occurrence. Partial inflorescences are
Huthia, and Loeselia) have fiber-tracheids or libri- often cymose, thus leading to thyrso-paniculate or,
form fibers. Rayless or near-rayless woods occur if such cymes are reduced to single flowers, to
in Cobaea, Eriastrum, Linanthus, and Phlox. botryoid inflorescences. These cymes are usually
Although wide vessels, vessel diameter dimor- dichasia. The occurrence of paired flowers is due
phism, and abundant ray parenchyma in Cobaea to unequal branching of such dichasia (e.g., in
are common in dicotyledonous vines, the absence Bonplandia geminiflora). Other modifications of
of rays appears to be unique (Carlquist et al.1984). paniculate inflorescences include condensation
Overall, ancestors of Acanthogilia, Cantua and to capitate clusters. The arrangement of parts is
Loeselia appear to have been woody. In contrast, sometimes obscured by metatopies (e.g.,
raylessness or predominantly erect ray cells Langloisia, Gilia). Both basipetal and acropetal
suggest secondary woodiness in Cobaea, Erias- development occurs in various parts of the
trum, Linanthus, and Phlox (Carlquist 1992). inflorescence. In some species of hummingbird- or
Leaves can be either bifacial or isolateral. The hawkmoth-pollinated Ipomopsis, flowers are dis-
cuticle varies from smooth to verrucose. In some placed to one side of thyrso-paniculate inflores-
species of Collomia, Gilia, and Phlox, the epider- cences. Inflorescences can be bracteate throughout
mal cells are mucilaginous. Stomata, sometimes or ebractetae distally. Floral bracts are often
present on both upper and lower surfaces, are reduced but can be foliaceous. In Loeselia the
anomocytic but approach paracytic in taxa with foliar bracts often have conspicuous "window-
linear leaves or leaflets (Metcalfe and Chalk 1950). like" hyaline or scarious areolae.
lsolateralleaves are associated with narrowly dis-
sected or linear leaves, especially in species of arid FLORAL MoRPHOLOGY AND ANATOMY. Polemo-
habitats. Large intercellular spaces are notably niaceae flowers, with few exceptions, are
absent in the meosphyll of bifacialleaves. actinomorphic and hypogynous and have a
Trichomes are either uniseriate eglandular or pentamerous perianth composed of fused sepals
uniseriate glandular with terminal, globose to and fused petals, and 5 alternipetalous stamens.
ovoid heads that are either unicellular or multicel- Tetramerous and hexamerous flowers are found in
lular and often glandular. Eglandular trichomes Linanthus. The ovary has 3, rarely 2, locules
predominate in Eriastrum, Ipomopsis, Langloisia, with axile placentation; the style has 3 stigmatic
Linanthus, and Phlox, whereas glandular tri- branches and is inserted on an entire to lobed
chomes predominate in Aliciella, Allophyllum, annular nectariferous disk. The sepals of Cobaea
Bonplandia, Cantua, Collomia, Gilia, Navarretia, and some species of Gilia are almost completely
and Polemonium. Although most species of Gilia free, but form a tube in other genera. The tube is
have unicellular glands, those of the related herbaceous in Bonplandia, Cantua, Cobaea, Collo-
Aliciella are multicellular. In some species of mia, and Polemonium. The tube of Collomia is
Allophyllum, Collomia, Gilia, Navarretia and notably plicate in late flower and fruit. In all other
Polemonium, glandular trichomes are the source genera the tube is composed of 5 herbaceous
of a distinctive mephitic odor. Langloisia is dis- costae that extend into the lobes and are separated
tinctive in its possession of branched trichomes. by hyaline portions proximal to the sinuses. The
Densely tangled trichomes in some genera (e.g., hyaline portions can be narrower or wider than
Eriastrum, Ipomopsis, Gilia) are uniseriate eglan- the herbaceous costae, and are often ruptured by
dular, repeatedly geniculate, and composed of the developing fruit.
"rectangular parallelepipeds, joined at right angles Corollas can be campanulate, salverform, fun-
to each other" (Grant 1959). Multicellular bristles nelform, or sometimes rotate (Fig. 100). Zygomor-
or setae occur on foliar organs in Langloisia and phic, often bilabiate, flowers occur in Bonplandia
some Loeselia. and some species of Ipomopsis, Langloisia and
Loeselia. The petal lobes are convolute in bud,
INFLORESCENCE STRUCTURE. A detailed morpho- although imbricate petals are reported in Cantua
logical analysis of the inflorescences of Polemoni- {Johnson et al. 1999). In most Polemoniaceae, the
302 D.H. Wilken
5 stamens are inserted at or above the middle of

the tube, sometimes between the lobes. In
Bonplandia, Cantua, and Loeselia, the stamens are
inserted in the lower tube, but those of Cobaea
and some Gilia are nearly basal. Filaments can be
equal or unequal in length. Anthers are basifixed
in most species, but are dorsifixed in Cobaea and
Eriastrum.
Ten vascular bundles diverge from the central
stele of the pedicel (Dawson 1936). Five alternate
bundles become the midribs of the sepals. The
other five branch several times distally, giving rise
sequentially to traces supplying the corolla and
stamens, the nectariferous disc, and the ovary. The
nectariferous disc in most Polemoniaceae is sup-
plied by 5 bundles, but 10 bundles supply those of
Cantua and Cobaea. In most Polemoniaceae, the
central bundle leading to each petal lobe trifur-
cates at the base of the tube; the 3 traces often
anastomose in the throat or distal portion of the
lobe but give rise to secondary branches (Day and
Moran 1986). Anastomoses are lacking in Linan-
thus, but the 3 primary traces can branch several
times. In Acanthogilia, Cantua, Cobaea, and Phlox,
some of the distal secondary branches can form
additional anastomoses. In Allophyllum, Erias-
trum, Gilia, Ipomopsis, and Navarretia the second-
ary branches often remain free. Small corollas in
several genera often have either 1 or 3 unbranched
veins.
EMBRYOLOGY. The pollen grains are 2-celled

when shed. The ovule is anatropous to hemianat-
ropous, unitegmic, and tenuinucellate. The arches-
poria} cell functions directly as the megaspore
mother cell and the chalazal megaspore of a linear
tetrad develops into a Polygonum-type embryo Fig. 99. Polemoniaceae, pollen. A Collomia biflora,
sac. Endosperm formation is nuclear, but the zono{6-8)colporate, REM. B Cantua buxifolia, pantoporate,
tissue later becomes cellular (Johri et al. 1992). sexine insulate, REM. Bars = IO!!m. Photo B.E. Herber
PoLLEN MoRPHOLOGY. The pollen grains ofPole-

moniaceae are oblate to spheroidal and mostly "lines offorce in a magnetic field" (Stuchlik 1967).
pantoporate; zonotreme (Fig. 99A), bizonotreme The sexine of pantotreme reticulate pollen varies
or anomotreme/porate (Fig. 99B) or colp(or)ate from homobrochate to heterobrochate.
figurations are found in a few taxa. Pollen grains
vary in size between 22 and 190 ~m, and the KARYOLOGY. In Polemoniaceae, chromosome
number of apertures vary in the range (4-)6-50 numbers are based on multiples of 6, 7, 8, and 9.
(Stuchlik 1967; Taylor and Levin 1975). Pollen size Grant (1959) hypothesized x = 9 as ancestral and
is more closely correlated with style length than suggested that taxa with lower numbers were
with taxonomic affinity (Plitmann and Levin derived, sometimes independently among line-
1983). The sexine can be striate, reticulate, pertec- ages. Chromosome numbers exclusively with x = 9
tate, or verrucate. Germinal apertures range from are reported in Acanthogilia, Cantua, Gilia, Loe-
circular to oval or elliptic in shape. The lirae in selia, Navarretia, Linanthus, and Polemonium.
pollen with striate sexine can be vermiform, but Chromosome numbers with x = 7 occur in Erias-
most often radiate from the apertures like the trum, Ipomopsis, Langloisia, and Phlox. Collomia
Polemoniaceae 303
and Gymnosteris have chromosome numbers and scent, promoting attractiveness to bumble-
based on x = 8 and x = 6, respectively. Both Aliciella bees while repelling nectar-thieving ants (Galen
and Allophyllum have diploid chromosome and Newport 1988).
numbers based on either x = 8 or x = 9. The most Most genera include two or more pollination
parsimonious origin for the 2n = 30 reported in syndromes. Cobaea includes taxa that are autoga-
Bonplandia may be tetraploidy involving ances- mous or chasmogamous and pollinated by hawk-
tors with n = 7 and 8. Notably, Giliastrum includes moths and bats. Although autogamy occurs in
species with 2n = 12, 18, 20, 24 and 36, with x = 6 some annual taxa of Ipomopsis, the genus also
as the most likely base number. The highest includes chasmogamous floral syndromes associ-
numbers are 2n = 54 in Cantua (hexaploid based ated with effective pollination by moths (Sphingi-
on x = 9) and 2n = 52 in Cobaea scandens, which dae), bees (Andrenidae, Apidae, Halictidae),
may be derived by an aneuploid reduction from butterflies (Hesperiidae), beetles (Melyridae), and
the hexaploid condition. Tetraploid and some hummingbirds (Trochilidae). Diversification also
hexaploid taxa occur in Aliciella, Gilia, Giliastrum, has occurred at the infraspecific level in Gilia
Ipomopsis, Linanthus, and Phlox. B-chromosomes splendens, which includes geographic races polli-
have been reported in Linanthus and Phlox (Smith nated by hummingbirds or flies, and one predom-
and Levin 1967; Patterson 1980). inantly autogamous race.
REPRODUCTIVE AND POLLINATION BIOLOGY. A FRUITS AND SEEDS. Dehiscence is either septici-
rich diversity of breeding systems is found in dal in Cobaea, or loculicidal in the remaining
Polemoniaceae and has been discussed at genera. Except for some species of Navarretia,
length by Grant and Grant (1965). Both self- in which fruits dehisce from the base, the fruits of
incompatibility and self-compatibility are found the family are capsules. Mature fruits vary from
within most genera. Most self-compatible taxa globose to ovoid. The number of seeds varies from
have mixed mating systems, but autogamy has 1 to many per locule. Seeds, which vary from less
been documented in members of at least 9 than 100 11m to over 2 em, are often ovoid to ellip-
genera (Plitmann and Levin 1990). Cleistogamy is soid and can be smooth, angular, or verrucose.
known in Collomia grandiflora and Polemonium Seeds of Acanthogilia, Cantua, and Cobaea are
micranthum. Distyly, accompanied by a weak self- thin and often prominently winged. The seeds are
incompatibility system, occurs in Gilia heterostyla albuminous and the embryo is straight or highly
and functional gynodioecy has been reported curved. The seed coats of many Polemoniaceae
in Ipomopsis rubra. The effective pollinators have cylindrical cells with prominent helical sec-
of many Polemoniaceae are Coleoptera, Diptera, ondary thickenings and contain mucilage (Grant
Hymenoptera, and Lepidoptera. Variously colored 1959). When wetted, imbibition causes the outer
corollas, some with distinctive nectar guides, walls to separate from the testa, accompanied
attract insects that collect either pollen, nectar, or by extrusion of the expanded thickenings and
both. The type of insect visitor is often correlated mucilage (Grubert and Hambach 1972). Collomia,
with flower size, corolla shape and color, and inflo- Ipomopsis, Phlox, and Polemonium each include
rescence structure. Taxa visited or pollinated by species with either copious (annuals) or little, if
Coleoptera (e.g., Ipomopsis congesta, Linanthus any, mucilage (perennials). Thus, copious mucilage
parryae) have broadly campanulate, often light- in the family appears correlated with life form
colored corollas or small flowers disposed in rather than taxonomic affinity.
congested or capitate inflorescences. Erect, often
fragrant flowers with small corollas with long, PHYTOCHEMISTRY. Polemoniaceae are particu-
narrow tubes are often pollinated by Diptera larly rich in flavonoids, including 6-methoxy-
(Bombyliidae, Cyrtidae) or relatively small Lepi- flavonols (patuletin, eupalitin, eupatoletin) and
doptera. Hawkmoth pollination is known in at C-glycosylfavones (Smith et al. 1977). Floral pig-
least 7 genera, associated with fragrant, vespertine ments include delphinidin, cyanidin, and pelar-
or nocturnal, white, long-tubed corollas. Spread- gonidin glycosides, seemingly correlated with
ing or pendent flowers with relatively large, pollination syndromes (Harborne and Smith
brightly colored, diurnal corollas, and copious 1978), with pelargonidin associated with hum-
nectar attract hummingbirds in at least 8 genera. mingbird pollination and cyandin or delphinidin
Large, nocturnal solitary flowers in some taxa of associated with insect pollination. The almost
Cobaea are bat pollinated. Polemonium viscosum exclusive presence of acylated anthocyanins in
displays an unusual polymorphism for floral size corollas of Linanthus, Loeselia, and Phlox is corre-
304 D.H. Wilken
lated with the presence of C-glycosyl flavones in have x = 8, but x = 9 occurs throughout Navarretia
the vegetative organs. Other constituents of sys- and the remaining species of Allophyllum.
tematic interest include triterpene saponins Eriastrum, Langloisia, and most species of
(Hiller et al. 1979; Jurenitsch et al. 1979) and Ipomopsis also have senescent basal rosettes,
ketose-isoketose fructose oligosaccharides gradually reduced cauline leaves, and terminal,
(Pollard and Amuti 1981). congested or capitate inflorescences. However, all
members of this group share x = 7 and have
SUBDIVISIONS AND RELATIONSHIPS WITHIN THE light-colored seeds. With the exception of bilabi-
FAMILY. Based on cladistic analyses of molecular ate species in Ipomopsis and Langloisia, most
and morphological data, Porter and Johnson (2000) have salverform corollas. The seeds of Eriastrum
subdivided Polemoniaceae into three subfamilies, and Ipomopsis are often ellipsoid and angled
the Acanthogilioideae, Cobaeoideae and Polemoni- to minutely winged, but those of Langloisia are
oideae. The monotypic Acanthogilia, an endemic ovoid and smooth or angled. Although some
shrub of xeric habitats in Baja California, is the sole Ipomopsis have reticulate pollen, most species in
member of Acanthogilioideae. It has dimorphic the 3 genera have a striate-reticulate sexine in
leaves, axillary inflorescences, large salverform which the lirae radiate from the apertures.
corollas, and verrucate, zonotreme pollen. The Ipomopsis minutiflora, with uniformly glandular,
primary leaves are pinnately divided and become puberulent indumentum, slightly unequal calyx
spinose, but the axillary short-shoot leaves are lobes, and bizonotreme pollen, may be better
simple, entire, and borne in fascicles. Cobaeoideae placed elsewhere, perhaps recognized as a distinct
are composed of genera that share a woody or genus.
semiwoody habit, herbaceous calyces lacking Although Grant (1998) placed Loeselia in
conspicuous hyaline intervals, pantotreme pollen, Cobaeoideae, molecular phylogenies suggest
and winged seeds. Cantua is composed of shrubs closer relationships to Eriastrum, Cilia, Ipomopsis,
and small trees occupying relatively mesic and Langloisia (Johnson et al. 1996; Porter and
montane and subalpine habitats in the Andes of Johnson 1998). Most Loeselia have a suffrutescent
South America. Leaves in most taxa are simple and habit, often congested inflorescence, intercostal
the flowers, with salverform to funnelform corol- calyx membranes that are slightly hyaline, pan-
las, are relatively large and disposed in relatively totreme, pertectate pollen, and minutely winged
loose, terminal clusters. Calyx tubes are either seeds. Although leaves and bracts are entire to
completely herbaceous or have narrow, relatively dentate, their margins bear setae similar to those
thin to hyaline intervals between the herbaceous of Langloisia. Floral bracts in most Loeselia are
costae. Bonplandia has an herbaceous habit, het- foliaceous, with hyaline or thinly membranous
erophylly, and paired, zygomorphic, axillary areolae.
flowers. Cobaea, with a distribution restricted to Aliciella, Cilia, and Giliastrum have flowers dis-
neotropical forests, is the most distinct genus posed in relatively open inflorescences. The sepal
within the family. Its species share a viny habit, lobes often have a well-developed hyaline or scar-
deeply compound leaves, terminal tendrils, broadly ious margin. Aliciella and Cilia have paniculate or
campanulate corollas, and septicidal capsules. thyrsoid inflorescences, salverform to funnelform
Polemonioideae are equally diverse but many corollas, zonotreme, striate to reticulate pollen,
genera do not have distinct combinations of and x = 9. Glands in Aliciella are multicellular, but
characters (Mason 1945). Polemonium appears they are unicellular in Cilia and Giliastrum. Gilias-
unique in the subfamily by having pinnately com- trum has either solitary flowers or flowers dis-
pound leaves, herbaceous calyces that remain posed in open dichasia, rotate to short-funnelform
unruptured by the developing fruits, and striate corollas, zonotreme pertectate pollen, and x = 6.
pollen with 30-100 apertures and mostly vermi- Several species in Cilia appear sufficiently distinct
form lirae. to merit recognition. They apparently lack basal
Most species of Allophyllum, Collomia, and rosettes and have entire cauline leaves, solitary
Navarretia have senescent basal rosettes, cauline pedicellate flowers, and short-funnelform to cam-
leaves gradually reduced upwards, congested panulate corollas. Cilia tenerrima, with short-
or somewhat capitate inflorescences, and dark- funnelform corollas, has anomotreme pollen. Cilia
colored ovoid seeds. Pollen grains may be either capillaris, G. sinistra, and G. leptalea have fun-
pantotreme or zonotreme, with 6-20 apertures. A nelform corollas but their pollen is pantotreme.
pertectate or pilate sexine occurs in Allophyllum Another group, including G. filiformis, G. inyoen-
and Navarretia. Collomia and some Allophyllum sis, and G. maculata, has mostly free sepals, cam-
Polemoniaceae 305
panulate to rotate corollas, and stamens inserted Most members of the Cobaeoideae occur in
near the corolla base. subtropical to tropical woodlands and forests of
Linanthus and Phlox have opposite leaves with Central and western South America. The majority
entire to palmatifid blades, terminal inflorescences of Polemonioideae occur in western North
often composed of dichasia or solitary flowers, American woodlands and shrublands, associated
mostly salverform corollas, pantotreme pollen primarily with summer-dry Mediterranean, Great
with reticulate to striate-reticulate sexine, and x = Basin, and Sonoran Desert climates. Some genera
7. The closely related Gymnosteris, with x = 6, (Polemonium, Phlox) include taxa adapted to the
salverform corollas, and pantotreme reticulate relatively moist summer and cool to cold winter
pollen, apparently lacks cauline leaves, but the climates of both eastern and western North
terminal capitate inflorescence is subtended by America, including the arctic. Polemonium is also
a whorl of photosynthetic bracts. represented by a few species in relatively cool
As an alternative to the above phylogenetic moist climates of Eurasia as far south as the north-
system of classification of the family, another clas- ern flank of the Himalayas. Initial diversification
sification based on traditional taxonomy has been may have occurred during the late Cretaceous or
proposed by Grant (1998). In a review of these early Tertiary in what is now Mexico, Central
classifications, Grant (2001) reviews and compares America, or northern South America. Dispersal to
both of them. Eurasia may have involved both amphi-Beringian
and amphi-Atlantic bridges. However, over 90% of
AFFINITIES. Based on morphology, the Polemoni- Polemoniaceae occur in western North America,
aceae have been traditionally treated as allied to the majority of which are annuals (Grant 1959;
Convolvulaceae (Takhtajan 1980) or Hydrophyl- Raven and Axelrod 1978). Thus, progressive
laceae (Cronquist 1981) within Asteridae. Thorne cooling and drying, successive orogenies, and the
(1983), among others, placed Polemoniaceae development of a Mediterranean climate during
nearest Fouquieriaceae, but with both in Asteri- the mid to late Tertiary may have played a sig-
dae, based on their pentamerous perianth and nificant role in radiation, especially within Pole-
androecium, gamopetaly, epipetalous stamens, monioideae. Some predominantly western North
axile placentation, loculicidal capsule, and well- American genera (Collomia, Gilia, Ipomopsis,
developed embryos. Fouquieriaceae and some Linanthus, and Navarretia) have 1-2 close rela-
Polemoniaceae also have thin, winged seeds and tives in western South America, and two species
fasciculate spinescent leaves (Acanthogilia). (Phlox gracilis, Polemonium micranthum) occur
Several cladistic analyses using diverse mole- disjunctly. These amphitropical relationships are
cular data have consistently suggested a closer best attributed to long-distance dispersal during
alliance with Fouquieriaceae and such families as the late Tertiary (Raven 1963).
Diapensiaceae, Ericaceae, and Primulaceae within
Ericales/Primulales (Downie and Palmer 1992; FossiL RECORD. Fossil pollen has only rarely
Olmstead et al. 1993; Johnson et al. 1996, Porter been attributed to the family (see Muller 1981). A
1997; Porter and Johnson 1998). A tricarpellate megafossil from the Eocene of Utah, USA, Gilise-
gynoecium characterizes Fouquieriaceae, Diapen- nium, which is close to Gilia, is important as an
siaceae, Roridulaceae, and some Ericaceae, but early megafossil record of a herbaceous plant (Lott
elsewhere is unusual in Asteridae. Ketose and et al. 1998). Pleistocene fruits attributed to Phlox
isoketose oligosaccharides also occur in some Eri- have been reported from Alaska (Chaney and
caceae, but not in Asteridae s.s. (Pollard and Am uti Mason 1936).
1981). Accordingly, Polemoniaceae have been
included in an expanded Ericales at the base of EcoNOMIC IMPORTANCE. Some species of
Asteridae by the Angiosperm Phylogeny Group Cobaea, Cantua, Collomia, Ipomopsis, Linanthus,
(APG 1998). In the five-gene sequence analysis by and Polemonium are cultivated for ornament, and
Anderberg et al. (2002), Polemoniaceae and cultivars have been developed from several species
Fouquieriaceae appear in close association, albeit of Phlox. Species of Ipomopsis, Loeselia, and Pole-
with only moderate support, which is understand- monium are used by native North Americans to
able in view of the strong morphological diver- produce a mild soap. In Central America, infusions
gence between the two families. of Cobaea flowers have been used as a cough
suppressant. Potentially useful anti-inflammatory
DISTRIBUTION AND HABITATS. Acanthogilia is and cyto-toxic antitumor compounds (triterpene
endemic to thorn shrublands of Baja California. saponins, cucurbitacins) have been isolated from
306 D.H. Wilken
Ipomopsis and Polemonium (Aurada et al. 1982; - Leaf segments rounded to mucronate or spinescent, setose
Arisawa et al. 1984). Cantua buxifolia, considered bristles lacking 18
18. Upper cauline leaves palmatifid 19
sacred by the Incas, is the national flower of Peru. - Upper cauline leaves entire to pinnately toothed or
pinnatifid 20
KEY TO THE GENERA 19. Terminal leaf segment often longer than lateral segments,
rounded 7. Allophyllum
1. Leaves pinnately compound 2 - Terminal leaf segment not markedly longer than lateral
Leaves simple, entire to pinnately or palmately divided 3 segments, spinescent 16. Linanthus
2. Vines; terminal leaflet a tendril 2. Cobaea 20. Corolla rotate to short-funnelform, lobes longer than the
- Annuals or herbaceous perennials; terminal leaflet foliar tube; stamens inserted in the lower tube 13. Giliastrum
5. Polemonium - Corolla salverform to long-funnelform, lobes equal to or
3. Axis above cotyledons unbranched and apparently leafless; shorter than the tube; stamens inserted in the upper tube
leaves reduced to a bracteate whorl subtending the termi- or throat 21
nal inflorescence 18. Gymnosteris 21. Indumentum predominantly glandular; trichomes with
3. Cauline nodes and leaves present; cauline leaves mostly multicellular glands 14. Aliciella
- alternate or opposite 4 - Indumentum eglandular to glandular; glandular trichomes
4. Leaves predominantly opposite 5 with unicellular glands 22
- Leaves predominantly alternate 6 22. Corolla campanulate, funnelform, or rotate, often with a
5. Stamens inserted at the same level in the tube or throat well-developed throat 15. Gilia
16. Linanthus - Corolla salverform, throat lacking 9. Ipomopsis
- Stamens inserted at different levels in the tube 17. Phlox
6. Stamens inserted below the middle of the tube 7
- Stamens inserted at or above the middle of the tube, some-
times between the lobes 10 I. SUBFAM. ACANTHOGILIOIDEAE J.M. Porter
7. Sepals mostly free; corolla lobes longer than tube and L.A. Johnson (2000).
15. Gilia
- Sepals mostly fused; corolla lobes shorter than tube 8 Shrubs; leaves dimorphic, the primary ones
8. Shrubs; corolla campanulate or salverform; seeds thin,
conspicuously winged 3. Cantua alternate, spinescent, and pinnate, the axillary ones
- Annuals, suffrutescent perennials, or subshrubs; corollas simple, in fascicles; corollas actinomorphic, salver-
more or less zygomorphic, seeds plump, wingless or incon- form; pollen zonotreme, verrucate; fruit dehiscing
spicuously winged 9 both loculicidally and septicidally; seeds flat,
9. Flowers axillary, in pairs; corolla tube conspicuously bent
4. Bonplandia
winged.
- Flowers terminal, solitary or in clusters, if axillary then
corolla actinomorphic; corolla tube straight 12. Loeselia
I. Acanthogilia A.Day & R.Moran
10. Plants woody; flowers axillary; seeds winged
1. Acanthogilia Acanthogilia A.Day & R.Moran, Proc. Calif. Acad. Sci. 44: 111
- Plants herbaceous, if woody then suffrutescent or sub- {1986).
shrubs; flowers not axillary; seeds wingless or in com-
pletely and minutely winged 11
11. Inflorescence capitate or corymbose 12 Spiny shrubs with dimorphic leaves; long-shoot
- Inflorescence paniculate or thrysoid, sometimes axillary leaves alternate, persistent, pinnate with spinescent
16 divisions; axillary short-shoot leaves in fascicles,
12. Calyx plicate, the intercostal membranes distended dis- linear; pubescence minutely glandular. Inflores-
tally, not ruptured by the developing fruit; leaf tips acute
to rounded 6. Collomia cence axillary, 1-3-flowered. Calyx tubular, often
- Calyx not plicate, the intercostal membranes hyaline, often with narrow hyaline intervals; corolla actinomor-
ruptured by the developing fruit; leaf tips mucronate to phic, salverform; stamens subequally inserted
spinulose 13 near the middle of the tube, filaments glabrous;
13. Inflorescence bracts spinulose, often pinnatifid; calyx lobes
anthers exserted; pollen zonotreme, verrucate;
subequal 14
- Inflorescence bracts acute to mucronate, entire, sometimes style exserted. Fruit loculicidally and septicidally
pinnatifid to palmatifid; calyx lobes equal 15 dehiscent; seeds 1-6 per locule, flat, winged.
14. Indumentum predominantly eglandular 10. Eriastrum 2n = 18. One sp., A. gloriosa (Brandegee) Day &
- Indumentum predominantly glandular; eglandular tri- Moran, thorn shrublands of N Baja California.
chomes not arachnoid 8. Navarretia
15. Corolla campanulate, funnelform, or rotate, often with a
well-developed throat 15. Gilia
- Corolla salverform, throat lacking 9. Ipomopsis II. SUBFAM. COBAEOIDEAE (D. Don)
16. Basal and cauline leaves mostly entire 15. Gilia Arnott (1832).
- Basal and cauline leaves pinnately lobed to pinnatifid or
palmatifid 17
17. Leaf segments with 1-3 terminal setose bristles
Vines, shrubs, small trees, or suffrutescent peren-
11. Langloisia nials; leaves monomorphic, simple to pinnately
Polemoniaceae 307
lobed or compound; corollas campanulate, salver-

form or somewhat bilabiate; pollen pantotreme,
reticulate, striate, or verrucate; fruit dehiscing
loculicidally or septicidally; seeds somewhat flat,
usually winged.
2. Cobaea Cavanilles Fig. lOOB-D

Cobaea Cavanilles, Icon. 1: 11. (1791 ); Prather, Syst. Bot.
Monogr. 57: 1-81 (1999).
Vines with alternate, pinnately compound leaves;

lateral leaflets 4-8, terminal leaflet a branched
tendril; plants glabrous to variously pubescent.
Inflorescence cymose, I-S-flowered; pedicels often
with 2 foliaceous bracts. Calyx with mostly free,
herbaceous sepals; corolla actinomorphic, broadly
campanulate, constricted near the base; stamens
inserted near the base of the tube, filaments
basally villous; anthers exserted, dorsifixed; pollen
large, pantotreme, reticulate; style exserted. Fruit
dehiscing septicidally; seeds 1-many per locule,
flat, often broadly winged. 2n = 52 reported in C.
scandens Cav. About 20 spp., forests of tropical
Central and NW South America.
3. Cantua Juss. ex Lam. Fig. lOOA

Cantua Juss. ex Lam., Encycl.1: 603 (1785); Infantes Vera, Lilloa
31: 75-107 (1962).
Huthia Brand (1908).
Shrubs or small trees, often with dimorphic leaves;

principal leaves alternate, sometimes also in fasci-
cles, thin to coriaceous, entire to crenate, rarely
lobed; plants glabrous or with eglandular to glan-
dular indumentum. Inflorescence of 3-10 flowers,
terminal, raceme-like/botryoid or somewhat
capitulate. Calyx tubular to campanulate, herba-
ceous throughout or thin and hyaline; corolla
actinomorphic, salverform to campanulate; Fig. 100. Polemoniaceae. A Cantua quercifolia, flowering
stamens inserted at different levels near or below shoot. B, C Cobaea scandens. B Flowering shoot with tendrils.
C Vertical section of flower with pollinating bat. D Cobaea pen-
the middle of the tube, filaments glabrous to duliflora, flower. E Gymnosteris parvula, whole plant. F Phlox
villous; anthers exserted to included; pollen pan- bryoides. G Phlox longiflorum. H Bonplandia geminiflora,
totreme, reticulate to verrucate; style included to flower. (Takhtajan 1981)
exserted. Fruit dehiscing loculicidally; seeds 3-8
per locule, flat, winged. 2n = 54. Ca. 20 spp.,
interandine region of Ecuador, Peru and Bolivia.
pinnately lobed leaves, margins entire to serrate;
indumentum minutely glandular puberulent,
4. Bonplandia Cav. Fig. IOOH sometimes eglandular. Inflorescence of terminal
and axillary sympodia. Flowers usually paired;
Bonplandia Cav., Anales Hist. Nat. 2: 131 (1800); Rzedowski et
al.,Acta Bot. Mex. 31:55- 61 (1995). perianth zygomorphic; calyx herbaceous through-
out; corolla somewhat bilabiate, geniculate at the
Suffrutescent perennial, sometimes flowering the throat; stamens inserted near the mid-tube, fila-
first year, with alternate, linear to ovate, simple to ments basally puberulent; anthers exserted; pollen
308 D.H. Wilken
pantotreme, striate-reticulate; style slightly membranes distended in fruit; corolla actinomor-

included to exserted. Fruit dehiscing loculicidally; phic, salverform to funnelform; stamens inserted
seeds one per locule, somewhat thick, narrowly at the same or different levels in the upper tube,
winged or wingless. 2n = 30. One species, B. gemi- filaments equal to unequal in length, glabrous or
niflora Cav., woodlands and forests of Mexico. basally puberulent; anthers exserted to included;
pollen pantotreme or zonotreme, striate to re-
ticulate; style slightly included to exserted. Fruit
Ill. SUBFAM. POLEMONIOIDEAE (Juss.) dehiscing loculicidally; seeds 1-3 per locule,
Arnott (1832). ovoid, smooth or somewhat angled, weakly to
strongly mucilaginous when wetted. 2n = 16. Ca.
Annuals, perennials, sometimes shrubs; leaves 15 species, shrublands, woodlands, and forests of
simple to pinnately or palmately lobed or com- W North America, 1-2 in W South America.
pound; corollas rotate to salverform, sometimes
bilabiate; pollen pantotreme or zonotreme, mostly
7. Allophyllum (Nutt.) A. & V. Grant
striate to reticulate, sometimes pertectate; fruit
dehiscing loculicidally; seeds ovoid to ellipsoid, Allophyllum (Nutt.) A. & V. Grant, Aliso 3: 99 (1955); Grant &
wingless. Grant, Aliso 3:93-110 (1955).
Annuals. Leaves sometimes in basal rosettes,

5. Polemonium L. basal leaves senescent at bolting, cauline ones
Polemonium L., Sp. Pl. 1: 162 (1753); Davidson, Univ. Calif. alternate, entire to twice pinnatifid, uppermost
Pub!. Bot. 23: 209-282 (1950), rev. leaves sometimes palmatifid, the lobes oblong, the
Polemoniella Heller (1904). terminal ones often broader than the lateral lobes;
indumentum predominantly glandular, with a
Herbaceous perennials, often from rhizomes, mephitic odor. Inflorescence terminal, mostly 2-3-
rarely annual; cauline leaves alternate, compound; flowered, somewhat congested. Calyx actinomor-
indumentum eglandular or glandular. Inflores- phic, hyaline between the herbaceous costae, the
cences mostly terminal, paniculate to capitulate. lobes equal; corolla actinomorphic to slightly
Calyx actinomorphic, herbaceous throughout, zygomorphic, funnelform; stamens inserted at
lobes equal; corolla actinomorphic, salverform, the same or different levels in the upper tube,
funnelform, or rotate; stamens inserted at the filaments equal to unequal in length, glabrous;
same level in the lower tube, filaments equal in anthers included to exserted; pollen pantotreme,
length, pubescent; anthers included to exserted; pertectate; style slightly included to exserted.
pollen pantotreme, mostly vermiform striate; style Fruit dehiscing loculicidally; seeds 1-3 per lo-
included to exserted. Fruit dehiscing loculicidally; cule, ovoid, smooth or somewhat angled, strongly
seeds 1-12 per locule, ovoid to ellipsoid, smooth mucilaginous when wetted. 2n = 16, 18. Five
or angled, strongly to not at all mucilaginous when species, shrublands and forests of California and
wetted. 2n = 18. Ca. 25 species, shrublands and Baja California.
mesic sites in woodlands and forests of North
America and Eurasia.
8. Navarretia Ruiz & Pav.
6. Collomia Nutt. Navarretia Ruiz & Pav., Fl. Peruv. Prodr.: 20 (1794); Crampton,
Madrofio 12: 225-256 (1954).
Collomia Nutt., Gen. Amer. 1: 126 (1818); Wilken et a!.,
Biochem. Syst. Ecol. 10: 239-243 (1982). Annuals; basal leaves often in a rosette, senescent
at bolting, cauline leaves abruptly to gradually
Annual or perennial herbs. Leaves often in basal reduced in size. Leaves entire or once to twice
rosettes; rosette leaves in annuals senescent at pinnatifid, the ultimate lobes entire to lobed;
bolting, cauline leaves alternate, entire to lobed, plant glabrous, or with eglandular to glandular
rarely palmatifid or pinnatifid; indumentum trichomes. Inflorescence congested or capitate
eglandular to glandular, sometimes with a strong panicle. Calyx tubular, hyaline between the
mephitic odor. Inflorescence terminal and capi- herbaceous costae, the lobes equal to subequal,
tate, rarely axillary. Calyx actinomorphic, herba- mucronate; corolla actinomorphic, funnelform to
ceous or narrowly hyaline between the herbaceous salverform; stamens 5, equally inserted at the same
costae, the lobes equal to subequal, the intercostal level in the upper tube or between the lobes; fila-
Polemoniaceae 309
ments mostly equal, glabrous; anthers included to equal to unequal in length, glabrous; anthers
exserted; pollen pantotreme, mostly pertectate to included to exserted, often basifixed; pollen pan-
pilate; style included to exserted. Fruit dehiscing totreme or zonotreme, striate to striate-reticulate;
loculicidally; seeds 1-many, strongly mucilagi- style included to exserted. Fruit dehiscing loculi-
nous when wetted, ovoid to subglobose, mostly cidally; seeds 1-3 per locule, ovoid to ellipsoid,
smooth, sometimes angled. 2n = 18.About 30 spp., angled to minutely winged, mucilaginous when
shrublands and woodlands of western North wetted. 2n = 14. Fourteen species, shrublands and
America. forests of W North America.
9. Ipomopsis Michx. 11. Langloisia Greene

Ipomopsis Michx., Fl. Bor. Am. 1: 141 (1803); Grant, Aliso 3: Langloisia Greene, Pittonia 3: 30 (1896); Timbrook, Madroiio
351-362 (1956). 33: 157-174 (1986).
Loeseliastrum (Brand) Timbrook (1986).
Annuals, long-lived monocarps, or suffrutes-
cent perennials. Leaves often in persistent basal Annuals. Leaves sometimes in basal rosettes, sen-
rosettes, cauline leaves alternate, entire to twice escent at bolting, cauline ones alternate, dentate
pinnatifid, gradually reduced in size, the ultimate to once pinnatifid, the teeth or segments with
segments often mucronate or spinulose; indumen- 1-3 terminal bristles; indumentum glandular
tum glandular to eglandular. Inflorescences mostly and eglandular, the trichomes often branched.
terminal, thyrsoid, paniculate, or capitulate, rarely Inflorescence mostly terminal, capitulate, flowers
with few flowers. Calyx actinomorphic to slightly sometimes axillary and solitary. Calyx actinomor-
zygomorphic, hyaline between the herbaceous phic, narrowly hyaline between the herbaceous
costae, the lobes equal to subequal; corolla actin- costae, the lobes equal to subequal; corolla actin-
omorphic to slightly zygomorphic, salverform to omorphic and funnelform or strongly zygomor-
funnelform; stamens inserted at the same or phic and bilabiate; stamens inserted at the same
different levels in the throat or upper tube, level or slightly different levels in the throat or
rarely lower; filaments equal to unequal in length, upper tube, filaments equal to subequal in length,
glabrous; anthers included to exserted; pollen straight to somewhat declinate, glabrous; anthers
pantotreme or zonotreme, bizonotreme in one included to exserted; pollen zonotreme, striate
species, striate to reticulate; style included to to striate-reticulate; style included to exserted.
exserted. Fruit dehiscing loculicidally; seeds 1-12 Fruit dehiscing loculicidally; seeds 1-10 per
per locule, ovoid to ellipsoid, angled to minutely locule, ovoid, smooth, mucilaginous when wetted.
winged, mucilaginous when wetted. 2n = 14, 28. 2n = 14. Three species, desert shrublands and
Ca. 30 species, shrub lands and forests of W North woodlands of SW North America.
America, SE North America (1), and W South
America (1). 12. Loeselia L.
Loeselia L., Gen. Pl. 276 (1754); Turner, Phytologia 77:318-337
10. Eriastrum Woot. & Standi. (1994).
Eriastrum Woot. & Standi., Contrib. U.S. Nat. Herb. 16: 160
(1913); Harrison, Brigham Young Univ. Sci. Bull., Bioi. Ser. Annuals, suffrutescent perennials, or subshrubs.
16: 1-26 (1972). Leaves alternate or opposite, simple, margins
entire to aristate; indumentum eglandular,
Annuals or suffrutescent to woody perennials. glandular, or glabrous. Inflorescences axillary or
Leaves in annuals often in basal rosettes, senescent terminal, somewhat congested to capitulate. Calyx
at bolting, alternate, entire to twice pinnatifid, actinomorphic or slightly zygomorphic, the lobes
cauline leaves gradually reduced in size, the ulti- and tube mostly herbaceous, the tube somewhat
mate segments often mucronate; indumentum hyaline or thinly membranous below the sinuses;
predominantly eglandular, often arachnoid. Inflo- corolla actinomorphic to zygomorphic, salver-
rescences mostly terminal, capitulate. Calyx actin- form to somewhat rotate or bilabiate; stamens
omorphic, hyaline between the herbaceous costae, inserted at the same level below the mid-tube,
the lobes slightly unequal; corolla actinomorphic, filaments equal in length, glabrous or pubescent
salverform to funnelform; stamens inserted at the basally; anthers often exserted; pollen pantotreme,
same level in the throat or upper tube, filaments pertectate; style included to exserted. Fruit dehisc-
310 D.H. Wilken
ing loculicidally; seeds 1-10 per locule, globose to lands, woodlands, and forests of W North
ellipsoid, angled to minutely winged, mucilagi- America.
nous when wetted. 2n = 18. Ca. 14 species, shrub-
lands and forests of S North America, Central
15. Gilia Ruiz & Pav.
America, and NW South America.
Gilia Ruiz & Pav., Fl. Peruv. Prodr. 4: 25 (1794); Grant & Grant,
Aliso 3: 59-91 (1954), 3: 203-287 (1956).
13. Giliastrum (Brand) Rydb.
Giliastrum (Brand) Rydb., Fl. Rocky Mts.: 699 (1917); Porter, Annuals or herbaceous to suffrutescent perenni-
Aliso 17: 83-85 (1998). als, rarely subshrubs; basal leaves often in a
rosette, often senescent at bolting, cauline leaves
Annuals or suffrutescent perennials, sometimes alternate, abruptly to gradually reduced in size;
flowering the first year. Leaves in basal rosettes, pubescence eglandular or glandular. Leaves entire
cauline ones alternate, entire to pinnatifid, gradu- or once to thrice-pinnatifid. Inflorescence panicu-
ally reduced in size; indumentum predominantly late, relatively open, sometimes congested and
glandular puberulent. Inflorescences open, panic- capitate, ultimately of 1-3 subsessile to pedicel-
ulate, sometimes solitary. Calyx actinomor- late flowers. Calyx tubular, hyaline between the
phic, broadly hyaline between narrow herbaceous herbaceous costae, the lobes equal; corolla actin-
costae, the lobes equal; corolla actinomor- omorphic, rotate, campanulate, funnelform, or
phic, rotate to short-funnelform; stamens equally salverform; stamens 5, equally inserted on the tube
inserted below the mid-tube, filaments equal in or throat, filaments equal to unequal in length,
length, glabrous or pubescent basally; anthers rarely declinate, glabrous; anthers included to
often exserted; pollen zonotreme, pertectate; style exserted; pollen zonotreme, striate to reticu-
often exserted. Fruit dehiscing loculicidally; seeds late; ovary glabrous to minutely glandular, style
3-15 per locule, ovoid to globose, smooth to ver- included to exserted. Fruit dehiscing loculicidally;
rucate or angled, mucilaginous when wetted. seeds 1-many per locule, mucilaginous when
2n = 12, 18, 20, 24, 36. Eight species, shrublands wetted, generally many per locule, ovoid to sub-
and woodlands of W North America and Central globose, angled to rough. 2n = 18, 36, 72. About
America. 50 spp., shrublands, woodlands, and forests of W
North America.
14. Aliciella A.Brand
16. Linanthus Benth.
Aliciella A.Brand, Helios 22: 78 (1905); Porter, Aliso 17: 23-46
(1998). Linanthus Benth., Edward's Bot. Reg. 19: 1622 (1833);
Patterson, Madrofio 24: 36-48 (1977).
Annuals and perennials, often monocarpic. Leaves Leptodactylon Hook. & Arn. (1839).
in a basal rosette, often senescent at bolting, Linanthastrum Ewan (1942).
cauline leaves abruptly (most annuals) to gradu-
ally reduced (perennials) in size; pubescence Annuals, herbaceous or suffrutescent perennials,
mostly of uniseriate trichomes bearing multi- or shrubs. Leaves opposite, sometimes alternate,
cellular glands. Leaves entire to once-pinnatifid. simple, entire to palmatifid, sometimes pinnatifid;
Inflorescence paniculate. Calyx tubular, hyaline indumentum eglandular, glandular, or glabrous.
between the herbaceous costae; corolla actin- Inflorescences axillary or terminal, somewhat
omorphic, salverform to funnelform; stamens congested to capitulate. Calyx actinomorphic, nar-
5(3), stamens inserted at the same or different rowly hyaline between the herbaceous costae, the
levels in the upper tube or in the sinuses of the lobes equal; corolla actinomorphic, campanulate
lobes, sometimes declinate, filaments glabrous, to funnelform or salverform; stamens inserted
sometimes papillose; anthers included to exserted; at the same level in the throat or tube, filaments
pollen zonotreme, striate-reticulate to reticulate; equal in length, glabrous or minutely puberulent
ovary glabrous, style included to exserted. basally; anthers exserted to included; pollen pan-
Fruit dehiscing loculicidally; seeds not or only totreme, striate-reticulate to reticulate; style
slightly mucilaginous when wetted, generally included to exserted. Fruit dehiscing loculicidally;
1-12 per locule, but 18-28 in A. latifolia and seeds 1-10 per locule, ovoid, smooth or angled,
A. ripleyi, ovoid to subglobose, smooth or angled. either mucilaginous or remaining unchanged
2n = 16, 18, 32, 34, 36, 50. About 20 spp., shrub- when wetted. 2n = 18, 36. About 55 species, shrub-
Polemoniaceae 311
lands, woodlands, and forests ofW North America APG (Angiosperm Phylogeny Group) 1998. See general
and W South America. references.
Arisawa, M., Pezzuto, J., Kinghorn, A., Douglas, A., Cordell, G.,
Farnsworth, N. 1984. Plant anticancer agents. 30. Cucur-
17. Phlox L. Fig. lOOF, G bitacins from Ipomopsis aggregata (Polemoniaceae). J,
Pharmaceut. Sci. 73: 411-413.
Phlox L., Gen. Pl. 1: 52 (1737); Wherry, Morris Arb. Monogr. 3: Aurada, E. Jurenitsch, J., Kubelka, W. 1982. Structure of tri-
1-174 (1955), rev. terpene sapogenins from Polemonium caeruleum L. Sci.
Microsteris Greene, Pittonia 3: 301 (1898). Phar. 50: 331-350.
Carlquist, S. 1992. Wood anatomy of sympetalous dicotyledon
families: a summary, with comments on systematic rela-
Annuals, herbaceous or suffrutescent perennials, tionships and evolution of the woody habit. Ann. Missouri
sometimes from rhizomes; indumentum pre- Bot. Gard. 79: 303-332.
dominantly eglandular. Leaves opposite, simple, Carlquist, S., Eckhart, V., Michener, D. 1984. Wood anatomy of
entire to toothed, ovate to linear, those of the inflo- Polemoniaceae. Aliso 10: 547-572.
Chaney, R., Mason, H. 1936. A Pleistocene flora from Fair-
rescence sometimes alternate. Inflorescences banks, Alaska. Am. Mus. Nov.: 887.
mostly terminal, paniculate to capitulate. Calyx Cronquist, A. 1981. See general references.
actinomorphic, narrowly hyaline between the Dawson, M.L. 1936. The floral morphology of the Polemoni-
herbaceous costae, the lobes equal; corolla aceae. Am. J. Bot. 23: 501-511.
actinomorphic, salverform; stamens inserted at Day, A., Moran, R. 1986. Acanthogilia, a new genus of Polemo-
niaceae from Baja California, Mexico. Proc. Calif. Acad. Sci.
different levels in the tube, filaments mostly equal 44: 111-126.
in length, glabrous; anthers mostly included; Downie, S., Palmer, J. 1992. Restriction site mapping of the
pollen pantotreme, reticulate; style included to chloroplast DNA inverted repeat: a molecular phylogeny of
exserted. Fruit dehiscing loculicidally, often the Asteridae. Ann. Missouri Bot. Gard. 79: 266-283.
explosively; seeds 1 per locule, ovoid to ellipsoid, Galen, C., Newport, M. 1988. Pollination quality, seed set, and
flower traits in Polemonium viscosum: complementary
smooth, angled, or minutely winged, either muci- effects of variation in flower scent and size. Am. J. Bot. 75:
laginous or remaining unchanged when wetted. 900-905.
2n = 14, 28. About 70 species, grasslands, shrub- Grant, V. 1959. Natural history of the phlox family. The Hague,
lands, woodlands, and forests of North America Netherlands: Martinus Nijhoff.
Grant, V. 1998. Primary classification and phylogeny of the
and NE Asia (1).
Polemoniaceae, with comments on molecular cladistics.
Am. J. Bot. 85: 741-752.
Grant, V. 2001. A guide to understanding recent classifications
18. Gymnosteris E.Greene Fig. 100£ of the family Polemoniaceae. Lundellia 4: 12-24.
Gymnosteris E.Greene, Pittonia 3: 303 (1898); Wherry, Am. Grant, V., Grant, K.1965. Flower pollination in the phlox family.
Midland Nat. 31: 230-231 (1944). New York: Columbia University Press.
Grubert, M., Hambach, M. 1972. Untersuchen iiber die ver-
schleimenden Samen von Collomia grandiflora Doug!.
Diminutive annuals with persistent cotyledons, (Polemoniaceae). Beitr. Bioi. Pflanz. 48: 187-206.
stems solitary or with 1 branch; indumentum Harborne, J.B., Smith, D. 1978. Correlations between antho-
glandular puberulent. Leaves reduced to a whorl of cyanin chemistry and pollination ecology in Polemoni-
basally connate bracts subtending the terminal aceae. Biochem. Syst. Ecol. 6: 127-130.
Hiller, K., Paulick, A., Doehnert, H., Franke, P. 1979. Saponins
inflorescence. Inflorescence of 1-several flowers, of Polemonium caeruleum L. Pharmazie 34: 565-566.
terminal, capitate. Calyx actinomorphic, broadly Johnson, L., Schultz, J., Soltis, D., Soltis, P. 1996. Monophyly and
hyaline to scarious between narrow herbaceous generic relationships of Polemoniaceae based on matK
costae, the lobes equal; corolla actinomorphic, sequences. Am. J. Bot. 83: 1207-1224.
salverform; stamens inserted at the same level in Johnson, L., Soltis, D., Soltis, P.1999. Phylogenetic relationships
of Polemoniaceae inferred from 18s ribosomal DNA
the upper tube, subsessile, filaments equal in equences. Plant Syst. Evol. 214: 65-89.
length, glabrous; anthers included; pollen pan- Johri, B.M. et a!. 1992. See general references.
totreme, reticulate; style included. Fruit dehiscing Jurenitsch, J., Haslinger, E., Kubelka, W. 1979. Structure of
loculicidally; seeds 1-3 per locule, ovoid, smooth, sapogenins from Polemonium reptans. Pharmazie 34:
strongly mucilaginous when wetted. 2n = 12. Two 445-446.
Kapil, R., Rustagi, P., Venkataraman, R. 1968. A contribution to
species, cold desert shrublands and woodlands of the embryology of Polemoniaceae. Phytomorphology 17:
W North America. 403-412.
Lott, T.A., Manchester, S.R., Dilcher, D.L. 1998. A unique and
complete polemoniaceous plant from the middle Eocene of
Utah, USA. Rev. Palaeobot. Palynol. 104: 39-49.
Selected Bibliography Mason, H. 1945. The genus Eriastrum and the influence of
Bentham and Gray upon the problem of generic confusion
Anderberg, A.A. et a!. 2002. See general references. in Polemoniaceae. Madrofto 8: 33-59.
Metcalfe, C., Chalk, L. 1950. See general references.
312 D.H. Wilken
Muller, J. 1981. See general references. Raven, P. 1963. Amphitropical relations in the flora of North
Olmstead, R., Bremer, B., Scott, K., Palmer, J. 1993. A parsi- and South America. Quart. Rev. Bioi. 29: 151-171.
mony analysis of the Asteridae sensu Jato based on rbcL Raven, P., Axelrod, D. 1978. Origin and relationships of the
sequences. Ann. Missouri Bot. Gard. 80: 700-722. California flora. Univ. Calif. Pub!. Bot. 72: 1-134.
Patterson, R. 1980. The occurrence of B chromosomes in Smith, D., Levin, D. 1967. Karyotypes of eastern North
Linanthus pachyphyllus. Caryologia 33: 141-149. American Phlox. Am. J. Bot. 54: 324-334.
Plitmann, U., Levin, D. 1983. Pollen-pistil relationships in the Smith, D., Glennie, C., Harborne, J.B., Williams, C. 1977.
Polemoniaceae. Evolution 37: 957-967. Flavonoid diversification in the Polemoniaceae. Biochem.
Plitmann, U., Levin, D. 1990. Breeding systems in the Polemo- Syst. Ecol. 5: 107-115.
niaceae. Plant Syst. Evol. 170: 205-214. Spencer, S., Porter, J. 1997. Evolutionary diversification and
Pollard, C., Amuti, K. 1981. Fructose oligosaccharides: possible adaptation to novel environments in Navarretia (Polemoni-
markers of phylogenetic relationship among dicotyledo- aceae). Syst. Bot. 22: 649-668.
nous plant families. Biochem. Syst. Ecol. 9: 69-78. Stuchlik, L. 1967. Pollen morphology in the Polemoniaceae.
Porter, J. 1997. Phylogeny of Polemoniaceae based on nuclear Grana Palynol. 7: 146-240.
ribosomal internal transcribed spacer DNA sequences. Takhtajan, A. 1980. Outline of the classification of flowering
Aliso 15: 57-77. plants (Magnoliophyta). Bot. Rev. 46: 225-359.
Porter, J., Johnson, L. 1998. Phylogenetic relationships of Takhtajan, A. (ed.) 1981. See general references.
Polemoniaceae: inferences from mitochondrial nadlb Taylor, T.N., Levin, D.A. 1975. Pollen morphology of Polemo-
intron sequences. Aliso 17: 157-188. niaceae in relation to systematics and pollination systems:
Porter, J., Johnson, L. 2000. A phylogenetic classification of scanning electron microscopy. Grana 15: 91-112.
Polemoniaceae. Aliso 19: 55-91. Thorne, R. 1983. Proposed new realignments in the angio-
Prather, L.A., Ferguson, C.J., Jansen, R.K. 2000. Polemoniaceae sperms. Nord. J. Bot. 3: 85-117.
phylogeny and classification: implications of sequence data Weberling, F. 1998. Die Infloreszenzen. Vol. 11/2. Jena: G.
from the chloroplast gene ndhF. Am. J. Bot. 87: 1300-1308. Fischer.
Primulaceae 313
Primulaceae
A.A. ANDERBERG
Primulaceae Vent., Tab!. Regne Veg. 2: 285 (1799), nom. cons. the hypocotyl shows an initial swelling but never
develops into a tuber. The leaves are typically alter-
Perennial or sometimes annual herbs, often nate but are sometimes opposite or whorled.
cushion-forming, and sometimes woody at the Most species have a basal leaf rosette and a
base; stem and leaves with long articulated hairs, scapose inflorescence terminating the short,
short glandular hairs, or sometimes glabrous. often rhizomatous stem. Stolons are formed in
Leaves usually alternate, often forming a basal some Androsace. The leaves are usually short-
rosette but sometimes scattered on the stem, petiolate or the blades gradually taper towards the
opposite or whorled, entire, dentate or rarely base. In some genera such as Cortusa, Soldanella
pinnatifid or pinnate; stipules absent; young leaves and part of Primula, the petioles are distinctly
involute or conduplicate, seldom revolute. Inflo- demarcated from the lamina which has a palmate
rescence terminal or axillary, racemose, spicate, venation. Most species have dentate or serrate leaf
umbellate, sometimes scapose and with one to margins and marginal veinlets ending in hyda-
many superimposed umbels, or flowers solitary. thodes. Ptyxis is important systematically: in most
Flowers perfect, hypogynous, actinomorphic or genera it is involute or conduplicate, but revolute
rarely slightly zygomorphic, with imbricate, some- in Dionysia, Cortusa, and most Primula.
times quincuncial corolla aestivation, 5(6-8)- Immersed capitate hairs, like in Myrsinaceae, are
merous, sometimes heterostylous; bracts present not known from Primulaceae, but long, articulated
or sometimes absent, prophylls lacking; calyx hairs are typical of virtually all genera, albeit
synsepalous, campanulate or cylindrical, with five characteristically absent from some (e.g. Dode-
herbaceous lobes; corolla sympetalous, campanu- catheon). Ordinary short glandular hairs are also
late to hypocrateriform; corolla-tube usually dis- common. In species of Androsace, Douglasia and
tinct, short to very long; corolla-lobes entire, Vitaliana, white stellate hairs are frequent but
emarginate, or fringed (Soldanella), sometimes otherwise not present in the family. Omphalo-
reflexed (Dodecatheon); stamens 5( -8), ante- gramma, Bryocarpum and certain Primula have
petalous, epipetalous; anthers tetrasporangiate, blackish, flattened gland-dots on the leaves. Many
dithecal, introrse, opening with longitudinal slits, Primula and Dionysia have a distinct farinose
completely distinct or connivent and forming a coating on young leaves, stems and parts of the
protruding anther cone; gynoecium syncarpous, inflorescence. This can be either powdery or
5-carpellate; ovary superior, unilocular, with a woolly in appearance, and its presence is often
stipitate, free central placental column, only rarely species-specific.
with rudimentary septa basally; ovules few to
many arranged in several series on the surface of VEGETATIVE ANATOMY. Secretory cavities and
or rarely (Dionysia) immersed in the placenta, calcium oxalate crystals in the leaves and flower
anatropous, bitegmic, tenuinucellate; style some- parts are absent from Primulaceae, and extra-
times exserted; stigma truncate to capitate. Fruit a xylary bundles of unbranched fibres, as in
capsule opening with apical valves, a circular lid, Threophrastaceae, are lacking. Irregularly shaped
or by irregular disintegration; seeds more or foliar sclereids are found in Dionysia and some
less angular, reticulate or papillose; endosperm other genera.
copious.
A predominantly north temperate family with 13 INFLORESCENCE STRUCTURE. The inflorescence
genera and c. 600 species. in Primulaceae is basically racemose and the
flowers are arranged in racemes, spikes, corymbs,
VEGETATIVE MORPHOLOGY. All species are herbs or umbels. In Hottonia and several Primula, the
but in the cushion-forming species the stem is flowers are arranged on scapes in superimposed
often more or less lignified. In some Dodecatheon umbels. The bracts which subtend the flowers may
314 A.A. Anderberg
sometimes be fused to the pedicel to a greater or with showy flowers with well-developed corollas
lesser extent. In Kaufmannia the bract is broad are often visited for nectar and nectaries situated
and deeply incised. Only a few taxa are ebracteate. on the gynoecium are found in many genera, e.g.
Prophylls are lacking throughout. Flowers are Primula, Androsace, and Soldanella. There are also
often nodding, particularly in species growing in a number of species with nodding flowers and
alpine or montane habitats, such as many Primula, an anther cone, as in Dodecatheon, Soldanella,
Dodecatheon, Soldanella, and Bryocarpum. and Cortusa. These flowers are likely to be buzz-
pollinated.
FLORAL STRUCTURE. Branching of the vascular
system in the corolla of Soldanella has been been FRUIT AND SEED. Most Primulaceae have many-
brought forward as support for a staminodial seeded capsules, although some genera may have
origin of parts of the corolla, but this needs to be few seeds, e.g. Douglasia and Vitaliana. The cap-
confirmed. In Androsace, Douglasia and Vitaliana, sules are often globose but in a few genera ovoid,
the corolla throat is characteristically constricted ellipsoidal, or even long cylindrical capsules are
and provided with scale-like outgrowths, which found. In most species the capsules dehisce with
are formed by swellings of the corolla tissue. apical valves; Pomatosace has operculate capsules
The heterostylous taxa have short-styled thrum- that dehisce with a circular lid. A number of other
type flowers with the style included in the corolla- taxa are often stated to have operculate capsules
tube, and long-styled pin-type flowers with long (Bryocarpum, Soldanella, Primula sect. Caro-
styles and the stigma presented at the mouth of the linella) but in these taxa capsules open through a
tube. The stigma is usually distinctly capitate. more or less irregular split of the basal portion of
The gynoecium is probably formed by five the style, leaving short, blunt valves and a part of
carpels, although it is initiated from a ring-shaped the style-base as a lid-like structure at the distal
primordium (Schaeppi 1937). end of the mature fruit. It is evident that so-called
operculate capsules comprise different fruit types
EMBRYOLOGY. The anther tapetum is secretory; that have arisen several times.
cells are uninucleate. Pollen is binucleate when The seeds are normally angular with a smooth,
shed. The ovules are anatropous, bitegmic, and reticulate or papillose surface and a hilum more or
tenuinucellate with a micropyle formed by both less flush with the surface. The testa is rather thick,
integuments, the inner of which has an in- usually distinctly two-layered and generally
tegumentary tapetum. In Soldanella the inner provided with rhomboid crystals. The seeds of
integument is partly resorbed. The archespore is Douglasia and Vitaliana are almost semiglobose
multicellular, and the embryo sac is of the Poly- with a fiat or slightly concave ventral surface. The
gonum type with ephemeral antipodals but lacks endosperm is copious and composed of cells with
endosperm haustoria. The seeds are albuminous smooth and evenly thickened walls except in Dou-
with nuclear endosperm formation. Embryogeny glasia, Vitaliana, and some Androsace which have
follows the caryophyllad type; the embryo is short irregularly thickened endosperm cell walls with
and provided with short, narrow cotyledons distinct, narrow constrictions.
(Davis 1966; Johri et al. 1992).
DISPERSAL. The mature fruit is usually borne on
PoLLEN MoRPHOLOGY. The pollen grains of a straight pedicel or scape, and the seeds are
the Primulaceae are basically 3-colporate. Many released ballistically through the openings of the
Primula have syncolpate pollen; stephanocolpate capsule. Some Primula are myrmecochorous, with
pollen is found in Primula (incl. Sredinskya) and a swollen funicle acting as an elaiosome. In the
Dionysia (Erdtman 1952; Wendelbo 1961; Nowicke aquatic Hottonia as well as in some wetland
and Skvarla 1977; Richards 1993). Primula, the seeds are dispersed by water (van der
Pijl1972).
KARYOLOGY. Primulaceae show a great varia-
bility in chromosome numbers but chromosome REPRODUCTIVE BIOLOGY. Heterostyly is known
numbers based on n = 9, 10, 11, and 12 are from Hottonia, Dionysia, Vitaliana, and Primula,
common. Polyploidy is frequent in Androsace and and in the latter genus more than 90% of the
Primula. species are heterostylous. Wendelbo (1961) con-
sidered the genus to be primarily homostylous
PoLLINATION. Most Primulaceae seem to be pol- and stated that other heterostylous genera had
linated by insects, chiefly bees and flies. Species evolved within Primula. Heterostyly in Primula
Primulaceae 315
has been studied extensively and it is known that Dodecatheon much resembles Primula sect. Parryi
the long-styled pin-type is homozygous, whereas with their involute leaves, and Cortusa comes close
the short-styled thrum-type is heterozygous to many species of Primula sect. Cortusioides.
for the heterostyly-gene. In heterostylous Primu- Sredinskya is closely related to the Primula veris
laceae the pin-type flowers also have longer stig- clade. Soldanella, Hottonia, and Omphalogramma
matic papillae and smaller pollen than the share many potential synapomorphies with
thrum-type flowers, except in Vitaliana in which species in Primula, but in the studies by Mast et al.
the pollen grains are of equal size in pin- and (2001) and Trift et al. (2002) the three genera form
thrum-type flowers (Richards 1993). a clade separate from Primula. The Androsace
complex is the sister group of the other genera in
PHYTOCHEMISTRY. Primulaceae often contain va- the family, and the genera Douglasia, Vitaliana,
rious saponins but lack compounds like iridoids and Pomatosace are derived within the Androsace
and ellagic acid. Many genera, including Primula, complex, in which Androsace itself is paraphyletic.
also produce quinoid compounds which are Detailed relationships between the Androsace
known to be allergenic. The flavonol gossypetin, complex and the Primula complex still need to be
which has a very restricted distribution in flower- clarified.
ing plants, has been found in Dionysia, Douglasia The studies by Kallersjo et al. (2000), Mast et al.
and several Primula species. The seeds store oil (2001), and Trift et al. (2002) have demonstrated
and amylose rather than starch (Hegnauer 1969, that a number of genera have their closest relatives
1973). in Primula or Androsace, and this will lead to a
number of new generic circumscriptions. In the
SUBDIVISION AND AFFINITIES. Pax (1889) and following taxonomic account, the traditional
Pax and Knuth (1905) subdivided Primulaceae generic circumscriptions are maintained, awaiting
into five tribes. Melchior (1964) hypothesized that a new generic classification.
the herbaceous growth habit in Primulaceae is
derived, and Judd et al. (1994) put forward that FosSIL RECORD. Seeds identified as Androsace are
Myrsinaceae are a paraphyletic group from which known from the Siberian Miocene, but other
Primulaceae have evolved. Anderberg and Stahl megafossils referred to Primulaceae, and particu-
(1995) performed a preliminary cladistic analysis larly to Primula, are less certain.
of family circumscription in Primulales, and the
work by Kallersjo et al. (2000), which was based DISTRIBUTION AND HABITATS. Primulaceae are
on morphological and DNA sequence data from mainly distributed in temperate and arctic regions
three chloroplast genes, demonstrated that not of the Northern Hemisphere; only few species
only Myrsinaceae but also Primulaceae in the grow in the Southern Hemisphere. The Himalayas
conventional circumscription are paraphyletic, are particularly rich in species of Primula and
since Samolus belongs to Theophrastaceae, and Androsace, and many small genera such as
genera such as Caris, Ardisiandra, Cyclamen, and Pomatosace, Bryocarpum and Omphalogramma
the genera of Primulaceae-Lysimachieae belong are also confined to this region. Many species are
to Myrsinaceae. Kallersjo et al. (2000) therefore confined to alpine habitats including meadows,
reduced Primulaceae to only one of the five rock crevices, and scree; other species grow in
primulaceous tribes proposed by Pax (1889), woodlands. Hottonia, with its submersed gill-
Primuleae, which is characterised by imbricate/ leaves, is adapted to life in freshwater.
quincuncial corolla aestivation. In this circum-
scription, Primulaceae, with few exceptions, are ECONOMIC IMPORTANCE. Many species of Primu-
herbaceous plants with a basal leaf rosette and a laceae are appreciated indoor plants, especially
scapose inflorescence and dry capsular fruits. A cultivars of Primula vulgaris and P. obconica,
later study by Anderberg et al. (2001) showed that which are grown commercially. Species of Cortusa,
also Stimpsonia is part of the herbaceous basal Androsace, and Primula are common as garden
complex of the Myrsinaceae, and that it should be ornamentals and widely used in rock gardens.
excluded from Primulaceae. Many Dionysia are very decorative but grown only
Wendelbo ( 1961) considered it likely that Sredin- by specialists.
skya, Hottonia, Dionysia, Cortusa and Dode-
catheon have their closest relatives within Primula, KEY TO THE GENERA
which is supported by the results of Kallersjo et al. 1. Submerged aquatic plants. Leaves pinnate 7. Hottonia
(2000), Mast et al. (2001), and Trift et al. (2002). - Terrestrial plants. Leaves simple or rarely pinnatifid 2
316 A.A. Anderberg
2. Corolla-lobes conspicuously reflexed 13. Dodecatheon lobate or crenate, flowers in umbels; sect. Chamae-
- Corolla-lobes spreading or erect 3 jasme Koch, caespitose perennials, leaves scarcely
3. Corolla-lobes 7, flowers solitary yellow; capsule narrowly
cylindrical, 3-4cm long 9. Bryocarpum petiolate, spathulate to linear, entire or minutely
- Corolla-lobes 5{6-8), flowers solitary or many; capsule dentate, flowers in umbels; sect. Aretia (L.) Duby,
globose or ellipsoid, not more than 2cm long 4 caespitose perennials, leaves entire or minutely
4. Throat of corolla constricted; corolla-tube short 5 dentate, flowers solitary.
- Throat of corolla not constricted; corolla-tube long or Androsace is paraphyletic if the three following
short 8
5. Capsule opening with an operculate lid; leaves pinnatifid genera are excluded.
4. Pomatosace
- Capsule opening with valves; leaves simple 6 2. Douglasia Lindley
6. Corolla tube generally short 1. Androsace
- Corolla tube longer 7 Douglasia Lindley, Quart. J. Sci. Lit. Arts 1827: 385 ( 1827), nom.
7. Homostylous; flowers pink 2. Douglasia cons.; Constance, Am. Midi. Nat. 19: 249-259 {1938), rev.;
- Heterostylous; flowers yellow 3. Vitaliana Kelso, Can. J. Bot. 70: 593-596 (1991).
8. Corolla six to eight-lobed, slightly zygomorphic
8. Omphalogramma
- Corolla five-lobed, actinomorphic 9
Perennial herbs, sometimes pulvinate. Leaves
9. Corolla-lobes fringed; leaves orbicular to reniform often in basal rosettes, entire or dentate, often
12. Soldanella with stellate hairs. Inflorescence scapose. Flowers
- Corolla-lobes usually entire; leaves of various shapes but if homostylous, in terminal umbels; calyx campanu-
the corolla-lobes are fringed, then the leaves are not entire late; corolla hypocrateriform, with a short tube,
and orbicular or reniform 10
10. Stamens inserted at the base of the corolla 11
constricted at the throat and with scale-like out-
- Stamens inserted in the corolla-tube 12 growths, violet or purple; stamens adnate to the
11. Flowers pink; bracts entire 10. Cortusa tube; stigma capitate. Capsule globose; seeds few,
- Flowers yellow; bracts deeply incised 11. Kaufmannia reticulate or papillose. 2n = 36, 38, 40. Eight spp.,
12. Corolla tube at least three times longer than the calyx. USA, Canada, E Siberia.
Ovules few 6. Dionysia
- Corolla tube not three times longer than the calyx. Ovules Douglasia is a derived part of Androsace.
many 5. Primula
3. Vitaliana Sesler
GENERA OF PRIMULACEAE
Vitaliana Sesler in Donati, Essai Hist. Nat. Mer Adriat.: 69
{1758); Constance, Am. Midi. Nat. 19: 249-259 {1938);
1. Androsace L. Ferguson, Fl. Europaea 3: 20 {1972).
Douglasia subg. Gregoria (Duby) Knuth {1905).
Androsace L., Sp. Pl.: 141 (1753); Smith and Lowe, Androsaces
{1977). Perennial herb, branched, prostrate and forming
A retia L. {1753 ).
mats. Leaves alternate, linear, with stellate hairs.
Flowers heterostylous, subsessile, solitary or
Suffrutices, perennial, or annual herbs, sometimes
paired; calyx tubular; corolla hypocrateriform,
densely pulvinate, or prostrate and forming mats.
with a tube about twice as long as calyx, somewhat
Leaves entire or dentate, alternate, sessile or peti-
constricted at the throat and with scale-like out-
olate, generally in basal rosettes, ovate, spathulate
growths, yellow; corolla-lobes linear, longer than
or linear, often with stellate hairs. Inflorescence
wide, imbricate when young; stamens adnate
generally scapose. Flowers homostylous, in ter-
to the tube; stigma capitate. Capsule globose,
minal umbels or solitary in the leaf-axils; calyx
opening with apical valves, seeds few, almost
campanulate; corolla hypocrateriform, with a
smooth. 2n = 32, 40, 80. One sp., V. primuliflora
short tube, constricted at the throat and with
Bertol., European Alps.
scale-like outgrowths, white, pink, or red; corolla-
Vitaliana is a derived part of Androsace.
lobes entire or emarginate; stamens adnate to the
tube; stigma capitate. Capsule globose, opening
with valves; seeds reticulate or papillose. 2n = 18, 4. Pomatosace Maxim.
20, 22, 30, 36, 38, 40, 48, 78. About 150 spp., north Pomatosace Maxim., Bull. Acad. Imp. Sci. Saint-Petersbourg 27:
temperate areas, many spp. in China and the 499 {1882).
Himalayas. Four sections: sect. Androsace (=
Andraspis (Duby) Koch), annual, leaves oblong or Annual herb. Leaves basal, pinnatifid. Inflores-
linear, entire or dentate, not clearly petiolate, cence scapose. Flowers homostylous, in terminal
flowers in umbels; sect. Pseudoprimula Pax, umbels; calyx campanulate with conspicuous tri-
perennial, leaves long petiolate, large, margin angular, persisting lobes; corolla white, with short
Primulaceae 317
tube, constricted at the throat and with scale-like Primula. Many of the subgenera are also in need of
outgrowths; corolla-lobes entire, erect to spread- a new circumscription.
ing; stamens adnate to the tube; stigma capitate.
Capsule globose, opening with a lid; seeds many, 6. Dionysia Fenzl
papillose. 2n = 20. One sp., P. filicula Maxim., NW
China. Dionysia Fenzl, Flora 26: 389 (1843); Wendelbo, Arb. Univ.
Pomatosace is a derived part of Androsace. Bergen, Mat. Nat. Ser. 3: 1-83 (1961); Grey-Wilson, The
genus Dionysia ( 1989 ); Al Wadi & Richards, New Phytol. 121:
303-310 (1992).
5. Primula L. Fig. lOlD
Perennial herbs, often woody at base and often
Primula L., Sp. Pl.: 142 (1753); Wendelbo, Arb. Univ. Bergen,
Mat. Nat. Ser. 11: 1-49 (1961), rev.; Wendelbo, Arb. Univ.
forming dense tufts. Leaves alternate or whorled,
Bergen. Mat. Nat. Ser. 7: 1-15 (1961), pollen morph.; Smith, revolute when young, often with articulated hairs
Forrest and Fletcher, Plant Monogr. Reprints 11: 1-835 and a farinose coating. Inflorescence generally
(1977); Richards, Primula (1993); Mast et al. Int. J. Plant Sci. scapose. Flowers heterostylous, in terminal simple,
162: 1381-1400 (2001), phylogeny; Trift et al. Syst. Bot. 27: or superimposed umbels, or solitary; calyx
396-407 (2002), phylogeny.
Sredinskya (Stein.) Fedorov (1950). tubular; corolla hypocrateriform, with a very long
slender tube, violet, pink or yellow; corolla-lobes
Perennial herbs. Leaves basal, revolute or involute spreading, entire or emarginate; stamens adnate to
when young, often with a white or yellow farinose the tube; stigma capitate. Capsule subglobose,
coating, entire, dentate or serrate, evenly attenuat- opening with apical valves; seeds few or many,
ing or with well demarcated blade and petiole, papillose. 2n = 20. 40 spp., Mountains of Central
sometimes pinnatifid, with articulated hairs or Asia.
glabrous, exceptionally with flattened gland-dots. Dionysia is a derived part of Primula, diagnosed
Inflorescence often scapose. Flowers heterostylous by longer corolla tube and fewer ovules.
or less often homostylous, in umbels, sometimes in
superimposed umbels, spike-like racemes, or soli- 7. Hottonia L.
tary; calyx campanulate or cylindrical; corolla cam-
panulate to hypocrateriform, with a short or long Hottonia L., Sp. Pl.: 145 (1753).
tube, white, yellow, pink, red, violet or blue; corolla-
lobes spreading or erect, entire, emarginate, bifid, Perennial aquatic herbs. Leaves submersed,
or fringed; stamens adnate to the tube; stigma cap- pinnate, alternate or verticillate, glabrous; leaf-
itate. Capsule globose or ellipsoid, opening with lobes linear, involute when young. Inflorescence
apical valves, rarely with a lid, or through irregular scapose, held above the water surface. Flowers in
disintegration; seeds reticulate or papillose. 2n = several whorls (superimposed umbels), hetero-
18,20,22,23,24,29,32,36,40,44,48,54,62,66,72, stylous; corolla white to pink, longer or shorter
126 (x = 8, 9, 10, 11, 12). About 400 spp., mainly N than the calyx; corolla-lobes spreading, often
Hemisphere. Six subgenera, often characterized by emarginate; stamens adnate to the tube; stigma
the following string of features: subg. Sphondylia capitate. Capsule opening laterally with apical
(Duby) Rupr. ( 1863), generally homostylous, leaves valve-slits; seeds few, reticulate. 2n = 20, 22. Two
involute, pollen 3-colporate; subg. Auriculastrum spp., North America, Eurasia.
Schott ( 1851 ), heterostylous, leaves involute, pollen
3-colporate or 3-syncolpate; subg. Primula, het- 8. Omphalogramma (Franch.) Franch.
erostylous, leaves revolute, pollen stephanocolpate;
subg. Auganthus (Link) Wendelbo {1961), het- Omphalogramma (Franch.) Franch., Bull. Soc. Bot. France 45:
178 (1898); Fletcher, Not. Roy. Bot. Gard. Edinburgh 20:
erostylous or homostylous, leaves revolute, pollen 125-159 (1949).
3-colporate, 3-syncolpate, or stephanocolpate; Primula subg. Omphalogramma Franch. (1885).
subg. Carolinella (Hemsley) Wendelbo {1961),
heterostylous, leaves revolute, capsule operculate; Perennial herbs. Leaves basal, involute when
subg. Aleuritia (Duby) Wendelbo (1961), het- young, ovate to elliptic, entire to denticulate,
erostylous or homostylous, leaves revolute, pollen with flattened gland-dots. Inflorescence scapose,
3-colporate, 3-syncolpate, or stephanocolpate. ebracteate. Flowers (5)6{ -8)-merous, homosty-
Primula is paraphyletic in its present circum- lous, somewhat zygomorphic, solitary; corolla
scription.At least the genera Dionysia and Cortusa, hypocrateriform, with a rather long tube, blue,
and Dodecatheon have their closest relatives within rose to dark purple; corolla-lobes spreading,
318 A.A. Anderberg
Fig. 101. Primulaceae. A Soldanella

montana, immature fruit. B Same,
1
dehisced, showing stylopodiallid. C Dode-
catheon meadia, flower. DPrimula denticu-
lata, flower, vertical section. E Cortusa
matthioli, habit. F Same, inflorescence.
Drawn by Pollyanna von Knorring
emarginate to dentate; stamens adnate to the tube; 11. Kaufmannia Regel

stigma capitate. Capsule cylindrical, opening with
Kaufmannia Regel, Trudy Imp. S.-Peterburgsk Bot. Sada 3: 293
valves; seeds reticulate with winged margin. 2n = (1875).
48, 96. 15 spp., Himalayan region.
Perennial herbs. Leaves basal, coarsely dentate,
9. Bryocarpum Hook. f. & Thoms. petiolate. Inflorescence scapose; bracts deeply
incised. Flowers homostylous, in umbels; corolla
Bryocarpum Hooker f. & Thomson, Hooker's J. Bot. Kew Gard. yellow; corolla-lobes entire; stamens connivent;
Misc. 9: 200 (1857).
style exserted; stigma capitate. Capsule elliptic,
opening with valves; seeds many, papillose. 1-2
Perennial herb. Leaves basal, involute when young,
spp. in Central Asia.
ovate with truncated base, with blackish flattened
Kaufmannia is very close to Cortusa, and like
gland-dots. Flowers 7-merous, nodding, homosty-
this probably a derived part of Primula.
lous, solitary; calyx campanulate; corolla narrowly
campanulate, yellow; corolla-lobes erect, emar-
ginate; stamens adnate to the tube; stigma capi- 12. Soldanella L. Fig. lOlA, B
tate. Capsule long cylindrical, opening with a lid
Soldanella L., Sp. Pl.: 144 (1753).
and with apical valves; seeds many, reticulate. One
sp., B. himalaicum Hook. f. & Thoms., E Himalayas.
Perennial herbs. Leaves ovate to reniform, entire,
petiolate, often with articulated hairs. Inflo-
10. Cortusa L. Fig. lOlE, F rescence scapose, bracteate. Flowers nodding,
homostylous, in terminal few-flowered umbels or
Cortusa L., Sp. Pl.: 144 (1753).
solitary; calyx campanulate; corolla blue, campan-
ulate; corolla-lobes erect, laciniate; stamens
Perennial herbs. Leaves basal, revolute when
connivent, forming a cone; style exserted; stigma
young, petiolate, coarsely dentate. Inflorescence
capitate. Capsule elliptic, opening with a lid and
scapose; bracts entire. Flowers homostylous, in
with apical valves; seeds many, reticulate. 2n =38,
umbels; calyx campanulate; corolla campanulate,
40. 10 spp., European Alps.
pink with darker blotches in the throat; cor-
olla-lobes erect, entire; stamens connivent; style
exserted; stigma capitate. Capsule elliptic, opening 13. Dodecatheon L. Fig. 101C
with valves; seeds many, papillose. 2n = 24. Eight
Dodecatheon L., Sp. Pl.: 144 (1753); Thompson, Contrib.
spp., mountains of Europe and Asia. Dudley Herb. 4: 73-154 (1953).
Cortusa is a derived part of Primula.
Perennial herbs. Leaves basal, linear to spathulate,
glabrous. Inflorescence scapose. Flowers many
Primulaceae 319
together in a terminal umbel, nodding, homosty- Kallersji:i, M. et a!. 2000. See general references.
lous; calyx campanulate; corolla campanulate, Mast, A.R., Kelso, S., Richards, J., Lang, D.J., Feller, D.M., Conti,
E. 2001. Phylogenetic relationships in Primula L. and related
white to pink often with darker blotches; corolla- genera (Primulaceae) based on noncoding chloroplast
lobes entire and distinctly reflexed; stamens con- DNA. Int. Y. Plant Sci. 162:1381-1400.
nivent, forming a protruding cone; style exserted; Melchior, H. 1964. Primulales. In: Melchior, H. (ed.) Engler's
stigma capitate or truncate. Capsule elliptic, Syllabus der Pflanzenfamilien 2. Berlin: Borntrager, pp.
389-394.
opening with apical valves or with a lid; seeds
Nowicke, J.W., Skvarla, J.J. 1977. Pollen morphology and
angular or somewhat winged, reticulate. 2n = 22, the relationships of the Plumbaginaceae, Polygonaceae,
44, 88. 13 spp. in E North America and Siberia. and Primulaceae to the order Centrospermae. Smithsonian
Dodecatheon is a derived part of Primula, diag- Contrib. Bot. 37: 1-64.
nosed e.g. by distinctly reflexed corolla lobes. Pax, F. 1889. Primulaceae. In: Engler & Prantl, Die natiirlichen
Planzenfamilien IV. 1. Leipzig: Engelmann, pp. 98-116.
Pax, F., Knuth, R. 1905. Primulaceae. In: Engler, A. (ed.) Das
Pflanzenreich IV, 237: 1-386.
Selected Bibliography Richards, J. 1993. Primula. Portland: Timber Press.
Schaeppi, H. 1937. Vergleichend-morphologische Unter-
Anderberg, A.A, Stahl, B. 1995. Phylogenetic interrelationships suchungen am Gynoecium der Primulaceen. Zeitschr.
in the order Primulales, with special emphasis on the family Gesamte Naturwiss. 3: 239-250.
circumscriptions. Can. J. Bot. 73: 1699-1730. Trift, I., Kallersjo, M., Anderberg, A.A. 2002. The monophyly of
Anderberg, A.A., Peng, C.-I., Trift, I., Kallersji:i, M. 2001. The Primula (Primulaceae) evaluated by analysis of sequences
Stimpsonia problem; evidence from DNA sequences of from the chloroplast gene rbcL. Syst. Bot. 27: 396-407.
plastid genes atpB, ndhF and rbcL. Bot. Jahrb. Syst. 123: Van der Pijl, L. 1972. Principles of dispersal in higher plants.
369-376. Berlin Heidelberg New York: Springer.
Davis, G. 1966. See general references. Wendelbo, P. 1961. Studies in Primulaceae III. On the genera
Erdtman, G. 1952. See general references. related to Pri;rzula with special reference to their pollen
Hegnauer, R. 1969, 1973. See general references. morphology. Arb. Univ. Bergen, Mat. Nat. Ser. 7: 1-15.
Johri, B.M. et a!. 1992. See general references. Zhang, L.-B., Comes, H.P., Kadereit, J.W. 2001. Phylogeny
Judd, W.S., Sanders, R.W., Donoghue, M.J. 1994. Angiosperm and quaternary history of the European montane/alpine
family pairs: preliminary phylogenetic analyses. Harvard endemic Soldanella (Primulaceae) based on ITS and AFLP
Papers Bot. 5: 1-51. variation. Amer. J. Bot. 88: 2331-2345.
320 D. Medan and C. Schirarend
Rhamnaceae
D. MEDAN and C. SCHIRAREND
Rhamnaceae Juss., Gen. Pl.: 376 (1789) ('Rhamni'), nom. cons. VEGETATIVE MORPHOLOGY. Architecture is
intermediate between Attim's and Raux's (Zizi-
Deciduous or evergreen, often thorny trees, phus), or between Koriba's and Raux's (Ziziphus,
shrubs, woody climbers or lianes, rarely herbs. Paliurus) models (Tourn et al. 1992). Dichasial
Leaves simple, petiolate, alternate or opposite, with sympodia are common: the apical meristem of
1 main vein or 3-5 veins, entire to serrate, some- orthotropic shoots ceases growth or produces a
times much reduced; stipules small, caducous spine, while buds of two distal nodes continue
or persistent, sometimes fused intrapetiolarly or the growth. Leaves usually subtend a single bud,
interpetiolarly, or transformed into spines, absent less commonly serial buds (Colletieae, Gouania,
in most Phylica. Inflorescence basically cymose, Ziziphus, Tourn et al. 1991). The lowermost serial
cymes mostly axillary, sessile or pedunculate, or bud can form vegetative long shoots, short -shoots,
reduced to many-few-flowered fascicles. Flowers or flowering shoots, and the distal bud often pro-
small, 3-6mm in diameter, regular, (3)4-5(6)- duces a spine that may ramify (Tortosa et al. 1996).
merous, bisexual or unisexual, plants sometimes Spinyness is absent only in Gouanieae and Venti-
dioecious, haplostemonous, hypogynous to lagineae but pervasive only in the Colletieae.
epigynous, yellowish to greenish, rarely brightly Twining shoots occur in Berchemia, Ziziphus, Ven-
coloured; hypanthium patelliform or hemispheri- tilago and Smythea, and circinate tendrils in
cal to tubular, sometimes absent, at the rim Gouania, Reissekia and Helinus. Rhizomes were
bearing calyx, corolla and stamens; sepals 4 or 5, reported for Colletia, and shoots adventitious on
valvate in bud, triangular, erect to more or less roots for Rhamnus, Phylica, and Ziziphus. Some
recurved during anthesis, often keeled adaxially; species are able of sprouting after fire (Zedler
petals 4 or 5, rarely 0, usually smaller than sepals, 1995; Tortosa et al. 1996).
concave or hooded, rarely almost flat, often shortly
clawed, often enfolding the stamens; stamens 4 or VEGETATIVE ANATOMY. Leaves are generally
5, antepetalous, filaments thin, adnate to the base dorsiventral, the hairs mostly simple (exceptions:
of the petals, anthers minute, versatile or not, 2( 4)- two-armed in Sageretia, stellate in Pomaderreae).
locular, dehiscing by longitudinal slits, usually The epidermis is often mucilaginous or includes
introrse; disc intrastaminal, nectariferous, thin to mucilaginous cells. Stomata are usually anomo-
more or less fleshy, entire or lobed, glabrous, rarely cytic, less commonly paracytic or anisocytic.
pubescent, free from ovary or tightly surrounding Stomata-bearing furrows or cavities are found in
it, or adnate to the hypanthium; combined Ceanothus and Condalia. A multiple epidermis is
pubescence of hypanthium and style sometimes unusual (Rhamnus); a hypodermis is reported
forming a secondary pollen presenter and pollen- for several genera. Palisade parenchyma is one
dosing structure; ovary superior to inferior, to several cells thick, often including isolated,
(1)2-4-locular, with 1(2) ovules in each locule, enlarged cells filled with mucilage, tannin, or both.
ovules anatropous, basal and erect. Fruit indehis- Secretory cavities and secretory channels occur
cent, winged or not, schizocarpic, capsular, rarely occasionally in Rhamneae and Paliureae. Lysige-
explosively dehiscent, or a more or less fleshy nous mucilage cavities have been recorded for the
drupe with 1-4 indehiscent, rarely dehiscent leaf vein parenchyma and petioles from all tribes.
pyrenes. Seeds with thin, oily albumen, sometimes Druses are common in the mesophyll; acicular
exalbuminous, embryo large, oily, straight or crystals are known only from Gouanieae. Small
rarely bent. lateral, vertically transcurrent veins are known
An almost cosmopolitan family of 52 genera and from several tribes. The petioles have usually a
about 925 species. single open or U-shaped bundle. Characters of
diagnostic value include type of leaf structure,
venation, thickness and striation of the cuticle,
Rhamnaceae 321
presence of a multiple epidermis, type of ing synflorescences and perennial short-shoots as

mucilaginous epidermis, presence of stomata- derived.
bearing cavities or furrows, type of stomata, dis-
tribution of indumentum, crystals and secretory FLOWER STRUCTURE. Claims for an axial respec-
cells, proportion of palisade tissue, and presence tively appendicular nature of the hypanthium
of collenchyma connecting the main vein with lack adequate support (Medan 1985). Flower size
the upper epidermis (Gemoll 1902; Herzog 1903; and hypanthium depth are correlated in Colletieae
Medan 1986). (Medan and Aagesen 1995). The nature of the
The wood is generally diffuse-porous to ring- petal-stamen complex is much disputed among
porous. Growth rings are ± distinct, marked by plant morphologists (Bennek 1958; Sattler 1973;
differences in vessel diameter, latewood fibres or Ronse Decraene et al. 1993).
marginal parenchyma bands. Vessels (2-220 per The disc is intrastaminal, either annular (secre-
mm 2 ) are solitary or in radial multiples or clusters tory surface a ring surrounding the ovary),
of 2-5(-15); vessel member length is from 110- adpressed (secretory surface lining the hypan-
700 J..Lm. Perforations are mostly simple, rarely thium, delimited distally by a rim), revolute (a
scalariform to reticulate; intervessel pits often coa- laminar projection of the flower tube above ovary
lesce and are rarely vestured. Vessel-ray and level) or indistinct (as the adpressed type but the
vessel-parenchyma pits are exclusively half-bor- boundary with non-secreting area only anatomi-
dered. Vascular tracheids intergrading with cally discernible) (Medan and Aagesen 1995).
narrow vessels are rather abundant, often with The development of the gynoecium starts with
spiral thickenings. Fibre elements are exclusively a unilocular primordium; plication and cen-
libriform, 650-IOOOJ..Lm long. Wood rays (10- tripetal growth of the fused carpellary flanks build
20 per mm) are 1-5 cells wide and 10-30 cells up the septa, which later fuse at the centre, often
high and distinctly heterogeneous to homoge- incompletely, thus leaving a compitum (Medan
neous. Axial parenchyma is usually scanty para- 1985, 1988). Carpels bear an ovule on one of their
tracheal to vasicentric, rarely diffuse apotracheal margins, the other margin being sterile (van
or the parenchyma cells are arranged in uni- to Tieghem 1875; Suessenguth 1953). In 3-carpellate
multiseriate, tangential to diagonal bands. Pri- ovaries the sequence is usually fertil-sterile-fertil-
smatic to diamond-shaped, rarely styloid crystals sterile-sterile-fertil, i.e. one septum bears two
are very frequent in the axial and ray paren- ovules, the second a single ovule, and the third is
chyma cells or in chambered strands of the sterile, but over 15 different septation patterns
axial parenchyma. The heartwood is often filled have been found (Suessenguth 1953; Tortosa 1984;
with brownish, gum-like deposits (Schirarend Medan 1985). The stigmata are terminal on the
1984). stylar branches; entire styles have various stigma
Ultrastructural data are scant. The mucilaginous types (Medan and Aagesen 1995). The style is tra-
epidermal cells of Rhamnus frangula were studied versed by as many stylar canals as locules (Medan
in detail by Jakovleva (1988); sieve-element plas- 1985); rarely a single canal is present, or the style
tids are of the S-type in all four genera investigated is solid (Hanackova and Pifteyro Lopez 1999).
by Behnke (1974); in Rhamnus frangula the Increased functional syncarpy is achieved in
mucilage-secreting cavities in shoot apices and genera with a single stylar canal or a solid style, or
flower buds have been found to be of lysigenous when a stigmatic compitum is formed (Colletia,
origin (Bouchet 1974). D'Ambrogio and Medan 1993).
INFLORESCENCE STRUCTURE. The inflorescences EMBRYOLOGY. Anthers are 2- or 4-locular; the

are generally described as cymes, or more specifi- development of anther wall follows the basic type.
cally as umbelliform, corymbiform, racemiform, The endothecium has fibrous thickenings; the two
spiciform cymes (Suessenguth 1953; Brizicky middle layers are ephemeral, and the cells of the
1964). Solitary terminal flowers have also been glandular tapetum become 2-nucleate. Cytokine-
reported. Detailed studies are only available for a sis in the microspore mother cells is simultaneous;
few species of Ceanothus, Noltea, Phylica, Paliurus, tetrads are tetrahedral or isobilateral. Pollen
Pomaderris and Rhamnus, and for Colletieae grains are 2-celled when shed. The ovules are
(Tortosa et al. 1996). Within this tribe, deciduous anatropous, bitegmic (outer integument thicker),
monotelic synflorescences (either thyrsoids with crassinucellate, and the micropyle is usually
cymes as paracladia, or botryoids with !-flowered formed by the outer integument. A hypostase is
paracladia) were considered basic, and proliferat- sometimes present, an epistase is rare (Ziziphus).
Fig. 102. Rhamnaceae pollen. A Ventilago viminalis, equato- pattern), with ± densely spaced perforations. The
rial view; sexine supra-reticulate. B Karwinskia rocana, polar boundary between tectum and subtectal elements
view; sexine rugulate. C Auerodendron glaucescens, equatorial
section showing paired exine thinnings parallel to colpi; all is rather indistinct; the subtectallayer is very dif-
SEM (x2000). Photo C. Schirarend ferent in thickness and mostly granular. Solid sub-
tecta} bacula are rare (Pomaderris); the foot layer
is of different thickness and has a distinct distal
Funicular vascular strands usually almost reach surface. The nexine is always well differentiated,
the tip of the outer integument. The archesporium thin at the mesocolpia, and± thickened at the colpi
is 1- to several-celled, each cell giving rise to a (costae transversales; Schirarend and Kohler
parietal derivative that produces up to 13 parietal 1993).
layers. The apical cells of the nucellar epidermis
form a nucellar cap 5 cells thick. Sometimes a KARYOLOGY. The basic chromosome number of
nucellar beak protrudes through the micropyle. Rhamnaceae appears to be x = 12; 2n = 24 is
The embryo sac is of the Polygonum type, less reported for most tribes. Other counts include
commonly of the Allium type. Antipodals are n = 11, known from Helinus and most Colletieae
generally ephemeral; in Ziziphus they persist or studied, and n = 10, 11, and 13 recorded for
become coenocytic. The polar nuclei fuse at about Rhamnus (Rhamneae). Polyploidy is apparently
the time of fertilisation. Multiple embryo sacs are restricted to cultivated species of Ziziphus (e.g. 24,
reported for several genera. Endosperm formation 48, 60, 72, 96 for Ziziphus jujuba) and to Pomader-
is nuclear; the tissue later becomes cellular from ris (e.g. 24, 36 and 48; Richardson et al. 2000a).
the micropylar pole, but cellularization does not
always reach the chalaza} end. Embryogeny follows REPRODUCTIVE BIOLOGY. Relevant data are avail-
the Asterad type, sometimes the Solanad type able for some 14 genera, but detailed studies have
(Ziziphus). Synergid polyembryony is occasionally been conducted for only 15 species (references in
found (Ziziphus; Davis 1966). Medan and D'Ambrogio 1998; Medan and Arce
2000). Overall, flowers of Rhamnaceae are small,
PoLLEN MoRPHOLOGY. Pollen grains are usually often inconspicuous, and receive visits of insects
isopolar, radially symmetrical, angulaperturate, that are rewarded with easily accessible nectar and
3{4)-zono-colporate, suboblate to subprolate, pollen.
11-40~m in polar diameter, 12-34~m in equato- Flowering is usually in spring and summer;
rial diameter, and angular to circular in polar view some taxa start blooming in late winter
(Fig. 102). Apertures are composed of compara- (Berchemia, Discaria, Retanilla, Rhamnus) or con-
tively narrow colpi and ± narrow, lalongate to tinue to flower into the cold season (Colletia,
elliptic, rarely lolongate endoapertures. The colpi Phylica). Individual flowers last 1.5-10 days, being
extend over about 4/5 of the length of the polar usually protandrous (a shorter male phase
axis. The exine is 1-2{-3.5) ~m thick, with con- generally overlaps with a longer female phase).
spicuous thinnings parallel to the colpi (Fig. Homogamy was reported for Colletia; a report on
102C). The sexine is mostly pertectate-perforate. protogyny (Maesopsis) needs confirmation.
The tectum is microreticulate, striate, rugulate, The flowers are generally white, greenish, less
fossulate, verrucate, pilate or ± psilate (LO- commonly yellow, rarely pink or blue (Ceanothus)
Rhamnaceae 323
or red (Colletia). They are weakly scented Protandry and SI are apparently maintained
(Rhamnus) to almost oppresively fragrant (Dis- when abortion of the stamens or the pistil give rise
caria), from foetid or nauseating (Discaria to pistillate or staminate flowers. Andromonoecy
americana, Ziziphus mucronata) to pleasant is reported for Alphitonia, Colubrina, and Trevoa,
(vanilla-, cinnamom-, or honey-like). the latter showing reduced incompatibility. The
Pollen is commonly exploited and may function pollen-ovule ratio is relatively high (Trevoa
as a subsidiary reward. Nectar is easily accessible quinquenervia: 14500: 1). True dioecy is known
except in species with relatively long hypanthia; from Rhamnus, where the pollen-ovule ratio is
amounts may be minute, but no true nectarless very high (ca. 32300: 1). Less well-known cases of
species has been reported. Sugar concentrations dimorphism occur in Noltea, Alphitonia, Phylica,
are usually lower than 20% but may reach 30% Gouania, Rhamnus, and Discaria.
(Ziziphus). Both fructose/glucose and saccharose- Under open pollination, the fruit!ftower ratio of
dominated nectars have been reported. SI species is usually below 10%. Fruit set is higher
The cucullate petals cover often the anthers dur- in species with reduced incompatibility (Trevoa
ing anthesis, which may prevent pollen robbing quinquenervia: 48.8%) and in self-compatible
and may protect pollen from desiccation (Medan species (Pomaderris kumeraho: 43.9%). Crop size
and Aagesen 1995). Centrifugal stamen move- varies from 300 to 8000 fruits per individual per
ments during anthesis, recorded for several genera, year (Rhamnus), and from 120 (Discaria) to over
possibly prevents pollen-stigma interference 17000 viable seeds per plant per year (Colletia).
(Medan and Hilger 1992). In Trevoa and Retanilla Reproductive success is pollen-limited at the
pollen becomes trapped in the hairs inside the flower level in some cases (Rhamnus, Ziziphus, and
floral tube; this hairy barrier acts as an exposed, Discaria).
secondary pollen presenter because mouthparts of
insects must traverse it to reach nectar. Both wet FRUIT AND SEED. Dry fruits may be indehiscent,
and dry stigmas have been reported. schizocarpous, or capsular; dehiscence can be slow
Generalized, unspecialised entomophily is the or explosive. Fleshy fruits are always drupaceous.
rule, with Hymenoptera and Diptera co-dominat- All types have a sclerenchymatous endocarp
ing nearly always. Increased flower tube length usually developed from the inner carpel epidermis
seems associated with a higher proportion of (Vikhireva 1952). Each carpel produces one stone,
Hymenoptera and Lepidoptera in some assem- or the fruit produces a single, compound stone.
blages (Colletieae, Medan and Aagesen 1995). Parthenocarpic fruits were recorded in Discaria,
Evidence of pollen transport by wind exists for Rhamnus, and Ziziphus.
Rhamnus (Aronne and Wilcock 1995) and Zizi- The thin-walled, xerochastic, 3-4-carpellate ex-
phus (Zietsman 1990). plosive capsules open through sudden aperture of
Monomorphic hermaphroditism combined ventricidal, dorsicidal and septifragal dehiscence
with protandry and self-incompatibility (SI) is lines in each of the 3-4 stones (endocarpids),
widespread in the family. Incomplete dichogamy, which also separate along septicidal planes. Dehis-
high flower number per individual and sequential cence is caused by oblique bending of the endo-
flowering favour selfing, which is counterbalanced carpids due to a distinct distribution of endocarpal
by gametophytic Sl. Pollen-ovule ratios under this cells. These occur in parallel layers with crossing
system are about 7000: 1. micellary structure, which leads to enhancing
Heterodichogamy occurs in Ziziphus. It implies stress during fruit dessication. A pedestal may
the occurrence of two floral morphs, both being persist on the plant after dehiscence (Colletieae,
protandrous, one starting flowering several hours Ceanothus: colletioid dehiscence; Medan 1985) or
after the other. While the flowers of one morph are not (Colubrina p.p., Noltea: colubrinoid dehis-
receptive, those of the other release pollen; later, cence; Johnston 1971; Medan and Aagesen 1995).
the situation is reversed, so that each individual The seeds are ejected up to 9 m away.
is alternately male and female (Galil and Zeroni Slowly dehiscing capsules are similar but the
1967; Zietsman and Botha 1992). Pollen-ovule endocarpids open slowly and often tardily and
ratios vary from 3300: 1 to 35000: 1. incompletely; the fruit disintegrates into one-
Self-compatibility occurs in some Pomaderris seeded endocarpids that dehisce later, or the seeds
spp. (Harvey and Braggins 1985). Their flowers fall off from the open fruits, or remain on the fruit-
are monomorphic and apparently protandrous, ing pedicel after pericarp breakdown.
but SI is largely suppressed (fruit set under self- Most Gouanieae have schizocarps that split into
pollination above 60%). three indehiscent, winged, one-seeded mericarps
which temporarily remain attached to a car- DISPERSAL. Entire fruits (samaras) are dispersed
pophore. Mericarps may be solid (Gouania, in Paliurus and Ventilago. In Smythea the disper-
Crumenaria) or inflated (Reissekia). Fruits sal by wind precedes dehiscence (Ridley 1930). In
transitional between capsules and schizocarps the Gouanieae the diaspores are winged mericarps
occur in the Gouanieae (Alvimiantha, Fig. 105, or hemimericarps. Undehisced Smythea fruits
p. 331), in which the septicidal capsule liberates may float in seawater for months (Guppy 1906;
winged, indehiscent hemimericarps (Medan Ridley 1930). The seeds of Colubrina asiatica may
1989). float in seawater over long time (Carlquist 1966).
Dry indehiscent winged fruits occur in Ventilago Transport of whole fruits by rainwater was
(Ventilagineae, with terminal wing) and Paliurus reported for Retanilla (Reiche 1907). All explosive
(Paliureae, with horizontal wing). A 3-seeded nut capsules effect ballistic seed dispersal (e.g. Cean-
with a fleshy, edible pedicel occurs in Hovenia othus, Evans et al. 1987).
(Paliureae). Trevoa (Colletieae) has a papery, Birds of over 35 families are known to disperse
usually one-seeded, rostrate nut partially covered whole fruits of Condalia, Berchemia, Maesopsis,
by the persistent floral tube (Fig. 103E, p. 328). Phylica, Rhamnus, Scutia, and Ziziphus, and
Typical drupes occur in Rhamneae, with either apparently also the arillate seeds of Alphitonia
a compound stone or 2-4 1-seeded individual (Sun and Dickinson 1996). Mammals including
stones, which may be tardily dehiscent. some 15 different groups from rodents to ele-
Seeds are exotestal, small to medium-sized, phants are involved in dispersal of Karwinskia,
rounded and somewhat dorsiventrally compressed Rhamnus, and Ziziphus, and perhaps Hovenia.
(rarely cordate: Alvimiantha, or dorsally furrowed: Ants transport arillated seeds of Phylica
Rhamnus p.p.). The testa is generally multiplica- (Kilian and Cowling 1992), several Pomaderreae
tive, smooth. The outer epidermis is a palisade of (Andersen and Ashton 1985), Ceanothus (Mills
thick-walled prismatic cells, often with a linea and Kummerow 1989), and (after dehiscence of
lucida. The mesophyll is aerenchymatous and ± bird-dispersed fruits) non-arillate seeds of
crushed, rarely with scattered sclerotic cells (Ven- Rhamnus (Aronne and Wilcock 1994).
tilago). The inner epidermis is unspecialised. The
tegmen is not or slightly multiplicative, soon PHYTOCHEMISTRY. The family is generally known
crushed; the inner epidermis is not lignified. The for the widespread occurrence of calcium oxalate,
vascular bundle extends into the raphe or has tannins and certain alkaloids. Phenolic com-
postchalazal extensions (Ventilago, Ziziphus). The pounds are accumulated in large quantities as
endosperm is nuclear, oily, generally as a thin fiavonols and leucoanthocyanins and the con-
layer, rarely well developed and ruminate densed tannins based on them. Derivatives of
(Reynosia), sometimes absent (Ventilagineae, anthraquinone are restricted to Rhamnus, where
several Rhamneae, Ampeloziziphus). The embryo they are pharmacologically used as laxatives (e.g.
is straight or rarely bent, invariably chlorophyllous Cortex Rhamni Frangulae, Fructus Rhamni
(Yakovlev and Zhukova 1980), oily; cotyledons are Cathartici). More widely distributed within the
comparatively large, usually fiat and reaching the family are the derivatives of naphtalin and lupeol
seed coat (thus dividing the endosperm in two and different types of saponins. For many genera
sectors), rarely curved (some Rhamnus). Funicu- the accumulation of alkaloids has been proved,
lar arils are often present, sometimes conspicuous which comprise peptid alkaloids and isochinolin
( Ceanothus, Colubrina, Phylica) and different in alkaloids (Jossang et al. 1996).
colour from the seed coat (Alphitonia), perhaps
overlooked in some genera (Medan and Aagesen SUBDIVISION AND RELATIONSHIPS WITHIN THE
1995). A tissue of raphal origin serves as an elaio- FAMILY. On the basis of a phylogenetic analysis of
some (some Rhamnus; Aronne and Wilcock 1994). rbcL and trnL-F sequences of the plastid genome
Heat may promote germination (Rhamnus, and morphological data, Richardson et al. (2000a,
Phylica, Ceanothus; Keeley 1992; Kilian and 2000b) revised the tribal classification of the
Cowling 1992). Embryo dormancy is overcome by family. Two of the five tribes recognised by Sues-
stratification in Discaria, Hovenia and Retanilla senguth ( 1953) were shown to be polyphyletic,
(Soriano 1960; Frett 1989; Keogh and Bannister whereas three appeared monophyletic. For three
1993). The stone hinders germination in Ziziphus of the six, well-supported further groupings, older
(Grice 1996). tribal names were taken up, and three were newly
named. The eleven tribes now recognised can be
characterised morphologically. At the next higher
Rhamnaceae 325
level, the tribes group together into three clades, the mid-Cretaceous of Nebraska (94-96MaB.P.;
for which no morphological characters can be Basinger and Dilcher 1984) has been ascribed to
indicated, possibly because they are relics of for- Rhamnaceae. The earliest known pollen records
merly much more coherent groups (Richardson are from the Oligocene (Muller 1981).
2000a).
ECONOMIC IMPORTANCE. Bark, leaves and fruits
AFFINITIES. Recent angiosperm classifications of several Rhamnus have been used as laxatives
placed Rhamnaceae in Rhamnales, either as the (notably R. frangula and R. catharticus). Diverse
sole family of the order (e.g. Takhtajan 1997), or Old World species of Rhamnus provide yellow and
together with Elaeagnaceae (Thorne 1992). Celas- green dyes as well as drugs. Several species of
trales, Urticales, and Euphorbiales had often been Karwinskia are highly toxic (Waksman et al. 1989;
considered as closely related groups. Analyses of Lux et al. 1998) and of potential use in human
DNA sequences, including those by Soltis et al. medicine. Maesopsis eminii is widely planted in
Africa and Asia for its timber, which is used in
(1997, 2000), Thulin etal. (1998), Richardson et al.
(2000b), and Savolainen, Chase et al. (2000), house and boat building; timber of Ziziphus,
support the placement of Rhamnaceae together Reynosia, Krugiodendron, Hovenia, Colubrina
with their closest relatives, Barbeyaceae and species are used for construction, fine furniture,
Dirachmaceae, in Rosales sensu APG II (2003), as carving, lathework and music instruments. Many
part of the eurosid I assemblage. Ziziphus have edible fruits; among them Z. jujuba
(Chinese jujube) and Z. mauritiana (Indian
DISTRIBUTION AND HABITATS. The family is of jujube) are cultivated at a commercial scale.
worldwide distribution, with habitat preferences Hovenia dulcis is also grown for the edible inflo-
ranging from tropical rain forest to moderately rescence stalks. Species of Ceanothus, Colletia,
arid areas and from the sea level to the treeline. Hovenia, Noltea, Paliurus, Phylica, Pomaderris
A basic preference for tropical and subtropical and Rhamnus are cultivated as ornamentals. Intro-
regions can be recognized. Many members of duced species may be invasive: Ziziphus mauri-
the family are genuine forest species, but several tiana in Australia (Grice 1996, 1998); Rhamnus
taxa occur in extratropical, xerophytic, open cathartica in Canada (Malicky et al. 1970); R. fran-
vegetation. gula in the U.S.A. (Possessky et al. 2000); the native
Discaria toumatou may turn invasive in New
SYMBIONTS AND PARASITES. Root nodules Zealand (Bellingham 1998).
forming perennial coralloid masses are found
in all genera of Colletieae and in Ceanothus. The CoNSERVATION. Studies specifically addressing
nodules are interpreted as modified lateral roots conservation are scarce (Hall and Parsons 1987,
and are inhabited by actinomycetes of the genus Discaria; Godt et al. 1997, Ziziphus). Almost 30
Frankia. Fixation of atmospheric nitrogen takes genera include endangered or vulnerable species
place in particular structures of the microsym- (World Conservation Monitoring Centre 1996). A
biont (Baker and Schwintzer 1990; Cruz-Cisneros few Australian species of Cryptandra, Spyridium
and Valdes 1991; Swensen 1996; Huss-Danell and Trymalium are considered extinct, and species
1997). Endomycorrhizae have been reported from of Gouania, Lasiodiscus and Rhamnus are also
Ceanothus, Colletia, Discaria and Trevoa (Gardner potentially extinct.
1986). Mites have been observed in domatia of
Rhamnus (Lundstrom 1887). Galls are produced CONSPECTUS OF RHAMNACEAE
on male flowers of Rhamnus ludovici-salvatoris by The following tribal classification follows that of
a cecidomyiid fly (Traveset 1999). Insects associ- Richardson et al. (2000a) with the addition of
ated with European Rhamnaceae have been used genera 25 and 50.
in biological control of Rhamnus cathartica in
Canada (Malicky et al. 1970). 1. Tribe Paliureae Reiss. ex End!. (1840).
- Genera 1-3.
2. Tribe Colletieae Reiss. ex Endl. (1840).
PALAEOBOTANY. One of the oldest fossils attrib- - Genera 4-9.
uted to Rhamnaceae are leaf impressions from the 3. Tribe Phyliceae Reiss. ex End!. (1840).
Upper Cretaceous (Hollick 1930), but the assign- - Genera 10-13.
ment of such fossils is often problematic, espe- 4. Tribe Gouanieae Reiss. ex End!. (1840).
- Genera 14-19.
cially before the Oligocene (Johnston 1977). A 5. Tribe Pomaderreae Reiss. ex End!. (1840).
bisexual, obhaplostemonous flower reported for - Genera 20-26.
6. Tribe Rhamneae Hook. f. (1862). 14. Filaments distally geniculate; nectary inconspicuous, ad-
- Genera 27-39. pressed to lower floral tube; floral tube persistent in fruit
7. Tribe Maesopsideae A. Weberb. (1895). 6.Adolphia
Genus 40. - Filaments curved distally; nectary forming a conspicuous
8. Tribe Ventilagineae Hook. f. (1862). disc; floral tube not persistent in fruit 15
- Genera 41-42. 15. Bases of opposite leaves united forming a line; stomata
9. Tribe Ampelozizipheae J.E. Richardson (2000). anomocytic; disc ring-like, encircling the ovary at bottom
- Genus 43. of floral tube 9. Discaria
10. Tribe Doerpfeldieae J.E. Richardson (2000). - Bases of opposite leaves not united; stomata paracytic; disc
- Genus 44. forming a revolute laminar projection of the floral tube
11. Tribe Bathiorhamneae J.E. Richardson (2000). located above ovary level 8. Colletia
- Genus 45. 16. Fruit fleshy 17
- Fruit dry, dehiscent 34
17. Leaf venation pinnate 18
The relationships of seven additional genera - Leaf venation palmate 31
(genera 46-52) are uncertain; therefore, these taxa 18. Ovary 1-locular; stigma mushroom-like. Maesopsideae
are not included in any of the above tribes. Generic 40. Maesopsis
boundaries within tribe Pomaderreae are cur- - Ovary 2- or 4-locular; stigma not as above. Rhamneae
rently under study and new genera will be proba- 19
19. Fruit a fleshy, one-stoned drupe 20
bly proposed in near future. - Fruit with 2-4 free pyrenes 29
20. Leaves alternate 21
KEY TO THE TRIBES AND GENERA - Leaves generally opposite 24
21. Armed shrubs or trees 36. Condalia
1. Fruit with apical appendages, longitudinal wings, or a - Unarmed shrubs or small tree 22
transversal membranous ring 2 22. Leaves leathery, margin entire 30. Berchemiella
- Fruit not as above 9 - Leaves papery, margin serrate 23
2. Shrubs; fruit with a transversal membranous ring. 23. Endosperm present 31. Rhamnella
Paliureae p.p. 1. Paliurus - Endosperm absent 32. Dallachya
- Climbers or herbs; fruit not as above 3 24. Petals present 25
3. Ovary 3-locular; fruit with longitudinal wings; tendrils - Petals absent 28
present; endosperm present. Gouanieae p.p. 4 25. Leaves never gland-dotted 33. Berchemia
- Ovary 2-locular; fruit with apical appendages; tendrils - Leaves often distinctly gland-dotted, pinnately veined and
absent; endosperm absent. Ventilagineae 8 with strongly parallel tertiaries 26
4. Perennial or annual herbs 18. Crumenaria 26. Ovules 2 per locule 35. Karwinskia
- Erect or climbing shrubs or Hanes 5 - Ovule 1 per locule 27
5. Erect, non-climbing shrubs 6 27. Seeds albuminous; cotyledons flat 37. Auerodendron
- Lianes or climbing shrubs with tendrils 7 - Seeds exalbuminous; cotyledons convex
6. Fruit a schizocarp, separating into 3-4 indehiscent, 34. Rhamnidium
winged, inflated mericarps 16. Reissekia 28. Leaves coriaceous 38. Reynosia
- Fruit longitudinally 2-4-winged, indehiscent - Leaves papery, margin undulate 39. Krugiodendron
19. Pleuranthodes 29. Flowers and fruits almost sessile, inflorescences
7. Fruit an explosive capsule 15. Helinus spike- or panicle-like, medifixed hairs present, disc
- Fruit not explosive 14. Gouania cylindrical 29. Sageretia
8. Fruit a samara 42. Ventilago - Characters not as above 30
- Fruit a capsule 41. Smythea 30. Petals emarginate or notched; flowers often unisexual; if
9. Infructescence axis succulent; disc usually hairy. Paliureae plants armed, seeds furrowed 27. Rhamnus
p.p. 3. Hovenia - Petals deeply obcordate or bilobed; flowers always bisex-
- Infructescence axis not succulent; disc not hairy 10 ual; plants mostly armed, but then seeds not furrowed
10. Trees or shrubs, usually armed; leaves decussate; roots with 28. Scutia
actinorhizal nodules. Colletieae 11 31. Ovary 3-locular 32
- Combination of characters not as above 16 - Ovary 2- or 4-locular 33
11. Inflorescences with a terminal flower; floral tube pubes- 32. Climber; endosperm absent. Ampelozizipheae
cent inside; fruit indehiscent or slowly splitting into 43. Ampeloziziphus
indehiscent endocarpids; arils remaining enclosed in the - Tree; endosperm present. Bathiorhamneae
endocarpids 12 45. Bathiorhamnus
- Inflorescences without terminal flowers; floral tube 33. Disc adnate to ovary only; endosperm present.
glabrous inside; fruit a explosive capsule; aril detached Doerpfeldieae 44. Doerpfeldia
from endocarpid and seed at dehiscence 13 - Disc adnate to ovary and floral tube; endosperm absent.
12. Anthers 4-locular; stigma exserted; fruit a papery rostrate Paliureae p.p. 2. Ziziphus
nut 4. Trevoa 34. Plant stellate-hairy. Pomaderreae 35
- Anthers 2-locular; stigma not exserted; fruit a barely fleshy - Plant glabrous or with simple hairs 41
drupe, indehiscent or slowly splitting into indehiscent 35. Flowers in pendent, terminal heads enclosed in an involu-
endocarpids 5. Retanilla cre of large brownish bracts; petals absent; leaves
13. Floral tube caducous at fruit maturity; rim of fruit pedestal opposite 21. Siegfriedia
sinusoid; margin of aril deeply lobed 7. Kentrothamnus - Flowers not as above; leaves alternate 36
- Rim of fruit pedestal smooth, or floral tube persistent at 36. Floral tube tubulate to campanulate 37
fruit maturity; margin of aril more or less smooth 14
Rhamnaceae 327
- Floral tube not as above 38 Evergreen or deciduous, often thorny shrubs or

37. Disc pubescent 25. Cryptandra small to medium-sized trees. Leaves distinctly
- Disc glabrous 24. Stenanthemum
38. Flowers sessile, in heads enclosed by persistent brownish triplinerved. Flowers in axillary cymes; hypan-
bracts 23. Spyridium thium hemispherical to dish-shaped; disc fleshy,
- Flowers pedicellate, solitary or in umbel-like cymes adnate to the intrastaminal surface of the hypan-
forming terminal racemes or panicles 39 thium; ovary half-inferior, 2-3-locular. Fruit dry,
39. Schizocarp often irregularly splitting; mericarps indehis- indehiscent, with a disc- to cup-shaped or hemi-
cent; stamens incurved, enclosed by the hooded petals at
early anthesis 22. Trymalium spherical wing around the apex. Four species in E
- Schizocarp splitting regularly; mericarps dehiscent; Asia, one species (P. spina-christi) in the Mediter-
stamens straight, never enclosed by the petals 40 ranean region.
40. Floral tube absent, sepals wide-spreading at anthesis, ovary
(half-)inferior 20. Pomaderris
- Floral tube present, ovary superior 26. Blackallia 2. Ziziphus Mill.
41. Roots with actinorhizal nodules; petals sometimes blue;
North America 47. Ceanothus Ziziphus Mill., Gard. diet., abr. ed. 4 (1754); Johnston, Am. J.
- Roots without actinorhizal nodules; petals never blue; Bot. 51: 1113-1118 (1964), West Indian spp.
rarely in North America 42 Condaliopsis (A. Weberb.) Suesseng. (1953).
42. Climber with tendrils. Gouanieae p.p. 17. Alvimiantha
- Tree or shrub without tendrils 43 Deciduous or evergreen, erect or straggling, often
43. Seed persisting on receptacle after dehiscence 44
- Seed not persisting on receptacle after dehiscence 45
climbing shrubs or small to medium sized trees.
44. Exocarp thin and leathery; ovary usually 2-locular Leaves distinctly triplinerved rarely pinnately
49. Emmenosperma veined; stipules often spinose. Flowers in axillary,
- Exocarp thick, spongy and crumbly at maturity; ovary corymb-like cymes or terminal or axillary thyrses;
usually 3-locular 46. Alphitonia hypanthium shallow, patelliform to hemispherical;
45. Leaves opposite 46
- Leaves usually alternate 47 petals usually present; disc flat, fleshy, 5-10-lobed;
46. Leaf margin not revolute; stipules interpetiolar ovary superior, 2-3( 4)-locular. Fruit a single-
51. Lasiodiscus stoned, (1)2-3-locular drupe. Almost pantropical,
- Leaf margin revolute; stipules not interpetiolar. Phyliceae about 100 spp., centred in tropical America and SE
p.p. 12. Nesiota Asia.
47. Branches clustered; stipules nearly always absent.
Phyliceae p.p. 48
- Branches not clustered; stipules present 49
48. Stipules absent; disc glabrous 10. Phylica
3. Hovenia Thunb.
- Stipules present; disc pubescent 11. Trichocephalus Hovenia Thunb., Nov. Gen. 7 (1781).
49. Ovary superior 52. Schistocarpaea
- Ovary (half-)inferior SO
50. Plant glabrous; leaf margins always dentate. Phyliceae p.p. Deciduous, unarmed trees or shrubs. Leaves long-
13. Noltea petiolate. Flowers in axillary or terminal, 3- to
- Plant sometimes pubescent; leaf margins not always several-flowered cymes; hypanthium obconical to
dentate 51 dish-shaped; disc fleshy, plane, tomentose, filling
51. Pericarp with outer mealy layer; Western Australia
50. Granitites
the hypanthium; ovary half-inferior, 3-locular.
- Pericarp without mealy layer; pantropical 48. Colubrina Fruit a 3-seeded nut, peduncles and pedicels
becoming fleshy and juicy at fruit maturity. Seven
GENERA OF RHAMNACEAE spp., restricted to E Asia.
1. TRIBE PALIUREAE Reiss. ex Endl. (1840).

2. TRIBE COLLETIEAE Reiss. ex Endl. (1840).
Armed or unarmed trees or shrubs with alternate
leaves. Disc sometimes hairy, adnate to ovary and Usually armed, often nearly aphyllous, erect
hypanthium. Fruit dry with a membranous, trans- shrubs or small trees. Leaves decussate, each sub-
versal ring (Paliurus), a drupe (Ziziphus), or dry tending 2 serial buds. Root nodules actinorhizal.
and indehiscent with fleshy pedicel (Hovenia). Disc either a ring-like, laminar projection of the
Three genera. hypanthium or an inconspicuous nectary adnate
to lower hypanthium. Fruit a papery nut, a barely
fleshy drupe or an explosive capsule. Six genera
1. Paliurus Tourn. ex Mill. (Aagesen 1999).
Paliurus Tourn. ex Mill., Gard. Diet. abr. ed. 4 (1754);
Schirarend & Olabi, Bot. Jahrb. Syst. 116: 333-359 (1994),
rev.
3-locular. Fruit an indehiscent, thin-walled nut

containing 1-2(3) seeds. One species in central
Chile.
5. Retanilla (DC.) Brongn.

Retanilla Brongn., Ann. Sci. Nat. (Paris) I, 10: 364 (1827);
Tortosa, Darwiniana 31: 223-252 (1992), rev.
Leafy to aphyllous, unarmed to spiny shrubs up to

4m tall. Leaves subtending 1-2 buds. Flowers
pubescent, 4-5-merous, terminal or in 3-5(7)-
flowered cymes; hypanthium campanulate to
urceolate, caducous to persistent in fruit, pubes-
cent inside; petals cucullate, partly covering the
pollen-presenting surface of the anthers; filaments
distally geniculate, anthers 2-locular; disc indis-
tinct, nectariferous surface lining the lower part of
hypanthium; ovary half-superior, 2-3(4)-locular.
Fruit somewhat fleshy, indehiscent or splitting into
indehiscent endocarpids. Four species, S South
America.
6. Adolphia Meisner
Adolphia Meisner, Pl. Vase. Gen. tab. 70, comm. SO (1837);
Tortosa, Darwiniana 32: 185-189 (1993), rev.
Subaphyllous shrubs up to 3m tall; the distal bud

in each node forming a spine and the proximal one
a vegetative shoot or a proliferating synftores-
cence. Flowers in 3-7-flowered cymes, pubescent;
hypanthium campanulate, persistent in fruit;
petals present; stamen filaments distally genicu-
late, anthers 4-locular; disc inconspicuous, adnate
to the lower hypanthium; ovary half-inferior, 3-
locular. Fruit an explosive capsule. One species in
Fig. 103. Rhamnaceae-Colletieae. Trevoa quinquenervia. A southern North America.
Branch. B Flower. C Same, vertical section. D Petal and stamen
seen from inside. E Fruit. (Tortosa 1992)
7. Kentrothamnus Suessenguth & Overkott
4. Trevoa Miers Fig. 103 Kentrothamnus Suessenguth & Overkott, Repert. Spec. Nov.
Regni Veg. 50: 326 (1941); Johnston, J. Arnold Arb. 54:
Trevoa Miers, Bot. Misc.1: 158 (1829); Tortosa, Darwiniana 31: 471- 473 (1973), rev.
223-252 (1992), rev.
Subaphyllous shrubs up to 4 m tall; distal bud in
Leafy shrubs up to 3m tall. Flowers pubescent, each node producing a spine, the proximal one a
perfect or staminate, solitary or in 1-7-flowered vegetative shoot or a proliferating synftorescence.
cymes; hypanthium slightly campanulate, persist- Flowers pubescent, 5-merous, in 1-3(4)-flowered
ent in fruit, pubescent inside; petals cucullate, each cymes; hypanthium urceolate-campanulate,
partly covering the pollen-presenting surface of caducous in fruit, leaving a sinusoid scar that
the anthers; stamen filaments distally geniculate, includes the filaments; petals present; stamen fila-
anthers 4-locular; disc inconspicuous, adnate to ments distally geniculate, anthers 4-locular; disc
the lower part of hypanthium; style persistent inconspicuous, adnate to the lower part of
in fruit, stigma exserted. Ovary half-inferior, hypanthium; ovary half-superior, 3-locular. Fruit
Rhamnaceae 329
9. Discaria Hook.
Discaria Hook., Bot. Misc. 1: 156 (1829}; Tortosa, Bol. Soc.
Argent. Bot. 22: 301-335 (1983}, rev.
Prostrate to erect, leafy or subaphyllous, often

spiny shrubs or trees up to 8 m tall. Leaf bases
united, forming a line. Flowers perfect, rarely uni-
sexual, 4-5(6)-merous, in 1-3(6)-flowered cymes;
hypanthium campanulate to urceolate, caducous;
petals present or absent; anthers generally 4-
locular; disc ring-like, more or less lobed, encir-
cling the ovary at bottom of hypanthium; ovary
half-inferior, 3-locular. Fruit an explosive capsule.
Eight species, 5 in South America, 2 in Australia
and 1 in New Zealand.
3. TRIBE PHYLICEAE Reiss. ex Endl. (1840).
Often ericoid shrubs or trees. Leaves alternate or

opposite, usually densely tomentose beneath,
leaf margins usually revolute. Stipules present
or absent. Disc epigynous or adnate to the
hypanthium. Fruit splitting into 3 one-seeded
endocarpids. Four genera.
10. Phylica L.
Phylica L., Spec. Pl.: 195 (1753}; Pillans, J. S. Afr. Bot. 8: 1- 164
(1942}, rev.
Fig. 104. Rhamnaceae- Colletieae. Colletia hystrix. A Fruiting
branch. B Flowering branchlet. C Flower. D Stamen. E Fruit.
Evergreen ericoid shrubs or rarely small trees.
F Seed. Drawn by Nora Mugaburu. (Dimitri 1972}
Leaves alternate; stipules absent. Flowers bisexual
(plants polygamous) in tight head-like thyrses
with extremely pubescent bracts; hypanthium
bell-shaped or cylindrical, surpassing the ovary;
petals usually present; disc present or absent;
an explosive capsule. One species in Bolivia and ovary inferior, 3-locular. Fruit explosive; seeds
northernmost Argentina. arillate. About 150 spp. in Southern Africa to
S Tanzania, Madagascar, southern Antarctic
islands.
8. Colletia ]uss. Fig. 104
Colletia Juss., Gen. Pl. 380 ( 1789}, nom. cons.; Tortosa, Parodi-
ana 5: 279-332 (1989}, rev. 11. Trichocephalus Brongn.
Trichocephalus Brongniart, Ann. Sci. Nat. (Paris) I, 10: 374
Virtually aphyllous shrubs up to 4 m tall. Leaves (1827}.
minute, early caducous; the distal bud in each
node producing a spine or a spiniform twig ram- Evergreen ericoid shrubs. Leaves alternate, stipu-
ified up to 3rd order, the proximal one producing late. Flowers bisexual in tight head-like thyrses
a flowering short-shoot. Flowers 4-5-merous; with extremely pubescent bracts; hypanthium
hypanthium urceolate, caducous; petals present; cylindrical, surpassing the ovary; petals present;
anthers usually 2-locular; disc on a revolute disc adnate to hypanthium, pubescent; ovary infe-
laminar projection of the hypanthium; ovary half- rior, pubescent, 3-locular. Fruit explosive; seeds
inferior, 3(4)-locular. Fruit an explosive capsule. arillate. One species, T. stipularis (L.) Brongn. from
Five species, South America. Southern Africa.
12. Nesiota Hook. f. Evergreen woody climbers; tendrils coiled, axil-

lary. Flowers in axillary, (1- )3-15-flowered umbel-
Nesiota Hook. f., in Bentham & Hook. f., Gen. 1: 380 (1862).
liform cymes; hypanthium broadly campanulate·
disc massive, thickened around and adnate t~
Evergreen small trees. Leaves opposite. Flowers in
ovary, never lob~d. Fruit an explosive capsule. A
few-flowered lateral cymes; hypanthium shallow
tropical genus with five species, three in Eastern
campanulate; petals deeply hooded; stamen
and Southern Africa, one in India, and one in
filaments stout; disc thin, intrastaminal, densely
Madagascar.
pube~cent around ovary at early anthesis; ovary
mfenor, 3-4-locular. One species, N. elliptica
(Roxb.) Hook. f., St. Helena Island. 16. Reissekia Endl.
Reissekia End!., Gen.: 1103 (1840).
13. Noltea Reichb.
Noltea Reichb., Consp.: 145 (1828).
Evergreen shrub~. Flowers in terminal and axillary
c~mes; h~panthmm shallow, cup-shaped, sepals
shghtl~ wmged laterally; petals distinctly hooded,
Evergreen shrubs _or small trees up to 4 m high.
enclosmg the stamens; disc thin, adnate to the
Leav:s alternate, discolorous. Flowers in axillary or
hypanthium, nev~r 5-lobed or appendaged; ovary
termmal: few-flowered cymes in small panicles;
~-4-l~cular. ~chizocarp separating into 3-4
hypanthmm cup-shaped; disc very thin, adnate to
mde~uscent, ":"mg_ed and inflated mericarps. One
t~e lower part of the hypanthium; ovary half-infe-
species, R. smzlacma (Smith) Endl., Brazil.
nor, 3-locular. Endocarpids dehiscent along inner
face. One species, N. africana (L.) Reichb. in South-
ern Africa. 17. Alvimiantha Grey-Wilson Fig. 105
Alvimiantha Grey-Wilson, Bradea 43: 287-290 (1978).
4. TRIBE GOUANIEAE Reiss. ex Endl. (1840).

Evergreen_ climbing shrubs. Leaves triplinerved.
~lowers bisexual or functionally male, in axillary
Unarmed climbers or herbs with tendrils. Leaves
I~florescences. Hypanthium cup-shaped, glabrous;
alternate, . petiolate. Ovary inferior, usually 3-
disc very thin. Schizocarp inflated, unwinged,
locular; disc fleshy, stellate or 5-angled. Fruit a
subglobose, hemimericarps 3, winged, with a cor-
longit~din_ally 3-win~ed schizocarp, a schizocarp
date seed each. One species, A. tricamerata Grey-
re!easmg mflated, wmged (hemi-) mericarps, a
Wilson, Brazil.
wmged nut, or an explosive capsule. Six genera.
18. Crumenaria Mart.

14. Gouania Jacq.
Crumenaria Mart., Nov. Gen. Sp. Pl. 2: 68, pl. 160 (1826).
Gouania Jacq., Select. Stirp. Am. Hist.: 263 (1763).
Annual or perennial herbs. Leaves often more

Evergreen climbing shrubs, or up to 20m high
o~ less reduced to completely absent. Flowers
woody vines; tendrils circinnate. Flowers bisexual
bisex_ual to polygamous, solitary or in axillary or
to polygam~us, in axillary small cymes, usually
termmal cymes; hypanthium obconical to
ag~regated mto ample, leafless panicles; hypan-
campan~late; disc very thin or absent. Schizocarp
thmm shallow obconical to campanulate; petals
dry, mencarps 3, winged, fruit axis persistent. Six
rarely absent; disc broadly annular, with five more
species in tropical Brazil and NE Argentina, one in
or less distinct, antesepalous appendages. Schizo-
Guatemala.
carp__dry, ~ongitudinally 3-winged, separating
septlCidally mto three 2-winged, indehiscent meri-
carps. A pantropical genus of about 50 species in 19. Pleuranthodes A. Weberb.
need of revision.
Pleuranthodes A. Weberb. in Engler & Prantl, Nat. Pflanzenfam.
III, 5: 424 ( 1896).
15. Helinus E. Mey. ex Endl.
Evergreen shrubs. Flowers in axillary, long-pedun-
Helinus E. Mey. ex End!., Gen.: 1102 (1840), nom. cons.
c~late_di~hasia; hypanthium shallow, dish-shaped;
disc distmctly 5-lobed, ovary (2)3-4-locular. Fruit
Rhamnaceae 331
20. Pomaderris Labill.

Pomaderris Labill., Nov. Hoi!. Pl. Sp. 1: 61, pl. 86-87 (1804).
Evergreen, unarmed shrubs or small, often multi-

stemmed trees up to 10m high. Leaves alternate,
the margins distinctly revolute. Flowers in small,
umbelliform cymes forming terminal racemes or
panicles, sometimes head-like clusters, rarely soli-
tary; hypanthium absent; petals present or absent;
disc very thin and annular or absent; ovary half-
inferior. Mericarps generally dehiscing ventrally
through a basal valve. About 40 species, Australia
and New Zealand.
21. Siegfriedia C.A. Gardner

Siegfriedia C.A. Gardner, Enum. Pl. Austral. Occ.: 76 (1931),
nomen; J. Roy. Soc. W. Austral. 19: 85 (1934), descr.
Evergreen, unarmed shrubs. Leaves decussate, the

margins distinctly revolute. Flowers congested in
head-like, pendent clusters, covered by a whorl of
10-12 reddish-brown bracts; hypanthium 0; petals
0; stamens exserted, with about 6mm long fila-
ments; disc thin and delicate; ovary inferior. Mer-
icaps dehiscing ventrally through a basal valve.
One species, S. darwinioides C. A. Gardner, Western
Australia.
22. Trymalium Fenzl

Trymalium Fenzl, Enum. Pl. Hueg.: 20 (1837).
Fig. 105. Rhamnaceae-Gouanieae. Alvimianth~ tricam.erata.
A Habit. B Functional male flower. C Same, vertical sectiOn. D Evergreen, usually unarmed shrubs. Leaves alter-
Hermaphrodite flower, vertical section. E Fruit. F D ehisced
fruit releasing seeds. G Half-fruit, showing position of seed
nate, flat or with revolute margins. Flowers pedi-
with the pericarp valve and the central c olumn in part. Drawn cellate, in raceme-like cymes or narrow panicles;
by C. Grey-Wilson. (Grey-Wilson 1978) hypanthium very short or 0; disc ring-like or dis-
tinctly lobed. Schizo carp often irregularly splitting
in dehiscent or indehiscent mericarps. 14 spp.,
dry, longitudinally 2-4-winged, indehiscent. Two most in Western Australia.
species, Hawaii.
23. Spyridium Fenzl

5. TRIBE POMADERREAE Reiss. ex Endl. {1840). Spyridium Fenzl, Enum. Pl. Hueg.: 24 (1837).
Unarmed or armed shrubs or small trees with stel- Small, evergreen, unarmed, densely pubescent
late hairs. Leaves stipulate, opposite or alternate. shrubs. Leaves alternate, comparatively small,
Disc surrounding the ovary and adnate to the coriaceous. Flowers in sessile heads or clusters,
hypanthium. Ovary 3(2)-locular, superior to infe- surrounded by persistent brown bracts, the heads
rior. Fruit schizocarpic, the mericarps usually often united into compound heads; hypanthium
dehiscent ventrally. Seeds conspicuously arillate. short or 0; disc more or less lobed; ovary inferior.
Seven genera (but ongoing morphological re- Mericarps ventrally dehiscing or indehiscent.
search will most probably result in the addition About 30 spp., Western and Southern Australia.
of five new genera to the tribe; Kevin Thiele, pers.
comm.).
24. Stenanthemum Reissek 27. Rhamnus L.

Stenanthemum Reissek, Linnaea 29: 295 (1858). Rhamnus L., Sp. Pl. 1: 193 (1753).
Frangula Mill. (1754).
Usually unarmed, pubescent shrubs. Leaves entire Oreorhamnus Rid!. (1920).
Oreoherzogia W. Vent (1962).
or toothed, the margins revolute or not. Flowers
aggregated in dense head-like clusters surrounded
Deciduous or evergreen, unarmed or thorny
by bracts and leaves; hypanthium short to long, the
trees or shrubs. Leaves alternate, rarely opposite.
glabrous disc adnate to it or inconspicuous. Meri-
Flowers perfect or unisexual (then usually dioe-
carps enclosed in persistent hypanthium, apically
cious), mostly in sessile or peduncled, axillary,
and ventrally dehiscent; aril 3-lobed. At least 28
umbelliform cymes or fascicles; hypanthium cam-
Australian species.
panulate to cup-like; petals usually +; disc con-
spicuous, adnate to the hypanthium, or apparently
25. Cryptandra Sm. 0; ovary superior. Drupe 2-4-stoned, the stones
indehiscent or ventrally deshiscing. Seeds unfur-
Cryptandra Sm., Trans. Linn. Soc. 4: 217 (1798).
rowed or with dorsal or lateral furrows. About 100
spp., widespread, absent only from Madagascar,
Evergreen, often spinescent shrubs. Leaves very
Australia and Polynesia.
small and narrow, often clustered, lower surface
usually whitish-tomentose. Flowers solitary or
in terminal, clustered spike- or head-like cymes; 28. Scutia Comm. ex Brongn.
hypanthium campanulate to tubular; disc annular,
Scutia Comm. ex Brongn.,Ann. Sci. Nat. I, 10:362 (1827), nom.
often slightly lobed, stellate pubescent; ovary infe- cons.; Johnston, Bull. Torrey Bot. Club 101:64-71 (1974),rev.
rior. Mericarps dehiscing apically and ventrally,
aril succulent, 3-lobed. About 40 spp., extratropi- Evergreen, thorny, rarely scan dent shrubs or small
cal Australia. trees with opposite to subopposite leaves. Flowers
solitary or in sessile, axillary, fasciculate cymes;
26. Blackallia C.A. Gardner hypanthium short, cup-like; disc fleshy, adnate to
the hypanthium; ovary half-inferior. Drupe with
Blackallia C.A. Gardner, J. Roy. Soc. W. Austral. 27: 183 (1942). 2-4 indehiscent pyrenes. Five species, one in
the Old World Tropics and 4 in tropical South
Evergreen, spinescent shrubs with small, alternate, America.
bilobate leaves. Flowers solitary or in fascicles,
long pedicellate; hypanthium tubular; sepals erect;
petals cucullate; disc annular; ovary superior. 29. Sageretia Brongn.
Mericarps dehiscent. One species, B. biloba C.A. Sageretia Brongn., Ann. Sci. Nat. I, 10: 359, pl. 18, fig. 2 (1827).
Gardner, Western Australia. Lamellisepalum Engl. (1897).
Evergreen or deciduous, spinescent or unarmed

6. TRIBE RHAMNEAE Hook. f. (1862). small trees or shrubs, rarely climbing by thorns
representing modified inflorescence axes. Leaves
Usually unarmed trees, shrubs, or climbers. Leaves alternate or subopposite. Flowers minute, sessile,
alternate or opposite, entire or serrate. Petals + or in glomerules grouped in terminal or axillary
0; disc free or adnate to hypanthium, style often spike-like thyrses; hypanthium short, campanulate
persisting in fruit. Ovary superior or half-inferior. to patelliform, or absent; disc thin to fleshy,
Fruit a 1-4-locular drupe, the pyrenes sometimes annular to cupular, sometimes adnate to the
secondarily releasing the seeds. Thirteen genera. hypanthium; ovary superior. Drupe ± fleshy with
Note that generic limits among Reynosia,Rham- 2-3 indehiscent pyrenes. About 35 spp. mostly
nidium, Auerodendron and Karwinskia are prob- from Asia, but also in Africa and America.
lematic and these genera are in need of critical
study.
30. Berchemiella Nakai
Berchemiella Nakai, Bot. Mag. Tokyo 37: 30 (1923).
Rhamnaceae 333
Evergreen shrubs or small trees. Leaves alternate, Evergreen shrubs or small trees. Leaves opposite,
glabrous. Flowers in axillary or terminal fascicles often± distinctly gland-dotted, blackish. Flowers
or cymes; hypanthium dish-shaped, persistent; in small, long-peduncled, umbelliform axillary
disc comparatively thick, filling the hypanthium; cymes; hypanthium hemispherical to obconical,
ovary half-inferior. Drupe 1-locular. Two species, persistent in fruit; disc very thin, adnate to the
China and Japan. hypanthium; ovary superior to half-inferior.
Drupe single-stoned, incompletely 2-locular with
1-2 exalbuminous seeds. About 12 species, Cuba,
31. Rhamnella Miq.
Jamaica, Panama and tropical South America, in
Rhamnella Miq., Ann. Mus. Bot. Lugd.-Batav. 3: 30 (1867). need of revision.
Chaydaia Pit. in Lecomte (1912).
Evergreen shrubs or small trees. Leaves alternate. 35. Karwinskia Zucc.

Flowers in axillary or terminal, contracted, umbel- Karwinskia Zucc., Nov. Stirp. 1: 349, t. 16 (1832).
liform cymes; hypanthium cup- to dish-shaped;
sepals adaxially keeled; disc conspicuous, filling, Evergreen shrubs or small trees. Leaves opposite,
or adnate to, the hypanthium; ovary half-inferior. often gland-dotted. Flowers solitary or in pedun-
Drupe single-stoned; endocarp very hard, 1- or cled, axillary umbel-like cymes; hypanthium
incompletely 2-locular. About 10 species, Central hemispherical to dish-shaped; disc thin, adnate
China, Japan and Korea, in the Himalayas up to to hypanthium; ovary superior, incompletely 2-
3000m alt. locular, with 2 ovules per locule. Drupe single-
stoned, 1-3-seeded. About 15 species, from
32. Dallachya F. Muell. southern North America to northern South
America and the Caribbean.
Dallachya F. Muell., Fragm. 9: 140, 1875.
Small tree. Leaves alternate. Flowers in sessile 36. Condalia Cav.

axillary umbel-like cymes; petals +; hypanthium Condalia Cav., Anal. Hist. Nat. Madrid 1: 39 (1799), nom. cons.;
short, with the disc adnate to it; ovary superior. Johnston, Brittonia 14: 332-368 (1962), rev.
Drupe single-stoned, the cartilaginous stone gen- Microrhamnus A. Gray (1852).
erally 1-locular. One species, D. vitiensis (Seem.) F.
Muell., northern Australia, Papua New Guinea and Evergreen shrubs or small trees often with numer-
the Pacific region. ous divaricate, thorn-tipped short shoots. Leaves
alternate or appearing fasciculate on short shoots.
Flowers solitary or fascicled on short shoots;
33. Berchemia Necker ex DC. hypanthium cup-shaped, persistent in fruit; disc
Berchemia Necker ex DC., Prodr. 2: 22 (1825), nom. cons. thin, adnate to hypanthium; ovary superior. Drupe
Phyllogeiton (A. Weberb.) Herzog (1903). single-stoned, the stone 1-2-locular. About 20 spp.,
North and South America.
Evergreen to deciduous, climbing or erect,
unarmed shrubs or small trees. Leaves mostly
37. Auerodendron Urb.
alternate. Flowers in thyrses composed of few-
flowered, peduncled to sessile, corymb-like cymes; Auerodendron Urb., Symb. Antill. 9: 221 (1924).
hypanthium dish-shaped to hemispherical; disc
fleshy, filling the hypanthium; ovary superior. Evergreen shrubs or trees up to 20 m high. Leaves
Drupe single-stoned, 2-locular. About 20 species, opposite, glabrous,± conspicuously gland-dotted.
most in SE Asia, one in New Caledonia and one in Flowers in sessile or peduncled cymes; hypan-
North America. thium hemispherical; disc comparatively thin,
adnate to the hypanthium; ovary superior. Drupe
single-stoned, 2-locular. Seven species, Greater
34. Rhamnidium Reissek
Antilles, in need of revision.
Rhamnidium Reissek in Mart., Fl. Brasil. 11(1): 94, pl. 31
(1861).
38. Reynosia Griseb.
Reynosia Griseb., Catal. Pl. Cub.: 33 (1866); Urban, Symbol.
Antill. 9: 225-228 (1924).
Evergreen shrubs or small trees. Leaves opposite.

Flowers solitary or in axillary, sessile, subumbel-
late, cymose fascicles; hypanthium short-campan-
ulate to hemispherical; petals + or 0; disc fleshy,
adnate to the hypantium; ovary superior. Drupe
single-stoned, usually 1-locular by abortion of the
second locule; seeds with copious, hard and rumi-
'/,·',\\
nate endosperm. About 15 species, southern North
111
America, Central America and the Caribbean, in
need of revision.
; ~c ' I •,, ; ~
39. Krugiodendron Urb. 0 ..
Krugiodendron Urb., Symb. Antill. 3: 313 (1902).
Evergreen shrubs or small trees. Leaves opposite.

Flowers in axillary, sessile or shortly peduncled,
umbelliform cymes; hypanthium short, broadly
obconical to dish-shaped; petals 0; disc broadly
annular, crenate, fleshy, surrounding the base of
the ovary; ovary superior. Drupe single-stoned,
pericarp thin, fleshy, endocarp bony, 1(2)-locular;
seeds without endosperm. One species, K. ferreum
(Vahl) Urban, West Indies, Southern Florida,
Mexico and parts of Central America, in need of
revision.
7. TRIBE MAESOPSIDEAE A. Weberb. (1895).
Unarmed, large trees. Leaves opposite or alternate, Fig. 106. Rhamnaceae-Maesopsideae. Maesopsis eminii. A
Flowering branch. B Flower. C Petals, dorsal and ventral view.
strongly toothed. Flowers in peduncled, axillary D Stamens, ventral and dorsal view. E Pistil. F Portion offruit-
cymes; hypanthium campanulate; petals sessile, ing branch. G Fruit, vertical section. (Johnston 1972)
deeply hooded; disc thin, adnate to the hypan-
thium; ovary superior, 1-locular, style thick, api-
cally expanded into a mushroom-like stigma. Fruit
41. Smythea Seem.
a 1(2)-seeded drupe. One genus.
Smythea Seem., Bonplandia 9: 255 (1861).
40. Maesopsis Engl. Fig. 106

Shrubs, often climbing. Flowers solitary or in axil-
Maesopsis Engl., Pflanzenw. Ost-Afr. C: 255 (1895); Johnston, lary glomerules, sometimes in spike-like inflores-
Fl. Tr. E. Afr. Rhamnaceae: 36 (1972). cences; hypanthium dish-shaped; disc thin, adnate
to the hypanthium. Capsule compressed, 1-locular,
Characters as for the tribe. One species, M. eminii 1-seeded, dehiscing with 2 valves. About 10 spp.,
Engl., tropical Africa. tropical SE Asia.
8. TRIBE VENTILAGINEAE Hook. f. (1862). 42. Ventilago Gaertn.

Ventilago Gaertn., Fruct. 1: 223, pl. 49, fig. 2 (1788).
Evergreen unarmed climbers or rarely small trees
without tendrils. Leaves alternate. Petals present or Climbing shrubs or strong lianes, rarely small
absent, disc thick, filling the short hypanthium; trees. Flowers solitary or in axillary glomerules,
ovary half-inferior. Fruit an apically-winged sometimes in spike-like inflorescences;
samara or a rostrate capsule; seeds exalbuminous. hypanthium dish-shaped; disc thick, adnate to
Two genera. the hypanthium; petals usually present, cucullate.
Rhamnaceae 335
Fruit a 1-seeded samara with an elongate terminal Characters as for the tribe. Two species,
wing. About 40 spp., Old World tropics. Madagascar.
9. TRIBE AMPELOZIZYPHEAE J.E. Richardson Genera incertae sedis

(2000).
46. Alphitonia Reissek ex Endl.
Unarmed climbers without tendrils. Leaves alter-
Alphitonia Reissek ex End!., Gen. Pl.: 1098 (1840); Braid, Kew
nate, distichous. Flowers in axillary, clustered Bull. 1925: 171-186 (1925), rev.
cymes, or composed of several cymes forming a
panicle; hypanthium shortly turbinate; petals Evergreen, unarmed shrubs or trees, sometimes
cucullate; disc thick, annular; ovary half-inferior. tall. Leaves entire, alternate. Flowers perfect and
Fruit a 3-locular, 3-seeded, stipitate, explosive staminate, in dichasial or trichasial axillary or ter-
capsule; seeds exalbuminous. One genus. minal cymes; hypanthium short; disc compara-
tively thick, filling the hypanthium; ovary inferior.
43. Ampelozizyphus Ducke Fruit a 2-3-seeded capsule. Seeds arillate, tem-
porarily persistent on the receptacle after the fall
Ampelozizyphus Ducke, Arch. Inst. Bioi. Veg. Rio de Janeiro 2: of the pericarp, albuminous. About 15 species,
157 (1935).
Malaysia, West Pacific islands, New Caledonia and
Australia.
Characters as for the tribe. One species, A.
amazonicus Ducke, Amazonian Brazil.
47. Ceanothus L.
Ceanothus L., Sp. Pl.: 195 (1753); van Rensselaer & McMinn,
10. TRIBE DOERPFELDIEAE J.E. Richardson Ceanothus. Sta. Barbara Bot. Gard. (1942), rev.
(2000).
Deciduous or evergreen shrubs or rarely small
Evergreen, unarmed trees. Leaves alternate, entire trees, sometimes thorny, mostly with nitrogen-
or emarginate. Flowers solitary or in axillary fixing actinorhizal root nodules. Leaves alternate
fascicles; hypanthium very short and shallow; or opposite. Flowers white, pink, or blue, borne
petals 0; disc thin, adnate to ovary; ovary superior, in terminal or axillary, raceme- or corymb-like
pseudobilocular, style bifid. Fruit a single-stoned, thyrses composed of few-flowered umbelliform
unequally 2-locular drupe with a single seed in the cymes; hypanthium shallow, cupulate to
larger locule; seed albuminous. One genus. hemispherical; petals +; disc annular, ovary
half-inferior, 3-4-locular. Capsule 3-4-seeded, ex-
44. Doerpfeldia Urban plosive; seeds albuminous, sometimes conspicu-
ously arillate. 55 species, North and Central
Doerpfeldia Urban, Symb. Ant. 9: 218 (1924). America.
Characters as for the tribe. One species, D. cuben-
sis Urban, Cuba. 48. Colubrina Rich. ex Brongn. Fig. 107
Colubrina Rich. ex Brongn., Mem. Pam. Rham. 61: (1826), nom.
cons.; Johnston, Brittonia 23: 2-53 (1971), rev.
11. TRIBE BATHIORHAMNEAE J.E. Richardson Hybosperma Urban (1899).
(2000).
Evergreen or deciduous, unarmed or spinescent,
Evergreen, unarmed trees. Leaves triplinerved, rarely scandent trees or shrubs. Leaves alternate,
entire to toothed, alternate. Flowers in axillary fas- rarely opposite, often glandular. Flowers in sessile
cicles; disc thick; ovary superior, 3-locular. Fruit or shortly peduncled, axillary cymes or small
splitting into 3 indehiscent mericarps; seeds exal- thyrses or fascicles, rarely solitary; hypanthium
buminous. One genus. cup- to dish-shaped; petals present; disc fleshy,
almost filling the hypanthium; ovary half-inferior,
3( 4)-locular. Capsule 3-seeded, explosively or
45. Bathiorhamnus Capuron
slowly dehiscent with the outer pericarp irregu-
Bathiorhamnus Capuron, Adansonia II, 6: 121 (1966). larly breaking away. Seeds albuminous, sometimes
leafy clusters; hypanthium short; disc prominent,

adnate to the hypanthium; petals +; ovary inferior,
3-locular. Fruit beaked, apparently a 3-seeded
capsule with a mealy layer in the outer pericarp.
Seeds arillate. One species, G. intangendus (F.
Muell. ex F. Muell.) Rye, Western Australia; related
to Alphitonia (Fay et al. 2001).
51. Lasiodiscus Hook. f.

Lasiodiscus Hook. f. in Bentham & Hook. f., Gen. Pl. 1: 381
(1862); Figueiredo, Kew Bull. 50: 495-526 (1995), rev.
Evergreen, unarmed, often climbing shrubs or

small trees. Leaves opposite; stipules comparatively
large and interpetiolar. Flowers± densely ferrugi-
neous pubescent, congested in long-pedunculated,
axillary, divaricate cymes or pseudo-umbels;
hypanthium broadly campanulate; disc massive,
fleshy, broadly annular; ovary inferior or half-infe-
rior, 3-locular. Fruit a velvety, elastically dehiscent,
3-seeded capsule; seeds exalbuminous. 12 spp.,
tropical Africa and Madagascar.
52. Schistocarpaea F. Muell.

Schistocarpaea F. Muell., Victor. Nat. 7: 182 (1891).
Evergreen, unarmed trees with alternate leaves.

Fig. 107. Rhamnaceae. Colubrina asiatica. A Flowering and
fruiting branch. B Flower. C Same, vertical section. D Trans- Flowers cream or pinkish, in loose panicles of
verse section of ovary. E Portion of fruiting branch. F Capsule. 3-many-flowered cymes in the upper axils; hypan-
G Seed. (Johnston 1972) thium cup-shaped, persistent in fruit; disc con-
spicuous, forming a narrow rim at the base of the
conspicuously arillate. About 33 species, Old World ovary; ovary superior, 3-locular. Fruit a subglo-
and New World tropics. bose capsule; seeds exalbuminous. One species, S.
johnsonii F. Muell., Australia.
49. Emmenosperma F. Muell.
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Roridulaceae 339
Roridulaceae
J.G. CONRAN
Roridulaceae Engl. & Gilg. in Engl., Syllabus ed. 9 & 10: 226 VEGETATIVE ANATOMY. Calcium oxalate druses
(1924), nom. cons. are present in the epidermal cells near the major
veins (Bruce 1907). Tannin cells occur in the
Perennial insect-trapping evergreen woody shrubs endosperm and seed integuments (Dahlgren and
from a taproot with limited lateral root develop- van Wyk 1988).
ment; stems perennial, erect and few-branched. Primary stems have a ring of fibres connecting
Leaves alternate, crowded apically, linear to taper- the vascular bundles (Solereder 1908). The wood
ing, exstipulate, margins entire or with short lateral anatomy of the family was described by Carlquist
linear lobes, with insect-trapping stalked glands. (1976) who observed that there was distinct
Flowers in terminal, few-flowered botryoids, her- secondary thickening in both species and clear
maphroditic, medium-sized and showy; calyx and growth rings in R. dentata. Vessels occur scattered
corolla 5-merous, well developed; perianth actin- and singly in the roots and stems, with spiral
omorphic, free, rotate, bright purple, reddish pink thickening, scalariform perforation plates and
or white; petals imbricate in bud; stamens 5, oppo- bordered bars in both species, as well as vessel
site the sepals; anthers hypogynous; filaments free, plates with a meshwork of rhomboid perforations
filiform; anthers sub-basifixed, 2-thecate, 4- in R. dentata. Xylem parenchyma rays are uniseri-
sporangiate, introrse, incurved in bud, subtended ate with uniquely upright ray cells. The tracheids
by a basal swelling containing a nectariferous have bordered pits (Carlquist 1976), not simple,
cavity, irritable and swinging up when touched at as reported by Solereder (1908). Behnke (1991)
anthesis to become erect, dehiscing by four short reported Ss-type phloem sieve-tube plastids.
apical pores or slits; gynoecium of three united The leaves show brochidodromous venation
carpels; ovary superior, 3-locular with axile pla- with a clear marginal vein. The cuticle is thin and
centation; ovules anatropous, unitegmic, solitary the anomocytic stomata are present on both sur-
or 2-4 per locule; style terminal, tapering with a faces and oriented along the leaf axis. The non-
small capitate stigma or expanding terminally with vascularised tentacles vary from uni- to 5-seriate
an obconical stigma, papillate, erect. Fruit a smooth stalks with six-celled to multicellular glandular
loculicidal, 3-valved cartilaginous capsule. Seeds heads. The larger hairs have a central head column
exotestal, small, ellipsoid and smoothly reticulate three cells wide onto which the otherwise free
or angular-trilete and prominently warty or resin-secreting cells are basally attached (Fenner
honeycomb-sculptured, dark reddish-brown; 1904; Dahlgren and van Wyk 1988). The mesophyll
endosperm copious; embryo linear. is uniformly spongy, with prominent abaxial
One genus, Roridula, with two species endemic lacunae. The vascular bundles are surrounded by
to the Cape Province of South Africa. Both species sclerenchyma (Solereder 1908).
are occasionally cultivated as ornamental
curiosities. INFLORESCENCE STRUCTURE. Although the inflo-
rescence in Roridulaceae superficially resembles a
VEGETATIVE MORPHOLOGY (Fig. 108). There is a raceme, it is determinate with a terminal flower,
well-developed taproot with relatively limited and is thus derived-paniculate (botryoid)
lateral root development (Bruce 1907). The few- (Dahlgren and van Wyk 1988).
branched woody stems show a more or less divar-
icate branching pattern with the leaves restricted FLORAL MORPHOLOGY. The rotate actinomor-
to clusters near the branch apices. The leaves phic corolla consists of five broadly obovate,
are linear and tapering. They are simple in R. entire, glabrous and free petals which are im-
gorgonias and finely laterally linear-lobed in R. bricate in bud. The sepals have three non-
dentata. Both species have densely glandular anastomosing vascular traces (Chrtek et al. 1989).
leaves. The anthers are strongly outwardly and ventrally
340 J.G. Conran
endosperm nucleus. Fertilisation is porogamous,

there are micropylar haustoria and endosperm
formation is cellular. Embryogeny conforms to the
Solanad type.
The mature endosperm is tanniniferous and the
hypostase and testa walls become thickened. The
outer epidermal cells of the testa are enlarged with
strongly thickened inner tangential and radial
walls (Vani-Hardev 1972).
POLLEN MORPHOLOGY. Pollen grains differ

markedly between the two Roridula species. R.
dentata has isopolar, actinomorphic tricolporate
prolate-spheroidal pollen 36.5 J.lm (polar) by
c. 36 J.lm (equatorial). The exine has a spinulifer-
ous perforated tectum and infratectal baculae. Ora
are indistinct and the colpi are marginally granu-
lar and centrally warty. R. gorgonias pollen is
apolar, asymmetrical, spheroidal, c. 50 J.lm in
diameter without obvious apertures. The exine is
insular with bacula-supported polygonal to
rounded segments (Erdtman 1952; Dahlgren and
van Wyk 1988).
KARYOLOGY. Chromosomes in Roridulaceae were

reported by Kress (1970) and Peng and Goldblatt
(1983) as 2n = 12 (n = 6).
PoLLINATION. Hemiptera of the genus Pameridia

(Miridae), one species for each Roridula species,
pierce and feed from the anther connective nec-
taries. The piercing causes the anthers to spring
up through 180°, dusting the insects with pollen
(Marloth 1925). Plants in cultivation are also
Fig. 108. Roridulaceae. Roridula gorgonias. A Floriferous visited by hover flies and are self-fertile (Conran,
branch. B Part of leaf. C Flower. D Same, vertical section. E pers. obs.)
Stamens and pistil. F Stamen. G Young fruits. H Seeds.
(Dahlgren and van Wyk 1988) FRUIT AND SEED. The fruit is a smooth loculici-
dal, 3-valved cartilaginous capsule. Seeds are
recurved in bud but, when disturbed at anthesis, small, ellipsoid and smoothly reticulate in R.
they swing upwards through 180° and dehisce by dentata, and angular-trilete and prominently
short, pore-like apical slits (Fig. lOSE, F). warty to honeycomb-sculptured in R. gorgonias,
dark reddish-brown; endosperm is copious; the
EMBRYOLOGY. Anther wall formation is of the embryo is linear. The seeds of both species become
basic type. There are no fibrous thickenings of the sticky and mucilaginous when wet.
endothecium and the glandular tapetal nuclei
exhibit both fusion and division. One of the EcoLOGY. The plants grow in areas of permanent
middle layers if crushed at maturity and the other moisture, although R. dentata can apparently
is tanniniferous. Microsporogenesis is simultane- inhabit slightly drier areas than R. gorgonias
ous and the mature pollen grains are binucleate. (Carlquist 1976). The rootstock is fire-resistant
The anatropous ovule is unitegmic and there and the roots are considered to be mycorrhizal
is a well-developed endothelium. The chalaza! (Marloth 1910), although this fungal association
megaspore follows Polygonum-type development. was questioned by Obermeyer (1970). The glan-
The polar nuclei fuse prior to fertilisation and the dular tentacles secrete a balsam-like resin which,
antipodals persist until the first division of the although it appears to trap insects, was not diges-
Roridulaceae 341
tive in function (Marloth 1903, 1925); however, A single genus:

these trapped insects are preyed upon by
Hemiptera of the genus Pameridia (Miridae: Het-
Roridula L. Fig. 108
eroptera), P. marlothii on R. dentata and P. roridu-
lae on R. gorgonias, and by a crab spider, Synema, Roridula L., Gen. ed. 6: 567 (1764).
all of which can move easily over the sticky leaves.
Ellis and Midgley (1996) demonstrated that the Description as for family.
excreta from these predators are absorbed by the
leaves of Roridula, resulting in carnivory by mutu-
alism. In addition, the foliage of Roridula is highly Selected Bibliography
UV-adsorptive (Midgley and Stock 1998), a feature
shared with other flypaper carnivorous plants Albert, V.A., Williams, S.E., Chase, M.W. 1992. Carnivorous
such as the Droseraceae, and which is thought to plants: phylogeny and structural evolution. Science 257:
make the plants attractive to potential insect prey 1491-1495.
(Joel et al. 1985). references.
Behnke, H.-D. 1991. See general references.
PHYTOCHEMISTRY. lridoids of unknown struc- Bentham, G., Hooker, J.D. 1865. Genera Plantarum vol. I.
ture have been reported for Roridulaceae by London: L. Reeve.
Jensen et al. (1975). Naphthoquinones are absent Bruce, A.N. 1907. On the distribution of the tentacles of
Roridula. Notes Roy. Bot. Gard. Edinburgh 17: 83-98.
(Zenk et al. 1969). Carlquist, S. 1976. Wood anatomy of Roridulaceae: ecological
and phylogenetic implications. Am. J. Bot. 63: 1003-1008.
AFFINITIES. Roridula has been associated vari- Chrtek, J., Slavikova, Z., Studnicka, M. 1989. Beitrag zur Leit-
ously with the Droseraceae (e.g. Bentham and biindelanordnung in den Kronblattern von ausgewahlten
Arten der fleischfressenden Pflanzen. Preslia 61: 107-124.
Hooker 1865), Ochnaceae (Planchon 1848),
Conran, J.G., Dowd, J.W. 1994. The phylogenetic relationships
Clethraceae (Hallier 1912), mostly on differently of the Byblis-Roridula (Byblidaceae-Roridulaceae) complex
emphasised features of gross morphology. More inferred from 18S rRNA partial sequences. Plant Syst. Evol.
recently, the genus has been included in the Bybl- 188: 73-86.
idaceae (e.g. Hutchinson 1959; Cronquist 1981) or Cronquist, A. 1981. See general references.
Dahlgren, R.M. T., van Wyk, A.E. 1988. Structures and relation-
treated as a monogeneric family (e.g. Takhtajan ships of families endemic to, or centered in Southern Africa.
1987; Dahlgren and van Wyk 1988; Juniper et al. Monogr. Syst. Bot. Missouri Bot. Gard. 25: 1-94.
1989), generally with affinities near the Ericales. Diels, L. 1930. Roridulaceae. In: Engler, A., Prantl, K. Die natiir-
Molecular studies of its position, using chloroplast lichen Pflanzenfamilien ed. 2, 18a. Leipzig: W. Engelmann,
rbcL (Albert et al. 1992) and mitochondrial 18 S pp. 346-348.
Ellis, A.G., Midgley, J.J. 1996. A new plant-animal mutualism
RNA sequences (Conran and Dowd 1994), con- involving a plant with sticky leaves and a resident
firmed that it was unrelated to the Byblidaceae. Hemipteran insect. Oecologia 106: 478-481.
In particular, Albert et al. (1992) placed it in the Erdtman, G. 1952. See general references.
Ericales as sister to the Northern Hemisphere car- Fenner, C.A. 1904. Beitrage zur Kenntnis der Anatomie,
nivorous pitcher plant family Sarraceniaceae. In Entwicklungsgeschichte und Biologie der Laubblatter und
Drusen einiger Insectivoren. Flora 93: 335-434.
the recent classification of the flowering plants Hallier, H. 1912. L' origine et le systeme phyletique des
based on combined multigene studies, Roridu- angiospermes. Arch. Neerl. Sci. Exactes Nat., III. B 1:
laceae are retained as a separate family in the Ed- 146-234.
cales (APG 1998). Hutchinson, J. 1959. The families of flowering plants. Vol. 2.
Monocotyledons, 2nd edn. Oxford: Clarendon Press.
Jensen, S.R., Nielsen, B.J., Dahlgren, R. 1975. Iridoid com-
DISTRIBUTION AND HABITATS. The family is pounds, their occurrence and systematic importance in the
endemic to the Cape Province region of South angiosperms. Bot. Not. 128: 148-180.
Africa in mountainous regions, with R. gorgonias Joel, D.M., Juniper, B.E., Dafni, A. 1985. UV patterns in the traps
found between Gordon's Bay and Hermanus to of carnivorous plants. New Phytol. 101: 585-594.
Juniper, B.E., Robins, R.J., Joel, D.M. 1989. Carnivorous plants.
the Genadenal region. R. dentata occurs in the
London: Academic Press.
Tulbagh and Ceres districts through to the Cedar- Kress, A. 1970. Zytotaxonomische Untersuchungen an einigen
burg mountains. The plants grow on sandstone Insektenfangern (Droseraceae, Byblidaceae, Cephalotaceae,
slopes in areas of seepage or near streams Roridulaceae, Sarraceniaceae ). Ber. Deutsch Bot. Ges. 83:
(Carlquist 1976). 55-62.
Lloyd, EM. 1934. Is Roridula a carnivorous plant? Can. J. Res.
10: 780-786.
ECONOMIC IMPORTANCE. Roridulaceae are of Lloyd, EM. 1942. The carnivorous plants, 2nd edn. Waltham,
interest to specialist carnivorous plant collectors. Mass.: Chronica Botanica Co.
342 J.G. Conran
Marloth, R. 1903. Some recent observations on the biology of Planchon, J.E. 1848. Sur Ia famille des Droseracees. Ann. Sci.
Roridula. Ann. Bot. 17: 151-157. Nat. Bot. III, 9: 79-99.
Marloth, R. 1910. Further observations on the biology of Solereder, H. 1908. Systematic anatomy of the Dicotyledons.
Roridula. Trans. Roy. Soc. S. Afr. 2: 59-61. Vol. 1, Introduction, Polypetalae and Gamopetalae. Oxford:
Marloth, R. 1925. Roridulaceae. In: The Flora of South Africa Clarendon Press.
vol. 2, part I. Cape Town: Darter Bros., pp. 26-30. Takhtajan, A.L. 1987. Systema Magnoliofitorum. Leningrad:
Midgley, J.J., Stock, W.D. 1998. Natural abundance of Delta Nauka (in Russian).
N-15 confirms insectivorous habit of Roridula gorgonias, Vani-Hardev, 1972. Systematic embryology of Roridula
despite it having no proteolytic enzymes. Ann. Bot. 82: gorgonias Planch. Beitr. Bioi. Pflanzen. 48: 339-351.
387-388. Zenk, M.H., Fiirbringer, M., Steglich, W. 1969. Occurrence and
Obermeyer, A.A. 1970. Roridulaceae. In: Codd, L.E., De Winter, distribution of 7-methyljuglone and plumbagin in the
B., Killick, D.J.B., Rycroft, H.B. (eds.) Flora of Southern Droseraceae. Phytochemistry 8: 2199-2200.
Africa, vol. 13. Pretoria: National Botanical Institute, pp.
201-204.
Peng, C.-I., Goldblatt, P. 1983. Confirmation of the chromo-
some number in Cephalotaceae and Roridulaceae. Ann.
Missouri Bot. Gard. 70: 197-198.
Rosaceae 343
Rosaceae
C. KALKMAN 1
Rosaceae Juss., Gen. Pl.: 334 (1789), nom. cons. !-several (often 2) per carpel, anatropous, ascend-
Spiraeaceae Maxim. (1879). mg or pendulous. Fruits various, fleshy or dry
Amygdalaceae (Juss.) D. Don (1825), nom. cons.
dehiscent or not; seeds 1-several, testa usuall;
Malaceae Small ex Britton & Small (1903), nom. cons.
firm, endosperm 0 or a thin layer, cotyledons fleshy
or flat.
Woody or herbaceous. Leaves usually alternate,
As recognized here, a moderately large, almost
sometimes distichous, rarely opposite, simple or
cosmopolitan family with 85 genera and c. 2000
compound; stipules on the twig or on the base of
sexual species. ~part from those, there are a large
the petiole, free or adnate to the petiole, rarely 0.
number of obhgately or facultatively apomictic
Inflorescences various, usually terminal, usually
(compound) racemes. Flowers actinomorphic, 'microspecies' in several genera.
mostly (4)5-merous, mostly bisexual, rarely uni-
sexual and then the plants monoecious or dioe- ~EGETATIVE ~ORPHOLOGY. Variation in vegeta-
~lve char~cters 1s ~ery great in this family and there
cious; hypanthium usually well-developed (not
evident in some staminate flowers), from saucer- 1s not a smgle trmt, not even the presence of stip-
shaped to tubular or campanulate, the epicalyx, ules, that is characteristic of the family as a whole.
Woody Rosaceae are mostly shrubs or small
sepals, petals, and stamens inserted on its rim its
inside usually lined by nectariferous tissue; disk trees, not more than 12m high. Small and low-
growing shrubs with creeping woody branches are
sometimes distinct, intrastaminal; epicalyx + in
present in several genera, while large forest trees
some genera; sepals free; petals free, from large
ove~ 25-30 m high are rare and occur in few genera
and showy to small and not or hardly distinct from
(Erwbotrya, Sorbus, Prunus). Woody species may
sepals, rarely 0; stamens few to numerous, often
be evergreen or leaf-shedding. Thorns as cauline
their number distinctly related to the number of
perianth parts; filaments free; anthers bilocular, m:tamorphoses are sometimes developed, and
pnckles (emergences) are common in some
dehiscing longitudinally; carpels 1-many, free or
genera (Rosa, Rubus). Buds are normally protected
variously connate with each other and/or adnate
by bud-scales in woody plants of climates with an
~o th~ hypanthi~m, forming l or more superior to
mfer.wr ovary(1es); stylodia (in monocarpellate unfavourable season. Woody genera not rarely
show a differentiation in long and short shoots.
ovanes styles) +,these sometimes (some Maleae)
Herb~ceous members of the family are mostly
fused into a common, branched style; ovules
perenmal and when growing in temperate climates
they perennate by means of a subterranean verti-
1 In his contribution to this series the late Dr. Kalkman
cal rhizome or horizontal rootstock. Annuals are
(tl9 January 1998) had included Quillaja in, and excluded extremely rare (Alchemilla, Potentilla). Herba-
Lyonothamnus from Rosaceae, although he had been aware of
the results of the rbcL analysis by Morgan et al. (1994) that
ceousness may with some confidence be seen as an
were in favour of the contrary. I have found no reason for apomorphic state.
rejecting these molecular findings and consequently have Many Rosaceae have simple leaves but in more
chan?ed Dr. Kalkman's manuscript and included Lyonotham- t~an 30 genera compound leaves, mostly of a
nus :n Rosaceae, altho~gh its position within the family pmnate type, are the rule. That raises the question
remams unclear for the time being, whereas Quillaja has been
removed from it and will be treated as a separate family in a
of what state is primitive in the family. Wolfe and
further volume. After completion of Dr. Kalkman's manu- Wehr (1988), in relation to their study of some
script, some important papers dealing with the morphology fossil North American genera, made a distinction
and II_IOl~cular systematics .of Rosaceae have appeared, which between true, pinnately compound leaves "formed
may md1cate that the mamtenance of tribe Exocordeae is by discrete laminar units that dehisce" (p.l78), and
probably not justified. I have inserted references to these
papers into the bibliography but in general have left )aracompound" leaves that look pinnate but are
untouched Dr. Kalkman's argument, which to a large degree is m f~ct deeply dissected simple leaves, detaching
supported by recent findings. (or, m Wolfe and Wehr's terminology, "dehiscing")
344 C. Kalkman
as a whole. Their contention is that in Rosaceae the Woon ANATOMY. A comprehensive account was
trend is from simple and pinnatifid to pinnatisect published by Zhang {1992), based on a survey of
leaves to paracompound leaves and probably (cer- about 80% of the shrubby or arborescent genera.
tainly, I would say) also to truly compound leaves. The following is a short summary of his family
As a descriptive term, 'paracompound' may be description, with the rare exceptions omitted.
useful and in some groups of the family the dis- Growth rings distinct except in tropical genera.
tinction may even have some morphological and Vessels vary substantially in number per sq. mm,
phylogenetic value but at the moment there is too in degree of grouping and pattern, as well as in
little evidence for making the choice between phy- diameter and length of the elements. In the tem-
logenesis going in either of the two directions - perate and shrubby taxa, vessel diameter is typi-
from simple to compound, or vice versa (or both!). cally small and vessel frequency is low; the reverse
That the majority of the woody genera have simple holds true for the tropical trees. In most maloids,
leaves, whereas the herbaceous genera have (with vessels are typically solitary (over 80%) while
few exceptions) compound leaves may probably be Pruneae have a high percentage (over 60%) of the
seen as an indication that the simple leaf is the ple- vessels in radial multiples. The other groups show
siomorphic state in the family. Troll's typological a variation range in between these two extremes.
approach of the compound leaf types in Rosaceae Perforations mostly exclusively simple, but rare
(Troll1935) does not give the solution but stresses scalariform or reticulate plates occur in some taxa.
the diversity. Intervessel pits alternate, vessel-ray and vessel-
Leaves are normally alternate, only in three parenchyma pits usually half-bordered. Vessel
genera (Coleogyne, Rhodotypos and Lyonotham- walls with or without helical thickenings. Tyloses
nus) are they opposite. In many woody genera absent. Ground tissue mostly of fibre-tracheids,
there are erect orthotropic shoots with spirally rarely mixed with libriform fibres in some
arranged leaves, and horizontally growing pla- Pruneae, non-septate. Parenchyma scanty to abun-
giotropic shoots, with distichous leaves. The dor- dant, differently patterned but mainly diffuse
siventral orientation of the plagiotropic shoots and scanty paratracheal. Rays 1-16-seriate, very
may already be visible in bud, as demonstrated variable within and between taxonomic groups.
by Charlton ( 1993) for Prunus laurocerasus (his Crystals abundant to absent, when present mostly
"rotated lamina syndrome"). prismatic but in the Pruneae druses may also be
Especially in the malo ids, the nervation type has present.
some taxonomic value at the generic or subgeneric Zhang (1992) recognized 12 phenetic groups,
level. Nervation may there be craspedodromous some of which clearly reflect taxonomic groupings
(secondary nerves going straight to the margin based on morphological characters. His groups
and terminating there) or camptodromous (sec- VI and VII are exclusively composed of maloid
ondaries not reaching the margin). The lowermost genera (with only Cercocarpus as an exception in
secondaries are often stout and pedately divided. group VII), and groups XI and XII consist only of
Extrafloral glands are present on the leaves of (split-)genera that in the present treatment have
some woody genera: in Prunus on the marginal been taken together as Prunus s.l. However, also
teeth, on the petiole, or on the blades, in several the other, more mixed groups are not without
maloids (Aronia and others) on the upper midrib. merit. In Zhang's words (p. 79): "... with few
Water pores and/or guttation have been demon- exceptions all woody tribes, if not characterised by
strated in a number of common, Northern Hemi- a single wood type, encompass few closely adja-
sphere Rosaceae. Well-developed hydathodes with cent wood anatomical groups".
their anatomical specialization have been reported
and described for only few species belonging INFLORESCENCES. According to Troll (1964, and
to Sanguisorbeae (Agrimonia, Sanguisorba), especially 1969) the basic synflorescence in the
Alchemilla, Geum, and Physocarpus (see Lersten Rosaceae is monotelic (=with a terminal flower).
and Curtis 1982). Rich-flowered inflorescences may have the appear-
Stipules are absent in one tribe, viz. Spiraeeae, ance of a panicle or a corymb. However, they are
and exceptionally in single species of normally better described as thyrsoids since their partial
stipulate genera. Often stipules are small and inflorescences are cymose. Thyrsoids are found in
caducous but in several groups they are relatively many Maleae and Crataegeae but also, e.g., in
large and persistent, more or less adnate with the Sorbaria (Gillenieae). The anthela of Filipendula is
petiole, and forming a sheath around the axillary a special case. Its shape results from the fact that
bud (Potentilleae, Rosa, etc.). the side-branches are crowded and of unequal
Rosaceae 345
length, overtopping the terminal flower of the hypanthium is constricted at top and closed over
mam ax1s. the carpel(s), so that only the stylodia emerge
Terminal flowers or even a larger terminal part through the central orifice. This is the case in
of the synflorescence may be more or less reduced, Alchemilla, Sanguisorbeae, Rosa, and in a large
either consistently or as one possibility within part of the maloids, where hypanthium and
the variation in a species. Holodiscus, Sibiraea ovary(ies) are fused so that the ovary becomes
and some Spiraea species (all Spiraeeae) are (semi-)inferior.
good examples, but possibly also Agrimonia In few genera (Aremonia, Spenceria), under the
(Sanguisorbeae). hypanthium, that is under the flower, an involu-
The axillary partial inflorescences of a thyrsoid crum is developed, consisting of bracts/prophylls
may represent dichasia or monochasia. On some (bracteoles). These organs must be distinguished
occasions, they are reduced to single flowers. well from an epicalyx which is, where present,
In such cases, the inflorescence is a botryoid always inserted at the apex of the hypanthium,
(= raceme with a terminal flower, e.g. in some with the sepals.
Rubus; Troll1969). These simple-looking inflores- The phyllomatic organs forming the epicalyx are
cences are not rare, occurring, e.g., in spiraeoid only present in genera of Potentilleae, the Geum
groups, Sanguisorbeae, maloid genera and in group, Sanguisorbeae, Kerrieae (see below), and
Prunus. Alchemilla (all rosoids). The segments are nor-
The occurrence of single terminal flowers can be mally present in the same number as the sepals.
interpreted as the result of a complete reduction The general opinion was, and is that the epicalyx
of lateral partial inflorescences. Inflorescences segments represent stipular appendages of the
reduced to one terminal flower occur in several sepals. Isomery of sepals and epicalyx originates
groups: Lindleya (Exochordeae), Kerria and in this vision by connation of the stipular
Rhodotypos (Kerrieae), Cowania and Dryas appendages of neighbouring, valvate sepals.
(Dryadeae), some species in Rosa and Rubus, Based on a teratological specimen of Geum rivale,
and in Cydonia. In Fallugia (Geum group?) and Bolle (1935) proposed another explanation of the
Mespilus (Crataegeae), there are often or always isomery: the outer 2 sepals have two stipules, the
one or few (up to 3 in Mespilus) lateral flowers or inner 2 none, the number-3 sepal one. Kania
at least buds under the terminal flower. (1973) investigated the ontogeny of the flower in
It must not be imagined that the first Rosaceae several species and saw that in Aremonia and
had a very elaborate thyrsus, evolving only by Potentilla the epicalyx primordia develop after
reductions to other inflorescence types. It seems those of the sepals, in Geum and Fragaria after the
more plausible to envisage the first Rosaceae as petal primordia, in Alchemilla after the stamen
having solitary terminal flowers, evolving by pro- primordia. In Rhodotypos (Kerrieae) the basal
gressive branching to more elaborate types (see appendages of the sepals are also interpreted as an
Parkin 1914). It must be recognized, however, epicalyx (Schaeppi 1953), but according to Kania
that the presence of solitary flowers is not always ( 1973) they are formed at the same time as the
a plesiomorphy: they certainly have also evolved marginal teeth of the sepals. More study of the
secondarily as reductions. ontogenetical development of epicalyces is needed
to elucidate their morphological nature. It is not at
FLOWERS. Although there are some genera with all certain that all these organs are homologous.
extreme reductions in the flowers, the generalized Sepals are usually rather small and often valvate,
image of a rosaceous flower is that of a discoid and protect the flowerbud only in a very juvenile
structure, displaying the petals, the stamens and stage. Sometimes, however, they are relatively
the nectar on the same level to passing insects. large, as in Rosa and Rubus, with an imbricate ver-
The only character uniting the family is the pres- nation also expressed in the unequal development
ence of a hollowed hypanthium (also called flower of the marginal lobes.
tube, floral cup, or calyx tube). A recognizable Petals are normally showy and the colour white
hypanthium is wanting only in staminate flowers predominates. Yellow petals are, however, most
of some genera like Bencomia and Sarcopoterium. common in some groups: Sanguisorbeae (insofar
In the majority of the Rosaceae the hypanthium as petaliferous), Potentilla and the Geum group.
is cupular, tubular, campanulate, etc., rarely The petals may be clawed and they are caducous
saucer-shaped. The inside of an open hypanthium after anthesis, as is normal for petals. The absence
is normally lined with nectariferous tissue (see of petals, as in some genera (Alchemilla, Neviusia,
below, under disk). In several genera, however, the Cercocarpus, several Sanguisorbeae), is obviously
346 C. Kalkman
a derived state, probably at least in part related to lature has been studied by Sterling (1964-1969)
the evolution of wind pollination. over a wide range of genera. Stylodia are usually
Unisexual flowers occur scattered through well developed and stigmas are generally of a
several of the tribes but hermaphroditic flowers simple shape, e.g. linear, capitate or bifid. In several
are by far most common. Plants with unisexual genera of Sanguisorbeae the stigmas are fimbriate
flowers are rarely dioecious (some Spiraeeae, to penicellate, probably in connection with wind
some Sanguisorbeae), more frequent is polygamy pollination.
with hermaphroditic and staminate flowers in one Ovules are anatropous, rarely hemi-anatropous.
plant or even in one inflorescence. In several In the groups usually considered to have main-
genera the situation has not been observed in tained primitive flower characters (spiraeoids),
living populations and remains uncertain. there are several ovules per carpel. In many other
Stamens are free from each other and their groups the number is reduced to 2 (then usually
number is often about 20. Depending on the only one developing to a seed) or 1. Where the
dimensions of the flower, a disposition in one row number is 1 or 2, the ovules are often basal,
or in several whorls may be evident. The ontoge- ascending, and collateral, more rarely apically
netic studies of Lindenhofer and Weber ( 1999a, inserted and pendulous; horizontally oriented
1999b, 2000) suggest a preponderance of a 10 + 5 ovules also occur. Descending ovules are always
+ 5 arrangement of the stamens, with antesepalous epitropous, ascending ones apotropous. An obtu-
stamen pairs in the outer cycle. This pattern is rator, guiding the pollen tube, may be present
interpreted as derived from a helical arrangement, (especially in Pruneae, but also in Exochorda
whereas the possibility of intercalation or splitting and Adenostoma); sometimes this function is per-
of stamens is refuted. Any evidence that the prim- formed by a swelling of the funiculus. The struc-
itive number of stamens should be 5 or 5 + 5 is ture of the ovary, style/stylodium, stigma, and
lacking. In small, reduced flowers, the number of ovules in several genera all through the family are
stamens may also be very low. In filled (double) treated extensively by Juel (1918, 1927). Not easily
flowers the extra petals are clearly petaloid explained is the large number of ovules in
stamens. what is here called the Cydonia group. Is this a
A disk is only mentioned as such in the generic plesiomorphic character state ('left over' from the
descriptions when it is structurally visible as a ancestral spiraeoids) or the result of a reversal
separate organ, ring-shaped or divided into parts. from 2-ovulate (as in other maloids) to multi-
It is probably always covered with nectariferous ovulate?
tissue, as may be the inside of the hypanthium.
When a disk is present, it is intrastaminal on the FRUITS AND SEEDS. The follicle, originating from
upper inside rim of the hypanthium (but see a monocarpellate, multi-ovulate pistil, occurs in
Alchemilla). Where the hypanthium is closed, it spiraeoids and may confidently be regarded as the
may be the disk that surrounds the orifice. plesiomorphic fruit type. Derived from this type
Five carpels is possibly the plesiomorphic state are the very common achene and the drupaceous
in the family and, if that is the case, there have achene with a slightly fleshy layer in the pericarp.
been evolutionary changes to larger numbers or The drupaceous achenes as in Kerrieae cannot be
reductions in number. When carpels are isomer- sharply separated from the true drupes with hard
ous, they are either antesepalous or antepetalous. endocarp and fleshy mesocarp as in Prunus and
When there is a large number of carpels, a more unrelated Rubus. So-called pomes are morpholog-
or less highly elevated torus is usually developed ically diverse, and the term should be restricted to
(Potentilleae ), the carpels either covering the the fruits derived from a flower with the carpels
whole torus or being absent in its basal part. adnate to the hypanthium forming a completely
Sometimes (in the Geum group, see Iltis 1913) the or incompletely inferior ovary, with a fleshy
carpels are placed on gynophores, obviously layer either entirely hypanthial or mixed
homologous with the torus but shaped as a rather hypanthial/carpellary, and with the endocarpal
thin stalk with all (or at least most) carpels at the layers not woody. The carpels may be partly or
top. Connation of carpels (with one another) and entirely free from one another. When so defined,
adnation (to the hypanthium) define structure the pome is restricted to the Cydonia group and
and shape of the fruit in the maloid groups. In most of the genera in the tribe Maleae. When
most groups, however, the carpels are free from a flower as described above develops woody
one another and from the hypanthium, each carpel endocarpal layers in the fruiting stage (as in
forming a separate pistil and developing into a Crataegeae), such a fruit may be called a multi-
fruit. The anatomy of the carpels and their vascu- pyrenous drupe but actually this is not a correct
Rosaceae 347
description (see note under tribe Crataegeae, usually relatively long. In some Rosaceae the
p. 380). ectoapertures are operculate. As far as is known,
Fleshy fruits may also be produced in other all Potentilleae display this character, also several
ways. In Rosa it is the closed hypanthium that Sanguisorbeae and some species of Rosa.
becomes fleshy and the fruits are achenes with a Transmission electron micrographs show a col-
hard pericarp. That reminds of the situation in umellate infratectal layer. The outside pattern of
quite unrelated genera with a fleshy hypanthium, ornamentation is variable but mostly striate.
like Margyricarpus and Sarcopoterium, and also of As far as present knowledge goes, the characters
the situation in some genera of Maleae, viz. of the pollen do not seem to be of much critical
Heteromeles and Dichotomanthes. In Fragaria the relevance for classification within the family.
fruits have a fleshy torus.
In the peri carp of the fruits of maloid genera, the KARYOLOGY. Summarizing the present knowl-
fleshy layer shows anatomical differences the sys- edge of chromosome (base) numbers, the follow-
tematic value of which has been partially explored ing picture emerges.
by Iketani and Ohashi (1991a). Heterogeneous
flesh contains either large pigmented cells or large Spiraeoids
sclereids, or both; homogeneous flesh does not Exochordeae Not sufficient evidence
contain these specialized cells. for establishing a base
Seeds from follicles have a thick testa but, when number. 2n = 16, 30,
the seeds remain enclosed in an achene or other 34
fruit type, the testa is usually thin, although some- Spiraeeae x=9 Counts for all 8 genera
times of a quite firm consistency. The embryo has Neillieae x=9 1 of the 3 genera
flat to thick cotyledons. In full-grown seeds there insufficiently known
is either nothing left of the endosperm or only a Gillenieae x=9 Counts for all 4 genera
very thin layer.
Rosoids?
Kerrieae x=9 1 out of 3 genera not
EMBRYOLOGY. Pollen grains are 2-celled when
counted
shed. The ovules are anatropous or hemi-
Dryadeae x=9 Counts for all 5 genera
anatropous and bitegmic (the primitive state) or by
connation unitegmic. In many genera the micro- Rosoids
pyle is (almost) absent by growth of the integu- Ulmarieae x=7 1 genus only
ments over the nucellus. Meiosis usually occurs in Sanguisorbeae x=7 6 out of 12 genera not
only one megaspore mother cell, leading to a tetrad counted
of which the chalaza! cell develops into an 8-celled Potentilleae x=7 Counts for all 3 genera
Polygonum-type embryo sac with one haploid egg Geum group x=7 2 out of 9 genera, incl.
cell. In some genera, however, the megaspore those of uncertain
mother cell is replaced by one or more secondary position, not counted
megaspore mother cells undergoing meiosis. Rubeae x=7 1 genus only
The polar nuclei in the embryo sac generally Roseae x=7 1 genus only
fuse prior to fertilization. The secondary
Rosoids?
endosperm is first free-nuclear, later becoming
Alchemilla x=8 1 genus only
cellular (Nuclear type). The embryo is mostly of
group
the so-called Asterad type. For a full account of
embryological data and literature, see Johri et al. Prunoids
(1992). Pruneae x=8 1 out of 4 genera not
counted
POLLEN MORPHOLOGY. No recent comprehensive
Maloids
account of pollen morphology in Rosaceae has
Cydonia x= 17 2 out of 4 genera not
been published, although some detailed regional
group counted
accounts do exist. The following short account has
Maleae X= 17 3 out of 11 genera not
been extracted from Van der Ham in Kalkman
counted
(1993: 231-232), where also the relevant references
Crataegeae X= 17 2 out of 7 genera not
may be found.
counted. See note
Pollen grains of Rosaceae do not display large
under Osteomeles,
variation. They are monads, more or less spher-
p.381.
oidal, and generally tricolporate. The colpi are
348 C.Kalkman
A number of unresolved problems arise: According to the classic work by Knuth (1898:
345-396; 1904: 334-347), pollinating insects are
a) In few cases aberrant chromosome counts (in Europe, and there is no reason to suppose
confuse the picture without sufficient evidence that it is different elsewhere) especially flies and
to consider the aberrant numbers as either short-tongued bees like Anthrena but, to a lesser
error or aneuploidy. degree, also long-tongued bees, beetles and even
b) In several of the tribes, establishing the base butterflies.
number rests on (too) little evidence, most Cross-pollination is mostly ensured by the
notably in the Lindleya group but also in more or less distinct, protogynous development
Neillieae and Sanguisorbeae. The natural char- of the flowers, rarely by protandry, dioecy or by
acter of these groups is, consequently, also more structural peculiarities (movement and place of
or less in doubt. stamens) that more or less impede self-pollination.
c) In some cases the chromosome base number is The latter is, however, by no means rare.
part of the evidence on which higher taxa are Anemogamy may be supposed to be present in
based (Kerrieae, Dryadeae, Alchemilleae). See several genera belonging to the Sanguisorbeae. It
also the paragraph on Classification, p. 349. can be deduced from the fimbriate or penicellate
stigma and, in some cases, by the scabrate pollen.
Much has been written on the origin of the high Experimental evidence is usually lacking.
base number x = 17 for the maloid groups. Well- Uncertainties are many-fold. For instance, the
known is the postulate (going back to Sax 1932) staminate panicles of Aruncus dioicus (Spiraeeae)
that the number and the group are the result of are visited by hundreds of insects that take away
one or more, old hybridization events between the pollen and feed on it, but whether this con-
plants with n = 9 and with n = 8, followed by poly- tributes to pollination of the nectarless pistillate
ploidization. The n = 9 parent is usually referred flowers is quite uncertain, the species being nor-
to as spiraeoid and the other parent as prunoid or, mally dioecious. However, there is not even una-
more prudently, as proto-spiraeoid and proto- nimity in the literature about the pistillate flowers
prunoid. Phipps et al. (1991) are strong supporters being nectariferous or not.
of this hypothesis. There are, of course, several
other ways to arrive at 2n = 34 and at the present APOMIXIS. In a number of genera scattered over
stage there is, in my opinion, not sufficient evi- several tribes, apomixis has been established.
dence to make a responsible choice. 1 That poly- Most common in Rosaceae is apospory where the
ploidization played a role in the history of the embryo sac develops from a somatic cell (with
family is obvious, but successful hybridization unreduced nuclei) of the nucellus. Diplospory,
between a member of the (primitive) spiraeoids where the embryo sac develops from a generative
and a member of the (much derived) prunoids is but unreduced cell derived from a megaspore
in my opinion not very likely. The counts of 2n = mother cell, has also been established. A third
34 in Lindleya do not make things easier, although type, nucellar (adventitious) embryony, where the
the taxonomic relationship of this genus with embryos develop from somatic cells, seems to be
maloids is not at all a wild guess. unknown in Rosaceae (this type is not included
into apomixis by all authors).
PoLLINATION. In the terminology of Faegri and The egg cell with its unreduced nucleus of the
van der Pijl (1979), the blossom class most aposporous or diplosporous embryo sac, stimu-
Rosaceae belong to is the dish- to bowl-shaped lated in some way by pollination or not, will form
class and the individual flowers are the pollinator- the embryo without syngamy, i.e. without fusion
attracting units. Some members of the family have with a microgamete from the pollen tube. In the
brush-shaped blossoms. The individual flowers words of Asker ( 1979: 233 ), "it may be questioned if
are then rather small, not individually conspicu- there exists any 100% obligatory apomicts",
ous, and the visually attractive unit is the head- or although this has been claimed for a few Rosaceae
spike-shaped inflorescence. Several Sanguisorbeae (Alchemilla). In most cases apomixis is certainly
belong to this blossom class. facultative and mixed with sexual reproduction.
SEED DISPERSAL. Four main modes of seed/fruit

1 Evans and Campbell (2002) have now discarded the wide
hybridisation hypothesis for Maloideae and instead suggest a dispersal are realized in Rosaceae, linked of course
polyploid origin involving only members of a lineage that con- to the different fruit types (dry and dehiscent, dry
tained the ancestors of Gillenia. and not dehiscent, fleshy).
Rosaceae 349
Endozoochory. The animals involved may be rhiza, it can be stated that knowledge 1s still
small ones (snails) in the case of low herbaceous incomplete.
plants like Fragaria. In the case of woody plants
mostly birds, bats and other mammals are the DISTRIBUTION. Three genera, Prunus, Alchemilla
active party. In most cases endozoochory is and Rubus, are cosmopolitan. Of these only Prunus
only surmised on the basis of morphology and with certainty occurs naturally in Asia, Europe, the
often there are no observations or experiments. Americas, Africa, and the Australian region. The
Myrmecochory may be present in Aremonia, present-day distribution of the two other genera is
where an extrafloral elaiosome seems to be grossly distorted by human influence, and special-
present. The fleshy part is never an arilloid or ist research would be needed to detect their orig-
sarcotesta. inal areas of distribution.
Sixteen genera are distributed over the entire
Epizoochory. This method relies on accidentally Northern temperate zone with species in North
passing animals. Specializations on the side of the America, Europe and Asia (Agrimonia*, Ame-
plant include hooks on the hypanthium or a lanchier*, Aruncus, Crataegus*, Dryas, Filipen-
hooked style. dula*, Fragaria*, Geum*, Malus, Parageum,
Potentilla*, Rosa*, Sanguisorba*, Sorb us*,
Ballistochory. The receptacle containing the dias- Spiraea*, Waldsteinia). Most of them (indicated
pores is composed of hypanthium and sepals (and by*) extend southward into regions with Mediter-
epicalyx), or it is the open follicle. ranean climate or even into tropical latitudes
(often montane). Geum and Agrimonia are excep-
Anemochory is rare. Hagenia and Leucosidea have tional in being distributed also in the Southern
a flying apparatus composed of the epicalyx, Hemisphere.
Cowania, Dryas and Geum have feathery, persist- Eight genera are Eurasian (Aremonia,
ent stylodia. A hypanthial wing is present on the Cotoneaster*, Eriolobus, Mespilus, Orthurus,
fruits of some Bencomia spp. Whether the wings Pyracantha, Pyrus*, Sibiraea). Their distribution
on the seeds of the Exocordeae are functional for ranges from rather small areas in Southeast
dispersal by wind is unknown. Europe and West Asia to wider areas, some (indi-
cated by*) extending to part(s) of Africa.
EcoLOGY. Habitats. Rosaceae range from semi- There are only two genera restricted to, but
desert to lowland rainforest and open, alpine veg- widely disjunct in, the Southern Hemisphere:
etation in a large number of different habitats. Acaena and Oncostylis.
Nevertheless, a large number of Rosaceae may Endemicity at the level of genus is high in Asia
be found on wooded mountain slopes at medium (20 genera) and North America (17 genera), but
altitudes and temperate latitude. less significant in South America, Africa (with 3
and 5 genera respectively) and Europe (1 genus);
Mycorrhiza. Vesicular-arbuscular mycorrhiza Australia has no endemic genus.
seems to occur as common and erratic in Rosaceae Croizat (1952), Goldblatt (1976), Thorne (1983)
as in Dicotyledons in general. Ectomycorrhiza has and Kalkman (1988) postulated a Southern
been demonstrated in only a small number of (Gondwanan) origin for the family, or at least
woody species belonging to Dryadeae (Cercocar- found it a hypothesis seriously to be considered. It
pus, Chamaebatia, Dryas), maloids (Crataegus, must be admitted, however, that most authors
Malus, Pyrus, Sorbus), Prunus and Rosa. This list favour a Laurasian origin for the family, and
was taken from Harley and Smith (1983) and it support for a Gondwanan origin is still weak.
may be regarded as quite incomplete. Genera from
areas outside Europe and North America have CLASSIFICATION. Subfamily level. Four sub-
rarely or not been checked. families are usually distinguished: Spiraeoideae,
Rosoideae, Maloideae (or Pomoideae), and
Nitrogen fixation. Root nodules with Frankia as Prunoideae (or Amygdaloideae). This subdivision,
bacterial component, in which nitrogen fixation which still is much used, obviously goes back to
has been demonstrated, also occur in Rosaceae, Focke (1888). At one time or another, these
especially in Dryadeae (see Newcomb and Heisey subfamilies have all been elevated to families:
1984). Surprisingly, also a tropical Asian species Spiraeaceae Maxim., Malaceae Small, Pomaceae
of Rubus has to be included in the list (Becking Lois.,Amygdalaceae (Juss.) D. Don, and Drupaceae
1979) and, just as is the case for ectomycor- DC. Several smaller groups have in the past also
350 C. Kalkman
been elevated to subfamily or family rank, either mentioned, either as accepted or excluded names
formally or only as a suggestion (e.g. Kerriaceae, or as a synonym.
Sanguisorbaceae, Dichotomanthes ).
While Prunoideae and Maloideae are probably Intergeneric hybrids. Not listed in the taxonomic
monophyletic groups, this cannot be said for the part are the intergeneric hybrids which sometimes
two other subfamilies. Spiraeoideae contain the have been named, especially in maloids. In the
genera with dry,multi-seeded,dehiscent fruits (fol- maloid tribes there are only few natural inter-
licles), and thus are characterized by a plesiomor- generic hybrids, most having been made artifi-
phic character. In other, derived groups, the fruits cially or having been formed spontaneously
evolved to different states. Spiraeoideae are clearly when plants were brought together in cultivation
linked with both Rosoideae and Maloideae, the (Robertson et al.1991; Kriigel1992b). It is obvious
delimitation of rosoids against spiraeoids being that the (in)ability to hybridize cannot be used as
uncertain (see Table in the paragraph on Karyol- an argument for splitting or lumping genera.
ogy). Prunoideae are most probably an offshoot
from some part of primitive Rosoideae. PHYLOGENY. A phylogenetic analysis of the
If one wants to intercalate a taxonomic rank family, considering only rather few morpholog-
between tribe and family, a logical and phyloge- ical characters and tribes as operational units
netically defendable distinction would be to rec- (Kalkman 1988), was not successful. Another
ognize two subfamilies, one covering the classical attempt, based on 36 characters in 96 species rep-
Spiraeoideae and Maloideae, the other including resenting almost all genera, was made by Phipps
the classical Rosoideae and Prunoideae. However, et al. (1991). A phylogenetic analysis on the basis
this classification is not formally proposed here, of wood anatomical characters was conducted by
since the phylogeny within the family has not Zhang (1992).
been worked out sufficiently. Some tribes such as A molecular analysis of rbcL sequence data
Kerrieae are difficult to place. Nevertheless, for (Morgan et al. 1994), based on 40 species repre-
the sake of orientation, the conventional groups senting nearly all tribes and groups recognized in
(spiraeoids, rosoids, maloids and prunoids) are the present survey, established the family as a
used, although the delimitation of the former two monophylum. The 'quillajoid' genera Kageneckia,
is not undisputed. Lindleya and Vauquelinia (but not Quillaja which
appeared together with Surianaceae and Poly-
Tribal level. Names for tribes were coined by galaceae and consequently here is excluded from
Maximowicz (1879) (for his Spiraeaceae), by Focke Rosaceae) were placed in one clade with the five
(1888) for Rosoideae, and by Koehne (1890) for maloid genera sampled (Amelanchier, Crataegus,
Pomaceae (Maloideae). The Prunoideae have Eriobotrya, 'Photinia' and Sorbus). This stresses
never been subdivided into tribes. Hutchinson the relationship between this part of the spi-
(1964) divided the family into some twenty tribes raeoids and the maloids. A subdivision into sub-
but did not distinguish subfamilies. The classifica- families of the traditional circumscription was not
tion given in the taxonomic part of the present supported by the resulting tree.
treatment is rather similar to Hutchinson's system In another molecular study, Campbell et al.
and contains 18 tribes or (if a formal name did not (1995) employed sequences from internally tran-
yet exist) 'groups'. I have tried to make them scribed spacers ITS and part of the 5.8 S rDNA of
homogeneous, but some problems remain and are 19 species of maloids, and included also mor-
discussed at the appropriate places. phological characters. Of the three subgenera of
Sorbus that were sampled, Aria is not nested with
Generic level. Generally, genera are taken in a Cormus and Sorbus, and the coherence of the
moderately narrow sense, and some of them may Cydonia group is not unequivocally supported
be united when more data are available (e.g. (Chaenomeles not nesting with Cydonia and
Purshia and Cowania). In Alchemilla and Poten- Pseudocydonia). Vauquelinia was, as in the analy-
tilla I have taken a wide view by lack of evidence sis by Morgan et al. (1994), included in the maloid
in favour of splitting, and Sorbus has been recog- clade in close proximity to Eriobotrya and
nized in a wide sense mainly for practical reasons Rhaphiolepis.
(many cultivated species). Synonymy for genera is
not exhaustive. However, all genus names cited FAMILY RELATIONSHIPS. Rosaceae have always
under the family in Brummitt (1992) and all been placed in an order Rosales but the con-
names recognized in Gunn et al. (1992) have been tents of the order varies much in different
Rosaceae 351
classifications. Families such as Calycanthaceae, many published ethnobotanical lists. Sometimes

Dichapetalaceae, Connaraceae, Cunoniaceae, this is easily understood when the region involved
Grossulariaceae, Saxifragaceae, Cunoniaceae, is poor in Rosaceae (see, e.g. Barret al. (1993) for
Leguminosae, Anisophylleaceae, Crossosomat- Australia, Morgan ( 1981) and Bhat et al. ( 1990) for
aceae, Neuradaceae, Rhabdodendraceae and Suri- parts of Africa, and Cambie and Ash ( 1994) for Fiji).
anaceae have been considered as relatives of the In these four lists Rosaceae either are completely
Rosaceae. In contrast, recent molecular studies absent or represented by only one species. Less
agree in placing Rosaceae in a clade together with understandable is the absence of Rosaceae in many
Barbeyaceae, Rhamnaceae and Elaeagnaceae, and published ethnobotanicallists relating to parts of
sister to Urticales families (see APG II 2003). South and Central America, where Rosaceae are
more diverse and more abundant. North American
PHYTOCHEMISTRY. The extensive literature has Indians, on the other hand, have found many uses
been reviewed and evaluated by Hegnauer (1973, (fruits or, in a single case, the taproot as food,
1990), to which the reader is referred for details several medicines, technical applications of wood
and references. or bark). For parts of Asia, ethnobotanicallists are
Phenolics are accumulated often in considerable usually also poor in Rosaceae, except as folk medi-
amount and often appear species- or genus- cine. Perry (1980) lists species in 14 genera in her
characteristic. Tannins of the condensed type are book on medicinal plants of East and Southeast
almost ubiquitous, while hydrolysable tannins Asia, the majority of the records being for China, a
seem to be restricted to some groups of genera country that is very rich in Rosaceae. The difference
(Okuda et al. 1992). between folk medicine and 'official' medicine
Sorbitol, a sugar alcohol, replaces part of the sac- becomes obvious when Perry's list is compared
charose in many of the Rosaceae. The absence of with the Chinese pharmacopoeia (Stager 1989),
it seems to be characteristic of some rosoids, espe- where only seven species are treated.
cially those with x = 7. Kerrieae and Adenostoma In Western civilization, the economic impor-
have sorbitol and in this respect are anomalous tance of the family rests mainly on the edibility of
among rosoids. the fleshy fruits of many of its members. The apple
Cyanogenic compounds (amygdalin, prunasin, is the second most cultivated fruit in the world,
dhurrin, and some unidentified glycosides) are after (wine-)grape. Also, pears, prunes, cherries,
well known from seeds and/or other parts in many peaches, apricots, loquats, strawberries, black-
Rosaceae but often show a patchy distribution. Of berries, raspberries, etc., are all more or less
the spiraeoids only some genera have been proven widely grown, traded, and consumed, either in
to contain prunasin. The rosoid genera are nor- their fresh state, or preserved in tins or bottles, or
mally devoid of cyanogenic compounds, with in the form of juices, jellies, or jams.
the exception of Kerrieae and some Dryadeae. Economically important is also the use of many
In maloids many genera and species have been species as ornamentals in gardens and parks or as
proven to be strongly cyanogenic, the seeds often street trees, mainly in temperate climates. The
containing amygdalin while other plant parts may beautiful flowers and often decorative fruits make
have prunasin. Pruneae are more or less the same many species useful for such purposes. Rosa, the
in this respect but the cyanogenic compounds in 'queen of flowers', may be economically the most
Oemleria and Prinsepia have not yet been identi- important genus in this category.
fied. Kageneckia contains prunasin, which may be As suppliers of timber the family is not impor-
interpreted as a support for the intermediate place tant. Only few species reach timber size, and a
of the genus between spiraeoids and maloids. minority of them may reach the world market.
Exochorda leaves and twigs are weakly cyanogenic Some tree species, however, have a local impor-
but the compound responsible is unidentified. tance for construction purposes.
Adenostoma and Coleogyne have small amounts of Other useful products include pectin from
an unidentified cyanogenic compound, which apples, rose oil from the flowers of Rosa species,
does not help finding a good place for them. and tool handles from the hard wood of Holodis-
Saponins, derivatives of pentacyclic triterpenes, cus and Cercocarpus. Except for the first-
are widespread in the family, normally as complex mentioned one, these products are not of high
mixtures. market value. For pharmaceutical products, see
the end of this paragraph.
UsEs. Ethnobotanical value (usefulness to non- The best-known medicine, originally extracted
Western societies) is generally low, as evidenced by from a rosaceous species, is aspirin, the brand
352 C.Kalkman
under which the German firm Bayer marketed - Ovules several, pendulous 7
acetylsalicylic acid in 1899. The brand name mem- 7. Leaves imparipinnate, leaflets several em long and wide.
Seeds with small wings at both ends 16. Sorbaria
orizes Spiraea ulmaria (now Filipendula ulmaria), - Leaves bipinnate or pinnate with deeply pinnatipartite
source of old European folk medicines (Balick and leaflets, the ultimate segments only some mm long. Seeds
Cox 1996: 32}. From the flowerbuds of this plant, without wings 17. Chamaebatiaria
salicylic acid was isolated and a synthesized 8. Epicalyx 0 (hooked or barbed spines at the top of the
derivate became world-famous, aspirin now being hypanthium are not considered to represent an epicalyx)
9
a household-name for any over-the-counter - Epicalyx +. Fruits achenes 27
febrifuge/ sedative. 9. Fruits dry achenes 10
- Fruits of other types (drupes or pomes) 24
CONSERVATION. The database 'Globally threat- 10. Herbs, stem base usually woody 11
ened species' of the World Conservation Monitor- - Shrubs or trees 17
11. Hypanthium open above. Pistils 5-15 12
ing Centre WCMC at Cambridge (U.K.) contains - Hypanthium closed above or almost so, achenes remain-
(May 1997) 475 records pertaining to species or ing enclosed in it. Pistils 1-51 14
infraspecific taxa of Rosaceae. Five species are 12. Leaves 1-3 times deeply tripartite. Hypanthium campanu-
considered to be extinct: two Crataegus from late, more or less clearly differentiated into a lower part
Canada, two Potentilla ( 1 from U.S.A., 1 from surrounding the ovaries and an upper tubular part.
Stylodia basal. Ovule 1 per pistil 41. Chamaerhodos
Russia}, and one Rubus from U.S.A. Under cate- - Leaves interruptedly pinnate. Hypanthium cupular, ovaries
gory E (endangered) or Ex/E (endangered or and achenes not enclosed in it. Stylodia terminal 13
maybe already extinct), there are more then 100 13. Pistils on bottom of hypanthium or on low torus, often
entries listed in c. 25 genera (my classification). shortly stalked. Ovules 2 per pistil 28. Filipendula
- Pistils on torus on top of distinct gynophore. Ovule 1 per
Cases are very diverse and I would not consider pistil 44. Geum (vernum)
the loss of c. 30 North American microspecies of 14. Petals+, yellow 15
the Rubus fruticosus complex (see p. 370) or an - Petals 0 16
apomictic hybrid species of Sorbus as alarming 15. Hypanthium with hooked spines at top 31.Agrimonia
as the loss of one of the eight South American - Hypanthium almost rudimentary, without spines
31.Aremonia
Margyricarpus species or the only Chamaemeles 16. Hypanthium in fruit 4-angled to 4-winged, without spines
species endemic to Madeira. 34. Sanguisorba
Species not present in the database but, in view - Hypanthium in fruit with 2-4 barbed spines at apex or
of their small range of distribution, probably at with many scattered spines 40. Acaena
least of the 'vulnerable' category will almost cer- 17. Flowers bisexual 18
- Flowers unisexual. Petals 0 22
tainly be added in the updating process. We may 18. Petals 0 19
think of the Asian Dichotomanthes tristaniifolia - Petals +, 3-5 20
and Spenceria ramalana, the North American 19. Flowers in inflorescences 38. Polylepis
Chamaebatiaria millefolium, and the Mexican - Flowers solitary 39. Margyricarpus
Lindleya mespiloides and Xerospiraea hartwe- 20. Sepals and petals 3 84. Potaninia
- Sepals and petals 5 21
giana, all species from monotypic genera, not (yet) 21. Leaves very finely compound, 3 times pinnate with very
in the WCMC listing. small ultimate leaflets. Pistil 1 27. Chamaebatia
- Leaves pinnate. Pistils many 54. Rosa
KEY TO THE GENERA 22. Flowers usually solitary. Often ericoid shrubs or shrublets,
rarely small trees, unarmed, leaves not in rosettes
1. Leaves compound 2 37. Cliffortia
- Leaves simple, lobed, sometimes deeply, or not - Flowers in inflorescences 23
(Lyonothamnus pinnate to pinnatifid) 42 23. Small thorny shrubs with small leaves. Disk in flower not
2. Fruit a dehiscent follicle with more than 2 seeds 3 evident 35. Sarcopoterium
- Fruit not dehiscent (achene, drupe or pome) 8 - Candelabra shrubs, leaves in rosettes at end of branches.
3. Plant herbaceous 4 Flower with thick disk 36. Bencomia
- Plant with woody branches, erect or prostrate 5 24. Carpels several to many, superior, free from hypanthium,
4. Leaves 2 or 3 times ternate to bi- or tripinnate, stipules 0. forming a collective fruit, composed of monopyrenous
Flowers mostly unisexual; bisexual ones sometimes + in drupes 53. Rubus
staminate inflorescences. Ovules inserted apically - Carpels up to 5, dorsally adnate to hypanthium and
12.Aruncus forming with it one spurious fruit (pome) 25
- Leaves trifoliolate, stipules +. Flowers bisexual. Ovules 25. Stones (endocarps) hard and woody 78. Osteomeles
inserted near base 19. Gillenia - Endocarp not woody 26
5. Prostrate shrublets. Leaves 2 or 3 times ternate 26. Leaves pinnate 64. Sorbus (subg. Cormus and Sorbus)
9. Eriogynia - Leaves 3-foliolate 72. Eriolobus
- Erect shrubs. Leaves (bi)pinnate 6 27. Trees or (small) shrubs 28
6. Ovules 2, collateral, inserted near base of cell. Leaves - Herbs with often stout, sometimes slightly woody caudex
imparipinnate 18. Spiraeanthus or rhizome 31
Rosaceae 353
28. Stylodia subterminal to subbasal 42. Potentilla 52. Seeds 1-2 per fruit 15. Stephanandra
- Stylodia termina 29 - Seeds several in each fruit 13. Neillia
29. Flowers solitary. Small shrubs 50. Sieversia 53. Shrubs, the old twigs thorny 10. Xerospiraea
- Flowers in inflorescences. Trees or shrubs 30 - Unarmed shrubs 54
30. Flowers in large, axillary panicles, normally unisexual, 54. Evergreen cushion plants. Hypanthium saucer-shaped
trees monoecious 29. Hagenia S.Kelseya
- Flowers in terminal spikes, bisexual 30. Leucosidea - Leaf-shedding shrubs. Hypanthium campanulate
31. Involucre under flower+ 32 14. Physocarpus
- No involucre under the flower 33 55. Only 1 ovule per carpel 56
32. Stamens 5-10 32. Aremonia - Two or more ovules per carpel 69
- Stamens 35-40 33. Spenceria (see Brachycaulos (p. 383), where the number of ovules
33. Stylodia subterminal, lateral or (sub)basal 34 is unknown)
- Stylodia terminal 36 56. Fruit a dry achene 57
34. Torus not swollen after anthesis, or torus 0 and pistils on - Fruit drupaceous, i.e. with woody endocarp(s) and thin,
bottom of hypanthium 42. Potentilla slightly fleshy mesocarp 67
- Torus swollen after anthesis 35 57. Petals 0 58
35. Petals white 43. Fragaria - Petals+ 61
- Petals yellow or red 42. Potentilla 58. Stipules adnate to petiole 55. Alchemilla
36. Stylodium hooked 37 - Stipules free 59
- Stylodium not hooked 38 59. Intrastaminal disk +, a tubular sheath around the ovary
37. Upper part of stylodium caducous, lower part persistent, 83. Coleogyne
hooked 44. Geum - Disk not evident 60
- Top of stylodium hooked in fruit 47. Oncostylus 60. Sepals 5. Hypanthium narrowly tubular in lower part, the
38. Stylodium articulate at base 39 upper part widened and open 26. Cercocarpus
- Stylodium not articulate at base 40 - Sepals 3(4). Hypanthium 0 in staminate flowers, ellipsoid
39. Leaves trifoliolate 52. Waldsteinia and closed at mouth in pistillate flowers 37. Cliffortia
- Leaves pinnate to pinnatisect 61. Hypanthium closed at top, fleshy when fruiting
49. Coluria (see also 42. Potentilla, where the stylodia are 54. Rosa (persica)
± always subbasal to subterminal. In very few species - Hypanthium open above, not fleshy when in fruit 62
only the stylodia could be called terminal) 62. Flowers (sepals and petals) 7-10-merous 23. Dryas
40. Pistils up to c. 15, most of them on top of stalk-like - Flowers 5-merous 63
gynophore, some lower or on bottom of hypanthium 63.Shrubs 64
46. Orthurus - Herbs 66
- Pistils many, not on a stalk-like gynophore 41 64. Epicalyx +,small 51. Fallugia
41. Terminal leaflet and uppermost lateral leaflets much larger - Epicalyx 0 65
than lower leaflets. Pistils on bottom of hypanthium or on 65. Pistils 1(-3) 25. Purshia
low torus 45. Parageum - Pistils 5-12 24. Cowania
- Terminal leaflet not much larger than lateral ones. Pistils 66. Stamens 5 41. Chamaerhodos
on bottom of hypanthium or on cylindrical torus - Stamens many 52. Waldsteinia
48. Acomastylis 67. Flowers solitary. Petals large 20. Kerria
42. Fruit a 5-celled, woody, loculicidal capsule 2. Lindleya - Flowers in terminal racemes 68
- Fruit of a different type: follicle, achene, drupe or pome 68. Carpels free from hypanthium. Petals small or 0
43 21. Neviusia
43. Fruit follicular. Woody plants 44 - Carpels dorsally adnate to hypanthium. Petals large
- Fruit of a different type, not dehiscent. Woody or herba- 80. Hesperomeles
ceous plants 55 69. Fruit a drupe (the mesocarp fleshy or leathery) or at least
44. Carpels 5, ventrally connate. Ovules 2 per carpel 45 drupaceous (the mesocarp thin and slightly fleshy). Fleshy
- Carpels 1-5, free from each other or at most basally shortly layer only of carpellary origin 70
connate 46 - Fruit otherwise, the fleshy part, if present, not (or not only)
45. Intrastaminal disk evident. Ovules pendulous of carpellary origin 74
4. Exochorda 70. Pistil normally 1 71
- Disk 0. Ovules ascending 3. Vauquelinia - Pistils 2 or more 72
46. Stipules +.Ovules 4-many per carpel. Seeds winged or flat 71. Style terminal
47 57. Prunus (see also 58. Maddenia, which is hardly or
- Stipules 0. Ovules 2-several per carpel. Seeds not winged not different from part of Prunus)
48 - Style becoming (sub )basal in fruit 59. Prinsepia
47. Follicles 5, each with many winged seeds 1. Kageneckia 72. Leaves opposite, rarely whorled in 3s 22. Rhodotypos
- Follicles 2, each with 4 flat seeds 85. Lyonothamnus - Leaves alternate 73
48. Intrastaminal disk well developed 49 73. Stipules 0 or rudimentary 56. Oemleria
- Disk inconspicuous or 0 51 - Stipules + 53. Rubus
49. Dioecious plants with unisexual flowers 6. Sibiraea 74. More than 2 ovules per carpel 75
- Flowers bisexual 50 - Two ovules per carpel 78
50. Low, prostrate, evergreen shrubs 7. Petrophytum 75. Axillary thorns +, collateral with short shoots
- Small or large, erect or prostrate shrubs, leaf shedding 60. Chaenomeles
5. Spiraea - Plants unarmed 76
51. Pistil1(2) per flower 52 76. Flowers in terminal umbel-like inflorescences with up to
- Pistils 2-5 53 five flowers 62. Docynia
354 C.Kalkman
- Flowers solitary, terminal on leafy shoots 77 98. Petals upright. Carpels 2. Pigment cells in flesh of fruit
77. Flowers bisexual or staminate. Stylodia connate at very 64. Sorbus subg. Chamaemespilus
base, at base surrounded by a cup-shaped disk on top of - Petals spreading. Carpels 2-5. Sclereids in flesh of fruit
the ovary 63. Pseudocydonia 66.Aronia
- Flowers bisexual. Stylodia free, not surrounded by a cup-
shaped disk 61. Cydonia TRIBES AND GENERA OF RosACEAE
78. Fruits dry achenes, indehiscent 79
- Fruits with fleshy parts of carpellary and/ or hypanthial
origin 81 1. TRIBE ExocHORDEAE Schulze-Menz (1964).
79. Pistil 1 82. Adenostoma
- Pistils 4 or more in each flower 80 Woody, unarmed. Leaves alternate, simple,
80. Intrastaminal disk+, rim-like. Pistils 4-5 11. Holodiscus
- Disk not evident. Pistils 5-15, rarely fewer
penninerved; stipules +. Hypanthium open at top.
28. Filipendula Epicalyx 0. Petals +. Carpels 5, free or ventrally
81. Carpel (pistil) always 1, free from the hypanthium or connate; stylodia terminal; ovules numerous-2
basally adnate to it 82 per carpel. Fruits with woody pericarp. Seeds
- Carpels 2 or more, exceptionally only 1 developed 83 winged; endosperm thin or 0. Chromosome base
82. Fruit a drupe with a woody stone 79. Chamaemeles
- Fruit an achene, entirely free from, but almost fully sur-
number(s) unclear.
rounded by the enlarged and fleshy hypanthium
69. Dichotomanthes
83. Carpel walls (or only the endocarps) forming woody 1. Kageneckia Ruiz & Pavon
pyrenes in the fruit, surrounded by fleshy tissue of hypan- Kageneckia Ruiz & Pavon, Prodr.: 145 (1794).
thial (and sometimes partly carpellary) origin 84
- Carpel wall not woody 87
84. Carpels completely connate with each other and adnate to Shrubs or small trees. Leaves serrate. Dioecious.
the hypanthium 81. Mespilus Flowers showy, male ones in poor racemes or
- Carpels ventrally and laterally free from each other, dor- corymbs, female ones solitary. Hypanthium cam-
sally adnate to hypanthium 85 panulate to obconical. Petals white to yellowish.
85. Leaves lobed. Ovules superposed, one-stalked
77. Crataegus Disk rim-shaped. Stamens or staminodes c. 15,
- Leaves not lobed. Ovules collateral 86 inserted on rim of disk. Pistils with obliquely
86. Thorns +. Carpels 5 76. Pyracantha dilated, 2-fid stigma; ovules many, 2-seriate, hori-
- Plants unarmed. Carpels 2-3 75. Cotoneaster zontal; pistillodes in male flowers small. Follicles
87. Inflorescence a simple raceme 88
- Inflorescence a compound raceme 92
spreading. Seeds many, winged at apex. 2n = 34.
88. Leaves deeply tripartite 72. Eriolobus About three species in South America (Bolivia,
- Leaves slightly or not lobed 89 Peru, Chile, Argentine).
89. Carpel apices free from hypanthium, exposed in flower 90
- Carpel apices adnate to hypanthium, not exposed 91
90. Flesh of fruit soft, core thin, sepals persistent. Carpels with 2. Lindleya H.B.K.
incomplete septs 74. Amelanchier
- Flesh of fruit hard, core cartilagnous, sepals ultimately Lindleya Humb., Bonpl. & Kunth, Nova Gen. Sp., 6 ed. fol.: 188;
caducous. No incomplete septs in carpels ed. qu.: 239 (1824), nom. cons.
70. Macromeles
91. Stylodia free, emerging through a central hole in the disk Trees, evergreen. Leaves with glandular-toothed
that covers top of ovary 73. Pyrus margins. Flowers solitary, terminal on axillary
- Stylodia connate at base 71. Malus
92. Leaves deeply lobed 64. Sorbus sub g. Torminaria
short shoots towards the ends ofbranchlets, bisex-
- Leaves unlobed 93 ual, 5-merous, showy. Hypanthium obconical.
93. Nerves going straight to the margin 94 Sepals imbricate. Petals white. Stamens 15-20.
- Nerves not going straight to the margin 96 Disk not evident. Carpels antesepalous, free from
94. Carpel apices not exposed. Sepals and part of hypanthium hypanthium, laterally connate, forming a 5-locular
caducous 64. Sorbus subg. Micromeles
- Carpel apices usually free from the hypanthium. Sepals ovary; stigmas 2-lobed, oblique; ovules 2 per cell,
persistent on the fruit 95 ascending. Fruit a 5-celled, woody capsule, loculi-
95. Flesh of fruit with large pigment cells cidally 5-valved. Seeds 2 per cell, fiat, with a thin
64. Sorbus subg. Aria wing. n = 17 (Goldblatt 1976). One species,
- Flesh of fruit without large pigment cells 65. Eriobotrya
Lindleya mespiloides H.B.K., in Mexico.
96. Carpels completely adnate to hypanthium. Hypanthium
and sepals caducous 67. Rhaphiolepis
- Carpel apices free from hypanthium. Sepals (long) persist-
ent on the fruit 97
3. Vauquelinia Humb. & Bonpl. Fig. 109
97. Spurious fruit consisting of leathery achenes surrounded Vauquelinia Correa ex Humb. & Bonpl., Pl. Aequin. 1: 140
by a fleshy hypanthium 68. Heteromeles (1807); Hess & Henrickson, Sida Contrib. Bot. 12: 101-163
- Fruit a true pome 98 (1987), rev.
Rosaceae 355
2. TRIBE SPIRAEEAE Maxim. (1879).
Woody (exception: Aruncus). Leaves alternate,

simple (exceptions: Aruncus, Eriogynia), pennin-
erved; stipules 0. Inflorescences terminal
(compound) racemes (exception: Kelseya).
Hypanthium open at the top, dry at maturity. Epi-
calyx 0. Sepals valvate. Petals +. Disk+ (exception:
Xerospiraea). Carpels 2-5, free; stylodia free;
ovules inserted in apical part of suture. Follicles
(exception: Holodiscus). x = 9.
5. Spiraea L.
Spiraea L., Sp. Pl.: 489 (1753); K.M. Purohit & G. Panigrahi,
Rosaceae in India 1: 48-138 (1991), regional rev.
Pentactina Nakai (1917).
Fig. 109. Rosaceae-Exocordeae. Vauquelinia californica. A
Fruiting branch. B Flower. C Flower, vertical section. D Fruit. Shrubs, leaf-shedding. Leaves simple, rarely lobed,
E Seed. (Takhtajan 1981)
margins incised at least in apical part or some-
times entire. Inflorescences terminal racemes,
umbels or compound racemes. Flowers 5(4)-
Large shrubs or small trees, evergreen. Leaves merous, bisexual, rarely unisexual and then plants
coriaceous, with serrate margins. Inflorescences dioecious. Hypanthium campanulate to obconic.
terminal, corymb-shaped compound racemes. Disk at the rim of the hypanthium, collar-shaped
Flowers bisexual, 5-merous. Hypanthium hemi- or divided into lobes, intrastaminal. Petals white,
spherical. Sepals valvate. Petals white. Stamens rarely yellow, red. Stamens 15-many. Pistils usually
15-20. Disk not evident. Carpels 5, free from 5, antepetalous, free; stylodia (sub )terminal;
hypanthium, ventrally connate; ovules 2 per cell, stigmas capitate; ovules 2-several, pendulous. Fol-
ascending, apotropous. Fruit separating into 5 follicles with dry pericarp. Seeds few, small, with
licles with woody pericarp, dehiscing ventrally and membranous testa, endosperm thin or 0. 2n = 18,
dorsally. Seeds 2 per follicle. 2n = 30. Three species 27, 36, 54; aneuploids also recorded. About 80
in Southern U.S.A. (Arizona, New Mexico, Texas) species in the northern temperate zone, richest in
and Mexico, in chaparral and other arid to semi- Asia, with a centre in the Himalayas (c. 30 spp.),
arid habitats, up to 2700 m altitude, often on 10 species wild in North America, and 8 species
limestone. wild in Europe. Usually in sunny places or forest
margins, more rarely in forest, in low altitudes and
montane.
4. Exochorda Lindl.
Exochorda Lind!., Gard. Chron.: 925 (1858). 6. Sibiraea Maxim.
Shrubs, leaf-shedding. Leaves with entire or Sibiraea Maxim., Acta Hort. Petropol. 6: 213 (1879).
serrate margins. Inflorescence a terminal raceme.
Flowers large, (4)5-merous, bisexual (unisexual?). Shrubs, more or less prostrate, unarmed, leaf-
Hypanthium campanulate to obconical. Sepals shedding. Leaves simple, entire. Inflorescence a
imbricate. Petals white. Stamens 15- 30, 3- 5 terminal, compound raceme. Flowers 5-merous,
together before each petal, the groups separated unisexual, plants dioecious. Hypanthium hemi-
by a larger space. Disk well developed as a sphe~ic. Petals white or pinkish. Stamens resp.
rim. Carpels 5, free from hypanthium and free at stammodes c. 20. Disk intrastaminal, lobed. Pistils/
top, but ventrally connate; stigma dilated; ovules pistillodes mostly 5, very shortly connate at base;
2, collateral, pendulous from apex. Fruit separat- stylodia terminal; stigma capitate; ovules 4-8,
ing as 5 follicles with woody pericarp. Seeds 1 or pendulous from ventral placenta. Follicles with
2 per follicle, fiat. 2n = 16 (one record of bony pericarp, opening ventrally, partly also
2n = 18). Some 4 species in Siberia, Korea, China, dorsally. Seeds few, linear, with thin endosperm.
Japan. 2n = 18. One species, S. laevigata (L.) Maxim., in
356 C. Kalkman
Central and East Asia and south-eastern Europe. bracts leaf-like. Flowers 5-merous, bisexual.
Rocky places in montane altitudes. Hypanthium obconic. Disk a shallow, intrastami-
nal rim. Petals white. Stamens c. 20. Pistils 4-6,
mostly 5, free, antepetalous; stylodia terminal;
7. Petrophytum (Torr. & A. Gray) Rydb.
ovules several, pendulous from upper part of
Petrophytum (Torr. & A. Gray) Rydb., Mem. New York Bot. ventral suture. Follicles opening ventrally and dor-
Gard. 1: 206 (1900), 'Petrophyton'. sally. Seeds fusiform, small, endosperm 0. 2n = 18.
Spiraea§ Petrophytum Nutt. ex Torr. & A. Gray (1840), rank not
One species, E. pectinata (Pursh) Hook., in the
indicated.
Eriogynia sect. Petrophytum S. Wats. (1890). nort.h-western part of North America, from Bering
Strait to Cascade Mts. Subalpine to alpine habitats.
Low, prostrate shrubs, unarmed, evergreen. Leaves
crowded, simple, 1- or 3-nerved,entire,coriaceous. 10. Xerospiraea Henr.
Inflorescence a terminal (compound) raceme.
Flowers 5-merous, bisexual. Hypanthium saucer- Xerospiraea Henrickson, Aliso 11: 206 (1986).
shaped. Disk intrastaminal, entire. Petals white.
Stamens c. 20. Pistils 3-5, free; stylodia terminal; Evergreen shrubs, with long and short shoots, old
stigmas minute; ovules 2-4, pendulous from top. twigs often thorny. Leaves alternate, small, simple,
Follicles opening dorsally and ventrally, coria- with (sub)entire margins. Inflorescences terminal
ceous. Seeds linear. 2n = 18. Three very similar (compound) racemes, often leafy at base. Flowers
species in the Western half of North America, pos- 5-merous, bisexual. Hypanthium obconic. Petals
sibly better combined into one. Montane, often on white. Stamens 15-20. Disk not evident. Pistils 2-5,
limestone. free; stylodia terminal; stigma punctiform; ovules
The relationships of this genus and the following 2, pendulous. Follicles thin-walled, opening ven-
three with Spiraea were discussed by Henrickson trally and in the upper part also dorsally. Seeds
(1986a). The reasons to keep them separate seem (not observed) 1 or 2, testa thin, endosperm
to be sufficiently compelling. 0. 2n = 18. One species, X. hartwegiana (Rydb.)
Henr., in Mexico. Montane, on limestone.
8. Kelseya (S. Wats.) Rydb.

11. Holodiscus (K. Koch) Maxim.
Kelseya (S. Wats.) Rydb., Mem. New York Bot. Gard. 1: 207
(1900). Holodiscus (K. Koch) Maxim., Acta Hort. Petropol. 6: 253
Eriogynia Hook. sect. Kelseya S. Wats. (1890). (1879), nom. cons., see also Taxon 45: 672 (1996).
Spiraea 'Gruppe' Holodiscus K. Koch, Dendrologie 1: 309
(1869), rank not indicated, basionym.
Shrublets, growing in cushions, unarmed, ever-
green. Leaves simple, entire, coriaceous. Flowers
Shrubs, unarmed, leaf-shedding. Leaves simple,
solitary, axillary, 5-merous, bisexual. Hypanthium
alternate, shallowly to deeply lobed, nervation
saucer-shaped. Disk inconspicuous, entire. Petals
pinnate, margins incised. Inflorescences terminal,
purplish pink. Stamens c. 10. Carpels 3-5, free;
panicle-shaped compound racemes. Flowers 5-
stylodia terminal; ovules 3-7, pendulous. Follicles
merous, bisexual. Hypanthium saucer-shaped.
opening along ventral and dorsal sutures, pericarp
Petals cream-coloured. Stamens c. 20. Disk intrast-
coriaceous. Seeds fusiform. 2n = 18. One species,
aminal, rim-like, not lobed. Pistils 4-5, free, ante-
K. uniflora (S. Wats.) Rydb. in some north-western
petalous; stylodia terminal; ovules 2, collateral,
states of U.S.A. Montane, on limestone.
pendulous from apex. Fruits achenes, follicle-like,
with a bony pericarp, but obviously not dehiscing.
9. Eriogynia Hook. Seed 1 (2?), endosperm thin. 2n = 18, 36. One
species, H. discolor (Pursh) Maxim., from NW
Eriogynia Hook., Fl. Bor. Am. 1: 255, pl. 88 (1832), pro parte,
incl. type (Spiraea pectinata Pursh). North America through Central America to
Luetkea Bong. (1833). Northern South America. Montane habitats, in
forest and on rocks.
Evergreen shrublets, unarmed, with prostrate
woody branches and erect herbaceous scape-like 12. Aruncus L.
flowering stems. Leaves crowded at base of stems
and smaller ones on the scapes, much divided, 2 or Aruncus L., Opera varia: 259 (1758).
3 times ternate. Inflorescence a terminal raceme, Pleiosepalum Hand.-Mazz. (1922); Symb. Sin. 7: 455, f. 13
(1933), reduction.
Rosaceae 357
Large perennial herbs with thick rhizomes. Leaves

in rosette and also cauline,2-3-ternate to -pinnate,
leaflets pinnatisect to toothed. Inflorescence
a much branched terminal panicle-shaped
compound raceme. Flowers 5-merous, usually
unisexual and plants dioecious. Hypanthium
saucer-shaped. Epicalyx 0. Petals white or yellow-
ish-white. Stamens 15-30. Disk rim-shaped, not
conspicuous. Pistils 3-5, free; stylodia terminal but
often oblique; ovules few to several, pendulous.
Follicles with coriaceous pericarp. Seeds few, very
small, endosperm thin or 0. 2n = 18, 36, but also 2n
= 14,16,42 have been recorded. Usually recognized
as one polymorphic species, A. dioicus (Walter)
Fernald, widespread over the northern temperate
zone. Usually in shaded, damp to wet places,
montane to subalpine.
3. TRIBE NEILLIEAE Maxim. (1879).
Unarmed, leaf-shedding shrubs. Leaves alternate,

simple, palmatinerved; stipules free, on the twig.
Flowers 5-merous, bisexual. Hypanthium open at
top, dry at maturity. Epicalyx 0. Petals white. Disk
not evident. Stylodia terminal. Follicles large.
Seeds with abundant endosperm. Probably x = 9,
see Note under Neillia.
13. Neillia D. Don Fig. 110

Neillia D. Don, Prodr. Fl. Nepal.: 228 (1825); J. Vidal, Adanso-
nia II, 3: 142-166 (1963), rev.; J. Cullen, J. Arnold Arbor.
52: 137-158 (1971), rev.
Fig. llO. Rosaceae- Neillieae. Neillia thyrsiflora. A Flowering

Leaves variously lobed; stipules rather large, branch. B Flower. C Hypanthium with fruits. (Kalkman 1993)
persistent or caducous. Inflorescence a terminal
raceme or leafy panicle. Hypanthium campanulate
to tubular, in fruiting stage often with long-stalked 14. Physocarpus (Cambess.) Raf.
glands outside and on pedicels. Sepals imbrica~e.
Stamens 5-many. Pistil1(2, free); ovules 2-12, bis- Physocarpus (Cambess.) Raf., New Fl. 3: 73 (838), nom. et orth.
cons.
eriate on ventral placenta, pendulous. Follicle cori- Spiraea L. sect. Physocarpus Cambess. (1824).
aceous, enclosed in to slightly protruding from the
enlarged hypanthium, dehiscing ventrally. Seeds Leaves more or less distinctly 3-lobed; stipules
several, testa hard and shiny, raphe distinct. n = 9, early caducous. Inflorescence a terminal raceme,
but see Note below. About 12 species, in continen- ± globular to corymb-shaped. Hypanthium cam-
tal Asia from India to Korea, China, and Vietnam, panulate. Sepals valvate. Stamens many. Pistils 2-5,
one species also in Sumatra and Java. Montane connate in basal part; ovules 2-4, in lower half of
thickets, grasslands and forest edges, up to 2500 m ventral suture, variously oriented. Follicles shortly
altitude. stalked, inflated, protruding from hypanthium,
Ratter and Milne (1973) reported n = 9 for opening along both sutures. Seeds few, testa hard,
Neillia sinensis; Sharma (1970, not seen) counted shiny. 2n = 18, 54. Few species in North America,
n = 10, a rather deviant and possibly aneuploid one in Northeast Asia. Mostly in moist places,
number. along streambanks, etc. The bark scales off in
narrow strips.
358 C. Kalkman
15. Stephanandra Siebold & Zucc. Small shrubs, ± evergreen, aromatic-glandular.

Leav:s. pinnate with pinnatipartite segments,
Stephanandra Siebold & Zucc., Abh. Math.-Phys. Cl. Kiin. Bay.
Akad. Wiss. 3: 739 (1843). or b1pmnate. Inflorescence a terminal panicle-
shaped compound raceme, with some leaves.
L.eaves 3-more-lobed; stipules rather large, per- Hypanthium hemispheric. Sepals valvate. Petals
Slstent. Inflorescences terminal, leafy racemes white. Stamens c. 60. Carpels 5, alternipetalous,
or panicles. Hypanthium obconic-campanulate. ventrally connate at very base, otherwise free but
Sepals slightly imbricate. Stamens 10-20. Pistil 1; closely adhering; stylodia terminal; stigmas capi-
ovules 2, nearly apical, pendulous. Follicle pro- tate; ovules several, pendulous. Follicles dehiscent
truding from hypanthium, not inflated, opening ventrally and at apex dorsally. Seeds few, fusiform.
alo-?g ventral suture. Seeds 1-2, filling the follicle 2n = 18 (16?). One species, C. millefolium (Torr.)
entlrely, testa hard. 2n = 18. About 3 species, Korea, Maxim., in the western part of the U.S.A., in
Japan, and China. In thickets at low altitudes and montane altitudes.
montane.
18. Spiraeanthus (Fisch. & C.A. Mey.) Maxim.
4. TRIBE GILLENIEAE Maxim. (1879). Spiraeanthus (Fisch. & C.A. Mey.) Maxim., Acta Hort. Petropol.
6: 226 (1879).
Spiraea subg. Spiraeanthus Fisch. & C.A. Mey. (1842).
Unarmed. Leaves alternate, compound; stipules on
junction twig-leaf. Flowers in terminal inflores- Shrubs, probably evergreen. Leaves imparipinnate;
cences, 5-merous, bisexual. Hypanthium open at leaflets numerous, small, entire, coriaceous;
top, dry at maturity. Epicalyx 0. Petals +. Disk not stipules small. Inflorescence a terminal, panicle-
evident. Carpels up to 5, free. Follicles. Seeds with shaped compound raceme. Hypanthium
thin layer of endosperm. x = 9 (?). campanulate. Petals pink. Stamens 20-25. Carpels
2-5, alternipetalous, connate at base; stylodia ter-
16. Sorbaria (Ser.) A. Braun minal, thickened towards apex; stigmas capitate;
ovules 2, collateral, inserted near the base. Follicles
Sorbaria (Ser. in DC.) A. Braun in Asch., Fl. Prov. Brandenb. 1: connate at base, ventrally dehiscent and dorsally
177 (1860),nom. cons.; Rahn,Nord. J. Bot. 8:557-563 (1989),
rev. at apex. Seeds 1-2, shortly winged at both ends.
Spiraea L. sect. Sorbaria Ser. in DC. (1825), basionym. 2n = 18. One species, S. schrenckianus (Fisch. &
C.A. Mey) Maxim., in Kazakhstan: Karatau Range
Shrubs, leaf-shedding. Leaves imparipinnate, and Golodnaya Step.
leaflets with incised margins. Inflorescence a ter-
minal panicle-shaped compound raceme. Hypan- 19. Gillenia Moench
thium hemispherical. Sepals valvate, persistent.
Petals white or creamy. Stamens 20-50. Carpels Gillenia Moench, Meth. Suppl.: 286 (1802).
Porteranthus Small ( 1894), nom. superfi., see Taxon 37: 139
usually 5, antesepalous, ventrally connate in lower
(1988).
half and at base also connate over a part of the
lateral walls, otherwise free but closely adhering,
Perennial herbs, with rhizomes. Leaves trifoliolate,
forming a superior ovary with deep longitudinal
shortly petiolate, in inflorescence subsessile,
grooves between the cells; stylodia (sub )terminal,
leaflets with incised margins; stipules large or
recurving after anthesis; ovules several, pendu-
small. Inflorescence a terminal, corymb- or
lous, in 2 rows. Follicles much exserted from
panicle-shaped compound raceme. Flowers on
hypanthium, pericarp cartilaginous. Seeds few,
long pedicels, rather large. Hypanthium tubular to
fusiform, with small wings at both ends. 2n = 36.
c~mpanulate. Sepals imbricate. Petals white to pale
According to the recent revision, 4 species in
pmk. Stamens 10-20. Pistils 5, alternipetalous,
Central and East Asia. From low to montane and
free; stylodia terminal; ovules 2-4, inserted near
alpine altitudes.
the base, ascendent, apotropous. Follicles soon
rupturing the hypanthium, pericarp bony. Seeds
17. Chamaebatiaria Maxim. 1-4, rather large. 2n = 18. Two species in North
America: Canada (Ontario) and U.S.A. (Eastern
Chamaebatiaria (Brewer & Watson) Maxim., Acta Hort.
Petropol. 6: 225 (1879).
part), in montane forests.
Spiraea§ Chamaebatiaria Porter ex Brewer & Watson (1876).
Rosaceae 359
5. TRIBE KERRIEAE Focke (1888). larger, directed more outwards and equivalent to
or simulating an epicalyx (see below), persistent.
Unarmed, leaf-shedding shrubs. Leaves simple; Petals large, white. Stamens many, some of them
stipules +, free. Flowers bisexual. Hypanthium adnate to the disk. Disk intrastaminal, large, cov-
open at top, dry at maturity. Sepals imbricate. ering the pistils, 4-lobed. Pistils usually 4 (2-6);
Stamens many. Pistils several, free; stylodia termi- stigmas capitate; ovules 2, collateral. Achenes
nal; ovules attached in the middle. Achenes dru- black, mesocarp slightly fleshy, endocarp woody.
paceous, i.e. with slightly fleshy mesocarp and Seed 1, with thin testa, endosperm a thin layer.
woody endocarp. x = 9 (few data). 2n = 18. One species, R. scandens (Thunb.) Makino
in Japan and Central China.
The opposite leaves are an exceptional character
20. Kerria DC.
in the family. A relationship with Kerria is
Kerria DC., Trans. Linn. Soc. Lond. 12: 156 (1818). assumed, mainly because of the similarity in fruits
and leaves. The two outer sepals and sometimes
Leaves alternate, with incised margins. Flowers also the inner ones have at one or both sides a large
solitary, terminal on short lateral branches, large, basal tooth, usually separated from other marginal
5-merous. Hypanthium shallowly cup-shaped. teeth, which was interpreted by Schaeppi (1953) as
Epicalyx 0. Sepals imbricate, large. Petals large, stipular organs. The so-called disk was ontogenet-
(orange-)yellow (white in a cultivar). Stamens ically studied by Schaeppi (1977); it is not inner-
many. Disk not evident. Pistils 5-8; stigma puncti- vated and does not produce nectar. It cannot be a
form; ovule 1. Achenes yellow, mesocarp thin, staminodial structure but seems to be homolo-
fleshy, endocarp woody. Seed 1, with thin testa, gous with nectariferous, annular, intrastaminal
endosperm rather thick. 2n = 18. One species, K. disks, as occur in other genera.
japonica (L.) DC., in China and Japan. In forest
and open places.
6. TRIBE DRYADEAE (Focke) Hutch. (1964).
21. Neviusia A. Gray
Woody. Leaves alternate, simple (exception:
Neviusia A. Gray, Mem. Am. Acad. Arts II, 6: 374 (1858). Chamaebatia); stipules +. Flowers solitary or in
fascicles, normally bisexual. Hypanthium open
Leaves alternate, with incised margins; stipules at the top, dry at maturity. Epicalyx 0. Petals +
small. Inflorescence a short, terminal raceme, (exception: Cercocarpus). Disk 0. Pistils with long,
flowers on long pedicels, 5-merous, showy terminal stylodia; ovule 1, basally attached, obtu-
although apetalous. Hypanthium saucer- to cup- rator 0. Achenes with persistent stylodia or beak.
shaped. Epicalyx 0. Sepals imbricate, large and x= 9.
leafy, green to greenish white, margins incised.
Petals small or 0. Stamens many, white. Disk not
23. Dryas L.
evident. Pistils 2-5; stigma linear, on one side of
stylodium; ovule 1. Achenes with thin fleshy meso- Dryas L., Sp. Pl.: 501 (1753).
carp, endocarp crustaceous. Seed 1, endosperm
very thin. x = ? Two spp., N. alabamensis A. Gray in Dwarf shrubs with creeping branches, evergreen.
south-eastern states of U.S.A., and N. cliftonii Leaves close together, margin usually incised but
Shevock, Ertter & D.W. Taylor, California. On entire in some forms. Flowers solitary, axillary,
slopes and streambanks, often on limestone, long-pedicelled, bisexual or one of the sexes
shaded and more open places. reduced, showy. Hypanthium cup- to saucer-
shaped. Sepals 7-10, imbricate or valvate. Petals
usually isomerous with sepals, white, whitish, or
22. Rhodotypos Siebold & Zucc.
yellow. Stamens many. Pistils many, free, on
Rhodotypos Siebold & Zucc., Fl. Japon. 1: 185 (1841). bottom of hypanthium or on low torus; stylodia
terminal, long-hairy; stigmas minute, terminal,
Leaves opposite (sometimes whorled in 3), with linear, longitudinally canaliculate. Fruits achenes
incised margins. Flowers solitary, terminal on terminated by persistent and elongate, hairy stylo-
short shoots, 4-merous, bisexual, showy. Hypan- dia. Seed with membranous testa, endosperm a
thium small, saucer-shaped. Sepals imbricate, thin layer. 2n = 18. Two species: D. octopetala L.,
leafy, rather large, margins incised, lower teeth circumpolar-alpine, and D. drummondii Rich. in
360 C. Kalkman
Northeast Asia and North America. On rocky

places, up to high altitudes.
24. Cowania D. Don

Cowania D. Don, Trans. Linn. Soc. Lond. 14: 574 (1825).
Shrubs, evergreen, glandular. Leaves small, simple,

pinnatipartite with 3-5 lobes to entire; stipules
small, adnate to petiole. Flowers solitary, terminal
on long and short shoots, sessile, 5-merous,
normally bisexual, fragrant. Hypanthium funnel-
shaped to obconical or ± campanulate. Sepals 5,
slightly imbricate. Petals 5, white or pale yellowish.
Stamens many. Pistils 5-12, free, on bottom of
hypanthium, shortly stalked; stylodia terminal,
long-hairy; stigma linear. Fruits achenes termi-
nated by persistent, long-hairy stylodia. Seed with
membranous testa, endosperm thin. 2n = 18. One
species, C. mexicana D. Don, in south-western
U.S.A. and Mexico. Mountain slopes, often on
limestone. Hybridizing with species of Purshia
(Koehler and Smith 1981) and merged by
Henrickson (1986b).
25. Purshia Poiret

Fig. Ill. Rosaceae-Dryadeae. Cercocarpus traskiae. A Branch
Purshia DC. ex Poiret in Lam., Enc., Suppl. 4: 623 (181 6).
with short-shoot. B Flowering branch. C Fruits. (Sudworth
1967)
Shrubs with erect or prostrate branches, leaf-
shedding, glandular. Leaves small, simple, wedge-
shaped, 3(5)-lobed from the top; stipules small, (pedicel-like) in lower part, the upper part
adnate to petiole. Flowers solitary, terminal on widened, caducous. Sepals continuous with upper
short shoots, (shortly) pedicelled, 5-merous, bisex- part of hypanthium, valvate. Petals 0. Stamens
ual. Hypanthium obconical. Sepals imbricate. 15-25( -40). Pis till (2), on bottom of hypanthium;
Petals yellowish. Stamens 20-50. Pis till (2, 3, free), stylodium terminal, long-hairy; stigma linear,
on bottom of hypanthium; stylodium terminal; on the ventral side of the stylodium; ovule with
stigma linear. Fruit achene, beaked by persistent enlarged chalazal part. Fruit achene, stylodium
stylodium, pericarp fibrous. Seed with dark testa, after anthesis growing into a persistent, very long,
endosperm thin. 2n = 18. Two closely related twisted, plumose 'tail'. Seed 1, testa fibrous, stout,
species (or varieties?) in western North America: the apex an empty appendage, endosperm 0. 2n =
British Columbia to California and New Mexico. 18. About 5 species in western North America. In
Open mountain slopes and also in forest. more or less arid habitats, especially montane.
26. Cercocarpus Humb., Bonpl. & Kunth Fig. 111 27. Chamaebatia Benth.
Cercocarpus Humb., Bonpl. & Kunth, Nova Gen. Sp. 6, ed. fol.: Chamaebatia Benth., Pl. Hartweg.: 308 (1849).
183, ed. quart.: 232 (1824); F.L. Martin, Brittonia 7: 91 - 111
(1950), rev.
Shrubs, aromatic-glandular, evergreen. Leaves
alternate, 3 times pinnate, very finely compound
Small trees or shrubs, evergreen, with long and and fern-like; stipules small, lower part adnate
short shoots. Leaves usually leathery, with entire to the petiole, tips free. Inflorescence terminal
or incised margins; stipules free from petiole, on leafy twigs, cymose, few-flowered. Flowers 5-
caducous. Flowers solitary or fasciculate, terminal, merous, bisexual. Hypanthium obconical. Epicalyx
5-merous, bisexual. Hypanthium narrowly tubular 0. Sepals imbricate. Petals white. Stamens many.
Rosaceae 361
Disk not developed. Pistil I, on bottom of hypan- Trees up to 20m, evergreen. Leaves interruptedly
thium; stylodium rather short, constricted and imparipinnate; stipules large, adnate to petiole,
articulate at its base; stigma linear; ovule 1, basal, forming a wing over its entire length. Inflores-
ascending. Fruits achenes, with coriaceous peri- cences panicles, axillary, pendulous. Flowers 4-5-
carp. Seed with thin testa, endosperm thin. 2n = 18. merous, unisexual and trees monoecious, rarely
One species, C. foliolosa Benth., in California. bisexual; pedicel with 2 large prophylls. Hypan-
Mountain slopes and open forest. thium in male flowers small, saucer-shaped, in
female flowers turbinate, constricted at throat and
closed by a lobed disk. Epicalyx segments large,
7. TRIBE ULMARIEAE Vent. nerved. Sepals smaller than epicalyx, folded back.
Petals small to minute, linear. Stamens c. 16-20, in
Herbs. Leaves pinnate; stipules +. Hypanthium female flowers staminodial and very small. Pistils
open at top. Epicalyx 0. Petals +. Pistils several; 2-3, included; stigmas spathulate, fimbriate. Fruits
stylodia terminal; ovules 2, pendulous. Fruits enclosed in hardened hypanthium which is
indehiscent, 1-seeded. x = 7. crowned by enlarged epicalyx. 2n = ? One species,
H. abyssinica (Bruce) J.F. Gmel., in mountains of
continental Africa, from Central African Republic,
28. Filipendula Mill. Sudan, and Ethiopia to Zimbabwe, up to 3500 m
Filipendula Mill., Gard. Diet., abr. ed. 4 (1754); Schanzer, J. Jap. altitude.
Bot. 69:290-319 (1994), rev., without descriptions, with key.
Perennial herbs. Leaves alternate, in rosette and 30. Leucosidea Eckl. & Zeyh.
cauline, interruptedly pinnate; stipules rather Leucosidea Ecklon & Zeyher, Enum.: 265 (1836).
large, with incised margins. Inflorescence an
anthela-shaped compound raceme. Flowers 5( -7)- Shrubs or small trees, probably evergreen. Leaves
merous, bisexual, rarely unisexual and plants densely, rarely interruptedly imparipinnate,
polygamous. Hypanthium low cupular. Sepals leaflets in few pairs, margins incised; stipules
valvate, reflexed and persistent after flowering. forming an amplexicaul sheath, tips free, intra-
Petals white, cream-coloured, pink, or red. petiolar. Inflorescences terminal spikes. Flowers 5-
Stamens 20-45. Disk 0. Pistils 5-15, rarely fewer, on merous, bisexual, bracteate. Hypanthium obconic,
bottom of hypanthium or on low torus. Fruits thick disk at the throat. Epicalyx shorter than
achenes with the appearance of follicles but sepals. Sepals valvate. Petals green, smaller than
indehiscent, pericarp tough. Seed 1, testa thin, sepals, early caducous. Stamens 10. Pistils 2(-4);
endosperm very scarce. 2n = 14, 28, 42, aneuploid stylodia terminal, curved, rarely hooked at apex;
numbers also reported. According to the recent stigmas linear on curved end. Fruits included in
revision 15 species, in Europe, temperate Asia (also hardened hypanthium, crowned by persistent epi-
Himalayas and Yunnan) and Eastern North calyx and sepals. 2n =? One species, L. sericea Eckl.
America. In forest and in open (grassy) vegetation, & Zeyh., in Zimbabwe and South Africa. Montane
often in damp places. grassy slopes, riverbanks.
8. TRIBE SANGUISORBEAE Focke (1888). 31. Agrimonia L.

Agrimonia L., Sp. Pl.: 448 (1753).
Leaves alternate, imparipinnate or 3-foliolate;
stipules, at least those of rosette leaves, adnate to Perennial rhizomatous herbs. Leaves interruptedly
petiole. Hypanthium almost closed at the throat, imparipinnate. Stipules large, free. Inflorescences
enlarging in fruiting stage and persistent around terminal and axillary, simple, often interrupted
the achene(s). Pistils few or only 1; stylodia termi- racemes. Flowers 5-merous, bisexual; pedicel at
nal; ovule 1, pendulous. Achenes. Endosperm 0. base with two lobed prophylls. Hypanthium
x = 7, but chromosome number of several genera almost closed at the throat by a dome-shaped disk,
unknown. the outside armed with hooked spines. Epicalyx
0. Sepals imbricate, after anthesis closing over
29. Hagenia J.F. Gmel. Fig. 112 the fruit. Petals yellow, rarely white. Stamens
5-10( -20). Pistils usually 2, free. Achene usually
Hagenia J.F. Gmelin, Syst. Nat. 2, 1: 613 (1791). one per flower developing, enclosed in armed
Brayera Kunth ex A. Richard (1822).
362 C. Kalkman
Fig. 112. Rosaceae-Sanguisorbeae. Hagenia abessynica. A petiole but tips free, triangular. Flowers solitary or
Branch with female inflorescence. BMale inflorescence. CPart few fascicled in the axils of reduced leaves in apical
of the same. D Male flower, partly open. E Male flower. F Part
of male flower halved to show pistillode. G F emale flower, seen part of flowering stems, 5-merous, bisexual.
from above. H Female flower, lateral view, with outer calyx Hypanthium surrounded by, but free from a 6-12-
lobes removed. I Same, with hypanthium opened to show sta- lobed involucre, almost closed at the throat by the
minodes and pistil. J Pistil. (Graham 1960) intrastaminal disk. Epicalyx much smaller than
sepals, almost rudimentary. Sepals imbricate.
Petals yellow. Stamens 5-10. Pistils 2; stigmas reni-
hypanthium, pericarp thin, coriaceous. Seed with form. Fruits achenes with thin, hard pericarp,
membranous testa. 2n = 28, 42 (in hybrid), 56. often only 1 per flower developed, enclosed in
Between 10 and 15 species, widespread in Europe hardened but thin hypanthium, involucre enlarged
and Asia southward to Sri Lanka and Java, few in fruiting stage. 2n = 42. One species, A. agri-
species in North America and Mexico, 1 species monoides (L.) DC., in southeast Europe and west
in Haiti, one or few species in Brazil (Skalicky Asia. In forest, in montane and subalpine altitudes.
1971; Fuks 1987). Some species introduced and After flowering, the axis between the involucre
sometimes naturalized outside the native area, e.g. and the fruit-bearing hypanthium develops into
in South Africa. Temperate, in tropical regions an elaiosome (Gams 1923) and the fruit (hypan-
montane. Mostly in open vegetation, some species thium plus achene) is dispersed by ants (Ridley
weedy. 1930: 521).
32. Aremonia Nestl. 33. Spenceria Trimen

Aremonia Necker ex Nestler, Monogr. Potentilla: IV, 17 (1816), Spenceria Trimen, J. Bot. 17: 97, t. 201 (1879).
nom. cons.
Perennial rhizomatous herbs. Leaves (interrupt-
Perennial, rosette-forming herbs. Leaves inter- edly) imparipinnate, most in basal rosette, few
ruptedly imparipinnate; stipules adnate to the reduced ones cauline; stipules forming narrow
Rosaceae 363
wings. Flowering axis erect, inflorescence a fimbriate. Hypanthium in male flowers minute, in
narrow, simple or compound raceme. Flowers female flowers large, constricted at apex, unarmed.
rather large, 5-merous, bisexual, bracteate. Involu- Epicalyx 0. Sepals 4, caducous. Petals 0. Stamens
cre under the flower 7-9-lobed. Hypanthium free many in male flowers, 0 in female ones. Intrasta-
from involucre, obconic. Epicalyx leaves with minal disk not evident. Pistils 2 or 3, free; stigmas
rounded apex, much smaller than pointed sepals. penicellate. Achene(s) remaining enclosed in
Petals large, yellow. Stamens 35-40. Intrastaminal smooth, slightly fleshy, red-coloured hypanthium.
disk rim-like, with entire margin, closely sur- 2n = 28. One species, S. spinosum (L.) Spach, East
rounding the stylodia. Pistils 2, included; stigmas Mediterranean area. In xerophytic shrubland.
punctiform. Fruits achenes, usually one devel- Dispersal, however, is often hydrochorous, see
oped, included in the persistent, unarmed hypan- Litav and Orshan (1971).
thium. 2n = ? Possibly only one species, S.
ramalana Trim., in China.
36. Bencomia Webb & Berthel.
Bencomia Webb & Berthelot, Hist. Nat. Iles Canaries 3: 10
34. Sanguisorba L. (1842).
Marcetella Sventenius (1948).
Sanguisorba L., Sp. Pl.: 116 (1753); Nordborg, Opera Bot. 11, 2:
Dendriopoterium Sventenius (1948).
1-103 (1966), partial rev.; Nordborg, Opera Bot. 16: 1-166
( 1967), rev. of sect. Poterium.
Poterium L. (1753). Candelabra shrubs, leaf-shedding. Leaves in
Poteridium Spach (1846). rosettes at the end of branches, imparipinnate;
stipules in lower part adnate to petiole, upper part
Perennial, rhizomatous herbs. Leaves in radical free. Inflorescences axillary spikes, or racemes of
rosettes and fewer cauline, imparipinnate; stipules spikes. Flowers unisexual, plants dioecious, incom-
of rosette leaves membranous, those of cauline pletely dioecious, or monoecious with both sexes
leaves free. Inflorescences terminal on erect, in one inflorescence; floral bract and prohylls
branched stems, dense spikes or heads. Flowers connate. Hypanthium in male flowers minute, in
hi- or unisexual, the plants normally monoecious. female flowers large, constricted and almost closed
Hypanthium constricted and almost closed at at apex, unarmed. Epicalyx 0. Sepals 3-5. Petals 0.
apex, 4-angled, unarmed. Epicalyx 0. Sepals 4, Stamens many in male flowers, 0 in female flowers.
petaloid, white, green, or reddish, caducous. Petals Disk a thick collar around the orifice of the
0. Stamens 2-4 or up to 30 or more. Intrastaminal hypanthium. Pistils 2( -4), entirely enclosed in
disk 0 but bases of sepals rather thickened. Pistils hypanthium; stigmas penicilliform. Achene(s)
1 or 2( -5), free; stigmas penicellate. Achene(s) remaining enclosed, fruiting hypanthium globose
remaining included in the hardened, 4-angled to or pear-shaped,± 4-angled, or with 2 wings. 2n =
4-winged, dry hypanthium. 2n = 14, 28, 56. About 28. About six species restricted to the Canary
15 species, if species limits are not drawn too nar- Islands, and one endemic to Madeira. In rocky
rowly. Most species in Eurasia and North Africa, 2 places, montane.
in North America. In grassy vegetation, shrubland, Recognized as 3 separate genera by D. & Z.I.
forest-edges, sometimes under marshy to damp Bramwell (1974) and D. Bramwell (1978). In Vieira
circumstances. (1992) and Press and Short (1994), Marcetella is
Most of the species seem to be wind-pollinated, kept separate from Bencomia. Nordborg (1966)
but insect pollination also occurs in some of the included Dendriopoterium and Marcetella in
species (see Sa'id 1979). Sanguisorba and kept Bencomia separate.
35. Sarcopoterium Spach 37. Cliffortia L.

Sarcopoterium Spach,Ann. Sci. Nat. Bot. III, 5:43 (1846); Nord- Cliffortia L., Sp. Pl.: 1038 (1753); Weimarck, Monogr. Genus
borg, Opera Bot. 11,2: 73-76 (1966). Cliffortia: 1-229 (1934); Bot. Not. 1948: 167-203 (1948),
survey.
Small shrubs, often cushionplants, branches ter-
minating as thorns. Leaves small, imparipinnate, Shrubs, erect or procumbent, or shrublets, rarely
coriaceous; stipules small, tooth-like. Inflores- treelets, often ericoid, evergreen. Leaves trifolio-
cences spikes, opposite the leaves. Flowers unisex- late, bifoliolate, or simple, usually strongly xero-
ual, plants monoecious, female flowers at the end morphic, blades of leaves/leaflets linear or wider;
of the spike, greenish; floral bract and 2 prophylls sheath amplexicaul, distinct or short, stipules on
364 C. Kalkman
the sheath, rarely 0, usually scarious, rarely green,

free. Flowers axillary, usually solitary, rarely in
small fascicles, rarely female ones in cone-like
inflorescences, inconspicuous, wind-pollinated,
unisexual, exceptionally bisexual, plants dioecious
or monoecious. Hypanthium 0 in male flowers,
ellipsoid in female flowers, ribbed or smooth
outside, closed at mouth. Epicalyx 0. Sepals 3(4),
caducous or persistent, often coherent. Petals 0.
Stamens 3-20( -30). Pistils 1 or 2, enclosed; stigma
feathery. Fruits achenes, enclosed in bony to
slightly fleshy, ribbed or winged hypanthium.
2n =?At least 100 species in South Africa, richest
in the West Cape Province. Only one species
extending to Angola and Kenya. Often xerophytic
and in dry places but some species preferring
moist habitats.
The striking parallelism of the leaves with those
of Aspalathus (Papilionaceae) in the same region
was discussed by Dahlgren (1971).
38. Polylepis Ruiz & Pavon Fig. 113

Polylepis Ruiz & Pavon, Fl. Peruv. Prodr.: 80 (1794); B.B.
Simpson, Smithson. Contrib. Bot. 43: 1- 62 (1979), monogr.;
Kessler, Candollea 50: 131-171 ( 1995), rev. for Bolivia, 9 spp.;
Romoleroux in Fl. Ecuador 56: 71-89 (1996), 7 spp.
Trees up to c. 25m, or shrubs. Leaves imparipin-

nate or trifoliolate; stipules forming a closed
sheath. Inflorescences racemes, pendent or erect.
Flowers bisexual, bracteate. Hypanthium urceo-
late, constricted at throat and intrastaminal disk
almost closing the orifice. Epicalyx 0. Sepals 3 or
4. Petals 0. Stamens 6 to many, anthers hairy. Fig. 113. Rosaceae-Sanguisorbrae. Polylepis quadrijuga. A
Pistil 1( -3 ), included; stylodium short or long; Flowering twig. B Flower with penicillate stigma. C Flower in
stigma peltate, fimbriate or penicellate. Achene(s) the male phase. D Fruit. (Simpson 1979)
included in and closely accumbent to the hard and
tough hypanthium with irregular spine- or knob-
linear; stipules adnate to the petiole over their
like protuberances or ridges, the latter sometimes
entire length, forming an amplexicaul sheath.
expanded into wings. 2n = ? About 20 species in
Flowers solitary, axillary, bisexual. Hypanthium
the Andean region of South America, from
urceolate, constricted at apex by a disk, mostly 4-
Venezuela to N Chile and N Argentina. In montane
angled. Epicalyx 0. Sepals (3)4(5). Petals 0. Stamens
and subalpine habitats, often up to the tree line, to
(1)2(3). Pistil 1, included; stigma penicelliform.
5000 m altitude.
Fruit achene, enclosed in hypanthium, the latter
in fruiting stage either dry, hard to leathery, with
39. Margyricarpus Ruiz & Pavon 3-5 distinct wings, or swollen into a fleshy
pseudoberry without wings. 2n =?About 8 species
Margyricarpus Ruiz & Pavon, Fl. Peruv. Prodr.: 7, t. 33 (1794).
Tetraglochin Poepp., Fragm. Syn. Phan.: 26 (1833); Rothmaler, but badly in need of a critical revision. In the
Darwiniana 3: 429-437 (1939), rev. Andean region of South America from Ecuador to
Chile and Argentina, and also in South Brazil and
Shrubs, erect or creeping, possibly evergreen. Uruguay.
Leaves imparipinnate, sometimes on short shoots Margyricarpus and Tetraglochin have often, e.g.
1-foliolate, petiole and rachis of long-shoot leaves by Rothmaler, been kept separate on the strength
in most species persistent as hard spines, leaflets of the fleshiness of the fruiting hypanthium in
Rosaceae 365
probably only one species, M. pinnatus (Lam.)

Kuntze. Tetraglochin was described as being
dioecious and it is certainly not impossible that
unisexual flowers are sometimes present.
The intergeneric hybrid x Margyracaena
skottsbergii Bitter was described from the Juan
Fernandez Archipelago. Using RAPD markers,
Crawford et al. (1993) produced evidence that it is
a hybrid between the indigenous Margyricarpus
digynus (Bitter) Skottsb. and the introduced,
weedy Acaena argentea Ruiz & Pavon.
40. Acaena L. Fig. 114

Acaena Mutis ex L., Mant. 2: 145,200 (1771); Bitter, Bibl. Bot.
74: 1-336,37 pl. (1911), monogr.
Ancistrum J.R. & G. Forst. (1776).
Perennial herbs, usually woody at base, with creep-

ing stems and erect flowering shoots. Leaves
imparipinnate; stipules adnate to petiole, with
small free tips. Inflorescences terminal or axill-
ary, in heads or interrupted spikes. Flowers small,
normally bisexual. Hypanthium narrowed at
the throat. Epicalyx 0. Sepals (3)4( -7), valvate,
caducous or persistent. Petals 0. Stamens often 4
(1-10). Disk 0. Pistil usually 1(2-4), free; stylodium
terminal; stigma usually plumose. Achene(s) in
hardened hypanthium, the latter with 2- 4 barbed
spines under the sepals, or with many spines scat-
tered over the surface. 2n = 42, c. 72, 84, 126. About
40 species when a not too narrow species concept
is followed. Southern hemisphere, richest in extra-
tropical South America, extending to Mexico and
California (1 species), few in South Africa, several
in Australia and New Zealand extending to Hawai'i
(1 species) and New Guinea (1 species), some in
subantarctic island groups. Bitter (l.c.) mentioned
c. 110 species. In forests and open vegetation from
the lowland up to 4500 m altitude.
Possibly two subgenera may be distinguished,
on the basis of the number of spines on the fruit-
ing hypanthium, 2-4 in one subgenus, many in the
other. Bitter, however, was of the opinion that
reductions in the number of spines have occurred
repeatedly, in connection with greater compact-
ness of the inflorescence.
Fig. 114. Rosaceae- Sanguisorbeae. Acaena poeppigiana. A

Flowering plant. B Leaf, the sheath with a rudimentary stipule.
C Apex of inflorescence. D Chasmogamic flower from inflo-
rescence. E Cleistogamic flowers hidden among the sheaths of
rosette leaves. F Cleistogamic flower. GLeaf sheaths with fruits
from cleistogamic flowers. H Fruit from cleistogamic flower.
(Grandoso 1964)
366 C. Kalkman
9. TRIBE POTENTILLEAE Focke (1888). saucer-shaped, cup-shaped, hemispheric or cam-

panulate, often with hairs on the inside. Epicalyx
Unarmed. Leaves alternate, compound. Stipules as large as or smaller than sepals. Sepals valvate,
adnate to petiole. Flowers normally 5-merous, usually entire. Petals yellow or white, rarely red
almost always bisexual. Hypanthium open at top, or purple. Stamens 10 to many. Disk mostly not
dry at maturity. Epicalyx +(exception: Chamaer- evident, sometimes + as a rim on which the
hodos). Petals+. Carpels several to many, in some stamens are inserted. Pistils many-few, rarely 1, on
species in reduced number; stylodia not terminal; the bottom of the hypanthium or on a low to ele-
ovules 1 per carpel. Achenes. Endosperm 0. x = 7. vated, usually hairy torus; stylodia subapical,
ventral or subbasal, sometimes partly thickened,
persistent or articulated and caducous. Achenes
41. Chamaerhodos Bunge
usually remaining enclosed by the persistent
Chamaerhodos Bunge in Ledeb., Fl. Alt. 1: 429 (1829). hypanthium with epicalyx and sepals closing
above them, torus dry, rarely after anthesis enlarg-
Perennial herbs, more or less woody at base. ing and becoming spongy to somewhat fleshy.
Leaves in dense rosette-cushions and few on flow- Seed with thin testa. 2n = 14 or a multiple,
ering stems, 1-3 times tripartite, finely dissected; recorded up to 2n = 112, aneuploidy obviously
stipules in rosette-leaves adnate to base of petiole rare. Probably more than 300 species in the north-
as a membranous widened part, in the ± sessile ern temperate zone, and with few species extend-
cauline leaves practically 0. Inflorescences termi- ing to northern and eastern Africa and Malesia
nal, composed of cymes. Flowers 5-merous, (New Guinea). Mostly in open habitats, many
bisexual. Hypanthium campanulate, its lower part species montane to alpine. In need of a compre-
surrounding the ovaries, the upper part ± tubular. hensive, genus-wide study.
Epicalyx 0. Sepals valvate. Petals white or purplish. Molecular studies (Eriksson et al. 1998) raise
Stamens 5, antepetalous. Disk not evident. Pistils doubt on the monophyly of the genus. Some
5-10 (+),on bottom of hypanthium; stylodia basal, Eurasian species complexes (Potentilla arguta, P.
articulated. Fruits dry achenes. Seed with firm argentea a.o.) have a facultative apomictic repro-
testa. 2n = 14. About five species in Central and duction mode, implying the usual taxonomic
East Asia, one also in the western part of North difficulties.
America (U.S.A. incl. Alaska, Canada). In open veg-
etation on mountain slopes, also in forest.
43. Fragaria L.
Fragaria L., Sp. Pl.: 494 (1753).
42. Potentilla L.
Potentilla L., Sp. Pl.: 495 (1753). Perennial rhizomatous herbs, often stoloniferous.
Comarum L. (1753). Leaves radical, trifoliolate, only in anomalous
Sibbaldia L. ( 1753 ).
Duchesnea Sm. (1811).
cases with 5 leaflets; stipules adnate to petiole.
Horkelia Cham. & Schldl. (1827). Inflorescences terminal on leafless erect stems,
Ivesia Torr. & A. Gray (1858). cymose, few-flowered (except in cultivars).
Comarella Rydb. (1896). Flowers 5-merous, bisexual or unisexual and
Stellariopsis (Bail!.) Rydb. (1898). then plants (polygamo-)dioecious. Hypanthium
Purpusia Brandegee (1899).
Sibbaldiopsis Rydb. ( 1901).
obconic to saucer-shaped. Epicalyx of± same size
Drymocallis Fourr. ex Rydb. (1908). as sepals. Sepals valvate. Petals white, rarely yel-
Horkeliella (Rydb.) Rydb. (1908). lowish. Stamens c. 20-numerous. Disk sometimes
visible as a rim on which the stamens are inserted.
Perennial herbs with a rhizome or caudex, some- Pistils many, free, on cylindrical torus; stylodia
times forming cushions, sometimes with stolons lateral to subbasal, short; stigma capitellate; ovule
(shrubs, annual herbs). Leaves all radical or also inserted in the middle, descendent, epitropous.
alternate on longer stems, (interruptedly) pinnate, Achenes dry, on the much-enlarged, fleshy to juicy
palmate, or tri(uni)-foliolate; stipules in their torus, forming an oblong, ovoid, or globose fruit.
lower parts adnate to the petiole, membranous or 2n = 14 or a multiple, reported up to 2n = 56.
the upper free parts leafy and green. Inflores- About 10 species, in temperate Eurasia and North
cences cymose or thyrsoid, terminal or axillary, or America, one species, F. chiloensis (L.) Mill., also in
flowers solitary and axillary or opposite the leaves. South America (Chile). In open vegetation and in
Flowers (4)5(6)-merous, bisexual. Hypanthium forest.
Rosaceae 367
Kept separate from Potentilla, although the case part elongating after anthesis, persistent and ter-
for reduction is almost as strong as in Duchesnea minating in a hook, upper part caducous; stigmas
and Comarum, which also have a torus enlarging punctiform. Achenes tipped by long persistent sty-
after anthesis, although there becoming more lodia. Seeds with membranous testa. Most species
spongy than fleshy. hexaploid, 2n = 42, but 2n = 28, 56, 70 and 84 are
Fragaria x ananassa (Duchesne) Guedes, the also reported. About 30 species, widely distributed
Garden strawberry, widely cultivated as table fruit in Europe (including the Mediterranean), Asia,
and for preserves, originated (maybe more than and North and South America, one species, G.
once) in French gardens as an octaploid hybrid capense Thunb., in South Africa. In open vegeta-
between the North American F. virginiana Mill. tion and forests, some weedy.
and F. chiloensis, both octaploid species.
45. Parageum Nakai & H. Hara
10. GEUM GROUP Parageum Nakai & H. Hara, Bot. Mag. Tokyo 49: 124 (1935);
Knil, Preslia 38: 151 (1966), emend.
? Taihangia Te T. Yii & C.L. Li (1980), see Note below.
Perennial herbs with caudex, rarely rhizome,
Sieversia auctt., pro parte, not incl. type.
unarmed. Leaves imparipinnate, in rosette and few Acomastylis auctt., pro parte, not incl. type.
often reduced ones cauline, alternate. Stipules
adnate to petiole, at least in rosette leaves. Flowers Perennial, rarely stoloniferous. Rosette leaves
5-merous, bisexual, in scapose, mostly few- imparipinnate, lyrate, with terminal leaflets much
flowered cymes. Hypanthium open at top, dry at larger than lateral ones, cauline leaves few,
maturity. Epicalyx + (one exception in Geum). reduced. Inflorescence terminal on erect or
Petals mostly yellow. Stamens many. Ovule 1, basal, ascending leafy stems in axils of rosette leaves,
ascending. Achenes. Endosperm 0 (not known in cymose, few( -I)-flowered. Hypanthium obconoid.
Coluria). x = 7 (but see Orthurus). Epicalyx lobes smaller than sepals. Sepals valvate.
The characters of three genera (nos. 50-52}, of Petals yellow. Stamens many. Intrastaminal disk
uncertain position but probably related to the low. Pistils many, on bottom of hypanthium or on
present group, have not been taken into account in low torus; stylodia terminal, not articulate with
the above summary of characters common to all ovary, variously hairy, upper part not caducous;
genera. See p. 368. stigma linear to punctiform. Achenes crowned by
Lack of material precludes a definite decision elongated stylodia. Seeds with thin testa. 2n = 28,
on the status of Novosieversia, Taihangia, and 42. About 6 species, Eurasia and North America, in
Woronowia. the montane and subalpine belt.
The Chinese Taihangia rupestris Yii & Li has uni-
44. Geum L. sexual and bisexual flowers. It is possibly well
accommodated in this genus, but no material was
Geum L., Sp. Pl.: 500 ( 1753 ); F. Bolle, Fedde Rep., Beih. 72: 1-119 seen.
(1933), monogr. of Geum in a slightly wider sense than
accepted here.
46. Orthurus Juz.
Perennial. Rosette leaves (interruptedly) imparip-
innate, end-lobe (and upper lateral lobes) larger, Orthurus Juz., Fl. USSR 10: 616 (1941, orig. ed.),458 (1971, Engl.
trans!.).
cauline leaves less divided. Inflorescence terminal Geum L. subg. Orthostylus (Fisch. & C.A. Mey.) F. Bolle, Fedde
on erect leafy stems in the axils of rosette leaves, Rep., Beih. 72: 43-46 (1933), monogr.
cymose, mostly few-flowered. Hypanthium ? Woronowia Juz. (1941), see Note below.
saucer-shaped to cup-shaped or obconical. Epica-
lyx +(lacking in G. vernum (Raf.) Torr. & A. Gray), Perennial. Rosette leaves interruptedly imparipin-
lobes smaller than sepals. Sepals valvate or slightly nate, lyrate, cauline leaves less divided but still
imbricate. Petals mostly yellow, also white, pink, large. Flowering stems in the axils of rosette leaves,
red, or purplish. Stamens 20-many. Intrastaminal laxly branched in upper part and forming a leafy
disk sometimes + but never prominent. Pistils inflorescence. Hypanthium narrowly obconical.
several to many, on the bottom of the hypanthium Epicalyx lobes not very different from sepals.
or on a conoid to cylindric torus, lower part of Sepals valvate. Petals yellowish. Stamens many.
torus in some species developed as a gynophore; Intrastaminal disk not evident. Pistils rather few,
stylodia terminal, not articulate with ovary, lower up to c. 15, most in cluster on top of gynophore but
368 C. Kalkman
always 1 or 2 lower on gynophore and/or on obconical. Epicalyx lobes smaller than sepals,
bottom of hypanthium; stylodia terminal, sometimes incised from apex. Sepals valvate.
lower part persistent after anthesis, top with stiff Petals yellow(ish). Stamens many. Intrastaminal
retrorse hairs, upper part shorter, caducous; disk evident or not. Pistils many, on bottom of
stigmas short-linear. Achenes tipped by persistent hypanthium or on a cylindric torus; stylodia ter-
parts of stylodia, on lengthening gynophore. Seeds minal, not articulate with ovary nor elsewhere,
with firm testa. 2n = 28, 30 (see below). One not elongating after anthesis, glabrous or hairy;
species, 0. heterocarpus (Boiss.) Juz. in France, the stigmas punctiform. Seeds with firm testa. 2n = 28,
Mediterranean, West and Central Asia. Montane, in 70. In this restricted circumscription about twelve
shaded places. species, most of them in North America, one also
Woronowia speciosa (Albov) Juz., based on in Northeast Asia. Two species are known from the
Geum speciosum Albov (not seen) from the Himalayas, but further research may reveal that
Caucasus Mts., also has a long gynophore with the these are better placed in Parageum.
pistils partly at its base but, according to the Bolle (l.c.) recognized two subgenera, subg.
descriptions, the stylodia are not retrorsely Megacomastylis, part of which is here placed in
barbate. Still, it may belong in this genus. Parageum (subg. Parageum sensu Kral). The other
part, together with the second subgenus, subg.
Micracomastylis, here united with Erythrocoma, is
47. Oncostylus (Schltdl.) F. Bolle
placed here in Acomastylis.
Oncostylus (Schltdl.) F. Bolle, Fedde Rep., Beih. 72: 27-32 The genus Novosieversia, with its only species N.
(1933), monogr. glacialis (Adams) F. Bolle, occurs in Northeast Asia
Geum L. subg. Oncostylus Schltdl., Linnaea 28: 465 (1856), and is probably congeneric with the present one.
basionym.
Perennial. Rosette leaves imparipinnate,lyrate with 49. Coluria R. Br.

terminal leaflet much larger than lateral ones,
Coluria R. Br., J. Voy. N.-W. Pass., Bot.: 276 (1824); F. Bolle,
cauline leaves reduced. Flowering stems in axils of Fedde Rep., Beih. 72: 90-93 (1933), monogr.
rosette leaves, stiffly branched in upper part and
forming a bracteate, few-flowered inflorescence, or Perennial. Rosette leaves interruptedly imparipin-
stems not branched and scapose with one flower. nate to pinnatisect, end-lobe and upper lateral
Hypanthium obconical. Epicalyx lobes narrower lobes distinctly larger than lower ones or all
and usually shorter than sepals. Sepals valvate. (primary) leaflets± equal in size. Inflorescence ter-
Petals yellow or white. Stamens 10-many. Disk minal on axillary branches, cymose, mostly few-
0. Pistils many, on bottom of hypanthium or on flowered. Hypanthium obconical. Epicalyx lobes
cylindrical torus; stylodia terminal, more or less smaller than sepals. Sepals valvate. Petals yellow
glabrous, after anthesis hooked at apex; stigmas (white in one species?). Stamens c. 15 to many, on
punctiform. Achenes crowned by hooked stylodia. often distinct rim (disk). Pistils (3-)12-25, on
Seeds with thin testa. 2n =?About 5 species in New bottom of hypanthium or on short torus; stylodia
Zealand incl. Auckland Islands, one in Tasmania, (sub )terminal, articulate with ovary and caducous.
one or two in southern South America. In grassland Seeds not seen. 2n = 14. About 4 species in Siberia
and on rocky places, at low to montane altitudes. and China. Mountain slopes at high altitudes.
48. Acomastylis Greene

Doubtfully belonging to the Geum group
Acomastylis Greene, Leaflets Bot. Obs. 1: 174 (1906); F. Bolle,
Fedde Rep., Beih. 72: 78-90 ( 1933 ), monogr. of the genus in
a wider sense than here. The three genera placed here have in common that
Erythrocoma Greene (1906). they seem to be related to the Geum group but
? Novosieversia F. Bolle (1933), see Note below. differ from it in having distinctly to slightly woody
Sieversia auctt., pro parte, not incl. type. stems. The differences between them are such that
they do not form a separate group.
Perennial. Rosette leaves imparipinnate, often
interruptedly, terminal leaflet not strikingly larger
than lateral ones. Flowering stems in axils of 50. Sieversia Willd.
rosette leaves. Inflorescence terminal, cymose, Sieversia Willd., Ges. Naturf. Fr. Berlin Mag. Neuesten Entd. 5:
few- or 1-flowered. Hypanthium campanulate to 397 (1811).
Rosaceae 369
Small, evergreen shrubs, branches mostly pros- 52. Waldsteinia Willd.

trate, unarmed. Leaves in rosettes at the tops of the Waldsteinia Willd., Ges. Naturf. Fr. Berlin Neue Schr. 2: 105
branches, pinnate; lower part of stipules adnate (1799); F. Bolle, Fedde Rep., Beih. 72: 93-96 (1933), monogr.
to the petiole, tips free, long and thin. Flowering
stems scapose, not branched. Flowers solitary, 5- Perennial herbs, unarmed, stems firm, mostly
merous, bisexual. Hypanthium obconical. Epicalyx creeping. Leaves in rosette at the top of the stem,
segments about as long as sepals but narrower, simple and 3-5(-7)-lobed or 3-foliolate; stipules
sometimes 2-topped. Sepals valvate. Petals white. of radical leaves completely adnate to the long
Stamens very many. Intrastaminal disk 0. Pistils petiole, forming a membranous wing. Inflores-
many (40 or more), free, on bottom of hypan- cence terminal on axillary stems, cymose, cauline
thium, in fruit on a short gynophore; stylodia ter- leaves reduced, stipulate; bracts in inflorescence
minal, not articulate, straight, hairy except tip; mostly small. Flowers 5-merous, bisexual. Hypan-
stigmas punctiform; ovule 1, basal, ascending. thium funnel-shaped to obconical. Epicalyx small
Fruits achenes, crowned by persistent and elongate or 0. Sepals valvate. Petals yellow. Stamens many.
hairy stylodia. Seeds with membranous testa, Intrastaminal disk+, thin. Pistils (1)2-6( -15), free,
endosperm 0. 2n = 14. One species, S. pentapetala on bottom of hypanthium or on short torus; sty-
(L.) Greene, in Northeast continental Asia, lodia terminal, constricted and articulate at base,
Japan and the islands north of it. Often in damp deciduous; stigmas punctiform; ovule 1, basally
situations. inserted. Fruits achenes. Seeds with membranous
Sieversia has in the past been considered as a testa, closely adhering to pericarp, endosperm 0.
much larger genus, including also herbaceous 2n = 14, 21, 28, 35, 42. About 5 species, 2 or 3 in
species now placed in Parageum and Acomastylis. North America, 1 in Southeast Europe, 1 (W.
ternata (Steph.) Fritsch) disjunct from Alps to
51. Fallugia Endl. Korea. Montane, in heath- and shrubland and
other sunny places, also in forest.
Fallugia Endl., Gen.: 1246 (1840).
Small shrubs, more or less evergreen, unarmed. 11. TRIBE RUBEAE (Focke) Hutch. (1964).
Leaves alternate, deeply 3-7-partite, rarely simple,
wedge-shaped in general outline, small; stipules at Stipules free. Epicalyx 0. Pistils many, free, on ele-
base shortly adnate to petiole, largest part free, vated torus; ovules 2, pendulous, one developing.
basal part of petiole together with stipules remain- Drupes. x = 7.
ing on the twig. Flowers solitary, terminal on the
long shoots (growth sympodial), 5-merous, nor-
mally bisexual, under the terminal flower often 53. Rubus L.
two lateral ones. Hypanthium cup-shaped. Epica- Rubus L., Sp. Pl.: 492 (1753); Focke, Bibl. Bot. 72: 1-223
lyx segments small. Sepals imbricate. Petals white. (1910-1911); ibid. 83: 1-274 (1914), monogr.
Stamens many, inserted on a low ring. Intrasta-
minal disk not developed. Pistils many, on conoid Shrubs (i.e. determinate flowering shoots develop-
torus, shortly stalked; stylodia terminal, not artic- ing on indeterminate woody stems of 1 or more
ulate, hairy; stigmas minute, terminal, longitudi- years old), usually climbing, straggling or creep-
nally canaliculate; ovule 1 (see Note below), basally ing, rarely erect; only few species herbaceous,
inserted. Fruits achenes, crowned by the persistent perennial. Twigs and other parts nearly always
and elongated, hairy stylodia. Seed with membra- with prickles. Leaves pinnately, palmately, or
nous testa, endosperm 0. 2n = 28. One species, Fal- pedately compound or simple, when simple
lugia paradoxa (D. Don) Torr., in North America usually lobed; stipules free, on the base of the
from Utah to Texas and Mexico. In montane petiole or at the junction of twig and petiole,
shrubland. persistent or fugacious, rarely 0. Inflorescences ter-
In habit this genus reminds of Cowania in the minal, determinate, elaborately branched, or
Dryadeae, but it differs conspicuously by the pres- little or not branched and in axillary bundles,
ence of an epicalyx. According to Juel (1918: p. 61, rarely strongly reduced and flowers (sub)solitary.
figs. 115-119) there are two ovules, one long- Flowers usually 5-merous, mostly bisexual, rarely
stalked and mostly not fertile, one sessile and unisexual and the plants dioecious. Hypanthium
fertile. I have investigated several specimens but saucer- to cup-shaped. Epicalyx 0. Sepals imbri-
have not found this situation. cate, often unequal, exposed margins often lobed.
370 C. Kalkman
Petals white, cream, pink, purplish or red, in few from the wild (e.g. cloudberry, Rubus chamae-
species partly or entirely 0. Stamens many, rarely morus L. of sub g. Chamaemorus) or grown on a
few. Disk 0. Pistils many, rarely few, free, on a small and often non-commercial scale (e.g. the
mostly elevated torus; stylodia (sub )terminal; wineberry, Rubus phoenicolasius Maxim. of subg.
stigmas capitate or bifid; ovules 2, inserted almost Idaeobatus).
at same height but situated one above the other, Nitrogen fixation in root nodules has been
usually only the lower one developing, descendent, established in Rubus ellipticus J.E. Sm. from Java,
epitropous. Fruits drupes, exocarp black, red, see Becking (1979), so Frankia symbiosis in
yellow, orange or white, mesocarp juicy or fleshy, Rosaceae seems not be to be restricted to the
endocarp hard, drupelets cohering and falling as a Dryadeae.
collective fruit without or together with the dry
torus, rarely separating individually. Seed with
thin testa, endosperm scarce. 2n = 14 and multi- 12. TRIBE ROSEAE Focke (1888).
ples, recorded up to 2n = 98. Apart from several
thousands of 'microspecies' in subg. Rubus (see Woody. Leaves once pinnate (exception: Rosa
below), possibly about 250 species. Distribution persica); stipules adnate to petiole (exception:
almost cosmopolitan in temperate climates. Rosa persica). Epicalyx 0. Stamens many. Pistils
Several species have (intentionally or accidentally) many, entirely enclosed in hypanthium, fleshy
been introduced to places outside their natural after anthesis; ovule 1, pendulous. Achenes. x = 7.
area; sometimes they have naturalized completely
and some have become noxious weeds. Originally
54. Rosa L.
in natural open habitats but also in forest, now
in many anthropogenic vegetations like hedges, Rosa L., Sp. Pl.: 491 (1753).
shrubberies, etc. Hulthemia Dumort. (1824).
The revision by Focke (l.c.) is now outdated and x Hulthemosa Juz. (1941).
especially the infrageneric subdivision should be
studied anew. There are three large subgenera: Shrubs, erect, climbing, straggling, or prostrate,
Rubus, Malachobatus (Focke) Focke, and Idaeoba- evergreen or leaf-shedding, often glandular-hairy,
tus (Focke) Focke. Several monotypic to species- branches and petioles nearly always armed with
poor subgenera have also been recognized, e.g. straight or curved prickles and often also with
Anoplobatus Focke, Chamaebatus (Focke) Focke, thinner, straight bristles, often paired prickles
Chamaemorus Focke, Cylactis (Raf.) Focke, Dal- under the leaves. Leaves alternate, imparipinnate,
ibarda (L.) Focke, of which the taxonomic and leaflets penninerved, in subg. Hulthemia Dumort.
phylogenetic status has to be reviewed. Sometimes leaves reduced to one leaflet and without stipules.
these subgenera were recognized as separate Stipules in lower part adnate to the petiole, upper
genera. parts free. Inflorescence a terminal thyrsus or
Subgenus Rubus is taxonomically the most raceme, or flowers solitary and terminal. Flowers
problematic one. It comprises a group, variously (4)5-merous, bisexual, large and showy, often fra-
called 'Eubatus', the 'Moriferi', or the 'Rubus grant, rarely functionally unisexual and plants
fruticosus/caesius complex', that consists of sexu- cryptically dioecious (see Kevan et al., Biol. J. Linn.
ally reproducing, self-sterile, diploid species and a Soc. 40: 229-243, 1990). Hypanthium globular to
larger number of facultatively apomictic poly- urceolate, throat usually almost closed by a thick-
plaids of hybrid origin. The complex is repre- ened annular disc. Epicalyx 0. Sepals imbricate, the
sented in Europe and North America and it has exposed margins usually pinnately incised, per-
been studied most extensively in Europe where c. sistent or caducous after flowering. Petals imbri-
2000 species have been described. See Nybom cate, in cultivars often more than 5, large, red,
(1988) for a recent discussion on apomixis versus purple, white, or yellow. Stamens very many, on
sexuality in the subgenus. Apomixis seems to be relatively short filaments. Pistils usually many, on
absent or very rare in the other subgenera. hairy bottom of hypanthium and on the lower part
The fruits are edible. Blackberries, dewberries of its wall, shortly stalked or subsessile; stylodia
and loganberries (selections from species and terminal, free or their upper parts coherent to
crosses in subg. Rubus), and raspberries (cultivars connate, protruding through the orifice in the
of Rubus idaeus L. and crosses with other species disk; stigmas capitate. Fruits achenes with bony,
in sub g. Idaeobatus) are commercially cultivated leathery or woody peri carp, remaining enclosed in
in many places. Other species are only harvested the accrescent, ± fleshy hypanthium (hip). Seeds
Rosaceae 371
with thin testa, endosperm 0. 2n = 14 or poly-

plaids, recorded up to 2n =56. More than 100 wild
species, in the temperate parts of the Northern
Hemisphere, richest in Asia, in some places
(Philippines, Ethiopia, Mexico) extending into the
mountains of the tropical zone. In forest edges,
shrubland, riverbanks, etc., usually not in closed
forest.
Two species (Rosa roxburghii Tratt. from China
and Japan, R. minutifolia Engelm. from North
America) have been considered the types of two
subgenera, resp. Platyrhodon Rehder (probably
monotypic) and Hesperhodos (Cockerel) Rehder
with maybe two species. They are rather ordinary
roses but differ conspicuously from the other
species by having the hypanthium armed with
stout, straight prickles. The two species do not
seem to be closely related. Recognition of these
subgenera does not seem taxonomically useful.
A third aberrant species is Rosa persica Juss.
(R. berberifolia Pall.) from the Iran-Afghanistan Fig. 115. Rosaceae. Alchemilla rothii. A Plant with stolon and
inflorescence. B Flower, seen from above, the stylodia drawn
region, Kazachstan and Uzbekistan. Apart from to one side. C Flower, halved to show pistil. D Pistils. (Graham
having armed hypanthia, it has peculiar and very 1960)
strongly reduced leaves: unifoliolate and without
stipules. It is the type species of subgenus
Hulthemia (Dumort.) Focke and the genus the latter that I prefer to give Alchemilla s.l. its own
Hulthemia Dumort., and there may be sufficient group. The distinction is stressed by the difference
reason for a subgeneric status in this case. See also in chromosome base number. In the tree generated
Zielinski (1980). by the molecular analysis of Morgan et al. (1994),
Subgenus Rosa is usually divided into some ten the one species of Alchemilla is associated with
sections but its taxonomy is difficult for several Potentilla (1 species) and Fragaria. Hutchinson
reasons: many species cultivated, others very vari- (1964) placed the genus in tribe Sanguisorbeae. A
able and split by some into too many species, poly- name on tribal level does not seem to exist.
ploidy, apomixis, deviating reproduction methods
(like in Caninae where the male gametes have
n = 7, the female gametes n = 21, 28, or 35). 55. Alchemilla L. Fig. 115
Many of the garden roses have an untraceable Alchemilla L., Sp. Pl. 123 (1753); S. Froehner in Hegi, Ill. Fl. Mit-
breeding history with lots of undocumented inter- teleuropa, ed. 2, IV, 2B: 13-248 (1990), regional rev.
specific hybridisation since times immemorial and Aphanes L. (1753).
still going strong. Most of the cultivars cannot be Lachemilla (Focke) Lagerh. (1894); L.M. Perry, Contrib. Gray
brought to one species and must bear only the Herb. 84: 1-57 (1929), rev.; W. Rothmaler, Trab. Mus. Nac.
Cienc. Nat. Madrid,Ser. Bot. 31:52 pp. (1935),survey Colom-
genus name and the cultivar epithet. Cultivar bian spp.; Romoleroux in Fl. Ecuador 56: 89-133 (1996), 24
groups have not yet been formalized. spp.
Zygalchemilla Rydb. (1908).
13. ALCHEMILLA GROUP Perennial herbs with woody caudex, annuals,

or small shrubs, unarmed, evergreen or leaf-
Leaves usually palmately structured; stipules shedding. Leaves often partly in a radical rosette,
adnate to petiole. Hypanthium closed by disk. palmately structured, entire or lobed, folded in
Epicalyx +, exceptionally 0. Petals 0. Stamens and bud; stipules adnate to the petiole, large, lower
pistils few to 1. Stylodium basal; ovule 1, ascend- part membranous and circling the stem, upper
ing. x = 8. part leaf-like. Inflorescences cymose, never truly
Alchemilla has often been placed in tribe Poten- terminal on the main stems, ranging from rich-
tilleae but the closed hypanthium of the former flowered thyrses to reduced bundles in the axils of
contrasts so much with the open hypanthium of cauline leaves. Flowers small, 4(5)-merous, bisex-
372 C. Kalkman
ual. Hypanthium ± urceolate, throat almost closed some characters in common, is not an obvious
by the disk. Epicalyx segments usually smaller solution because of their distribution. Lacking a
than sepals, rarely epicalyx 0. Sepals valvate. Petals thorough modern phylogenetical analysis to lay
0. Stamens 1-4{5), ante- or alternisepalous, pollen the foundation for a definitive classification, it
in many species sterile and reproduction apoga- seems the best solution to keep the three groups
mous. Disk intra- or extrastaminal, nectariferous. together in one genus.
Pistils 1-4{-12), free, more or less stalked. Fruits When the genus is thought to be related to the
achenes, remaining enclosed in the dry, more or Potentilleae, the palmately structured leaf is con-
less hardened hypanthium. Seeds with thin testa, sidered to be homologous with the apical part of
endosperm 0. 2n = 16 or higher, mostly high- the pinnate leaves of genera in Potentilleae. One
polyploid, reported up to 2n = c. 224, aneuploid species, Alchemilla pinnata Rydb., has distinctly
numbers common. A genus of more than 1000 pinnate leaves and is the type species of mono-
species, many of which in cold- to warm- typic Zygalchemilla Rydb. The latter genus is
temperate Eurasia including the Mediterranean usually synonymized with either Lachemilla or
region, few reaching Sri Lanka and Java, East Alchemilla s.l.
Africa, South Africa and Madagascar; few in In Alchemilla s.str., and possibly also in part of
northern North America, many in South America, Lachemilla, the species are apomictic and have
one indigenous in Australia. Usually in open, stony sterile pollen. Recently Froehner (l.c.) studied
or grassy places, also (high) montane, some weedy. the European species and recognized 137 species
The genus has not been monographed in the last in Central Europe, stating that there would be
century. Rothmaler published extensively on it and certainly more than 1000 over the entire area.
first recognized the genus in the widest sense, According to him, hybridisation between the
with three subgenera Alchemilla, Aphanes, and (micro)species does never occur (with only one
Lachemilla. Later (Rothmaler 1937) he elevated established exception!), although it played a major
them to their original generic rank. They seem to role in the history of the group and in establishing
be well recognizable: the multitude of forms. Other authors also see
apomixis in this genus as obligatory, which would
Subg. Alchemilla. Perennial herbs or shrubs; contrast with the facultative apomixis in other
stamens 4, alternisepalous; disk intrastaminal; up genera.
to 4 pistils. More than 1000 (micro)species (see
Note below). Temperate Northern Hemisphere,
few spp. in montane tropical Asia, East Africa, 14. TRIBE PRUNEAE Hutch. {1964).
Madagascar, South Africa.
Woody plants. Leaves alternate, simple, pennin-
Subg. Lachemilla. Perennial herbs or shrubs; erved. Hypanthium open at top. Epicalyx 0. Disk 0.
stamens 2, antesepalous; disk extrastaminal; pistils Pistil(s) on bottom of hypanthium, free from it;
1-12. About 80 species. Only in Mexico, Central ovules 2, descendent and epitropous, over a con-
and South America, in Andes up to 5000 m siderable length attached to the ventral wall, obtu-
altitude. rator distinct. Fruit drupaceous. Seed usually 1 per
pistil developing, endosperm 0. x = 8. Exochorda is
Subg. Aphanes. Annual herbs; stamen 1, ante- often considered to be closely related, see there.
sepalous; disk extrastaminal; pistil 1. About 20
species. Remarkably scattered area: Mediter-
56. Oemleria Rchb.
ranean area, adventitious in large part of Europe,
Atlantic islands, North America (Southeast and Oemleria Rchb., Deutsche Bot., Herb. Buch: Index 195, Emend.
West), South America (Pacific & Atlantic), Aus- 236 (1841).
Nuttalia Torr. & A. Gray (1840), nom. illeg., non Raf. (1817).
tralia (1 species), Ethiopia (1 species). Osmaronia Greene (1891).
Distinction as three genera seems at first sight
indicated, but there remains considerable doubt
Shrubs or small trees, leaf-shedding. Leaves with
about Aphanes. Its area suggests that this is an old
entire margins; stipules 0 or rudimentary. Inflores-
residual group but its distinguishing characters cence a short raceme, axillary(?) on the short
are certainly apomorphic. So, it may be an unnat-
shoots, with large bracts. Flowers fragrant,
ural grouping of species derived from different 5-merous, unisexual, plants dioecious. Hypan-
lineages within Alchemilla s.str. or Lachemilla. A
thium campanulate. Sepals valvate. Petals shortly
fusion of Aphanes and Lachemilla, which have
Rosaceae 373
clawed, white. Stamens 15, inserted in 3 series, small Fig. 116. Rosaceae-Pruneae. Prunus arborea. A Branch with
and sterile in female flowers. Pistils 5 in female leaves. B Stipules with glands. C Fascicled racemes. D Halved
flower. E Infructescences. Drawn by R. van Creve!. (Kalkman
flowers, free, antesepalous, none in male flowers; 1993)
stylodia (sub )terminal; stigmas dilated. Fruits
drupes, usually only one or two developing in one
flower, exocarp black-blue, mesocarp thin, fleshy, Trees or shrubs, rarely with thorns, evergreen
endocarp leathery, smooth. Seed with membra- or leaf-shedding. Leaves with incised or entire
nous testa. 2n = 16 (only one record seen: Moffett margins, usually with glands in the margin or on
1931: 443, footnote). One species, Oe. cerasiformis the underside of the lamina or on the petiole; stip-
(Hook. & Arn.) J.W. Landon, in the western part ules on the twigs at the junction with the petioles,
of North America from British Columbia to Cali- free, rarely connate. Inflorescence basically a
fornia. Thickets, etc., from sea level to montane raceme, rarely branched, often reduced to a few-
altitudes, see Antos and Allen (1990). flowered umbel or to only 1 or 2 flowers. Flowers
normally 5-merous and bisexual. Hypanthium
57. Prunus L. Fig. 116 usually partly or entirely caducous after flowering.
Sepals imbricate, usually caducous. Petals white
Prunus L., Sp. Pl.: 473 (1753); Ghora & Panigrahi, Fam. Rosac. or pink. Stamens c. 15-many. Pistil1, sessile; sty-
India 2: 5-171 (1995), regional rev.; Bortiri et al., Syst. Bot.
26: 797-807 (2001), molec. analysis. lodium terminal; stigma capitate. Fruit a drupe,
Pygeum Gaertn. (1788); Kalkman, Blumea 13: 1-115 (1965), mesocarp in wild species thin, fleshy or juicy to
reduction. rather dry, sometimes dehiscent (almonds), endo-
374 C. Kalkman
carp forming a bony to woody, smooth or rugose stigma capitate. Exocarp purple, mesocarp pulpy
pyrene. Seed with thin testa. 2n = 16, 24, 32, 48, 56, to fleshy, leathery when dried, endocarp (slightly)
64, and aneuploid numbers. At least 200 spp., cos- woody. Seed subbasally attached, with membra-
mopolitan (in Africa only very few species indigenous testa. 2n = 32, once reported as 2n = 28 (see
nous). In forests and open vegetation, often in Missouri Bot. Gard. Monogr. Syst. Bot. 30, 1990).
montane or even subalpine altitudes. Three (or maybe four) species, in China, Mongo-
Subdivision is controversial; a distinction of lia, Himalayas, Taiwan. Thickets on slopes and in
five subgenera seems the most useful one to valleys, montane, up to c. 3000 m altitude.
me: Prunus, Amygdalus (L.) Focke, Cerasus (Mill.) Note: As to generic delimitation in tribes/groups
Focke, Padus (Mill.) Focke, and Laurocerasus 15-17 ('maloids') I fully agree with Weber (1964)
(Tourn. ex Duhamel) Rehd. who had stated: "The genera of the Maloideae are
Many species and species hybrids were domes- thus delimited mostly for the sake of convenience,
ticated long ago and are cultivated for the edible and there will always be disagreement among tax-
fruits. Several species and hybrids are grown as onomists as to how many genera and which ones
ornamentals. Also used in folk-medicine and should be recognized".
phytotherapy.
15. CYDONIA GROUP

58. Maddenia Hook. f. & Thoms.
Maddenia Hook. f. & Thoms., Hook. J. Bot. Kew Gard. Misc. 6: Woody. Leaves alternate, simple, penninerved;
381 (1854). stipules on junction of twig and petiole. Flowers
5-merous. Hypanthium with free rim above the
Small trees, leaf-shedding, unarmed. Leaves with pistil. Epicalyx 0. Carpels 5, dorsally adnate to
glandular-dentate margins; stipules large, on the hypanthium; stylodia terminal; ovules more than
junction of the petiole with the twig, caducous. 2 per carpel. Fruit a pome, flesh (meso carp)
Inflorescences terminal, short racemes, dense- hypanthial and carpellary, core (endocarp) not
flowered. Flowers bisexual. Hypanthium obconi- woody. Seeds with thin layer of endosperm. x = 17,
cal, circumscissile above base after anthesis, upper but chromosome number of two genera unknown.
part deciduous. Perianth segments c. 10, subequal This group of maloid genera is here distin-
or some more petal-like. Stamens 20-40. Pistil1 (2, guished as a separate unit characterized by the
0); stylodium terminal; stigma widened; ovules high number of ovules per carpel, although its
collateral, pendulous. Fruits drupes, mesocarp monophyly is not certain (see the general part
fleshy, endocarp woody. Seed with membranous under Phylogeny).
testa. 2n = ? Possibly only one species. Eastern part
of Himalayas, China, in montane and subalpine
altitudes. 60. Chaenomeles Lindl.
Chaenomeles Lind!., Trans. Linn. Soc. Lond. 13: 96, 97 (1821),
'Choenomeles', orth. cons.; C. Weber, J. Arnold Arbor. 45:
59. Prinsepia Royle 161-205,302-345 (1964), rev.
Prinsepia Royle, Ill. Bot. Himal. Mts. 1: 206 (1834), descr.; 2: t.
Shrubs, leaf-shedding, thorns axillary on long
38,1 (1839); Baronov, Taiwania 11: 99-112 (1965), rev.
Plagiospermum Oliv. (1886). shoots, collateral with the short shoots. Leaves
with incised margins; stipules large on long
Shrubs, leaf-shedding, thorny with 'stem-spines' shoots, very small on short shoots. Flowers few in
serially above the leaves and their axillary shoots. axillary fascicles or solitary, bisexual or staminate.
Leaves with distinctly incised margin to entire; Hypanthium campanulate, free upper part lined
stipules minute. Inflorescences racemes, some- with a nectary of which the rim is disk-like. Sepals
times only 1-flowered. Flowers 5-merous, bisexual. unequal, slightly imbricate when very young,
Hypanthium widely obconical to campanulate, deciduous with the upper part of the hypanthium.
persistent with sepals after flowering. Sepals Petals large, red (white in some cultivars), clawed.
imbricate or valvate. Petals narrowed at base, Stamens c. 20-60. Carpels 5, completely connate
white or yellow, fugacious. Stamens 10-30. Pistil and adnate to the hypanthium, forming an infe-
1, sessile; stylodium by unequal development of rior, 5-celled ovary; stylodia connate in basal part;
ovary becoming lateral and ultimately subbasal; stigmas thickened; ovules more than 20 per cell,
Rosaceae 375
axile; pistillode in staminate flowers minute. Pome 63. Pseudocydonia (C.K. Schneid.) C.K. Schneid.
with thick, hard flesh with sclereids, endocarp
Pseudocydonia (C.K. Schneid.) C.K. Schneid. in Fedde, Rep. 3:
forming one core, rather thin, bony. Seeds many, 180 (Nov. 1906).
dark brown, with firm testa. 2n = 34. Three species Chaenomeles sect. Pseudocydonia C.K. Schneid., Ill. Handb.
in the mountains of East Asia. Ch. japonica Lindl. Laubholzk. 1: 729 (May 1906), basionym.
is much cultivated as ornamental garden plant,
sometimes escaped and established. Shrub or small tree, unarmed, leaf-shedding.
Leaves with glandular-serrate margins; stipules
rather large. Flowers solitary on leafy short shoots,
61. Cydonia Mill. bisexual or staminate. Hypanthium in bisexual
Cydonia Mill., Gard. Diet., abr. ed.: 4 (1754). flowers narrowly campanulate. Sepals slightly
imbricate, caducous. Petals large, pink. Stamens 20
Shrub or small tree, unarmed, leaf-shedding. or more. Carpels 5, completely connate and also
Leaves entire; stipules large, long persistent. adnate to the hypanthium, forming an inferior, 5-
Flowers solitary, terminal on leafy shoots, showy, celled ovary; stylodia connate at the base and there
bisexual. Hypanthium narrowly campanulate, surrounded by a cup-shaped 'disc' on top of the
upper free part lined with a nectary. Sepals imbri- ovary; ovules many per cell. Pome not seen,
cate, glandular-dentate, soon reflexed, persistent. according to descriptions very hard with many
Petals white or pink. Stamens 15-25. Carpels 5, clusters of sclereids and with brown seeds. 2n = ?
dorsally adnate to the hypanthium, laterally and One species, P. sinensis (Thouin) C.K. Schneid., in
ventrally free from each other, forming an inferior, China.
5-celled ovary with a central cavity; stylodia free;
stigmas oblique; ovules many, in 2 rows, horizon-
tal, pleurotropous. Po me large, flesh with sclereids, 16. TRIBE MALEAE Schulze-Menz (1964).
endocarps forming one core, coriaceous. Seeds
several in each cell, with firm testa. 2n = 34. One Woody. Leaves alternate, penninerved. Inflores-
species, C. oblonga Mill., probably indigenous in cence terminal. Flowers 5-merous, bisexual.
the mountain areas of Caucasus and Kurdistan, Hypanthium adnate to carpels in lower part, upper
but since long in many places cultivated as a fruit part free (entirely free in Dichotomanthes). Epica-
tree (quince), sometimes naturalized. lyx 0. Stylodia terminal (exception: Dichotoman-
thes), sometimes fused to form a branched style.
Ovules 2 (exceptions occur, see Malus, Rhaphi-
62. Docynia Decne
olepis, Sorbus), normally basal and collateral. Fruit
Docynia Decne, Nouv.Arch. Mus. Hist. Nat. 10: 125,131 (1874). a pome (exception: Dichotomanthes, and see Note
Eriolobus sect. Docynia (Decne) C.K. Schneid. (1906). under Heteromeles), endocarp not woody. Seeds
with thin layer of endosperm. x = 17.
Trees, unarmed, leaf-shedding (sometimes semi-
evergreen?). Leaves sometimes lobed; stipules
small, fugitive. Inflorescences umbel-like with 64. Sorbus L.
(1)2-5 flowers, terminal on new shoots. Flowers Sorbus L., Sp. Pl.: 477 (1753); Aldasoro et a!., Syst. Bot. 23:
bisexual. Hypanthium tubular. Sepals valvate, 189-212 (1998), rev. Europ. & N Afr. spp.
persistent. Petals large. Stamens 30-50. Carpels Chamaemespilus Medik. (1789).
Torminalis Medik. (1789).
5, completely connate with each other and adnate Aria Host (1831).
with basal part of hypanthium, forming an almost Cormus Spach (1834).
inferior, 5-celled ovary; stylodia connate at base; Micromeles Decne (1874).
stigmas oblique; ovules 3-10 per cell. Pome with
hard and gritty flesh, with many sclereids, endo- Trees or shrubs, mostly leaf-shedding, unarmed.
carps bony, separated by flesh. Seed with firm Leaves simple or imparipinnate (in hybrids
testa. x = ? One species, D. indica (Wall.) Decne, in also intermediary), stipulate. Inflorescence a
eastern Himalayas, eastwards to North Thailand compound raceme, corymb- or panicle-shaped.
and South China. In montane forest. See also Hypanthium obconical to cupular, free upper part
under Macromeles in tribe Maleae, p. 378. lined with a ± obvious nectary. Sepals persistent
or deciduous. Petals white or pink. Stamens c. 20.
Carpels 2-5, dorsally adnate to hypanthium, apical
376 C. Kalkman
parts exposed or not, ventrally and laterally at least Subg. Chamaemespilus (Medik.) K. Koch. Leaves
in upper part usually free from each other, lower simple, nerves not running straight to the margin.
down completely connate; stylodia free or connate Sepals persistent. Carpels 2, carpel apices exposed,
in lower part; stigmas obliquely truncate; ovules stylodia free or slightly connate. Flesh of pome
2( -4) per cell. Po me variously coloured, flesh without sclereids, with large pigment cells. Only
(hypanthial and carpellary) with or without scle- species S. chamaemespilus (L.) Crantz in southern
reids, with or without large pigment cells; endo- and eastern Europe.
carp firm-membranous, leathery, bony or slightly
woody. Seeds with firm, leathery to membranous Subg. Torminaria (DC.) K. Koch. Leaves simple,
testa, endosperm thin. 2n = 34, 51, 68. About 150 deeply lobed, primary nerves running straight to
species, in Europe, Asia, and North America. In the margin. Sepals caducous, part of hypanthium
forests and more open habitats, from low altitudes persistent. Carpels 2 or 3, completely adnate to
to high montane. hypanthium, apices not exposed, stylodia basally
Sorbus presents a taxonomic problem of long connate. Flesh of pome with sclereids and large
standing, the two extreme possibilities being one pigment cells. Only species S. torminalis (L.)
genus or six, and several intermediary classifica- Crantz in the Mediterranean region, a second
tions also proposed by different authors. As many species described from Iran deserves no more
former authors had done, Hutchinson (1964) saw than varietal rank.
it as one genus. More recently, Robertson et al.
(1991) took an opposite position and recognized Subg. Micromeles (Decne) Phipps et al. Leaves
five genera. After careful consideration I decided simple, primary nerves running straight to
to recognize Sorbus as an inclusive genus. That so margin. Sepals caducous, free part of hypanthium
many species are of horticultural value and are often also. Carpels 3 or 4, carpel apices not
in the pertinent literature, mostly called by their exposed, stylodia basally connate. Flesh of pome
Sorbus name, has played a role in my decision. with sclereids and large pigment cells. 12-15
Distinction of six subgenera seems indicated, species, in Southeast and East Asia.
but with misgivings about some of them. They are
enumerated here, with the characters that have
65. Eriobotrya Lindl. Fig. 117B
been used as differential.
Eriobotrya Lind!., Trans. Linn. Soc. Lond. 13: 102 (1821).
Subg. Sorbus. Leaves imparipinnate, primary
nerves not running straight to the margin. Sepals Trees or shrubs, unarmed, evergreen. Leaves
persistent. Carpel apices and stylodia free. Flesh of simple, penninerved; stipules free, rarely intrapeti-
pome usually without sclereids and large pigment olarly connate. Inflorescence a compound raceme.
cells. The number of 'true' species is difficult to Hypanthium obconoid, open at top. Sepals persist-
assess because of extensive hybridization and ent. Petals white. Stamens 15-20( -40). Carpels 2-5,
apomixis in some complexes. The c. 80 species ventrally and laterally connate (in upper part ven-
names, enumerated in Phipps et al. (1990), may be trally free), dorsally adnate to the hypanthium, the
considered a maximum. Temperate Eurasia and hairy apex exposed; stylodia usually connate at
North America (south to Mexico). S. aucuparia L., base, hairy; stigmas capitate. Pome yellow, orange
grown as an ornamental. or red, flesh mostly of hypanthial origin, sclereids
0 or+, endocarps thin, membranous. Seeds 1-3,
Subg. Cormus (Spach) Duch. Differing from subg. large, with a thin but firm testa; endosperm 0,
Sorbus in: sepals deciduous but hypanthium per- cotyledons thick. 2n = 34. About 15-20 species,
sistent. Flesh of pome with sclereids, without large from the Himalayas throughout continental south-
pigment cells. S. domestica L. from warmer parts east Asia to Japan and the islands of western
of Europe, the fruits edible when very old. Malesia. Montane or in tropical lowland forest. E.
japonica (Thunb.) Lindl. widely cultivated (loquat).
Subg. Aria (Pers.) Beck. Leaves simple, primary
nerves running straight to the margin. Sepals
66. Aronia Medik. Fig. 117A
usually persistent. Carpels 2 or 3, carpel apices
usually free from hypanthium, stylodia free. Flesh Aronia Medik., Philos. Bot. 1: 155 ( 1789), nom. cons.; Robertson
of pome without sclereids, with large pigment eta!., Syst. Bot. 16: 391 (1991), united with Photinia.
Photinia Lind!. (1821).
cells. About 50 species, in Europe, North Africa and Stranvaesia Lind!. (183 7).
temperate to tropical Asia. Pourthiaea Decne (1874).
Rosaceae 377
A B c D
Trees or shrubs, unarmed, evergreen or leaf- Fig. 117. Rosaceae-Cydonia group, flowers and fruits. A
shedding. Leaves penninerved; stipules free, small Aronia sp. (= Photinia davidiana). B Eriobotrya bengalensis.
C Rhaphiolepis philippinensis. D Macromeles corymbifera.
and caducous. Inflorescence a panicle- or corymb- (Kalkman 1993)
shaped compound raceme. Hypanthium obconoid
to campanulate, open at top. Sepals long persistent,
but ultimately often (partly) weathering off. Petals
white to pinkish. Stamens 15-25, often 20. Carpels Asian disjunction. They correctly state that flowers
2-5, ventrally and laterally completely connate, and fruits of the two genera are indistinguishable,
dorsally adnate to the hypanthium but the apical but they kept the genus under the younger name
part free and protruding as a usually hairy dome; Photinia. Their taxonomic decision is followed
stylodia entirely free or connate at base; stigma here but the generic name Photinia had to be rel-
slightly broadened, truncate. Pome red or black, egated to synonymy. I am still uneasy about the
flesh (almost) entirely of hypanthial origin, inner monophyly of the enlarged genus Aronia/ Photinia
layer with sclereids, carpel wall rather thin, core and a worldwide monograph is badly needed. The
bony. Seeds usually several per fruit, rather small, Asiatic species of Photinia will generally not have
testa firm, endosperm thin or sometimes 0. 2n = a synonym under Aronia.
34, 68. Some forty species, in East Asia, and North
& Central America. In forest and thickets, in low as
67. Rhaphiolepis Ker Gawl. Fig. 117C
well as montane and subalpine altitudes. In China
40 and more species have been recognized but not Rhaphiolepis Lindley ex Ker Gawl., Bot. Reg. 6: t. 468 (1820),
all of them will survive closer taxonomic scrutiny. nom. et orth. cons.
Some species cultivated as ornamentals.
Photinia was described in 1821 from Asia, and Small trees or shrubs, unarmed, evergreen. Leaves
subsequently enlarged by description of several simple, penninerved; stipules on extreme base of
Asian and North and Central American species. petiole, free. Inflorescence a compound, rarely
Robertson et al. (1991) merged the small Ameri- simple raceme. Hypanthium obconoid, the free
can genus Aronia (indigenous inEast Canada and part at inside lined with an intrastaminal disk.
the eastern half of U.S.A.) with Photinia, thereby Sepals caducous after flowering, together with
creating a new example of the East-American/East the upper rim of the hypanthium. Petals white to
378 C. Kalkman
pinkish. Stamens 15-20. Carpels 2, completely lobed; ovules 2, basal, collateral, erect, apotropous.
connate with each other and adnate to the hypan- Fruit an achene, slightly exserted from enlarged
thium; stylodia free or basally connate; stigmas and fleshy hypanthium, pericarp coriaceous to
truncate; ovules normally 2 per carpel (rarely woody. Seed not observed. 2n = 34 (Gladkova, l.c.).
more, Sterling 1965c). Pome (sub)globular, with a One species, D. tristaniifolia Kurz, in tropical parts
distinct circular scar at top, purplish or bluish of China. Montane on open slopes, forest and
black, flesh rather thin, with many sclereids, carpel forest margins.
walls thin, membranous. Seeds 1 or 2 per fruit, This, of course, is a deviating genus but the rela-
large, globose; testa thin but firm, endosperm 0, tionship with Heteromeles seems obvious to me.
cotyledons thick. 2n = 34. Some 5 species, in The latter has two carpels in the flower, basally
Southeast and East Asia, see Ohashi (1988). In adnate to the hypanthium, while Dichotomanthes
forest, low and montane altitudes. has only one, entirely free carpel. In both cases the
hypanthium becomes fleshy and the fruit an
achene. A separate subfamily certainly does not
68. Heteromeles M. Roem.
seem justified, and a likeness to Prunus is only
Heteromeles M. Roem., Fam. Nat. Syn. Monogr. 3: 100, 105 superficial. Juel (1927) placed the genus in a sepa-
(1847); Phipps, Can. J. Bot. 70: 2138-2162 (1992), rev. rate tribe near his Quillajeae and considered
Vauquelinia (here in the Exochordeae) as the
Small trees or shrubs, unarmed, evergreen. Leaves nearest relative.
simple. Inflorescence a panicle-shaped compound
raceme. Hypanthium obconical. Sepals persistent.
Petals white. Stamens 10. Carpels 2, free from each 70. Macromeles Koidz. Fig. 117D
other except at entire base, the lower half or less Macromeles Koidz., Fl. Symb. Or. As.: 53 (1930).
adnate to hypanthium, the free tops conical, long- Malus sect. Docyniopsis C.K. Schneid. (1906), validly
hairy; stylodia free; stigmas capitate. 'Pome' (see published?
Note below) red or yellow, crowned by apical rim Docyniopsis (C.K. Schneid.) Koidz. (1934).
of hypanthium and persistent sepals, carpel walls
leathery, surrounded by and in the lower half Trees, leaf-shedding, sometimes thorny? (not
adnate to the fleshy hypanthium, no sclereids in observed). Leaves simple; stipules caducous. Inflo-
the flesh. Seeds 1 or 2 (maybe sometimes more), rescence a few-flowered raceme. Hypanthium
testa firm, endosperm 0. 2n = ? One species, H. obconical. Sepals long persistent but ultimately
salicifolia (C. Presl) Abrams (usually incorrectly deciduous. Petals white. Stamens 30-50. Carpels 5,
called H. arbutifolia (Lindl.) M. Roem.), in western dorsally adnate to hypanthium but apical part
states of U.S.A. and Northwest Mexico. In sclero- forming a small, free cone, laterally connate, ven-
phyllous scrub and (pine) woods at montane alti- trally with open sutures; stylodia connate only at
tudes up to c. 2000 m. very base, cohering by hairs in lower part; stigmas
The inadequacy of the term pome is well illus- not broadened. Pome globose, yellowish (always?),
trated in the fruits of this species: actually they are flesh hard, with a layer full of sclereids round the
'spurious' fruits consisting of a fleshy hypanthium core, the core (en do carps) cartilaginous, thin.
and two practically free, indehiscent achenes with Seeds 1 or 2 per cell, large, with thick and soft testa,
a leathery pericarp. endosperm very thin. 2n = ? Three species of
which the Japanese M. tschonoskii (Maxim.)
Koidz. is sometimes cultivated as an ornamental.
69. Dichotomanthes Kurz Other species include one or two Chinese species
Dichotomanthes Kurz, J. Bot. 11: 194 (1873); Gladkova, Bot. and the East Asian Pirus doumeri Bois, which does
Zhurn. 54: 431-436 (1969), new subfamily. not have a valid name in Macromeles. It was
treated as Malus doumeri (Bois.) Chev. by Vidal in
Trees or shrubs, leaf-shedding, unarmed. Leaves Fl. Camb., Laos &Vietn. 6:99-101 (1968). Growing
simple, entire. Inflorescence a compact compound in forest at montane altitudes up to 2000 m.
raceme, the last branches dichasial. Flowers
scented. Hypanthium cup-shaped, open. Sepals 71. Malus Mill.
persistent. Petals cream-coloured to white.
Stamens 20, on rim of disc. Pistil 1, at base of and Malus Mill., Gard. Diet., abr. ed.: 4 (1754).
free from hypanthium; ovary 1-locular; stylodium
slightly lateral, often sigmoid; stigma capitate, Trees or shrubs, leaf-shedding, rarely evergreen
and/ or thorny. Leaves simple, lobed or not; stipules
Rosaceae 379
caducous. Inflorescences few-flowered racemes thium obconical. Sepals reflexed, persistent. Petals
terminal on short shoots. Hypanthium obconical white. Stamens c. 20. Carpels 5, dorsally adnate to
to campanulate, upper part persistent. Sepals per- hypanthium and their apical part not exposed, lat-
sistent or deciduous. Petals white, pink or red. erally fully connate with each other, ventrally with
Stamens 15-20. Carpels 3-5, dorsally over their open sutures; stylodia connate at base, the connate
entire length adnate to hypanthium and also the part persistent; stigmas capitate. Pome ± globose,
apical part not exposed, laterally connate, ventrally red or yellow, rather hard and gritty, with a layer
with open sutures (see Note below); stylodia rich in sclereids around the cartilaginous core
connate in lower part; stigmas slightly broadened (en do carps). Seeds 1 or 2 per fertile cell, often only
to capitate; ovules 2 (4?) per carpel. Po me ± 1 cell fertile, testa thin, endosperm thin. 2n = ?
globose, green to red or yellow, rarely lenticellate, Monotypic, containing only E. trilobatus (Poir.)
flesh soft and mealy to rather hard, without scle- Roem., in Southeast Europe and Southwest Asia. In
reids, the core (endocarps) cartilaginous to bony forest and open places, at altitudes up to c. 1500 m.
(even slightly woody). Seeds with thin but firm, This remarkable species has, apart from having
dark coloured testa, endosperm a thin layer. Chro- been recognized as a genus, been moving around
mosome number 2n = 34, 51, 68, 85. Between 30 in Crataegus (basionym), Pyrus, Sorbus, Cormus
and 50 species, northern temperate. In woods and and Malus. When not recognized as a genus, it will
thickets. be best in place in Malus from which it differs
The circumscription of this genus is difficult and chemically by not possessing the flavonoid phlo-
much disputed. See Notes under Macromeles, Eri- ridzin in the leaves but trilobatin (Williams 1982).
olobus, and Pyrus, genera with which Malus has The most important difference with Macromeles
often been united. Langenfeld (1971) considered seems to be that in Eriolobus the apical part of the
the Macromeles spp. (as sect. Docyniopsis) and Eri- carpels is not free from the hypanthium.
olobus spp. (as sect. Eriolobus) to be relicts of the
original Tertiary Malus from which the other sec-
73. Pyrus L.
tions evolved. Merging Malus with Sorbus, as has
also been advocated, does not seem very sensible Pyrus L., Sp. Pl.: 479 (1753); Browicz, Arbor. Korn. 38: 17-33,
or enlightening. conspectus, distr.
In the present circumscription two or three sub-
genera or sections may be distinguished: subg. Trees or shrubs, sometimes thorny, leaf-shedding.
Malus with unlobed leaves in Eurasia, and subg. Leaves simple, rarely lobed; stipules caducous.
Sorbomalus with often lobed leaves (at least on the Inflorescence a simple raceme, corymb-shaped,
long shoots) in Asia. A third subgenus or section, with few flowers. Hypanthium urceolate, free
Chloromeles, is North American. Its species have upper part with a cushion-like outgrowth (disc)
usually lobed leaves and, according to Robertson covering the top of the carpels, leaving a central
et al. (1991), "a dense layer of sclereids around the hole for the stylodia, ± persistent. Sepals persistent
core" of the pome, but I did not have the opportu- or caducous. Petals white, rarely pinkish. Stamens
nity to observe this myself in the limited material 15-30. Carpels 2-5, dorsally adnate to hypan-
at my disposition. If confirmed, it would be a suf- thium, laterally and ventrally at least in the apical
ficient reason to separate the group as a genus, or part often free from each other, the ventral sutures
it may lead to fusion of Macromeles and Brio lobus open; stylodia free; stigmas truncate to 2-partite.
with Malus. Pome pyriform to globose, brownish to yellow,
sometimes lenticellate, flesh containing many scle-
reids, these sometimes compacted to a woody
72. Eriolobus (DC.) Roem. layer, core (endocarps) membranous to cartilagi-
Eriolobus (DC.) Roem., Faro. Nat. Syn. Monogr. 3: 104, 216 nous. Seeds with thin testa, endosperm a very thin
(1847). layer. 2n = 34, 51, 68, and aneuploid numbers also
Pyrus sect. Eriolobus DC., Prodr. 2: 636 (1825), basionym; reported. Between 10 and 20 species, but an exact
Browicz, Arbor. K6rn. 14: 5-23 (1969), taxonomic survey,
determination of their number is difficult because
distribution.
of extensive natural hybridization and establish-
ment of escapes from cultivation. Eurasian, also
Trees, leaf-shedding, unarmed. Leaves simple,
in North Africa. In woods, thickets, hedges, also
deeply 3-partite, the terminal and lateral lobes
on sunny slopes. Pyrus communis L., P. pyrifolia
more or less deeply lobed again, rarely the leaf 3-
(Burm.f.) Nakai and some other species are culti-
foliolate; stipules caducous. Inflorescence a simple
vated as fruit trees.
raceme, corymb-shaped, with few flowers. Hypan-
380 C. Kalkman
74. Amelanchier Medik. Leaves simple, usually leathery; stipules rather

pe~sistent. Inflorescences cymes, rarely flowers
Amelanchier Medik., Philos. Bot. 1: 155 (1789).
Peraphyllum Nutt. ex Torr. & Gray (1840). solitary. Flowers rather small. Hypanthium obcon-
Nagelia Lind!. ('Nagelia') (1842), nom. illeg. non Rabenh. ical to campanulate. Sepals persistent, incurved
(1844). after anthesis. Petals erect or spreading, white or
Cotoneaster § Malacomeles Decne, Nouv. Arch. Hist. Nat. 10: pink. Stamens (6-)10-20(-30). Carpels (1)2 or
177 (1874), rank not indicated.
Malacomeles (Decne) Engl. in Engl. & Prantl, Nat. Pflanzen-
3( -5), ventrally and laterally entirely free from
fam., Nachtr. 1: 186 (1897). one another or very shortly connate at base, with
exposed, hairy apex, dorsally adnate to hypan-
Shrubs or small trees, leaf-shedding, unarmed. thmm; ovules 2 per carpel. Fruit yellow, red or
Leaves simple; stipules small, caducous. Inflores- black, flesh rather dryish, without sclereids; stones
cence a simple raceme. Hypanthium obconic to mostly 2 or 3, their upper parts free from hypan-
campanulate, free upper part saucer- to cup- thium. Seed with thin testa, endosperm 0. 2n =34,
shaped. Sepals persistent. Petals white, rarely pink. 51, 68, rarely 85, 102. Variously recorded as having
Stamens usually c. 20, sometimes fewer. Carpels c. 60 or up to c. 300 species, with widespread fac-
2-5, ventrally (partly) free from each other, later- ultative apomixis and hybridization. Distributed
ally more or less connate, dorsally more than half over Asia, with few species in Europe and few in
adnate to hypanthium, top dome-shaped and North Africa and Ethiopia. Tetraploids are far
glabrous or hairy, cells of the ovary usually divided more numerous than diploids, the latter are
by an incom~lete sept between the ovules; stylo- mainly in the Himalayas and part of China, see
dta free to htghly connate; stigma(s) truncate to Kriigel (1992a). In open, often dry vegetation, and
capitate. Fruit a small pome, red to black, rarely in thickets and forests, up to c. 4000 m altitude.
yellow, flesh thin, uniform of consistency, with Many species and hybrids are cultivated as orna-
sclereids. Seeds several, testa rather thick, mentals or as landscaping elements. For the dif-
endosperm a very thin layer. 2n = 34, 68, and ane- ference with Pyracantha, see there.
uploid numbers also reported. Maybe around 20
spp., northern temperate: North America, East 76. Pyracantha M. Roem.
Asia, Europe (one species), in America south to
Mexico and Guatemala. Extensive hybridization Pyracantha M. Roem., Fam. Nat. Syn. Monogr. 3: 104, 219
(1847).
between species and apomictic reproduction
established. In rocky places of montane altitude,
Shrubs, evergreen, mostly with thorns. Leaf-
and open woods. Edible fruits.
margins incised or entire; stipules small,
caducous, but on long shoots longer persistent and
17. TRIBE CRATAEGEAE Koehne (1890). sometimes leafy. Inflorescence a terminal corymb-
s?aped thyr~e. Hypanthium obconical. Sepals per-
Woody. Buds perulate. Leaves alternate, simple Sistent and mcurved after flowering. Petals white.
(pinnate in Osteomeles); stipules on petiole-base. Stamens c. 20. Carpels 5, ventrally and laterally free
Flowers 5-merous, bisexual (sometimes male from each other, with exposed, hairy apex, dorsally
flowers in Mespilus). Hypanthium open, lower part adnate to hypanthium; ovules 2 per carpel. Fruit
red, orange or yellow, flesh rather thin, without
adnate to carpels. Epicalyx 0. Carpels 2-5 (excep-
sclereids; stones 5, free, upper part exposed. Seed
tions: 1 in Chamaemeles, rarely 1 in some
with thin testa. 2n = 34. About three species (up to
Cotoneaster and some Crataegus ), dorsally adnate
10 reported in horticultural literature). Distrib-
to hypanthium; stylodia terminal. Fruit a fleshy
uted from southern parts of Europe to East Asia.
hypanthium containing 5-l pyrenes (stones);
Pyracantha coccinea M. Roem. much cultivated
pyrenes 1-seeded; endosperm a thin layer. x = 17.
with many cultivars.
The differences with Cotoneaster are slight and
75. Cotoneaster Medik. probably the number of carpels per flower and the
presence/absence of thorns are the only decisive
Cotoneaster Medik., Phil. Bot. 1: 154 (1789); Klotz, Beitr. Phy-
totax. Univ. Jena 10: 7-82 (1982), synopsis. characters. Up to 5 carpels have been reported as
occurring in some species of Cotoneaster but I
Shrubs, sometimes prostrate, rarely small trees, myself have not observed that number.
leaf-shedding or more or less evergreen, unarmed.
Rosaceae 381
77. Crataegus L. for two Japanese species. Usually cited to have 3

to 5 species but rather a single variable species,
Crataegus L., Sp. Pl.: 475 (1753); Phipps, Ann. Missouri Bot.
Gard. 70: 667-700 ('1983' 1984), E As. and N Am. spp.
0. anthyllidifolia Lindl., distributed in China
(Sic~uan to Guangdong), Taiwan, Ryu Kyu Islands,
Shrubs or small trees, often with thorns, with short Bonm Islands, and Hawai'i. Often indicated from
and Ion~ shoots, evergreen or leaf-shedding. Myanmar but I have not seen any specimen from
Leaves pmnately lobed or pinnatifid or 3-lobed, there and it is not present in Thailand or Indo-
rarely unlobed, those on flowering branches often China. Forest margins, mountain slopes, also on
u~like those on strictly vegetative branches;
dry places, up to 3000 m altitude but also near the
stlpules usually small, mostly early caducous. coast. Sometimes cultivated as an ornamental.
Inflorescence terminal, a corymb-shaped thyrse or
cyme. Hypanthium obconical to campanulate. 79. Chamaemeles Lindl.
Sepals persistent or caducous. Petals orbicular to
Chamaemeles Lindl., Trans. Linn. Soc. Lond. 13: 96, 104
obovate, spreading, white, pink or red. Stamens
(1821).
mostly c. 20, inserted on a disk-like rim, well
visible on the fruit. Carpels 1-5, when more than
Evergreen shrubs, unarmed. Leaf margins entire to
1 ventrally free from each other, laterally free, at
serrulate; stipules small, caducous. Inflorescence
least pa:tly, dorsally adnate to hypanthium, apex
an axillary compound raceme. Flowers small.
free, hairy; ovules 2, basal. Fruit ± globose, red,
Hypanthium obconical. Sepals persistent. Petals
black or yellow, crowned by the apical part of
clawed, small, pink to white. Stamens 10-18. Pistil
th.e hypanthium, flesh mostly thin and rather dry,
1, .monocarpellat~, lower half adnate to hypan-
without sclereids; stones 1-5, with thick, hard and
thmm; style termmal, rather thick, broadened at
woody walls, forming one more-celled stone. Seed
apex; stigma oblique; ovules 2, basal, collateral.
compressed, with thin but firm testa. 2n = 34, 51,
Fruit with thin flesh. Seed 1, cotyledons convo-
68. Between 100 and 200 species, although more
luted. 2n = ? One species, C. coriacea Lindl.,
than 1000 have been described, northern temper-
endemic to Madeira, on inaccessible places on
ate zone and in Central America, richest in eastern
cliffs and ravines.
North America. According to some sources also in
the Andes but most probably in error. In woods,
forest margins, thickets, in low and montane alti- 80. Hesperomeles Lindl.
tudes. Often planted as ornamental or in hedges.
Hesperomeles Lindl., Bot. Reg. 23: t. 1956 (1837).
The taxonomy is complicated by hybridization and
apomixis.
Shrubs or small trees, probably evergreen, with
th.orns or unarmed. Leaves only on long shoots;
78. Osteomeles Lindl. stlpules small, caducous. Inflorescence a terminal
leafy compound raceme. Hypanthium obconical:
Osteomeles Lindl., Trans. Linn. Soc. Lond. 13: 96,98 (1821).
Sepals persistent. Petals white or cream-coloured.
Stamens c. 20. Carpels 4-6, dorsally adnate to
Shrubs, often prostrate, rarely treelets, evergreen
hypanthium, ventrally free, laterally ± free from
(?), unarmed. Leaves imparipinnate; leaflets in
e~ch other, apex exposed; stylodia terminal, free;
5-15(-20) pairs; stipules small. Inflorescence a
stigma truncate; ovule 1, basal. Fruit red, flesh
termi~al corymb-shaped thyrse. Hypanthium
with sclereids; stones c. 5, walls thick, hard and
obcomcal to campanulate, covering the carpels
woody, upper parts exposed and free from upper
at ~pex and a~most entirely adnate to them. Sepals
rim of hypanthium, covered by sepals. Seeds not
pomted, persistent and erect on the fruit. Petals
observed. 2n = ? Probably about 10 species, in
spreading, white. Stamens c. 20. Carpels 5, ven-
South America (Andes from Colombia to Bolivia)
trally and laterally connate, dorsally adnate to
and Central America (Costa Rica, Panama). In
hypanthium, forming an inferior, 5-celled ovary;
cloud forest, up to 4000 m alt.
stylodia mostly long-hairy at base; ovule 1 per
By its single ovules and other characters con-
carpel, basal, ascending. Fruit white, flesh thin,
nected with Osteomeles and formerly included in
hard; stones 5, hard and woody, rather easily loos-
that genus. However, the leaves are quite different
ened from each other. Seed with membranous
and recognition as a separate genus seems
testa. 2n = 34; according to Goldblatt in Missouri
indicated.
Monogr. Syst. Bot. 8 (1984), 2n = 32 was reported
382 C. Kalkman
81. Mespilus L. Adenostomateae (Adenostomeae). In Hutchin-

son's description the pistil is said to contain only
Mespilus L., Sp. Pl.: 478 ( 1753 ).
one ovule, but Juel (1918) correctly described the
genus as having two collateral ovules. The combi-
Trees or shrubs, with thorns (in the wild) or
nation simple leaves/one pistil/two ovules is found
unarmed (mostly in cultivation), leaf-shedding.
in all Pruneae except Oemleria, in some maloids
Leaf-margins serrate; stipules caducous. Flowers
and in part of Neillieae. Purshia (Dryadeae) has
solitary, terminal on short shoots, large, some-
some resemblance but differs in having a basal
times male, beneath them rarely with 1-3 later
ovule and no intrastaminal disk. Zhang (1992)
or not developing lateral flowers. Hypanthium
placed the genus in his wood-anatomical phenetic
obconical. Sepals large, persistent. Petals white.
group V, together with Hagenia (Sanguisorbeae)
Stamens 30-40, on the rim of a disk covering the
and Sorbaria (Gillenieae), and also in his dado-
free part of the hypanthium. Carpels (4)5, com-
gram the three genera are closely associated.
pletely connate and adnate to the hypanthium,
Morgan et al. (1994), in their molecular-
forming an inferior, 5-celled ovary, apex hairy; sty-
phylogenetical analysis, found Adenostoma
lodia free (to halfway connate: Gams 1922); ovules
together with Sorbaria and Chamaebatiaria
2(3) per cell, only the basal one fertile. Fruit
(Gillenieae), so this option must not be neglected.
brownish, flesh mealy with many sclereids; stones
Juel (1918) placed the genus in a group derived
with thick, hard, woody wall. Seed 1 per cell, testa
from Quillajeae and consisting of Exochorda,
thin but firm. 2n = 34. One species, M. germanica
Lindleya and the Pruneae. I could not find any ref-
L., from southeast Europe to Iran, now widely cul-
erence to a published chromosome number.
tivated outside its homeland; the fruit edible when
overripe. Wood very tough.
From Arkansas in U.S.A. a second species has 83. Coleogyne Torr.
been described by Phipps (1990). From a phyto-
Coleogyne Torr., Smiths. Contrib. Know!. 6: 8 (1853).
geographical standpoint this is unlikely. I wonder
whether this single population could have origi-
Shrubs, leaf-shedding, branches spinescent. Leaves
nated from a crossing between a cultivated medlar
opposite, crowded on the short shoots, simple,
and some indigenous Crataegus species.
small, linear, entire, coriaceous and grooved below,
sessile; stipules small, persistent on twig. Flowers
solitary, terminal, bisexual, with 2-3-lobed pro-
Rosaceous genera of unclear tribal position
phylls close under the hypanthium. Hypanthium
low obconical. Epicalyx 0. Sepals 4, large, persist-
82. Adenostoma Hook. & Arn.
ent, yellow inside. Petals 0 (see below). Stamens
Adenostoma Hook. & Arn., Bot. BeecheyVoy. 3: 139 (1832). 20-40. Intrastaminal disk forming a tubular
'sheath' around the ovary. Pistil1, monocarpellate,
Shrubs, evergreen, resinous. Leaves simple, rigid; on bottom of hypanthium; style lateral, twisted,
stipules basally adnate to leafbase, minute on short exserted from disc, persistent; stigma linear,
shoot leaves. Inflorescence a panicle with short on ventral side of style; ovule 1, attached in the
spike-like branches. Flowers many, very small, middle, descending, epitropous. Fruit achene,
sessile, 5-merous, bisexual. Hypanthium funnel- pericarp coriaceous. Seeds not seen, no data in lit-
shaped, open at the top, 10-ribbed. Epicalyx 0. erature. 2n = 16, two counts, recorded by McArthur
Sepals mucronate, imbricate. Petals orbicular, and Sanderson (1985). One species, C. ramossis-
white. Stamens 10-15. Intrastaminal disk lobed. sima Torr. in North America (California, Nevada,
Pistil 1, monocarpellate, on bottom of hypan- Arizona, Colorado). Open and dry mountain
thium; ovary oblique with 'cushion' on one side; slopes, sometimes in pure stands.
style terminal but put aside by the dorsal cushion; The tubular disk is obviously a much derived
stigma capitate; ovules 2, inserted below the apex. character and of no help in finding the closest rel-
Fruit achene, covered by indurated hypanthium. ative. Kearney and Peebles c.s. (1960: 391) record
2n =? a specimen with two pale yellow petals. Because
Three species, in western North America. Dry of the opposite leaves and the absence of petals,
mountain slopes, often in pure vegetation, see Hutchinson (1964) placed the genus in Cerco-
Jepson (1936). carpeae but differences with Cercocarpus are large.
Hutchinson (1964) and Schulze-Menz in Mel- Dryadeae and especially Kerrieae are often men-
chior (1964) placed the genus in a separate tribe, tioned as possible places to accommodate the
Rosaceae 383
genus. Juel (1918), for example, saw Rhodotypos 85. Lyonothamnus A. Gray
as the closest relative and detected a likeness
Lyonothamnus A. Gray, Proc. Am. Acad. Arts 20: 291 (1885).
with Neviusia. Zhang (1992) put the genus in his
wood anatomical group I, together with Kerria,
Evergreen tree with exfoliating bark. Leaves oppo-
Petrophytum and Stephanandra (Neillieae), Rosa,
site, petioled, simple and entire to pinnatifid or
Rubus, and Sibiraea (Spiraeeae). The chromosome
pinnately compound. Stipules deciduous. Flowers
number suggests a relationship with Alchemilla,
many, perfect, in large terminal corymbose com-
also with sepals 4 and petals 0, but otherwise very
pound panicles, with short pedicels. Flower tube
different.
campanulate, free from ovary, subtended by 1-3
bractlets. Sepals 5, persistent. Petals 5, clawless.
84. Potaninia Maxim. Stamens c. 15, inserted on a wooly disk lining the
flower tube. Pistils 2, distinct, 1-loculed. Stylodia
Potaninia Maxim., Bull. Acad. Imp. Sci. St. Petersb. 27: 465
(1881). stout, with subcapitate stigma. Ovules and seeds 4
in each ovary, the ovules bitegmic. Fruit a pair of
Much branched, small shrub, up to 40 em. Leaves small woody follicles. Seeds flat. n = 27. One sp., L.
crowded in short shoots, alternate (opposite floribundus Gray, with 2 subspecies, on Santa
according to Hutchinson), imparipinnate with (1)3 Catalina Isl. and adjacent islands off the coast of
or 5 small leaflets with entire margins; petiole California.
persistent, spiny; stipules adnate to the petiole, This genus is mostly recognized as spiraeoid,
relatively large. Flowers solitary, axillary, minute, sometimes placed in Sorbarieae (Juel 1927), by
3-merous, bisexual. Hypanthium funnel-shaped. others in Quillajeae (Hutchinson 1964). In the
Epicalyx 0. Sepals acuminate. Petals white or molecular-based tree published by Morgan et al.
pinkish. Stamens 3, inserted on a swollen disk. (1994), it associates with Cercocarpus and Purshia
Pistil 1, monocarpellate, on the bottom of the (Dryadeae), although the molecular support for
hypanthium; style basal, stigma capitate; ovule 1, this relationship is not strong, and morphologi-
basal, ascending. Fruits achenes, pericarp light cally it has little in common with these genera
yellow. 2n = ? One species, P. mongolica Maxim., either. Morgan et al. (1994) therefore conclude that
in Mongolia and China (Inner Mongolia). In Lyonothamnus may be considered as an isolated
sandy desert. Locally used as cattle feed (camels descendant of the ancestral spiraeoid complex; it
a.o.). certainly would merit a more profound study.
There are some mystifications about the flower
structure. Hutchinson (1964) described the flower
as having an epicalyx as well as sepals and petals, Doubtfully Rosaceous genus
but Yi.i Te-tsun et al. in Fl. Reip. Pop. Sin. 37:
455-456 (1985) do not mention an epicalyx in Brachycaulos Dikshit & Panigrahi
their elaborate description (translated for me by Brachycaulos Dikshit ('Dixit') & Panigrahi, Bull. Mus. Natl.
Dr. Ding Hou) nor is it drawn in their plate. The Hist. Nat. Paris IV, 3, sect. B. Adansonia 1: 57-60 (1981).
place of the stamens also is reported in different
ways: antesepalous according to Hutchinson, ante- Compact dwarf herbs, all parts glabrous (but
petalous according to Yi.i Te-tsun et al. I have seen see stipules). Leaves simple, entire, small; stipules
only one very insufficient herbarium specimen membranous, fully adnate to petiole, with shortly
myself, and my interpretation of the flower frag- ciliated margins, brown, clothing the stems.
ments available is in agreement with the descrip- Flowers solitary, terminal, bisexual. Hypanthium
tion in Yi.i et al.: epicalyx 0, sepals 3, petals 3, flat. Epicalyx 0. Sepals 5, free, entire. Petals 5,
stamens opposite the petals. clawed, slightly exceeding the sepals. Stamens 5,
Hutchinson placed the genus together with Cer- alternipetalous. Disk not evident. Pistils 2, free, on
cocarpus and Coleogyne in a separate tribe Cerco- bottom of hypanthium; stylodia terminal; stigmas
carpeae but in my opinion the three genera are inconspicuous; ovules unknown. Fruit achene,
not closely related. The best place for Potaninia smooth. x = ? One species described: B. simplici-
may be tribe Potentilleae but more evidence is folius Dikshit & Panigrahi, from Sikkim, India, in
needed. the alpine belt (4575m alt.).
This plant is only known from one collection,
W.W. Smith 3993 (CAL). The authors ascribed it to
tribe Potentilleae and compared it with Chamaer-
384 C.Kalkman
hodos. I am not at all sure that it is rosaceous and Bowden, W. 1945. A list of chromosome numbers in the higher
if so, it almost certainly does not belong to Poten- plants II. Am. J. Bot. 32: 191-201.
Bramwell, D. 1978. The endemic genera of Rosaceae
tilleae. Dr. G. Panigrahi (in litt.) kindly informed (Poterieae) in Macaronesia. Bot. Macar. 6: 67-73.
me that other collections of this species have not Bramwell, D., Bramwell, Z.l. 1974. Wild flowers of the Canary
come to his knowledge and that duplicates do not Islands, x + 261 pp. London: Thornes.
seem to exist in other herbaria. Dr. K. Brummitt Brummitt, R.K. 1992. Vascular plant families and genera. Kew:
(K) suggested (pers. comm.) that it may belong to Royal Botanic Gardens.
Cambie, R.C., Ash, J. 1994. Fijian medicinal plants. Canberra:
Saxifragaceae. For the time being I consider it as a CSIRO.
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Samolaceae 387
Samolaceae
B. STAHL
Samolaceae Raf., Ann. Gen. Sci. Phys. Bruxelles 5: 349 (1820). VEGETATIVE MORPHOLOGY. Some species are
more or less woody at base and rhizomatous; the
Perennial or sometimes annual herbs, rarely sub- Mexican S. cinerascens is a dwarf shrub. The leaves
shrubs, often woody at base, sometimes rhizoma- are simple and have entire margins that usually are
tous; secretory system usually appearing as a fine, decurrent or sometimes expanded on the petiole,
purple or dark brown striation on inflorescence which thus is poorly demarcated from the blade.
axes or short irregular lines or dots on calyx or In many species, all or most of the leaves are con-
bracts; indumentum mostly lacking, when present densed into a rosette at base of the plant. Samolus
then consisting of stalked or sessile glandular junceus and S. porosus have stems without leaves
hairs. Leaves alternate, often in a basal rosette, or with reduced, scale-like leaves.
sometimes lacking on aerial parts, rarely reduced
to scales, simple, petiolate but with petiole often VEGETATIVE ANATOMY. Although mostly glab-
poorly differentiated from blade; blades spathulate rous, short protrusive or immersed glandular
or oblong, sometimes linear or reniform, mar- hairs occur on reproductive and vegetative parts
gins entire. Inflorescences racemes, corymbs or in some species. Samolus ebracteatus is glandular-
sparsely branched compound racemes; pedicels puberulous and glandular-punctate throughout a
often long and slender, often sub tended by a single large part of its distribution (Henrickson 1983).
bract inserted at base or midway on the pedicel, The protrusive hairs have usually one or two stalk
sometimes ebracteate; prophylls always lacking. cells only, and a few-celled head. In this and a few
Flowers 5-merous, hermaphrodite, the perianth other species, the corolla mouth and petal bases
perigynous, actinomorphic; calyx herbaceous, the are densely puberulous from stalked glandular
tube ± fused to the ovary, the lobes narrowly to hairs with two to four stalk cells and a single-
broadly ovate or triangular, persistent; corolla celled head.
white, sometimes pink or purplish, campanulate Although not always evident, a secretory system
or urceolate, lobes broadly ovate, imbricate in bud; seems to be present in all species.
stamens 5, opposite the corolla lobes, included or
sometimes somewhat exserted; filaments largely INFLORESCENCE STRUCTURE. The inflorescence
fused with the corolla tube, free parts as long as in Samolaceae is basically racemose, being devel-
the anthers or shorter; anthers narrowly sagittate oped as a raceme, corymb or irregularly and
to oblong, dithecal, opening introrsely by longitu- sparsely branched compound raceme, sometimes
dinal slits, the connective sometimes apically on a long peduncle. The pedicels are often long
produced; staminodes often present, linear to and slender. Prophylls are lacking. In some species,
narrowly triangular, inserted at the mouth of the the bract is inserted near the middle of the pedicel.
corolla tube, alternating with the lobes; ovary
semi-inferior, globose to ovoid; style well devel- FLOWER STRUCTURE. In several species, ligulate,
oped, shorter than to somewhat longer than the linear or obtrullate staminodes are inserted
ovary; stigma truncate to capitate; ovules numer- between the lobes at the mouth of the corolla.
ous, hemitropous, inserted in several series on a These structures are provided with vascular tissue
free central placenta. Fruit a ± globose, loculici- (Thenen 1911); vestigial anthers are lacking.
dally dehiscent capsule. Seeds small, angular; seed Floral development in S. valerandi was studied by
coat pale to dark brown, reticulate, two-layered, Dickson (1936).
with rhomboid crystals; endosperm abundant, cell
walls smooth; embryo short, straight, cotyledons PoLLEN MoRPHOLOGY. The pollen of one exam-
narrow. x = 12, 13. ined species, S. valerandi, is triocolporate, elliptic,
A monogeneric family of worldwide distribu- relatively small (12-17 )lm diam.), and has a psilate
tion, comprising about 12 species. ornamentation (Punt et al. 1974).
388 B. Stahl
FRUIT AND SEED. The fruit is a globose capsule

opening loculicidally by five equal valves. The
seeds are numerous, obconic or more or less
angular, light to dark brown, and reticulate; the
testa is two-layered and has rhomboid crystals.
The seeds lack starch but contain amyloid and
have a short embryo with narrow cotyledons.
AFFINITIES. The family consists of a single genus,

Samolus, which usually has been referred to Prim-
ulaceae. Within that family, Pax and Knuth (1905)
placed it in a separate tribe, Samoleae. Because of
similarities in floral characters, Bartling (1830)
and Mez (1902) suggested a close relationship
between Samolus and Maesa (Maesaceae). How-
ever, phylogenetic analyses (Anderberg and Stahl
1995; Anderberg et al. 1998) did not support a
close relationship with Maesaceae, although its
true position within primuloid taxa remained
obscure. However, in a study combining mor-
phological and DNA sequence data (Kallersjo c
et al. 2000), Samolus is placed as sister to
Theophrastaceae.
DISTRIBUTION AND HABITATS. Unlike other pri-

muloid taxa, Samolaceae are well represented in
the Southern Hemisphere, with three species in
temperate South America, four in Australia and
Tasmania, and one in South Africa. Four or five
species are confined to southern North America (S
USA, Cuba and Mexico). One species, S. valerandi,
is cosmopolitan.
Most species grow in damp, temporarily inun-
dated areas near lakes, ponds or rivers. Some
species grow in more or less saline habitats such
as small, brackish ponds near the seashore or in
salt marshes.
Fig. 118. Samolaceae. A, B Samolus ebracteatus. C, D S.
valerandi. A Habit, flowering plant. B Flower. C Flower in ver-
ECONOMIC IMPORTANCE. At least one species, S. tical section showing ovary, stamens and staminodes. D Fruit
valerandi, is a popular aquarium and terrarium with pedicel and bract. (Orig., A redrawn from Pax and Knuth
plant (Crusio 1981), but the family is otherwise of 1905)
no economic importance.
A single genus: Selected Bibliography
Anderberg, A.A., Stahl, B.1995. Phylogenetic interrelationships

Samolus L. Fig. 118 in the order Primulales, with special emphasis on the family
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Pflanzenreich IV. 237: 336-344 (1905); Lourteig, Lilloa 8: Anderberg, A.A., Stahl, B., Kallersjo, M. 1998. Phylogenetic
241-252 (1 942 ), Argentinean spp.; Crusio, Mededel. W.A.P. relationships in the Primulales inferred from rbcL sequence
2: 13-25 (1982), 6 :13- 16 (1984). data. Plant Syst. Evol. 211: 93-102.
Bartling, F.G. 1830. Ordines naturales plantarum. Gottingen:
Dietrich.
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Wasserpfi. 11: 3- 8.
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385-393.
Samolaceae 389
Henrickson, J. 1983. A revision of Samolus ebracteatus (sensu Punt, W., de Leeuw van Weenen, J.S., van Oostrum, A.P. 1974.
lato) (Primulaceae). SouthWestern Nat. 28: 303-314. Primulaceae. The Northwest European pollen flora, 3. Rev.
Kallersjo, M. et al. 2000. See general references. Palaeobot. Palynol. 17: 31-70.
Mez, C. 1902. Myrsinaceae. In: Engler, Pfianzenreich IV. 236. Then en, S. 1911. Zur Phylogenie der Primulaceenbliite. Studien
Leipzig: W. Engelmann, pp. 1-437. tiber den Gefassbiindelverlauf in Bliitenachse und Perianth.
Pax, F., Knuth, R. 1905. Primulaceae. In: Engler, Pfianzenreich Jena: G. Fischer
IV. 237. Leipzig: W. Engelmann, pp. 1-386.
390 T.D. Pennington
Sapotaceae
T.D. PENNINGTON
Sapotaceae Juss., Gen. Pl.: 151 (1789) ("Sapotae"). fused with it, or absent; ovary superior, 1-15( -30)-
Sarcospermataceae H.J. Lam (1925). locular, loculi usually uniovulate, rarely 2-5-
ovulate, placentation axile, basi-ventral or basal;
Trees or shrubs, rarely geoxylic suffrutices or style simple, included or exserted; style-head
lianas, sometimes spiny; branching usually sym- simple or minutely lobed. Fruit a berry or rarely a
podial; latex nearly always present in trunk, drupe, or tardily dehiscent by a single lateral valve;
branches and fruits, usually white, rarely yellow or pericarp fleshy or less frequently leathery or
blue; indumentum nearly always of malpighia- woody; seeds 1-many, globose, ellipsoid, oblong,
ceous hairs (simple in Delpydora). Leaves alternate, often strongly laterally compressed, testa usually
spirally arranged or distichous, less frequently smooth and shining, free from the pericarp, less
opposite or verticillate, simple, entire or very rarely frequently roughened, wrinkled or pitted and then
spinous-toothed; petiole rarely bearing a pair of often adherent to the pericarp; hilum adaxial, basi-
minute stipels; stipules +or 0. Inflorescence fasci- ventral or basal, narrow or broad, sometimes
culate or flowers occasionally solitary, axillary, extending to cover most or all of the seed; embryo
ramiflorous or cauliflorous; fascicles occasionally vertical, oblique or horizontal, with thin foliaceous
arranged along short leafless axillary, panicle-like or thick flat or plano-convex, usually free, cotyle-
shoots; fascicle base sometimes developing into dons; radicle included or exserted; endosperm+ or
short, densely scaly brachyblasts. Flowers bisexual 0. X= 10, 11, 12, 13, 14.
or unisexual (plants monoecious or dioecious), A pantropical family of 53 genera and about
actinomorphic; calyx a single whorl of 4-6 free or 1100 species, mostly in humid forest, but some
partly fused, imbricate, sometimes quincuncial genera (e.g. Sideroxylon, Argania) extending into
sepals, or 6-11 sepals in a closely imbricate spiral, semi-arid and arid regions.
or with 2 whorls of 2-4 sepals and then the outer
whorl valvate or only slightly imbricate; corolla MORPHOLOGY. All Sapotaceae are woody plants,
rotate, cyathiform or tubular, sympetalous; tube the great majority trees or shrubs. They are pre-
shorter than, equalling or exceeding the petals; dominantly plants of moist lowland forest below
petals 4-18, entire, lobed or partly divided or 1000 m altitude, and are commonly canopy trees or
divided to the base into 3 segments and then emergents. They are also well represented in semi-
median segment entire, 2 lateral or dorsal seg- arid zones where they occur as small trees or
ments entire, laciniate or shallowly or deeply dense multi-branched shrubs often equipped with
divided; stamens 4-35( -43 ), fixed in lower or sharp axillary spines derived from modified axil-
upper half of corolla tube or at the base of the lary shoots.
lobes, rarely free, in a single whorl opposite Bole section (fluted or cylindrical), presence and
the corolla lobes, or, when more numerous than the type of buttressing, bark type (fissured, scaling,
corolla lobes, some opposite and some alternate smooth) are often specifically constant, but less
with the corolla lobes, or sometimes several useful at higher levels of classification. Nearly all
stamens clustered opposite each lobe, or arranged species of Sapotaceae have fissured or scaling
in 2-3 alternating whorls within the corolla tube, bark, and only a few examples are known of
exserted or included; filaments often geniculate large Sapotaceous trees with smooth bark (e.g.
in bud, free, rarely fused into a staminal tube, or Micropholis guyanensis and Pouteria laevigata in
partially fused to the staminodes; anthers often tropical America.)
extrorse; staminodes 0-8(-12) in a single whorl The growth and branching pattern of the main
alternating with the stamens or fixed in the corolla axis and branches gives rise to the characteristics
lobe sinuses, simple or variously lobed, toothed or of the crown, and several architectural models rec-
divided, sometimes petaloid; disk annular or patel- ognized by Halle and Oldeman (1970) occur in the
liform, surrounding the ovary base and sometimes Sapotaceae.
Sapotaceae 391
1. Model of Aubreville. Branched trees in which but less frequently large, conspicuous and some-
the main axis is monopodia! with rhythmic times persistent (e.g. Madhuca spp.). Elsewhere
growth producing subverticillate branches and a in the family stipules are generally absent, with a
layered crown. The branches are plagiotropic and few notable exceptions, viz. Sarcosperma, Chro-
sympodial with a series of long, bare horizontal molucuma, Ecclinusa, Synsepalum, Englerophy-
shoots each terminated by a short, vertically ori- tum, Capurodendron and Tsebona. In these latter
entated section bearing densely clustered spirally cases the stipulate condition is well correlated with
arranged leaves. The vertically orientated part of other generic characters. The petiole is generally
a branch continues to grow slowly, and axillary unappendaged, but in a small number of species
inflorescences are produced indefinitely. The (some Sarcosperma and Pradosia) it bears a pair
most frequently occurring architectural type in of small, undivided appendages near the base or
Sapotaceae, found in Autranella, Baillonella, apex. They have been called auricles or stipels.
Vitellaria,Manilkara,Neolemonniera, Tieghemella, Their origin and function is not understood.
Palaquium, Madhuca, Aubregrinia, Englerophy-
tum, Pouteria, Synsepalum, Omphalocarpum. VEGETATIVE ANATOMY. The reader is referred to
Metcalfe and Chalk (1950). A family characteristic
2. Model of Corner. Unbranched treelets with is the presence of sticky, slow flowing latex in the
lateral inflorescences and indeterminate growth of cut bark, branches and fruit; its colour is usually
the shoot apex. Delpydora, Niemeyera. white, but in a few species it may be yellow
(Pouteria congestifolia) or blue-green (Niemeyera
3. Model of Raux. Branched trees with the trunk acuminata). The laticiferous ducts occur in the
monopodia! and orthotropous, with continuous leaf, where they usually accompany the veins, but
growth producing spirally arranged branches. The sometimes are also interspersed in the mesophyll.
branches are plagiotropic with distichous phyl- In the stem they can be found in the cortex,
lotaxis, and they can be either monopodia! or sym- phloem, and pith. They are also present in flowers
podial. This type follows taxonomic lines closely and fruits. An indumentum of malpighiaceous
and is usually consistent for a genus or section of hairs is present in all genera except Delpydora,
a genus. Micropholis, Payena, Sarcaulus. which has simple hairs. A common condition is a
mixture of appressed 2-branched hairs with erect
4. Model of Troll. In this type all axes are pla- or spreading simple hairs. In a majority of species
giotropic from early in their development. Certain the malpighiaceous hairs are more or less sessile,
axes, however, especially in the juvenile state, may with one long branch and one short branch, but
have a short orthotropic part. All branches have in others both branches may be well-developed.
horizontal growth and distichous phyllotaxis, and Less frequently the 2-branched hairs have a long
growth in height is achieved by the straightening stalk. Some species of Manilkara and Pouteria
out of the basal part of certain shoots after leaf fall. sect. Pouteria have a minute, closely appressed
Documented for Chrysophyllum cainito and prob- indumentum which forms a pellicle on the lower
ably in other species of this genus. leaf surface. The mesophyll often includes scle-
Phyllotaxis is usually spiral or distichous; op- renchymatous fibres; stomata are ranunculaceous
posite leaf arrangement is typical of Sarcosperma, and nearly always confined to the lower leaf
Pichonia and Leptostylis, but occurs sporadically surface. Cork in the axis arises superficially. Soli-
in many other genera, e.g. Pouteria, Pradosia, tary crystals are always present and often abun-
Sideroxylon. dant in the cortex. In the wood, vessels occur in
The leaves of all Sapotaceae are simple. The leaf loose, radial or oblique lines and often in multiples
margin is entire in all species except Chrysophyl- of 4 or more cells. The perforations are simple, and
lum imperiale and C. subspinosum (both South the intervascular pitting is alternate and usually
American), which are spinous-serrate. Leaf vena- small. The vessel elements are usually of medium
tion provides many characters of use at generic length, but sometimes are moderately or very
and sectional level. The principal venation types, long. The axial parenchyma is apotracheal and
which are defined in terms of the primary tends to be banded. The rays are heterogeneous
(midrib), secondary, tertiary and quaternary and 1-6 cells wide. The fibres have simple pits, but
veins, are craspedodromous, eucamptodromous pits may occasionally be bordered. Vasicentric tra-
and brochidodromous (Hickey 1973). Stipules are cheids are present in some genera. Silica is often
commonly found in tribes Mimusopeae and Iso- present in elements other than the vasicentric
nandreae. They are usually small and caducous, tracheids.
392 T.D. Pennington
INFLORESCENCE MORPHOLOGY. The inflores- sepals (Figs. 120C, 121D), but in Isonandreae,
cence is a simple fascicle of several to many pedi- which also have a biseriate calyx, the number of
cellate flowers arising directly from the leaf axil or corolla lobes is often 2-3 times that of the sepals
rarely reduced to a solitary axillary flower (e.g. (e.g. Madhuca) and occasionally more (e.g. Bur-
Manilkara zapota). ckella, Pouteria sect. Antholucuma). The division
Scattered throughout the family are species in of the corolla lobes into a median and two lateral
which the inflorescence is an axillary shoot of segments is characteristic of many members of
limited growth bearing several lateral fascicles, Mimusopeae and Sideroxyleae. In Mimusops,
each sub tended by a leaf or bract, and the fascicles Manilkara and related genera, the two lateral seg-
may themselves be reduced to a few or only one ments equal the median segment and are them-
flower. This type of compound inflorescence may selves sometimes subdivided to the base. The
be found together with the more usual simple arrangement of the various corolla lobe segments
axillary fascicle in the same species (e.g. Pouteria in the complex flowers of Mimusops and
ramiflora) or it may be constant within a species Manilkara is distinctive. The central segment
(e.g. Sideroxylon racemosum, Pouteria macahen- remains erect and clasps the opposing stamen,
sis ), and in Sarcosperma it is found in every species but the lateral segments spread horizontally
of the genus. In Sarcosperma the inflorescence axis (Figs. 120C, 121D). As the anthers are generally
may itself be further branched. extrorse, this arrangement ensures that they
Another modification of the simple axillary dehisce into the clasping corolla segment. In
fascicle occurs sporadically in several genera, e.g. members of Sideroxyleae, the lateral corolla lobe
Micropholis. Here the fascicle produces flowers segments are generally poorly developed.
over a long period, during which time it develops All genera of Sapotaceae have the stamens
a short stout and densely scaly axis (brachyblast opposite the corolla lobes inserted within the
terminated by a single, few-flowered fascicle. corolla tube or occasionally (e.g. some Pradosia)
at the base of the corolla lobes, or rarely (e.g.
FLORAL STRUCTURE. The calyx provides some of Pouteria bangii) free. The position of the stamens
the best taxonomic characters at generic and tribal relative to the corolla (exserted or included)
levels of classification. There are two major calyx follows tribal, generic or sectional lines. Exserted
types, the uniseriate and the biseriate. stamens are associated with the short-tubed,
The uniseriate calyx has a single whorl of (4)5( 6) rotate or spreading corolla and are typical of
free or partially fused imbricate sepals (Fig.123D), genera in Sideroxyleae, and of such genera as
as in most Pouteria or Chrysophyllum, or 5-12 Elaeoluma, Pradosia and Niemeyera in Chryso-
sepals in a closely imbricate spiral (e.g. Pouteria phylleae. The arrangement in Mimusopeae is
sect. Aneulucuma). essentially the same, except that the stamens
The biseriate calyx, which is characteristic of are exserted relative only to the lateral corolla
tribes Isonandreae and Mimusopeae, has two segments, while they are clasped by the median
whorls of 2, 3 or 4 free sepals, the outer whorl segment. Included stamens, found in many
being valvate (Fig. 121G). The number of sepals in genera of Chrysophylleae, are associated with the
each whorl is very consistent and closely follows tubular or cyathiform corolla with short erect
generic lines. Isonandra and Madhuca have 2 lobes.
whorls of 2 sepals, Palaquium and Manilkara have The number of stamens relative to the number
2 whorls of 3 sepals, and Mimusops has 2 whorls of corolla lobes is diagnostic for the tribes. In
of 4 sepals. Chrysophylleae, Mimusopeae, Sideroxyleae there
The corolla is always actinomorphic and is nearly always one stamen opposite each corolla
gamopetalous. It may be rotate (e.g. Pradosia, lobe, whereas in Isonandreae and Omphalo-
Niemeyera), with the tube very short in relation to carpeae there are 2 or more stamens opposite each
the lobes, or the tube may be shortly cyathiform, lobe. The stamens may be grouped opposite the
with spreading lobes (e.g. Sideroxylon), or it is corolla lobes as in Omphalocarpum, or they
short or long tubular usually with the tube exceed- may be in a single continuous whorl, or in 2-3
ing the lobes, and with the lobes erect or only alternating whorls within the corolla tube (e.g.
slightly spreading (Chrysophylleae). The number Madhuca).
of corolla lobes relative to the number of sepals is The increase in number of stamens in Isonan-
a character of primary importance. In Mimuso- dreae is correlated with an increase in size of
peae, which have a biseriate calyx, the number of the anthers, which is possibly associated with bat
corolla lobes is usually the same as the number of pollination.
Sapotaceae 393
The stamens are generally free, but in a few 1. Mimusops type. Occurs in Mimusops,
genera become fused to the adjacent staminode Manilkara and other genera of tribe Mimusopeae.
(e.g. Autranella), or to the adjacent stamen, thus This is the most complex floral structure of the
forming a partial or complete stamina! tube family. The calyx is biseriate, with 2 whorls of 3 or
(e.g. Aulandra, Englerophytum). Magodendron is 4 sepals. The corolla, which has a short tube
unique in having locellate anthers. exceded by the lobes, opens widely, exposing the
Petaloid or scale-like structures alternating with erect staminodes and stamens. The corolla lobes
the fertile stamens (opposite to them in Gluema) are subdivided into 3 segments, the median
are widespread, and their presence or absence is segment is erect and clasping the opposing
highly correlated with other structures of generic stamen, while the 2 lateral segments spread hori-
importance. They are apparently derived from zontally. The lateral corolla lobe segments are
stamens, by loss of the anther, and then by further often subdivided to the base, so that each corolla
development or reduction of the filament. Stamin- lobe is subdivided into 5 equal-sized petaloid seg-
odes are occasionally seen to bear apical anthers ments. The well-developed and variously shaped
(Fig. 122D), and genera such as Manilkara, which staminodes alternate with the stamens, and are
typically have a whorl of stamens alternating with held erect to form a closed sheath around the
a whorl of sterile staminodes, occasionally (e.g. ovary. The majority of species in this group have
M. valenzuelana) present two whorls of fertile equal numbers of sepals, corolla lobes, stamens
stamens. Sometimes the staminode whorl is and staminodes, but modification and reduction
incomplete by reduction in number and the sta- of corolla lobe appendages and staminodes occur
minodes present are variable in size. They reach in all the main geographical regions. Two genera,
their greatest development in Mimusops and Labourdonnaisia and Letestua, have 2-3 times as
related genera, where they are large, hairy, hooded many corolla lobes and stamens as sepals. Bat pol-
structures alternating with the stamens, erect or lination has been recorded for this floral type
incurved, forming a closed receptacle around the (Manilkara).
ovary. The structure so formed acts as a nectar
receptacle in the newly opened flower. Elsewhere 2. Madhuca type. Characterized by a biseriate
in the family they are smaller or absent, and in calyx of2 whorls of2 or 3 sepals, and a short-tubed
some of the simple-flowered 5-merous genera open corolla with spreading lobes. The number of
such as Pouteria, they are inserted in the corolla lobes may equal the number of sepals or be
corolla lobe sinuses, above the level of the fertile 2-3 times as many, but the lobes are undivided.
stamens. There are 2-3 times as many stamens as corolla
Although the flowers secrete nectar, the nectar- lobes, usually with large exserted anthers. Stamin-
secreting area is morphologically poorly dif- odes are absent. The style is usually very long and
ferentiated, and represented only by a small exserted. This floral type is characteristic of most
ring-shaped disk around the ovary base. species of Isonandreae, and it also appears in the
The ovary is 1- to multilocular, with uniovulate Omphalocarpeae (Tsebona). The latter differs,
loculi; exceptions are Diploon with 1 loculus and however, in the presence of well-developed sta-
2 basal ovules, and some Sideroxylon which have minodes. This floral type is also associated with
lost some or all of the ovary septa, resulting in up bat pollination in Madhuca.
to 5 basal ovules. Placentation is axile or less fre-
quently basal or basi-ventral. 3. Sideroxylon type. Present in most species of
Members of Mimusopeae, Isonandreae and tribe Sideroxyleae, and less complex than the
Omphalocarpeae have ovary loculi equalling the 2 previous types, with 5-merous flowers, a unise-
number of sepals, twice as many or even more (up riate calyx of usually 5 imbricate sepals, and a
to 30 in Omphalocarpum) whereas in Sideroxyleae short-tubed open corolla with wide-spreading
and Chrysophylleae the trend is generally towards lobes. The corolla lobes of many species are sub-
reduction in the number of loculi towards the divided into 3 segments, but the two lateral
unilocular condition, which is reached in some segments are greatly reduced and much smaller
Pouteria species (sect. Franchetella). than the median segment, and in many cases
Five floral types are recognized in the family; they have disappeared altogether. The stamens
they are based on the number and degree of com- are attached at the top of the corolla tube
plexity of the floral parts and the precise arrange- and exserted and alternate with 5 variously
ment of the organs relative to each. They correlate developed staminodes. The style is generally
well with the tribal divisions. short.
394 T.D. Pennington
4. Pradosia type. This and the Pouteria type rep- Unisexual flowers are more common in Chryso-
resent the simplest floral arrangement. It occurs phylleae, especially in the large genus Pouteria,
only in several genera of Chrysophylleae, and is where they reach levels of around 50%. The true
most commonly found in pentamerous flowers percentage of unisexual species may be much
(e.g. Pradosia, Pichonia, Elaeoluma, Synsepalum). higher owing to the lack of observations, and to
The flowers have a uniseriate calyx of 5 imbricate the fact that functionally male flowers have well-
sepals, short-tubed rotate or spreading corolla developed rudiments of the opposite sex and may
with 5 simple corolla lobes and 5 exserted stamens be mistaken for hermaphrodite flowers. Within
fixed at the apex of the corolla tube. Simple sta- Chrysophylleae both monoecious and dioecious
minodes alternating with the stamens are some- conditions are known, but dioecy seems to
times present. The style is short. No observations predominate.
on the pollination of this type of flower have been The simplest form of sexual dimorphism seen
recorded, but it is probably effected by short- in this group involves the loss of anthers in the
tongued insects. female, the filaments remaining. In others the
anther may be converted into a flattened
5. Pouteria type. Confined to genera in Chryso- appendage. Further reduction involves the com-
phylleae such as Pouteria, Chrysophyllum, plete loss of stamens in the female flower (e.g.
Micropholis and Ecclinusa. The flowers have a Pouteria nudipetala), and loss of stamens may
uniseriate calyx and are usually pentamerous but, also be accompanied by reduction in size of the
in some sections of Pouteria, the calyx is of only 4 staminodes (e.g. Pouteria rufotomentosa). Finally,
imbricate sepals, and the corolla lobes may loss of stamens may be correlated with differences
number 6-8 or occasionally more. The corolla in flower size, with generally the female much
lobes are simple, and staminodes if present are smaller than the male. At the same time, differ-
small and undivided. It differs from the Pradosia ences in flower size may also be accompanied by a
type in the shape of the corolla, which is tubular change in the relative length of corolla tube and
or cyathiform with small, more or less erect lobes, corolla lobes. The female flowers of Pouteria have
and included stamens. The stamens may be fixed corolla lobes free almost to the base, whereas in
at any level in the corolla tube, but if at the apex, the male the corolla tube is only slightly shorter
then the filaments are poorly developed, so that than the lobes.
the anthers remain within the erect corolla lobes.
The corolla tube is often longer than the corolla EMBRYOLOGY. The anther has a secretory
lobes. The style is short and included, or some- tapetum with multinucleate cells. The pollen
times slightly exserted beyond the apex of the grains are 2-celled when shed. The ovule is erect,
corolla tube. Pollinators for this floral type are pos- anatropous, unitegmic and tenuinucellate. Pres-
tulated to be short- or long-tongued insects. ence of a hypostase is contentious. The archespo-
Bisexual flowers predominate in tribes Mimuso- ria! cell functions as megaspore mother cell and
peae, Isonandreae and Sideroxyleae. Only 1-2% of the embryo sac is of the Polygonum type.
the species of these tribes have unisexual flowers, Endosperm development is of the nuclear type;
but among the latter are several species which the endosperm is thin-walled or with thick, often
exhibit the strongest dimorphism of any in the amyloid walls and oily (Corner 1976; Johri et al.
family. These are Sideroxylon puberulum (Mas- 1992).
carenes), Neohemsleya usambarensis (Tanzania),
and Nesoluma polynesicum (Hawaiian Islands), all POLLEN MORPHOLOGY (by M.M. HARLEY). With
of which are dioecious. The functionally male few exceptions, Sapotaceae pollen is easily recog-
plants bear well-developed rudiments of the oppo- nised based on its morphology. Exine stratification
site sex, the pistillode containing small ovules, but together with shape and apertures characterise the
at least where field observations have been made, pollen of the family. Pollen grains are single, isopo-
they never set fruit. Female flowers of these species lar, angulaperturate or, less frequently, planaper-
are generally smaller than the male, and corolla, turate (in 4-aperturate grains). The amb is more
stamens and staminodes are strongly modified. or less circular, or governed by the number of
The corolla lobes are reduced in size, almost free apertures, for example, 3-triangular (Fig. 119H),
in Neohemsleya, and are represented by small, free 4-square (Fig. 1191) or 5-pentagonal (Fig. 119J).
vestigial petals in Nesoluma, or they are absent. The grains are predominantly prolate-spheroidal,
The stamens are reduced to small sterile vestiges, subprolate or prolate, occasionally spheroidal or,
or they are absent. rarely, oblate-spheroidal. Within single collections
Sapotaceae 395
the colporate apertures may be 3 only, 3 and 4, 4 Two major pollen groups, A and B, were defined
only, 4 and 5 or, rarely, 5 only in number. Very for Sapotaceae (Harley 1986a, 1990a), and 12
rarely some 6-colporate grains are recorded in pollen types (Harley 1990b, 1991}, based on
samples which have mainly 4- or 5-colporate presence or absence of continuous endexinous
grains. Average polar length is between 16 and thickening in the equatorial region, number of
93!-Lm. The colpi are usually narrow, occasionally apertures, surface patterning and colpus length.
broad, and range from vestigial (Fig. 119A) to The pollen morphologies of Chrysophyllum mar-
c. 4/5 1h (Fig. 119B) or 5/61h of the polar length ginatum, C. inornatum and Englerophytum stele-
(Fig. 119C). They are provided with a tough, gran- cantha are more or less anomalous within the
ular membrane (Fig. 119B) that does not rupture family (Harley 1990b).
during acetolysis, except in the endoapertural
region where there is no underlying endexine. The KARYOLOGY. For a review of chromosome
endoapertures are narrowly or broadly lalongate numbers in Sapotaceae, counts for 95 species have
(Fig. 119F), less frequently circular or, extremely been evaluated (Johnson 1991 ), all based on x = 10,
rare, broadly lolongate. The tectum in the central 11, 12, 13 or 14, nearly all at diploid level. The
apertural region in many species is protrudent higher numbers (x = 13, 14) predominate in tribes
(Fig. 119A, K) or semi-protrudent. This is usually Chrysophylleae and Omphalocarpeae, whereas
associated with narrow endoapertures and vestig- numbers in Mimusopeae, Isonandreae and
ial or short colpi. Wall thickness at the poles Sideroxyleae are, with one exception, based on x =
(Fig. 119G, K) is 1-3(-5)!-Lm; at the equator 10 or 11. Johnson (1991) hypothesised that these
1-5( -8) !liD. The ektexine is nearly always com- numbers may reflect a series of descending dys-
posed of a thick tectum, a very narrow infratectum ploidy, but the reverse may also be a possibility,
('columellar layer'), which is either granular or particularly in view of the correlation with floral
with very reduced columellae, that are sometimes structure.
interspersed with granulae, plus a foot layer of
similar width, or thicker than, the tectum PoLLINATION (from notes kindly provided by S.
(Fig. 119G). The endexine is usually absent in the VoGEL to the editor). Although the majority of
polar region but greatly thickened, especially Sapotaceae certainly is entomophilous, in the lit-
in the area surrounding the endoaperture and erature only instances of bat pollination seem to
underlying the colpus (Fig. 119D, E, K). In some have been recorded. Madhuca indica(= Bassia lat-
species it is absent or very thin in the mesocolpial ifolia) has been observed being visited and polli-
areas, while in others it may be slightly thinner, or nated by flying foxes in India (Cleghorn 1922); the
as thick, as in the endoapertural areas, resulting in trees start blooming there in the leafless state and
a continuously thickened band of exine in the the pendent flowers produce a strong, unpleasant
equatorial region. There is, however, a gradual odour. The animals feed on the fleshy petals, get
reduction in the thickness of this band towards the dusted with pollen when exploiting the protan-
poles. The tectum is usually sparsely punctate or drous flowers and, in turn, deposit the pollen on
perforate (Fig. 119B ). The perforations, which may receptive stigmas. The same tree species was
occur in shallow pits, may be more dense and/or found to be bat-exploited in Mauritius and Java,
coarse, in either the mesocolpia or the apocolpia. where the Megachiroptera Pteropus niger and
Microfossulae are recorded for a few species. Cynopterus sphinx, respectively, were feeding on
In some species the entire surface, seen with the petals (van der Pijl 1936; Cheke and Dahl
SEM, appears psilate, subpsilate (Fig. 119C), 1981). Petropus bats were also observed as visitors
scabrate, finely or coarsely granular (Fig. 119L), to the flowers of Palaquium gutta and P. querci-
anastomosed granular, finely striate-rugulate, folium and Madhuca macrophylla on Java, where
striate (Fig. 119M), low-relief angular rugulate they ate the petals of the bad-smelling flowers,
(Fig. 119N), or weakly to coarsely rugulate those of Madhuca macrophylla being reportedly
(Fig. 1190). In other species there is a clear differ- nectariferous in the evening (van der Pijl1936).
entiation between the more or less smooth
apocolpia and a weakly or distinctly rugulate FRUIT AND SEED. The fruit is typically a !-many-
mesocolpia (Fig. 119A). Rarely, Diploon cuspidata seeded berry, with a fleshy, leathery or rarely
and Micropholis retusa, the exine has supratectal woody outer pericarp. Differentiation of the
spines (Harley 1990b). In Sarcaulus the exine is endocarp to form a distinct cartilaginous layer
spinulose. occurs in Pradosia, the fruit of which is therefore
396 T.D. Pennington
Fig. 119. Sapotaceae, pollen. A Pouteria caimito, equatorial elii, equatorial plane xlOOO LM. J Palaquium quercifolium,
view x l200 SEM. B Payena endertii, equatorial view x610 SEM. equatorial plane xlOOO LM. K Sideroxylon glanduliferum, polar
C Vitellaria paradoxa, equatorial view xl310 SEM. D Pouteria plane x 1200 TEM. L Madhuca aristulata, tectum surface,
ucuqui, equatorial plane x2330 TEM. EMimusops schimperi, mesocolpium, x lOOOO SEM. M Sideroxylon celastrinum,
equatorial plane x ll60 TEM. F Pycnandra aff. vieillardii, polar tectum surface, mesocolpium, x lOOOO SEM. N Pouteria bail-
orientation, fracture to show interior of grain x l085 SEM. G lonii, tectum surface, mesocolpium, x lOOOO SEM. 0 Madhuca
Pouteria multiflora, wall fracture, pole x lOOOO SEM. H Poute- pal/ida, tectum surface, mesocolpium, x lOOOO SEM. Photos
ria laurifolia, equatorial plane xlOOO LM. I Tieghemella heck- M.M.Harley
Sapotaceae 397
considered to be a drupe. Four genera of the base. The width of the hilum is to some extent cor-
Mimusopeae (Lecomtedoxa, Neolemonniera, related with seed shape. The strongly laterally
Gluema, Eberhardtia) have a dry fruit which compressed seed of Chrysophyllum nearly always
dehisces tardily to form a 1-several-seeded loculi- has a narrow adaxial hilum, and conversely, the
cidal capsule. broad ellipsoid seed of Pouteria usually has a
The general appearance of the sapotaceous seed broad adaxial hilum, but there are numerous
is so distinctive as to be one of the family charac- exceptions to the latter.
teristics. It has a smooth, hard, shining brown testa The orientation of the embryo within the seed is
which strongly contrasts with the pale, rough generally vertical, with the cotyledons superior
"scar" area (hilum) by which the seed is attached and the radicle inferior. The only variation is
to the fruit. The smooth shining testa is normally found in Sideroxylon where the arrangement
free from the endocarp, but several modifications varies from vertical, through oblique to fully
of this arrangement are found. In some species, horizontal.
e.g. Pouteria gongrijpii, the testa, although still There are two principal embryo types, strongly
smooth, is adherent to the fruit over its whole correlated with seed shape and with the presence
surface. In this case the seed can still be removed or absence of endosperm:
from the fruit, but with difficulty. In others, such
as in species of Chrysophyllum sect. Prieurella, the a) Embryo with thin flat foliaceous cotyledons
testa becomes roughened and is strongly adherent and exserted radicle, which is generally associ-
to the fruit. The fruit and seed are thus effectively ated with a laterally compressed seed, and with
fused and cannot be separated. In other species, the presence of endosperm. On germination
such as Pouteria glomerata, fusion of the seed to the cotyledons emerge from the seed and are
the fruit is achieved by the extension of the hilun photosynthetic (phanerocotylar). This type
over almost all the seed surface. The seed shape, is well represented in Mimusops, Manilkara,
position of the hilum, and its extent relative to the Sideroxylon and Chrysophyllum.
smooth, shining area of testa all provide taxo- b) Embryo with thick plano-convex cotyledons
nomic characters useful at the generic and species with radicle included or only extending to the
level. surface. This is generally associated with a
The position of the hilum is a most important broader seed which is not laterally compressed,
character at the generic level and above, as it is and with the absence of endosperm. The cotyle-
highly correlated with other floral features. dons of this type are retained within the seed
Sideroxyleae are typified by a basal hilum, derived coat on germination and do not photosynthe-
from the basal or basi-ventral ovule position. A sise (cryptocotylar). Well represented in
basal or basi-ventral hilum is also characteristic of Pouteria.
Mimusops and some Manilkara species. In the
majority of Chrysophylleae, Omphalocarpeae and All species of Chrysophyllum have an embryo with
Isonandreae the hilum is adaxial (derived from thin foliaceous cotyledons, with the presence of
axile ovule placentation). endosperm, correlated with the simple 5-merous
Within the two general hilum positions, there is flower lacking staminodes. Many of the smaller
considerable variability in the extent and shape of genera of Sapotaceae have only one embryo type,
the hilar area. Thus, the basal hilum in members but the majority of the larger genera have a sig-
of the Sideroxyleae is generally as broad as long, nificant number of exceptions, e.g. Palaquium is
or broader than long, whereas the basi-ventral predominantly non-endospermous, with thick
hilum of Manilkara tends to be elongate. The plano-convex cotyledons, but a minority of
lateral adaxial hilum may be short and narrow species have thin foliaceous cotyledons with
(confined to the basal half of the seed), as in some endosperm. Similar exceptions occur in Sideroxy-
Pouteria species, or long and narrow (extending lon and Madhuca. Pouteria is predominantly non-
the full length of the seed), as in many Chryso- endospermous with thick cotyledons, but about
phyllum. In other genera such as Niemeyera, the one third of the total species have the other
adaxial hilum extends to cover up to half or more embryo type, and between the two is an extensive
of the seed surface, and in extreme cases covers the series of intermediate conditions, in which the
whole seed surface except for a thin abaxial strip cotyledons are flattened but thick and fleshy, and
which still retains the smooth, shining testa (e.g. the endosperm is reduced.
Pouteria speciosa). In Ecclinusa, the narrow hilum Corner (1976) found the testa multiplicative in
extends down the adaxial face and around the most layers, including the outer epidermis; the
398 T.D. Pennington
outer part, 8-25 cells thick or more, forming a have about 80% polyisomere content. Balata is a
heavily lignified, sclerotic layer, the inner part product of tropical America; it contains more
thin-walled and eventually crushed. resinous material and its source are the genera
Manilkara and Mimusops. Chicle is a polymer con-
DISPERSAL. There is great variation in the size and taining both trans- and cis-isoprene in a ratio 2: 1;
texture of the sapotaceous berry, correlated with the classical source is Manilkara zapota. The dis-
the dispersal agent. Primates are the major distribution of these polymers within the family is
persers, taking both small-fruited and large- still unknown.
fruited species. They are attracted to species Further characteristic compounds are saponins
with both soft fleshy pericarps or hard leathery and pentacyclic triterpenes. Saponins are fre-
pericarps, and the seeds are spat out, or pass quently found in bark, wood and seed; in the
unharmed through the gut. Some species such latter they may act as defence and storage com-
as Manilkara zapota are bat-dispersed, and pounds. The pentacyclic triterpenes are based on
these have large berries with a soft pericarp. Birds widespread skeleta, to which unusual esterifying
(especially parrots) are also seen to take Sapotaceae groups are attached at the C-3 alcohols.
fruit, although they often destroy the seed. Alkaloids are of limited occurrence in
Sapotaceae and are represented by the
SEEDLINGS. Bokdam (1977) studied the seedlings pyrrolizidine group, which is widely distributed
of African Sapotaceae and recognized two major in other plant families. Phenolic compounds
groups within them. Type 1 has enlarged photo- are represented by common flavonols including
synthetic foliaceous cotyledons and develops from myricetin, proanthocyanidins (procyanidin and
endospermous seeds; Type 2 has unenlarged and prodelphinidin) as constituents of condensed
often non-photosynthetic fleshy cotyledons devel- tannins, and gallic acid and the hydrolysable
oping from seeds with scant or no endosperm. tannins based upon it.
Bokdam subdivided these groups into phanero-
cotylar and cryptocotylar, and also on the thick- AFFINITIES. Sapotaceae have usually been
ness of the cotyledons, and on the development of included in Ebenales but, as Corner (1976)
cotyledonary nervation. He found correlations remarked, in Ebenaceae the ovule is bitegmic and
between seedling types and other morphological suspended, and seeds are exotestal in contrast with
characters including the level of insertion of Sapotaceae, where the ovule is erect, unitegmic
stamens in the corolla tube, number of ovary and the seeds are mesotestal. Moreover, molecular
locules, number of seeds in ripe fruit and shilum studies have revealed Ebenales as an artificial con-
position, shape and size, but the relationships are struct, and place Sapotaceae in a well-supported
imperfect, even on the basis of African species clade, the expanded Ericales. In a five gene-
alone. As the loss of endosperm has apparently analysis of this clade (Anderberg et al. 2002),
occurred several times in all the major groups of Sapotaceae appear sister to Lecythidaceae,
the family, the associated seed, hilum and seedling although with very low support (52%) and backed
characters have also changed, and therefore none by weak apo(rather plesio?)-morphies such as tri-
of these characters is of use in defining them. They lacunar nodes, stipules and nuclear endosperm, so
are, however, useful at the lower level of genus, that their affinity appears to remain obscure.
section and species.
DISTRIBUTION AND HABITATS. The distribution
PHYTOCHEMISTRY. Data are from Hegnauer of Sapotaceae extends across the humid tropics of
(1973, 1990 ), and Waterman and Mahmoud (1991}. Central and South America, Africa, Madagascar,
By far the most characteristic compound synthe- and Asia east to the Pacific. Generic diversity is
sised by Sapotaceae is the latex, a mixture of poly- greatest in Africa and Asia, while the greatest
isoprenes and resins, from which the commercial concentration of species is in Amazonian South
coagulation products gutta percha, balata and America and Malesia. In South America the family
chicle are derived. Sapotaceous latex is a mixture occupies a broad swathe from the Guianas to the
of trans-polyisoprenes and resins, in which the eastern foothills of the Andes, with very high levels
polyisoprene content, according to quality, ranges of species diversity and individual abundance,
from 80 to about 20%, whereas ruber is a pure cis- equalling or exceeding that of any other tree
isomer of isoprene. Gutta percha is a product of family. Madagascar and New Caledonia are also
Malakka, Indonesia and the Philippines, mostly exceptionally rich in generic and species diversity
derived from Palaquium species; good qualities and endemism.
Sapotaceae 399
The majority of species are confined to lowland CoNSPECTus oF SAPOTACEAE

rainforest below 1000 m altitude, and they vary in
I. Tribe Mimusopeae
habit from small understorey treelets to large 1. Subtribe Mimusopinae
canopy emergents. Most occupy non-flooded Genera 1-6
forest, but a significant number are confined to 2. Subtribe Manilkarinae
periodically or permanently flooded forest. A few Genera 7-12
3. Subtribe Glueminae
genera extend into savannah, as Ecclinusa in the Genera 13-17
Guianas, Vitellaria in West Africa, and Sideroxylon II. Tribe Isonandreae
occurs widely in tropical dry forest in the West Genera 18-24
Indies and Central America, and also extends into III. Tribe Sideroxyleae
North America as far north as Illinois. Genera 25-30
IV. Tribe Chrysophylleae
Genera 31-49
PoLLEN FossiL RECORD. The fossil record of V. Tribe Omphalocarpeae
Sapotaceae-like pollen is extensive. It has been Genera 50-53
summarised and discussed in Harley (1990b,
1991}. It is clear from this record that by the lower KEY TO THE TRIBES OF SAPOTACEAE
Eocene Sapotaceae were represented in all conti-
1. Sepals in 2 whorls, outer whorl valvate 2
nents and, by this time, pollen similar to all the - Sepals in 1 imbricate whorl 3
major modern groups was in existence. 2. Stamens as many as corolla lobes, corolla lobes often
divided into 3 or more segments I. Mimusopeae
ECONOMIC IMPORTANCE. The family provides - Stamens 2-3 times as many as the corolla lobes, corolla
four important economic products: latex, oil, lobes undivided II. Isonandrae (p. 407)
3. Stamens 2-6 times as many as the corolla lobes
fruit and timber. The latex of Palaquium and V. Omphalocarpeae (p. 418)
Manilkara is used to make gutta percha and - Stamens as many as the corolla lobes 4
balata, which was formerly used on a large scale 4. Corolla lobes often divided, stamens exserted, staminodes
for insulating marine cables and for chewing gum, +,hilum basal or basiventral III. Sideroxyleae (p. 409)
- Corolla lobes undivided, stamens exserted or included,
but these products are now largely replaced by staminodes + or 0, hilum usually adaxial
synthetic materials. However, gutta percha is still IV. Chrysophylleae (p. 411)
important in dentistry, where it is used for root
fillings.
The most important oil-producing species is I. TRIBE MIMUSOPEAE Hartog (1878).
Vitellaria paradoxa, the Shea Butter Nut, which
replaces the oil palm as the source of cooking Unarmed trees or shrubs. Leaves spirally arranged,
oil in N Nigeria (Hall 1996). Argania spinosa sometimes clustered. Inflorescences axillary.
(Morocco) is also used for cooking oil, and several Flowers bisexual (unisexual in Vitellariopsis).
American species of Pouteria have similar oily Calyx usually 2 whorls of 3 or 4 sepals, the outer
seeds, which have yet to be investigated. whorl valvate, less frequently (subtribe Gluemi-
Virtually all species of Sapotaceae have edible nae) a single whorl of 5 imbricate, sometimes
fruit, and some of the better ones have been pro- quincuncial sepals; corolla lobes, stamens and sta-
tected and improved by man over many centuries. minodes usually same number as sepals; corolla
Among the most well-known species are Pouteria rotate or cyathiform; corolla lobes usually divided
caimito (western Amazonia), Pouteria lucuma into 3 segments, a median one and 2 lateral or
(Andes) and Manilkara zapota (Mexico), and the abaxial ones; lateral segments exceeding, equalling
latter is now grown commercially in Asia. Many or smaller than the median segment, sometimes
others are sold locally throughout the range of the subdivided; stamens exserted, usually 8, inserted
family. in a single whorl at the top of the corolla tube;
Several Malesian genera, mainly Palaquium, anthers extrorse, usually glabrous; staminodes
Madhuca and Payena, provide good-quality usually well-developed, usually 8, alternating with
medium-density timber, used for plywood and the stamens; ovary pubescent; hilum usually basal
veneer (Soerianegara and Lemmens 1993). Other or basi-ventral, less frequently adaxial.
species of the same genera produce very dense,
durable timber for heavy construction. Elsewhere, KEY TO THE SUBTRIBES OF MIMUSOPEAE
in Africa and tropical America, the timber-
producing species are used principally for heavy 1a. Calyx of 2 whorls of 4 sepals, the outer whorl valvate;
corolla lobes, stamens, staminodes, and ovary loculi
construction and parquet.
400 T.D. Pennington
usually 8; staminodes usually hairy; fruit indehiscent; the style; lateral segments widely spreading,
hilum usually small and basal 1. Mimusopinae often equalling or slightly exceeding the median
1b. Calyx of 2 whorls of 3 sepals, the outer whorl valvate,
corolla lobes, stamens usually 6, less frequently 12-18; sta- segment, entire or deeply divided or laciniate.
minodes 6 or 0, glabrous; fruit indehiscent; hilum usually Stamens (7}8; filaments free or partially fused to
elongate, basi-ventral 2. Manilkarinae (p. 402) the staminodes; anthers hairy or glabrous. Sta-
1c. Calyx a single whorl of 5 imbricate or quincuncial sepals; minodes 8, inflexed and often forming an envelope
corolla lobes, stamens, staminodes and ovary loculi usually round the gynoecium, lanceolate, hairy. Ovary
5; staminodes hairy or glabrous; fruit dehiscent or not,
hilum long, usually narrow, adaxial 3. Glueminae (p. 405) (7}8-locular, placentation basi-ventral or basal;
style exserted or not. Fruit 1-6-seeded, fleshy.
Seed laterally compressed, usually with a smooth
1. SUBTRIBE MIMUSOPINAE Aubreville (1964). shining, rarely rugose woody testa; hilum small,
often circular or elliptic, basal or basi-ventral.
Calyx of 2 whorls of 4 sepals, the outer whorl Embryo with foliaceous cotyledons and exserted
valvate; corolla lobes, stamens, staminodes, and radicle; endosperm copious. About 20 species
ovary loculi usually 8; staminodes usually hairy; in Africa, 15 in Madagascar, 4 in the Mascarenes,
fruit indehiscent; hilum usually small and basal. 1 in the Seychelles, and 1 in Asia and the
Pacific.
KEY TO THE GENERA OF MIMUSOPINAE
1. Stipules large and conspicuous 2 2. Vitellariopsis Baillon Fig. 120
- Stipules 0 or minute 3 Vitellariopsis Baillon ex Dubard, Ann. Mus. Bot.-Geol. Colon.
2. Corolla tube much longer than lobes, filaments united with Marseille III, 3: 44 (1915); Aubreville, Adansonia II, 3: 41
staminodes for most of their length; hilum basi-ventral, (1963); Baehni, Boissiera 11: 112 (1965); Kupicha, Candollea
embryo with thin foliaceous cotyledons, endosperm 33: 29 (1978).
copious 3. Autranella Mimusops sect. Vitellariopsis Baillon (1891).
- Corolla tube shorter than lobes, filaments free; hilum
adaxial, embryo with plano-convex cotyledons, endosperm
0 Trees or shrubs. Leaves spirally arranged, clustered
5. Baillonella
at the shoot apex. Venation brochidodromous; ter-
3. Corolla lobes entire; inflorescences clustered at shoot apex
in axils of scale leaves; venation craspedodromous tiaries forming a distinctive finely areolate reticu-
6. Vitellaria
lum. Stipules small, caducous. Flowers bisexual,
- Corolla lobes divided into 3 segments (entire in Mimusops
rarely unisexual, solitary or fasciculate. Corolla
antsiranensis ); inflorescences axillary or at defoliated
nodes, venation brochidodromous glabrous; tube much shorter than the lobes; lobes
4
(6-}8(-10}, divided to the base into 3 segments;
4. Hilum small, basal or basi-ventral, embryo with thin foli-
aceous cotyledons and copious endosperm 1. Mimusopsmedian segment erect, clasping a stamen, lateral
- Hilum large, adaxial, embryo with thick plano-convex
cotyledons, endosperm 0
segments widely spreading or reflexed, equalling
5
or slightly exceeding the median segment, undi-
5. Corolla tube well-developed, central corolla lobe segment
vided. Stamens (6-}8(-10}; filaments free; anthers
narrowed or reduced to a filament; staminodes glabrous;
seed coat thick glabrous. Staminodes (6-}8( -10 ), inflexed and
4. Tieghemella
- Corolla tube very short; corolla lobe segments more or less
forming an envelope around the gynoecium,
equal in size; staminodes hairy; seed coat thin, crustaceous
or membranous lanceolate, hairy. Ovary 7-9-locular, placentation
2. Vitellariopsis
axile; style exserted or not. Fruit 1-several-seeded,
fleshy or leathery. Seed broadly ellipsoid to
1. Mimusops L. oblong-ellipsoid, or plano-convex when several in
Mimusosps L., Sp. Pl.: 349 (1753).
a fruit, not laterally compressed, with a rather thin,
dull, membranous or crustaceous testa; hilum
Trees or shrubs. Leaves clustered or not. Venation large, broad, adaxial, covering up to half the seed
brochidodromous, tertiaries usually parallel to the surface. Embryo with thick fleshy plano-convex
secondaries and descending from the margin, or cotyledons; radicle extending to the surface, not
reticulate. Inflorescence axillary or in the axils of exserted; endosperm 0. Six species in East Africa.
leaf scars. Stipules minute, caducous, sometimes 0.
Flowers solitary or fasciculate. Corolla hairy or 3. Autranella A. Chev.
glabrous; tube much shorter than the lobes; lobes
8, nearly always divided to the base into 3 seg- Autranella A. Chev., Veg. Ut. Afr. Trop. Franc. 9: 271, fig. 29
(1917).
ments; median segment slender, usually erect and
clasping the stamen, sometimes inflexed against
Sapotaceae 401
Trees. Leaves clustered at the shoot tip. Venation Fig. 120. Sapotaceae-Mimusupeae. Vitellariopsis kirkii. A
brochidodromous; tertiaries parallel to the sec- Flowering branch. B Inflorescence. C Flower. D Trifid corolla
lobe, viewed from outside. E Same, viewed from inside, with
ondaries or reticulate. Stipules large, caducous. stamen and staminodes. F Stamen and staminodes, viewed
Corolla tube glabrous, lobes hairy, tube 2-3 times from outside. GPistil. H Ovary, schematic transverse section.
as long as the lobes; lobes 8, divided to base into I Fruit. J Seed. (Hemsley 1968)
3 segments; median segment erect, clasping the
stamen; lateral segments 1.5-2 times the length of
the median segment, widely spreading or reflexed, dons and exserted radicle; endosperm copious. A
undivided. Stamens 8; filaments united with the single species, A. congolensis (De Wild.) A. Cheva-
staminodes for most of their length, the free lier, in West Africa.
portion strongly reflexed; anthers glabrous. Sta-
minodes 8, fused to the filaments for most of their
length, free portion truncate or irregularly lobed, 4. Tieghemella Pierre
glabrous; stamen-staminode tube narrowed at the Tieghemella Pierre, Not. Bot. Sapot.: 18 (1890); Aubreville, Fl.
throat. Ovary large, filling the corolla tube, and Gabon 1, Sapotacees: 45 (1961).
gradually tapering from near the base, 8-locular,
placentation basi-ventral; style short, included. Trees. Leaves weakly clustered. Venation brochido-
Fruit 1( -2)-seeded, fleshy. Seed broadly obovoid, dromous, tertiaries reticulate or parallel to the
slightly laterally compressed, with a thick shining secondaries. Stipules 0. Corolla glabrous, tube
woody testa; hilum moderately large, rectangular, well-developed, only slightly shorter than the
basi-ventral. Embryo with thin foliaceous cotyle- lobes; lobes 8, partly or completely divided into 3
402 T.D. Pennington
segments; median segment narrow or reduced to by a filiform point, subglabrous. Ovary (5)6-
a small filamentous rudiment; lateral segments locular, placentation axile; style slightly exserted.
longer and broader than the median segment, Fruit 1-2-seeded, fleshy. Seed globose or broadly
undivided. Stamens 8; filaments free; anthers, ellipsoid, not laterally compressed, with a rather
glabrous. Staminodes 8, erect, carnose, narrowly thin shining testa; hilum broad, adaxial. Embryo
lanceolate, glabrous. Ovary 8-locular, placentation with thick, fleshy, fused cotyledons; radicle not
basi-ventral; style included. Fruit !-several- exserted; endosperm 0. Possibly two species in
seeded, fleshy. Seed broadly ellipsoid, slightly lat- western tropical Africa and Cameroon.
erally compressed, with a thick, woody, shining
testa; hilum broad, adaxial, covering up to half the
seed. Embryo with thick, fleshy, plano-convex 2. SUBTRIBE MANILKARINAE H.J. Lam (1938).
cotyledons and exserted radicle; endosperm 0.
Two species in West Africa. Unarmed trees or shrubs. Leaves spirally arranged,
clustered at the shoot apex, secondaries parallel.
Inflorescence axillary or in the axils of fallen
5. Baillonella Pierre
leaves. Flowers bisexual (few Manilkara dioe-
Baillonella Pierre, Not. Bot. Sapot.: 13 (1890); Aubreville, Fl. cious). Calyx of 2 whorls of 3 sepals, the outer
Gabon 1, Sapotacees: 51 (1961). whorl valvate, corolla lobes, stamens usually 6,
Mimusops sect. Baillonella (Pierre) Engler (1897). less frequently 12-18, inserted at the top of the
corolla tube (within: some Manilkara), filaments
Unarmed trees. Leaves clustered at the shoot apex. free (partly fused with staminodes in some
Venation brochidodromous, tertiaries oblique. Manilkara), anthers extrorse, glabrous (exception-
Stipules large, persistent. Inflorescences densely ally pubescent in Manilkara and Faucherera); sta-
clustered at shoot apex. Corolla hairy (tube only), minodes 6 or 0, often small, glabrous, alternating
tube shorter than the lobes; lobes 8, divided to with stamens; fruit indehiscent, fleshy, usually 1-
base into 3 segments; median segment erect; seeded; hilum usually elongate, basi-ventral.
lateral segments spreading, exceeding the median
segment, undivided. Stamens 8; filaments free; KEY TO THE GENERA OF MANILKARINAE
anthers, glabrous. Staminodes 8, erect, narrowly
lanceolate, hairy. Ovary 8-locular, placentation 1. Corolla lobes, stamens and staminodes usually 6; stamin-
basi-ventral; style slightly exserted. Fruit 1-2- odes well-developed, vestigial or rarely 0 2
- Corolla lobes and stamens (10-)12-18, staminodes 0 5
seeded, fleshy. Seed broadly ellipsoid, with a thick, 2. Corolla lobes nearly always divided into 3 segments 3
hard, shining testa; hilum broad, covering the - Corolla lobes entire (vestigial lateral segments present in
adaxial surface of the seed. Embryo with thick Northia) 4
fleshy, plano-convex cotyledons and exserted 3. Seed laterally compressed, hilum narrow, nearly always
basi-ventral; staminodes usually well-developed
radicle; endosperm 0. A single species in West 7. Manilkara
Africa (B. toxisperma Pierre). - Seed not or only slightly laterally compressed, hilum
broad, covering the adaxial surface; staminodes usually
vestigial 8. Labramia
6. Vitellaria Gaertner 4. Venation craspedodromous; corolla tube equalling lobes;
seed c. 7 em long, with large adaxial hilum, endosperm 0
Vitellaria Gaertner, Fruct. 3: 131, t. 205 (1807); Hepper, Taxon
lO.Northia
11: 226 (1962).
- Venation brochidodromous, leaves striate; corolla tube
Butyrospermum Kotschy (1865).
much shorter than lobes; seed 1-2 em long, with small
basi-ventral hilum, endosperm + 9. Faucherea
Trees or shrubs. Leaves in a dense terminal cluster. 5. Corolla lobes entire, ovary loculi 5-10; hilum basal or basi-
Venation craspedodromous with a prominent ventral, hollowed 11. Labourdonnaisia
marginal vein; tertiaries parallel to the secondar- - Corolla lobes divided into 3 segments; ovary loculi 16-18;
hilum adaxial, narrow 12. Letestua
ies or reticulate; quaternaries finely areolate.
Stipules small, caducous. Inflorescences densely
clustered at the shoot apex. Corolla glabrous; tube 7. Manilkara Aubreville Fig. 121
much shorter than the lobes; lobes (6-)8, entire,
Manilkara Aubreville, Adansonia II, 11: 251-300 (1971).
contorted in bud, spreading. Stamens (6-)8; Achras L. (1753).
anthers glabrous. Staminodes (6-)8, erect or
inflexed and forming an envelope round the Trees, rarely shrubs, nearly always with sympodial
gynoecium, lanceolate, margin erose, terminated branching, rarely dioecious. Venation nearly al-
Sapotaceae 403
1 ~~
~
HI
'I '
I
' E~
'I o'
ways brochidodromous, secondaries often looping Fig. 121. Sapotaceae-Mimusupeae. Manilkara discolor. A
to form a submarginal vein (eucamptodromous Branch. B Inflorescences. C Young flower. D Flower at anthe-
sis. E Corolla opened out, viewed from inside to show stamens
and then with convergent secondaries); ter- and staminodes. FPistil. G Female flower. H Part of corolla of
tiaries often descending from the margin and female flower. I Pistil of female flower. J Section of ovary. K
parallel to the secondaries; higher order venation Young fruit, attached. L Seed. (Hemsley 1968)
usually reticulate or areolate. Stipules small,
caducous, or 0. Flowers solitary or fasciculate.
Corolla nearly always glabrous, occasionally tially divided or 2-3-lobed at the apex, or rarely
carnose; tube usually much shorter than the lobes, entire. Stamens 6( -12 ), very rarely fixed within the
rarely equalling or exceeding them; lobes 6( -9), tube; filaments sometimes fused with staminodes.
usually spreading, usually divided to the base into Staminodes (0- )6( -12), bifid, laciniate, truncate or
3 segments; median segment usually erect, often irregularly divided or vestigial, erect or rarely
clawed, clasping the stamen; two lateral segments incurved, but not forming an envelope round the
spreading, shorter than, equalling or exceeding the gynoecium, nearly always glabrous. Small annular
median segment, entire or deeply divided or disk occasionally+. Ovary 6-14-locular, hairy or
laciniate, or less frequently corolla lobe only par- glabrous, placentation axile or basi-ventral; style
404 T.D. Pennington
exserted. Fruit 1-several-seeded. Seed ellipsoid to Trees. Venation craspedodromous with a promi-
obovoid, laterally compressed, with a hard shining nent marginal vein; tertiaries obscurely reticulate.
woody testa; hilum nearly always narrowly elon- Stipules 0. Flowers in small fascicles. Corolla hairy;
gate, basi-ventral or less frequently extending tube equalling the lobes, carnose; lobes 6, con-
along most of the adaxial face, very rarely broad. torted in bud, median segment erect or slightly
Embryo with foliaceous cotyledons and exserted spreading, abruptly narrowed at base, two lateral
radicle; endosperm copious. Pantropical, 30 spe- segments reduced to small irregularly shaped ves-
cies in America, c. 20 in Africa and Madagascar tiges at the base of the median segment or 0.
and c. 15 in Asia and the Pacific. Stamens 6. Staminodes 0 or minute. Ovary 6-
locular, hairy, placentation axile; style slightly
exserted. Fruit 1-seeded. Seed broadly ellipsoid,
8. Labramia A. de Candolle
not laterally compressed, with a woody testa;
Labramia A. de Candolle, Prodr. 8: 672 (1844). hilum broad, adaxial, covering about a third of the
seed surface. Embryo with thick plano-convex,
Trees or shrubs. Venation brochidodromous, partially fused cotyledons; radicle extending to the
higher order venation obscure. Stipules small, surface; endosperm 0. A single species, N. seychel-
caducous or 0. Flowers fasciculate. Corolla lana J.D. Hooker, in the Seychelles.
glabrous; tube shorter than the lobes; lobes 6( -8),
divided to the base into 3 segments; median
11. Labourdonnaisia Bojer
segment erect, clasping the stamen; two lateral
segments erect or spreading; almost equalling or Labourdonnaisia Bojer, Mem. Soc. Phys. Geneve 9: 295 (1841).
much shorter than the median segment, often
deeply divided or laciniate. Stamens 6( -8). Sta- Trees with sympodial branching. Venation brochi-
minodes 6( -8), usually reduced to a small fleshy dodromous, secondaries often joining to form a
vestige. Ovary 8-12-locular, glabrous, placentation submarginal vein; tertiaries parallel to the sec-
axile; style exserted. Fruit 1-seeded. Seed ellipsoid, ondaries. Stipules 0. Flowers solitary or in small
not or only slightly laterally compressed, with a fascicles. Corolla glabrous; tube much shorter than
hard shining woody testa; hilum broad, covering the lobes; lobes (10-)12-18, imbricate in bud,
the adaxial surface. Embryo with foliaceous spreading or refl.exed, entire or with a few small
cotyledons and exserted radicle; endosperm irregular lateral teeth. Stamens 11-18(-21). Sta-
copious. Eight species in Madagascar. minodes 0 or a few small irregular vestiges. Ovary
5-1 0-locular, hairy, placentation basi-ventral; style
included or slightly exserted. Fruit 1 (-2)-seeded.
9. Faucherea Lecomte
Seed narrowly ellipsoid, not or only slightly later-
Faucherea Lecomte, Bull. Mus. Hist. Nat. (Paris) 26:245 (1920). ally compressed, with a shiny woody testa; hilum
basal or basi-ventral, broad, often strongly
Trees. Venation brochidodromous; secondaries concave, embryo with foliaceous cotyledons and
numerous, higher order venation parallel to the long exserted radicle; endosperm copious. Three
secondaries, obscure, leaves appearing striate. imperfectly known species in Madagascar.
Stipules 0. Flowers fasciculate. Corolla glabrous;
tube much shorter than lobes; lobes 6( -11 ), entire.
12. Letestua Lecomte
Stamens 6(-11), anthers hairy or glabrous. Sta-
minodes 6( -11), usually very short, irregular or Letestua Lecomte, Notul. Syst. (Paris) 4: 4 (1920).
dentate. Small annular disk sometimes + around
base of ovary. Ovary (5-)6(-10)-locular, hairy; Trees. Venation eucamptodromous with widely
placentation axile; style exserted. Fruit 1 (-4)- spaced secondaries; tertiaries horizontal; higher
seeded. Seed slightly laterally compressed, with a order venation reticulate. Stipules 0. Flowers fasci-
hard shining testa; hilum basi-ventral, less than culate. Corolla glabrous; tube much shorter than
half as long as the seed. Embryo with foliaceous lobes; lobes 12-18, erect or slightly spreading,
cotyledons and exserted radicle; endosperm divided to the base into 3 segments; median
copious. Eleven species in Madagascar. segment slightly exceeding the lateral segments, all
petaloid. Stamens 12-18. Staminodes 0 (occasion-
ally a few stamens lack anthers). Ovary 16-18-
10. Northia J.D. Hooker
locular, hairy, placentation basi-ventral; style
Northia J.D. Hooker in Hooker's Icon. Pl. t. 1473 (1884). included. Fruit 1-seeded. Seed ellipsoid, laterally
Sapotaceae 405
compressed, with a shining woody testa; hilum well-developed, erect, lanceolate, glabrous. Ovary
long, narrow, adaxial. Embryo with foliaceous hairy; style included. Fruit 1-seeded, indehiscent,
cotyledons and exserted radicle; endosperm fleshy. Testa smooth or rugose; hilum long and
copious. A single species in West Africa, L. duris- rather broad, adaxial. Embryo with thinly plano-
sima (A. Chevalier) Lecomte. convex cotyledons; endosperm 0. Two species, one
in East Africa, one in West Africa.
3. SuBTRIBE GLUEMINAE Baehni ex

14. Neolemonniera Heine
T.D. Pennington (1991).
Neolemonniera Heine, Kew Bull. 16: 301 {1960).
Unarmed trees. Leaves spirally arranged. Inflores-
cences axillary or in the axils of fallen leaves; Leaves densely clustered at shoot tip. Venation
flowers fasciculate, bisexual. Corolla tube shorter eucamptodromous-brochidodromous with
than the lobes (± equal in some Eberhardtia). slightly convergent secondary veins, the tertiaries
Calyx a single whorl of 5 imbricate or quincuncial forming a lax reticulum; numerous fine minute
sepals; corolla lobes, stamens, staminodes and striations perpendicular to the mibrib. Stipules +.
ovary loculi usually 5; stamens in a single whorl Corolla hairy; lobes 5, sub erect, divided to near the
inserted at the corolla tube, filaments free; sta- base into 3 segments, median segment exceeding
minodes hairy or glabrous; ovary 5-locular, pla- the 2 lateral ones. Stamens 5; anthers extrorse,
centation axile; fruit dehiscent or not, seed pubescent or glabrous. Staminodes 5, alternating
ellipsoid, laterally compressed, radicle exserted; with the stamens, lanceolate, hairy, inflexed and
hilum long, usually narrow, adaxial. forming an envelope around the gynoecium.
Ovary hairy; style slightly exserted. Fruit a 1-
KEY TO THE GENERA OF GLUEMINAE seeded, leathery capsule dehiscent by a single
lateral valve. Seed somewhat asymmetric; testa
1. Fruit indehiscent 13. Inhambanella
- Fruit dehiscent 2 smooth, shining; hilum long, narrow, adaxial.
2. Stipules + 3 Embryo with thick flat cotyledons; endosperm a
-%~~0 4 thin layer. Five species in West Africa.
3. Leaf without fine parallel striations, corolla & staminodes
glabrous, staminodes with terminal sagittate appendage;
fruit 3-5-seeded 17. Eberhardtia 15. Lecomtedoxa (Pierre ex Engler) Dubard
- Leaf with numerous fine parallel striations perpendicular
to midrib; corolla and staminodes hairy, staminodes Lecomtedoxa (Pierre ex Engler), Notul. Syst. (Paris) 3: 46
without terminal appendage; fruit 1-seeded {1914).
14. Neolemonniera Mimusops subgen. Lecomtedoxa Pierre ex Engler {1904).
4. Staminodes alternating with stamens, free
15. Lecomtedoxa Leaves clustered at shoot apex. Venation eucamp-
- Staminodes opposite the stamens, partially fused
16. Gluema
todromous with convergent secondary veins, and
a lax tertiary reticulum, or brochidodromous and
then secondaries parallel and tertiaries parallel to
13. Inhambanella (Engler) Dubard Fig. 122 the secondaries and descending from the margin.
Inhambanella (Engler) Dubard, Ann. Inst. Bot.-Geol. Colon.
Stipules 0. Inflorescence axillary or ramiflorous.
Marseille II, 3: 42 (1915). Corolla hairy or glabrous; lobes 5, erect or slightly
Mimusops sect. Inhambanella Engler {1904). spreading, deeply divided into 3 segments; median
Kantou Aubreville & Pellegrin {1957). segment exceeding the lateral segments. Stamens
5; anthers extrorse, hairy. Staminodes 5, free, alter-
Leaves spaced or clustered at the shoot apex. Vena- nating with the stamens, lanceolate, often ter-
tion eucamptodromous with slightly convergent minated by a fine point, erect, glabrous or
secondary veins; tertiaries variable. Stipules small, occasionally hairy. Ovary hairy; style exserted.
caducous. Corolla glabrous, lobes (4- )5( -6), erect Fruit 1-seeded, dehiscent by a single lateral valve,
or slightly spreading, subdivided to halfway or leathery. Seed asymmetric, strongly laterally com-
more into 3 segments; median segment generally pressed; testas smooth, shining, woody; hilum
longer and broader than the lateral segments. long, narrow, adaxial. Embryo with thick flat
Stamens (4- )5( -6); anthers extrorse, glabrous. Sta- cotyledons; endosperm a thin layer. Five species in
minodes (4- )5( -6), alternating with the stamens, Gabon.
406 T.D. Pennington
G
.
0
. ....
~
'~\~// 'p
. F
' I '
E'· _..
Fig.122. Sapotaceae- Mimusupeae. Inhambanella henriquesii. tiaries finely areolate. Stipules 0. Corolla hairy;
A Flowering branch. B Flower. C Part of corolla, viewed from lobes 5, divided to the base into 3 segments;
outside. D Corolla, opened out, viewed from inside, showing
stamens and staminodes. E Stamen. F Staminode. G Flower median segment erect, clasping the stamen; lateral
with two sepals and corolla removed to show pistil. H Fruit. segments equalling the median segment, widely
I Seeds. (Hemsley 1968) spreading. Stamens 5; anthers extrorse, hairy. Sta-
minodes 5, fixed inside and opposite the stamens,
fused at the base, lanceolate, densely hairy, inflexed
16. Gluema Aubreville & Pellegrin and forming an envelope around the gynoecium.
Gluema Aubreville & Pellegrin, Bull. Soc. Bot. France 81: 797 Ovary hairy; style exserted. Fruit 1-seeded, dehis-
(1935). cent by a single lateral valve, leathery. Testa
smooth, shining, woody; hilum long, narrow,
Leaves clustered at the shoot apex. Venation adaxial. Embryo, with thinly plano-convex cotyle-
eucamptodromous-brochidodromous with dons; endosperm 0. A single species in West Africa
slightly convergent or parallel secondaries; ter- (G. ivorensis Aubreville & Pellegrin).
Sapotaceae 407
17. Eberhardtia Lecomte - Leaves alternate and distichous; tertiary venation parallel
to secondaries and descending from the margin; embryo
Eberhardtia Lecomte, Bull. Mus. Hist. Nat. (Paris) 26: 345 with thin foliaceous cotyledons, endosperm copious
(1920). 22.Payena
6. Fascicles densely clustered at shoot apex forming a
Venation eucamptodromous, with parallel second- pseudo-terminal inflorescence; sepals 4; corolla lobes 8(9),
imbricate; ovary (3)4(5)-locular 23. Burckella
aries, tertiaries oblique, higher order venation - Fascicles axillary or ramifiorous; sepals usually 4-5 or 5;
finely reticulate-areolate. Stipules well-developed. corolla lobes 8-16, contorted; ovary (5)6-15-locular
Corolla glabrous, tube equalling or slightly shorter 24. Diploknema
than the lobes; lobes 5, divided for c. 2/3 of their
length into 3 segments; median segment linear or
18. Palaquium Blanco
narrowly subulate, lateral segments slightly longer
and much broader than the median segment. Palaquium Blanco, Fl. Filip.: 403 (1837); ed. 2: 282 (1845).
Stamens 5, included; anthers dehiscing laterally,
glabrous. Staminodes 5, alternating with the Leaves spirally arranged; venation usually
stamens, exceeding the stamens, narrowly lanceo- eucamptodromous with oblique to horizontal ter-
late, glabrous, bearing at the apex a large sagittate, tiaries or rarely brochidodromous and then with
versatile, caducous appendage. Ovary hairy or tertiaries descending from the margin and paral-
glabrous; style included. Fruit a 3-5-seeded loculi- lel to the secondaries. Stipules +,usually small and
cidal capsule, leathery-fleshy, slightly constricted caducous, sometimes 0. Flowers usually bisexual,
between the seeds. Testa smooth, shining; hilum rarely unisexual and plant dioecious, in !-many-
narrow or broad, adaxial, extending the length of flowered fascicles, axillary or in axils of fallen
the seed. Embryo with thin foliaceous cotyledons; leaves. Sepals 3 + 3, free or slightly united, the outer
endosperm copious. Three poorly defined species ± valvate. Corolla lobes (5)6, usually contorted, less
of montane forest in South China, Vietnam and frequently imbricate, usually spreading or
Laos; recently collected in Sabah. reflexed, nearly always exceeding the tube, rarely
only equalling it; corolla tube usually glabrous
inside. Stamens usually 12, less frequently 10, 13,
II. TRIBE ISONANDREAE Hartog (1878). 18,24 or c. 30, exserted, in a single whorl or rarely
in 2 whorls, inserted near top of corolla tube or at
Calyx usually 2 whorls of 2 or 3 sepals, the outer base of corolla lobes; filaments free or occasionally
whorl valvate, less frequently a single whorl of 4-5 a few partly fused, tapering to an acute apex, hairy
imbricate or quincuncial sepals; corolla cyathi- or glabrous, 0 in female flowers. Nectary 0 (repre-
form, lobes as many as or 2-3 times as many as sented by a small annular swelling fused to ovary).
sepals, undivided; stamens 2-3( -5) times as many Ovary hairy, rarely glabrous, (5)6( -10)-locular;
as corolla lobes, in 1-3 whorls; staminodes 0; style long exserted. Fruit 1-2( -several)-seeded.
hilum adaxial. Seed usually broadly oblong or ellipsoid, with
broad adaxial hilum covering up to 2/3 of the
KEY TO THE GENERA OF ISONANDREAE surface and then without endosperm and embryo
with plano-convex cotyledons, or less frequently
1. Calyx biseriate, 2 whorls of 3 sepals 2
- Calyx uniseriate or if biseriate then with 2 whorls of 2 seed laterally compressed with a narrow adaxial
R~ 3 hilum and then usually with endosperm and
2. Stamens free; inflorescences axillary or just below the embryo with foliaceous cotyledons, radicle usually
leaves, fascicles sessile 18. Palaquium extending to the surface, less frequently exserted.
- Filaments partially united in a stamina! tube; plant
caulifiorous or ramifiorous, fascicles produced on scaly
About 110 species from India, through SE Asia to
brachyblasts 19. Aulandra the Pacific Islands.
3. Calyx biseriate with 2 whorls of 2 sepals 4
- Calyx uniseriate with 4 or 5 imbricate sepals 6
4. Flowers subsessile; corolla lobes 4( -5); stamens 8( -10); 19. Aulandra H.J. Lam
ovary loculi 4( -5); seed with copious endosperm
Aulandra H.J. Lam, Bull. Jard. Bot. Buitenzorg III, 8:415 (1927).
20. Isonandra
- Flowers distinctly pedicellate; corolla lobes (6)7-12(-17);
stamens (12)13-36( -43); ovary loculi (4-)6-9( -15); Leaves spirally arranged, venation eucamptodro-
endosperm + or 0 5 mous with oblique tertiaries. Stipules small,
5. Leaves nearly always spirally arranged; tertiary venation caducous. Flowers bisexual, cauliflorous or rami-
usually oblique or reticulate; embryo usually with plano-
convex or thick fiat cotyledons, endosperm thin or 0 florous, in few-flowered fascicles borne terminally
2l.Madhuca
408 T.D. Pennington
on densely scaly, simple or divaricately branched in a single whorl on the same individual, aestiva-
brachyblasts up to 3 em long. Sepals 3 + 3, free or tion of outer pair valvate, imbricate or open.
united at base, the outer ± valvate. Corolla lobes 6, Corolla lobes (6-)8-12( -17), imbricate, usually
imbricate or contorted, spreading or reflexed, spreading or reflexed, nearly always equalling or
exceeding tube; corolla glabrous. Stamens 18-19, exceeding the tube; corolla tube usually hirsute or
exserted, in a single whorl inserted at the top of barbate at the throat. Stamens (12-)14-36(-43),
the corolla tube; filaments united for more than exserted (uppermost whorls only), in 1-3 alternat-
half their length, free portion usually geniculate in ing whorls inserted in the throat of the corolla
bud; anthers extrorse, hairy. Nectary 0. Ovary tube, or occasionally the lowermost whorl inserted
hairy, 6(-7)-locular; style exserted. Berry 1- near the base of the tube; filaments usually free,
seeded. Seed broadly ovoid, with a broad adaxial rarely partially or completely fused in pairs or into
hilum covering of seed surface. Embryo with a short tube, not geniculate, or anthers sessile.
plano-convex cotyledons, radicle extending to the Anthers extrorse or laterally dehiscent, usually
surface; endosperm 0. Three species in Borneo. large and tapering gradually to the acute apex,
hairy or glabrous. Nectary sometimes +, poorly
developed, annular. Ovary usually glabrous,
20. Isonandra Wight
less frequently hairy, (5-)8-9( -15)-locular, style
Isonandra Wight, Ic. Pl. 2:4, t. 359,360 (1840); 4: 9, t.1219, 1220 exserted, often long, tapering gradually to the
(1848). apex. Fruit 1-seeded. Seed broadly ellipsoid,
oblong, plano-convex or laterally compressed;
Leaves spirally arranged. Venation eucamptodro- hilum usually long narrow adaxial, less frequently,
mous with oblique or horizontal tertiaries. Stip- broader and covering up to of the seed surface.
ules often small and caducous. Flowers bisexual, in Embryo with plano-convex, thick but flat,
small dense-flowered subsessile fascicles, some- rarely foliaceous cotyledons; radicle exserted.
times ramiflorous. Sepals± free, 2 + 2 (2 + 3), the Endosperm + or 0. About 100 species from India
outer pair open or imbricate. Corolla lobes 4(5), through Malesia and South China to New Guinea.
imbricate, rarely contorted, erect or slightly
spreading, equalling or exceeding the tube; corolla
tube glabrous within. Stamens 8( -10), inserted in 22. Payena A. de Candolle
a single whorl at the top of the tube; filaments free Payena A. de Candolle, Prodr. 8: 196 (1844).
or only slightly fused at base, strongly geniculate
in bud; anthers extrorse, usually with an apical tuft Leaves distichous, sometimes spirally arranged on
of hairs. Nectary 0. Ovary hairy, 4( -5)-locular; vertical shoots. Venation brochidodromous, inter-
style slightly exserted. Berry 1-seeded. Seed later- secondary and tertiary venation descending from
ally compressed, with a long narrow adaxial hilum. the margin and more or less parallel to the sec-
Embryo with foliaceous cotyledons and exserted ondaries. Stipules usually caducous. Flowers bis-
radicle; endosperm copious. About ten species in exual, in axillary fascicles. Sepals 2 + 2, ± free, the
South India, Sri Lanka, Malay Peninsula and outer valvate or open. Corolla lobes 7-9, imbricate,
Borneo. erect or slightly spreading, exceeding tube; tube
usually glabrous inside. Stamens 13-20( -30),
21. Madhuca Hamilton ex Gmelin inserted in 1(2) whorls at the top of the tube; fila-
ments free or fused in pairs or bundles, often
Madhuca Hamilton ex Gmelin, Syst. 2: 773,799 (1791). geniculate in bud. Anthers extrorse or laterally
Ganua Pierre ex Dubard (1908). dehiscent, large, tapering gradually to an acute
apex. Nectary 0, or a poorly developed annulus
Leaves spirally arranged, very rarely distichous on around the base of the ovary. Ovary hairy or
horizontal shoots; venation eucamptodromous or glabrous, (4-)6-8( -9)-locular; style exserted, often
brochidodromous, with oblique, horizontal or long, tapering gradually to the apex. Fruit 1-2-
reticulate tertiaries, rarely intersecondaries and seeded. Seed slightly to strongly laterally com-
tertiaries descending from the margin and paral- pressed, with a long narrow adaxial hilum. Embryo
lel to the secondaries. Stipules +, sometimes large with foliaceous cotyledons and exserted radicle;
and persistent. Flowers bisexual in 2-many- copious endosperm. About 15 species in western
flowered fascicles, usually on young branches in Malesia from the Andaman Islands and Burma to
the axils of fallen leaves or axillary. Sepals 2 + 2, the Malay Peninsula, Sumatra, Java to Borneo and
free, rarely a mixture of 2 + 2 and 5 imbricate sepals Mindanao.
Sapotaceae 409
23. Burckella Pierre a broad or narrow adaxial hilum. Embryo with

plano-convex or thin foliaceous cotyledons,
Burckella Pierre, Not. Bot. Sapot.: 3 (1890).
radicle extending to the surface or exserted;
endosperm+ or 0. About 10 species from north-
Leaves spirally arranged. Venation brochidodro-
ern India, Nepal and Bhutan to Burma, Thailand,
mous, tertiaries often descending from the margin
Cambodia and Vietnam; also m Borneo,
and parallel with the secondaries, or reticulate or
Philippines and Amboina.
oblique. Stipules + or 0. Flowers bisexual, the fas-
cicles clustered densely in the axils of scale leaves
to form a pseudo-terminal inflorescence. Sepals 2
III. TRIBE SIDEROXYLEAE (Engler)
+ 2, strongly imbricate. Corolla lobes 8(9), imbri-
cate, erect or slightly spreading, exceeding the
H.J. Lam (1938).
tube; corolla tube nearly always barbate in
Unarmed or spinose trees or shrubs. Sepals 5, in a
the throat. Stamens 16-18(-30) in 1-2 whorls at
single whorl, imbricate, sometimes quincuncial;
the top of the corolla tube; filaments free, not
corolla lobes, stamens and staminodes as many as
geniculate, or anthers sessile. Anthers extrorse.
sepals; corolla rotate or cyathiform; corolla lobes
Nectary annular or patelliform, often enveloping
often divided into 3 segments, a larger median one
the ovary, rarely 0. Ovary hairy or glabrous,
and 2 smaller lateral ones; stamens in a single
(3)4(5)-locular, style long, exserted. Fruit 1-
whorl, exserted, fixed at top of corolla tube; sta-
seeded. Seed broadly ellipsoid or dorso-ventrally
minodes usually well-developed, rarely 0; disk 0;
compressed, hilum covering at least 1/2 the surface
seed hilum basal or basi-ventral.
of the seed and frequently nearly all of it and then
leaving only a narrow adaxial strip; hilum area
KEY TO THE GENERA OF SIDEROXYLEAE
often rugose or ruminate, sometimes with promi-
nent protuberances. Embryo with plano-convex 1. Leaves usually opposite; stipules +; inflorescence a raceme
cotyledons, radicle extending to the surface, not or panicle 29. Sarcosperma
exserted, endosperm 0. About 14 species in the - Leaves usually spirally arranged or fascicled on short
shoots; stipules 0; inflorescence nearly always a fascicle 2
western Pacific from the Moluccas and New 2. Corolla lobes and stamens usually 7-10, staminodes 1-2
Guinea to Fiji, Samoa and Tonga. 27. Nesoluma
- Corolla lobes and stamens usually 4-5, staminodes
equalling the number of corolla lobes or 0 3
24. Diploknema Pierre 3. Staminodes 0 30. Diploon
- Staminodes + 4
Diploknema Pierre, Arch. Neerl. Sci. Exact. Nat.19: 103 (1884). 4. Ovary 2-3-locular; fruit containing several seeds com-
pletely fused to form a single pyrene, hilum 0 28. Argania
Leaves spirally arranged. Venation eucamptodro- - Ovary usually 5-locular; fruit containing 1 or more free
mous, rarely brochidodromous, tertiaries oblique seeds with usually basal hilum 5
5. Flowers unisexual and plant dioecious, female flowers with
or reticulate. Stipules often small and caducous. vestigial corolla, and without stamens and staminodes; fil-
Flowers bisexual or unisexual (plant dioecious), in aments in male flowers short, anthers included or only
axillary fascicles or ramiflorous. Sepals a single slightly exserted; hilum almost full-length, adaxial
whorl of (4)5( 6) free or partly united, imbricate or 26. Neohemsleya
quincuncial sepals. Corolla lobes 8-16, contorted - Flowers nearly always bisexual, rarely unisexual but then
female flower lacking only anthers; filaments in bisexual or
(not known in some species), usually spreading or male flowers well-developed, anthers clearly exserted;
reflexed, exceeding tube; tube hairy or glabrous. hilum nearly always basal or basi-ventral 25. Sideroxylon
Stamens (10-)16-30( -80), inserted in 1 or 2, often
dense whorls at the top of the corolla tube; fila-
ments free or partially united in pairs or small 25. Sideroxylon L.
bundles, usually geniculate in bud. Anthers Sideroxylon L., Gen. Pl. ed. 5: 89 ( 1754).
extrorse, large, tapering to an acute apex, hairy or Bumelia Swartz, Prod. Veg. Ind. Occ.: 49 (1788), nom. cons.
glabrous; stamens in female flowers converted into Monotheca A. de Candolle (1844).
flat narrow staminodes. Nectary 0 or a small Dipholis A. de Candolle (1844), nom. cons.
Mastichodendron (Engler) H.J. Lam (1939).
annulus at base of ovary. Ovary hairy or glabrous, Sinosideroxylon (Engler) Aubreville (1963).
(5)6-15-locular; style often exserted, apex taper- Spiniluma Baillon ex Aubreville (1963).
ing and simple or truncate and then with minute
stigmatic lobes. Fruit 1-4-seeded. Seed ellipsoid or Spinous or unarmed trees or shrubs. Leaves spi-
laterally compressed (known in only 4 spp.), with rally arranged, rarely opposite, often fascicled on
410 T.D. Pennington
short lateral shoots. Venation variable. Stipules 0. minodes (4)5, much shorter than corolla lobes,
Inflorescence axillary or in the axils of fallen subulate, 0 in female flower. Ovary 5-locular, pla-
leaves, sessile or very rarely pedunculate. Flowers centation axile; style short, stout, included; stigma
solitary or fasciculate, bisexual or rarely unisexual minutely 5-lobed, apex more or less truncate.
(?dioecious). Sepals one whorl of 5( -8), quincun- Berry 1-2-seeded. Seed ellipsoid, not or slightly
cial, free. Corolla cyathiform, usually glabrous; laterally compressed, or plano-convex (when fruit
tube nearly always shorter than the lobes, rarely 2-seeded); testa smooth, shining; hilum almost full
equalling or exceeding it; lobes (4)5( -8), imbricate length, adaxial, almost as wide as seed. Embryo
or quincuncial, spreading, entire or divided into a vertical, with thin foliaceous cotyledons and
larger median segment and two smaller lateral long-exserted radicle; endosperm copious. A
segments. Stamens (4)5( -8), exserted; filaments single species in Tanzania (N. usambarensis
well-developed; anthers extrorse, usually glabrous; Pennington).
stamens sometimes converted into sterile stamin-
odes in male flowers. Staminodes (4)5( -8), usually
well-developed, often lanceolate, erose, infolded 27. Nesoluma Baillon
and incurved against the style, usually glabrous, Nesoluma Baillon, Bull. Mens. Soc. Linn. Paris 2: 964 (1891).
less frequently hairy. Ovary (1- )5( -8)-locular,
hairy or glabrous; placentation basi-ventral or Unarmed trees or shrubs. Leaves spirally arranged.
basal; style exserted or included. Fruit 1(-2)- Venation brochidodromous, secondaries parallel,
seeded, fleshy, usually glabrous. Seed globose, higher order venation finely areolate. Stipules 0.
ovoid, oblong or ellipsoid, not laterally com- Inflorescence axillary and in the axils of leaf scars,
pressed, rarely plano-convex when 2 seeds in a fasciculate. Flowers bisexual or unisexual (plant
fruit; testa smooth, shining, free from the pericarp, dioecious). Sepals 4-5, strongly imbricate or quin-
often thick and woody, often sculptured on the cuncial. Corolla glabrous, shortly tubular, tube
adaxial surface with several prominent thickened shorter than lobes; lobes (5-)7-10(-12), erect or
plates; hilum nearly always basal or basi-ventral, spreading, entire or sometimes with a small lateral
small, circular, lanceolate or elliptic, rarely adaxial segment, unequal, in female flower reduced to free
and then broad. Embryo vertical, oblique or hori- vestigial scales or 0. Stamens 7-10(-12), often
zontal, with thin foliaceous cotyledons, and more than 1 opposite each corolla lobe, exserted
copious endosperm, or with plano-convex cotyle- on well-developed filaments, with prominent
dons and then with a thin sheath of endosperm, or traces to the base of the tube; sometimes a
endosperm 0; radicle exserted. Forty nine species few stamens converted into sterile petaloid or
in the Neotropics, about 25 elsewhere (6 Africa, staminode-like structures; anthers extrorse,
6 Madagascar, 8 Mascarenes, 4 Asia, 1 in NW glabrous; stamens 0 in female flower. Staminodes
Pakistan, Afghanistan, Oman, Somalia, Ethiopia, irregular, usually only 1-2, well-developed, some-
Djibouti). times petaloid, glabrous; 0 in female flower. Ovary
3-5(6)-locular, slightly hairy, placentation axile;
style slightly exserted. Berry usually 1-seeded.
26. Neohemsleya T.D. Pennington
Seed obovoid or ellipsoid, not laterally com-
Neohemsleya T.D. Pennington, The genera of Sapotaceae: 175 pressed; testa smooth, shining, thick, woody;
(1991). hilum large, rounded, basi-ventral or basal.
Embryo vertical, oblique or horizontal, with thin
Unarmed dioecious trees. Leaves laxly spirally foliaceous cotyledons and exserted radicle;
arranged, not clustered or fascicled, simple. Vena- copious endosperm. Three poorly defined species,
tion eucamptodromous-brochidodromous with Hawai'i, Henderson Island, Rapa and Tahiti.
convergent secondaries, tertiaries few, reticulate to
horizontal. Stipules 0. Flowers axillary, solitary or
paired, unisexual. Sepals 5, free or slightly fused, 28. Argania Roemer & Schultes
quincuncial. Corolla (male) tube much shorter Argania Roemer & Schultes, Syst. Veg. 4: XLVI, 502 (1819),
than lobes; lobes (4)5, imbricate, entire, spreading, nom. cons.
(female) reduced to 5 minute, almost free petals,
persisting in fruit as slightly accrescent membra- Spinous shrubs or small trees. Leaves spirally
nous structures. Stamens (4)5, included or slightly arranged, becoming fascicled on short lateral
exserted; filaments short; anthers extrorse or lat- shoots. Venation eucamptodromous to brochido-
erally dehiscent; stamens 0 in female flower. Sta- dromous, with convergent secondaries, higher
Sapotaceae 411
order veins forming an open reticulum. Stipules 0. aries joining below the margin to form a sub-
Inflorescence axillary or in the axils of leaf scars, marginal vein, intersecondaries long, usually
fasciculate. Flowers bisexual. Sepals 5, quincuncial. extending to near the margin, giving the leaves a
Corolla glabrous; tube shorter than lobes; lobes 5, slightly striate appearance. Stipules 0. Flowers
imbricate, entire, spreading. Stamens 5, exserted bisexual, in axillary fascicles. Sepals 4-5, free.
on well-developed filaments, anthers extrorse, Corolla rotate, tube very short, greatly exceeded by
glabrous. Staminodes 5, subulate or toothed at lobes; lobes 4-5, widely spreading, simple. Stamens
base, glabrous. Ovary 2-3-locular, hairy; placenta- 4-5, exserted; filaments thickened basally. Stamin-
tion basi-ventral; style exserted. Fruit 1-3-seeded, odes 0. Ovary glabrous, 1-locular with 2 basal
fleshy. Seeds completely fused by their adaxial ovules. Fruit a 1-seeded berry. Seed with small
surfaces, the resultant woody pyrene ellipsoid broad basal or basi-ventral hilum; embryo with
to ovoid; testa smooth, shining, thick, woody, plano-convex, free cotyledons, radicle extending
attached to the pericarp along the lines of fusion to the surface; endosperm 0. One species, D.
of the seeds. Embryo often solitary by abortion, cuspidatum (Hoehne) Cronquist, South America.
vertical, with thin foliaceous cotyledons and
exserted radicle; endosperm abundant. A single
species in Morocco, introduced into Libya; natu- IV. TRIBE CHRYSOPHYLLEAE Hartog (1878).
ralized in Southern Spain.
Calyx a single whorl of 4-5( -11) imbricate or
quincuncial sepals; corolla lobes and stamens
29. Sarcosperma J.D. Hooker
usually same number as sepals; corolla tubular,
Sarcosperma J.D. Hooker in Bentham & J.D. Hooker, Gen. Pl. 2: cyathiform or rotate; corolla lobes undivided;
655 (1876). stamens exserted or included; staminodes small,
in a single whorl alternating with stamens or 0;
Unarmed trees. Leaves usually opposite, less fre- hilum adaxial or rarely basi-ventral.
quently spirally arranged. Venation eucamptodro-
mous, with convergent or parallel secondaries, KEY TO THE GENERA OF CHRYSOPHYLLEAE
tertiaries horizontal. Hollow pits sometimes
present on lower surface in axils of secondary 1. Corolla rotate or tubular with spreading lobes; stamens
fixed at the top of the corolla tube and exserted 2
veins. Petiole sometimes bearing 2 small scales - Corolla tubular or cyathiform with more or less erect
(stipels). Stipules +. Inflorescence axillary or in lobes; stamens fixed within or at the top of the corolla tube,
axils of leaf scars, a small raceme or panicle. included 18
Flowers bisexual. Sepals 5, quincuncial. Corolla 2. Staminodes +, seed endospermous or not 3
- Staminodes 0; seed without endosperm or rarely with a
glabrous; tube shorter than lobes; lobes 5, imbri-
thin sheath of endosperm 12
cate or quincuncial, entire, sometimes auriculate 3. Seed nearly always endospermous; embryo with thin foli-
at base, widely spreading. Stamens 5, slightly aceous cotyledons and exserted radicle 4
exserted, with very short filaments; anthers - Seed without endosperm; embryo with plano-convex
introrse or laterally dehiscent, glabrous. Stamin- cotyledons, radicle usually included 6
4. Leaves alternate and distichous; secondary venation
odes 5, small, glabrous. Ovary (1)2-locular, closely parallel, secondary and higher order venation
glabrous, placentation basal; style short, included; indistinguishable, the leaf appearing finely striate
stigma minutely 2-4-lobed. Berry 1 (-2)-seeded. 34. Micropholis (sect. Exsertistamen)
Seed ellipsoid or oblong, not laterally compressed - Leaves spirally arranged; venation not closely parallel,
(plano-convex in 2-seeded fruit); testa smooth, leaves not finely striate 5
5. Plant often spiny; flower buds slender, acute; large anthers
thin; hilum small, round, basal or basi-ventral. with connective produced at apex, usually closely pressed
Embryo vertical, with plano-convex, usually fused, against corolla lobes; staminodes often aristate and hairy;
cotyledons, radicle included or extending to the disk 0; style long-exserted 48. Xantolis
surface, endosperm 0. About 8 species from India - Plant not spiny; flower buds short and rounded; anther
to South China and Malesia. connective not produced, anthers not pressed against
corolla lobes; staminodes not aristate, glabrous; annular
disk often +; style included or slightly exserted
30. Diploon Cronquist 31. Pouteria (sect. Pierrisideroxylon)
6. Leaves usually alternate and distichous; corolla tube, fila-
Diploon Cronquist, Bull. Torrey Bot. Club 73: 466 (1946). ments and staminodes carnose; seed laterally compressed
with narrow adaxial hilum 40. Sarcaulus
- Leaves spirally arranged, opposite or verticillate; corolla
Leaves spaced, distichous or weakly spirally tube, filaments and staminodes not carnose; seed not !at-
arranged, venation brochidodromous, the second- erally compressed, hilum nearly always broad 7
412 T.D. Pennington
7. Usually stipulate; secondary veins close, parallel; higher 23. Flowers bisexual; stamens inserted near the top of the
order venation parallel to the secondaries, the leaf appear- corolla tube; hilum adaxial and extending around the base
ing coarsely striate; filaments often partially or completely 33.Breviea
fused into a staminal tube 47. Englerophytum - Flowers unisexual; stamens inserted halfway up the corolla
- Stipulate or not; secondary veins not closely parallel, leaves tube; hilum adaxial 32. Aubregrinia
not striate; filaments free 8 24. Stipules + 37. Ecclinusa
8. Small caducous stipules +; corolla lobes with contorted - Stipules 0 25
aestivation; staminodes densely woolly, incurved, forming 25. Seed without endosperm 26
a cap above the ovary; anthers closely applied to corolla - Seed endospermous 36. Chrysophyllum
lobes 49. Capurodendron 26. Indumentum of long, stiff, simple hairs; leaves glandular-
- Stipulate or not; corolla lobes with imbricate or valvate aes- striate 38. Delpydora
tivation; staminodes not densely woolly, not in curved over - Indumentum of short malpighiaceous hairs; leaves not
the ovary; anthers not closely applied to the corolla lobes glandular-striate 31.Pouteria (sect. Oxythece)
9
9. Stipules +; leaves spirally arranged 46. Synsepalum
- Stipules 0; leaves often opposite or verticillate 10 31. Pouteria Aublet
10. Higher order venation nearly always finely areolate;
annular disk often + 39. Pichonia Pouteria Aublet, Hist. Pl. Guiane 1: 85, pl. 33 (excl. fruit)
- Higher order venation not finely areolate; disk 0 11 (1775).
11. Leaves spirally arranged; hilum covering at least half the For generic synonymy see Pennington (1991).
seed surface 46. Synsepalum
- Leaves often opposite or verticillate; hilum covering less Trees or shrubs, rarely geoxylic suffrutices. Leaves
than half the seed surface 31. Pouteria (sect. Gayella)
12. Leaves opposite; calyx of 4 sepals; long filiform exserted spirally arranged, rarely opposite. Venation
style 44. Leptostylis eucamptodromous or brochidodromous, usually
- Leaves spirally arranged; calyx of 5( -6) sepals; style without a submarginal vein, never finely striate
exserted or not 13 (except P. keyensis). Stipules 0 (+in P. congestifo-
13. 2-4 stamens opposite each corolla lobe 45. Pycnandra lia). Inflorescence axillary or ramiflorous, fascicu-
- 1 stamen opposite each corolla lobe 14
14. Stipules + 15
late, fascicles single or occasionally arranged along
- Stipules 0 16 short leafless shoots. Flowers often unisexual
15. Secondary veins close, parallel; higher order venation par- (plant dioecious). Sepals 4-6, free, imbricate or
allel to the secondaries, the leaf appearing coarsely striate; quincuncial, or 6-11 in a closely imbricate spiral.
filaments often partially or completely fused into a stami- Corolla cyathiform to tubular, rarely rotate, tube
na! tube 47. Englerophytum
- Secondary veins not closely parallel, leaves not striate; fil- shorter to longer than the lobes, lobes 4-6( -9),
aments free 46. Synsepalum usually erect, rarely spreading, simple, sometimes
16. Lower leaf surface usually minutely punctate; higher order fringed-ciliate or papillose. Stamens 4-6( -9), fixed
venation obscure; ovary 2-3(4)-locular, hilum covering inside corolla tube, or rarely at base of lobes, rarely
less than half the seed surface; embryo with exserted
radicle; thin sheath of endosperm +
free, usually included, less frequently exserted; fil-
41. Elaeoluma
- Lower leaf surface not punctate; higher order venation not aments generally short; anthers usually extrorse or
obscure; ovary (2)3-locular; hilum often covering more laterally dehiscent, usually glabrous. Staminodes
than half the seed surface; embryo with exserted or usually same number as corolla lobes, less fre-
included radicle; endosperm usually 0 17 quently lacking, inserted in the corolla sinus or
17. Corolla lobes and stamens (4)5-10; fruit a berry; hilum
often covering or more of seed surface; radicle included
inside the tube, sometimes fringed-ciliate or papil-
42. Niemeyera lose. Disk+ or 0. Ovary 1-6(-15)-locular, placen-
- Corolla lobes and stamens 5; fruit a drupe; hilum not cov- tation axile; style included or exserted. Fruit a
ering more than of the seed surface; radicle often exserted 1-several-seeded berry. Seed broadly ellipsoid,
43. Pradosia plano-convex, shaped like the segment of an
18. Staminodes + 19
- Staminodes 0 24
orange or laterally compressed, testa smooth,
19. Seed without endosperm 20 wrinkled or pitted; hilum adaxial, usually full-
- Seed endospermous 21 length, narrow, broad or sometimes covering
20. Plant with large stipules 35. Chromolucuma almost all the seed surface. Embryo vertical, with
- Stipules 0 (+ in Pouteria congestifolia) 31. Pouteria plano-convex or thin foliaceous cotyledons,
(sects. Rivicoa, Aneulucuma, Antholucuma, Pouteria,
Oxythece, Franchetella) radicle exserted or included; endosperm + or 0.
21. Leaves alternate and distichous; secondary venation About 200 species in the Neotropics, c. 120 species
closely parallel; leaf appearing striate 34. Micropholis in Asia, Malesia, Australia and the Pacific, c. 5
(sect. Micropholis) species in Africa. Nine sections distinguished by
- Leaves spirally arranged; secondary venation not closely
parallel; leaf not striate
Pennington (1991); see there for sectional descrip-
22
22. Ovary 7-9-locular 23 tions and synonymy.
- Ovary 5-locular 31.Pouteria (sect. Oligotheca)
Sapotaceae 413
32. Aubregrinia Heine appearing finely striate. Stipules 0. Inflorescence

axillary, ramiflorous or cauliflorous, fasciculate.
Aubregrinia Heine, Kew Bull. 14: 301 (1960). Flowers often unisexual. Sepals (4)5, free, im-
bricate or quincuncial. Corolla campanulate to
Dioecious trees. Leaves spirally arranged. Venation short- or long-cylindrical, the tube nearly always
eucamptodromous. Inflorescence axillary, fascicu- exceeding the lobes, rarely equalling them, lobes
late. Stipules 0. Flowers unisexual. Sepals 5, free, (4)5, erect to reflexed, simple. Stamens (4)5, fixed
quincuncial. Corolla broadly tubular, tube exceed- near the top of the corolla tube, included or
ing the lobes, lobes 5, erect, simple. Stamens 5, exserted; filaments short and straight or long and
fixed about halfway up the corolla tube, included; geniculate (at least in bud); anthers extrorse in
anthers extrorse, glabrous, 0 in female flower. Sta- bud, glabrous. Staminodes (4)5, in the corolla
minodes 5, inserted in the corolla sinuses. Disk +? sinuses, alternating with the stamens, usually
(according to Aubreville & Pellegrin present and lanceolate or subulate, or rarely petaloid. Disk+ or
fused to ovary). Ovary 7-8-locular, placentation 0. Ovary (4)5-locular, placentation axile; style
axile; style included. Berry large, several-seeded. included or exserted. Fruit 1-several-seeded. Seed
Seed laterally compressed, testa smooth; hilum laterally compressed, testa smooth or often
adaxial, full-length, narrow. Embryo vertical, with minutely transversely wrinkled, shining or dull;
thin foliaceous cotyledons, radicle exserted; hilum adaxial, extending the length of the seed,
endosperm copious. A single species, A. taiensis usually narrow. Embryo vertical, with thin folia-
(Aubreville & Pellegrin) Heine, in West Africa. ceous cotyledons and exserted radicle, surrounded
by thick endosperm. Thirty eight species in
33. Breviea Aubreville & Pellegrin Central and South America and the West Indies.
Two sections, 1. sect. Micropholis, corolla less
Breviea Aubreville & Pellegrin, Bull. Soc. Bot. France 81: 792
(1934); Aubreville, Fl. For. Cote d'Ivoire, ed. 2, 3: 130 (1959);
than lOmm long, lobes erect or only slightly
Heine, Kew Bull. 14: 302 (1960). spreading; stamens included stamens; thirty
species throughout tropical America; 2. sect.
Trees. Leaves distichous. Venation eucamptodro- Exsertistamen T. Pennington, corolla usually more
mous-brochidodromous. Stipules 0. Inflorescence than lOmm long; corolla lobes spreading or
axillary, fasciculate. Flowers bisexual. Sepals 5, reflexed, stamens exserted; nine spp. centred on
free, quincuncial. Corolla tubular, tube greatly the Guianas and extending across Brazilian
exceeding the lobes; lobes 5, erect, simple. Stamens Amazonia to Peru.
5, fixed near the top of the corolla tube, included;
anthers laterally dehiscent, glabrous. Staminodes 35. Chromolucuma Ducke
5, fixed in the corolla sinuses. Disk annular and
slightly lobed, fused to the ovary. Ovary 8-9- Chromolucuma Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 160
(1925); Trop. Woods 71: 20 (1942); Pennington, Fl. Neotrop.
locular, placentation axile; style included. Berry 52: 229-232 (1990).
large, several-seeded. Seed strongly laterally com- Pouteria sect. Chromolucuma (Ducke) Baehni (1942).
pressed, testa smooth; hilum adaxial and extend-
ing around the base, narrow. Embryo vertical, with Latex yellow. Leaves spirally arranged, venation
thin foliaceous cotyledons and exserted radicle; eucamptodromous. Stipules large. Flowers unisex-
endosperm copious. A single species, B. sericea ual (plants monoecious or dioecious ), pedicellate.
Aubreville & Pellegrin, in West Africa. Sepals 5, imbricate. Corolla cyathiform or shortly
tubular, tube usually equalling or slightly longer
34. Micropholis (Grisebach) Pierre than lobes (slightly shorter in male flowers of C.
rubriflora). Stamens fixed in the upper half or at
Micropholis (Grisebach) Pierre, Not. Bot. Sapot.: 37 (1891); the top of the corolla tube, included; filaments
Pierre & Urban, Symb. Ant. 5: 111 (1904); Pennington, Fl.
Neotrop. 52: 172 (1990).
short. Staminodes +, vestigial in female flower of
Sapota sect. Micropholis Grisebach (1861). C. baehniana. Disk 0. Ovary broadly truncate to
ovoid, 2-5-locular. Seed with dull rough testa, and
Trees or shrubs. Leaves spaced, distichous or spi- broad adaxial hilum covering up to two thirds of
rally arranged. Venation brochidodromous with a seed; embryo with plano-convex, free cotyledons,
submarginal vein, or craspedodromous, secondary radicle slightly exserted; endosperm 0. Two species
veins closely parallel, often not differentiated from in the Guianas, southern Venezuela and central
the higher order venation, and then the leaf Amazonian Brazil.
414 T.D. Pennington
36. Chrysophyllum L. simple. Stamens 5( -7) included, usually fixed near

Chrysophyllum L., Sp. Pl.: 192 (1753).
halfway or in the upper half of the corolla tube,
rarely m the lower half; filaments well-developed,
free; anthers extrorse, glabrous. Staminodes 0.
Trees or shrubs, very rarely Hanas. Leaves distic-
Disk 0. Ovary (3-)5( -9)-locular, placentation axile
hous or spirally arranged. Venation brochidodro-
or basi-ventral; style included. Fruit !-several-
mous or eucamptodromous, tertiary veins often
seeded, often thin-walled and constricted between
parallel to the secondaries and descending from
the margin, or oblique and closely parallel, or the seeds. Seed globose, ellipsoid, sometimes
reticulate. Stipules 0. Inflorescence axillary, rami- slightly laterally compressed or shaped like the
s~gment of.an orange, testa smooth, thin, shining;
florous or cauliflorous. Flowers unisexual or bisex-
ual, fasciculate or rarely solitary. Sepals (4)5(6), hilum adaxial and nearly always extending around
th~ base of the seed, usually narrow. Embryo with
imbricate or quincuncial, sometimes accrescent in
thick plano-convex cotyledons, radicle not
fruit, frequently ciliate. Corolla globose, campanu-
late or .cylindrical, tube shorter than, equalling or exserted, ex~ending to the surface; endosperm 0.
exceedmg the lobes, lobes (4)5( -8), simple. Eleven species, from Panama throughout tropical
South America.
Stamens (4)5( -8), fixed in the lower or upper part
of the corolla tube, included; anthers extrorse in
bud, hairy or glabrous. Staminodes rarely + as 38. Delpydora Pierre
small lanceolate or subulate structures in the
Delpydora Pierre, Bull. Mens. Soc. Linn. Paris 2: 1275 (1897).
corolla lobe sinuses, alternating with the stamens.
Disk 0. Ovary (4)5( -12)-locular, placentation axile,
Small trees. Indumentum of long stiff simple hairs.
style included. ~ruit 1-many-seeded. Seed laterally
Leaves spirally arranged, pellucid-striate, venation
compressed, with a narrow adaxial hilum, some-
times extending around base of seed, or not later- eucamptodromous or brochidodromous, ter-
tiari~s oblique. S~ipules 0. Inflorescence axillary or
ally compressed and then the hilum broader
ramiflorous, fasciculate. Flowers bisexual. Sepals 5,
basi-ventral or adaxial; testa smooth and shining:
free, quincuncial. Corolla shortly tubular, tube
or rough and then adherent to the pericarp.
Embryo vertical, with thin foliaceous or thick flat much ~onge~ than lobes; lobes 5, erect or slightly
cotyledons and exserted radicle, endosperm abun- spreadmg, simple. Stamens 5, fixed in the lower
half of the corolla tube, included; filaments well-
dant or about equalling the thickness of the cotyle-
developed, free; anthers extrorse, closely applied to
dons. Forty three species in the Neotropics, c.15 in
Africa, c. 10 in Madagascar, and 2-3 extending the style and sometimes laterally connivent,
glabrous. Staminodes 0. Disk 0. Ovary 5-locular,
from India to Malesia and Australia.
placentation axile; style included. Berry several-
Six sections; for descriptions, keys and syn-
onymy see Pennington (1991). seeded. Seed broadly oblong or shaped like the
segmen~ of an orange, not laterally compressed,
testa thm, smooth; hilum adaxial and extending
37. Ecclinusa Martius around the base, very narrow. Embryo with plano-
convex cotyledons and included radicle; endo-
Ecclinusa Martius, Flora 22, Beibl. 1: 2 (1839); Pennington, Fl.
Neotrop. 52: 622-639 (1990). sperm 0. Two species in West Africa.
Trees or rarely shrubs. Leaves spirally arranged, 39. Pichonia Pierre

usually loosely clustered at the shoot apex; vena-
tion usually eucamptodromous or rarely brochi- Pichonia Pierre, Not. Bot. Sapot.: 22 (1890).
dodromous, intersecondaries usually 0; tertiaries
us~ally oblique, numerous, close, parallel, rarely Trees .. Leaves opposite, rarely spirally arranged.
reticulate or areolate. Stipules caducous, leaving a Venation eucamptodromous-brochidodromous
conspicuous hilum. Inflorescence axillary or in the higher order venation finely reticulate-areolate:
axils of fallen leaves. Flowers sessile, subtended by Stipules 0. Inflorescence axillary, fasciculate, rarely
small persistent bracts, usually unisexual (monoe- on leafless short shoots. Flowers bisexual. Sepals
cious or dioecious). Sepals (4)5, free, quincuncial. (4)5, free, quincuncial. Corolla cyathiform to
Corolla small (usually less than 5mm long), cam- rotate, ~ube ~horter than the lobes; lobes 5, widely
panula~e or shortly tubular, the lobes usually spreadmg, Simple. Stamens 5, inserted at the top of
exceedmg the tube, rarely equalling it; lobes 5( -7), the corolla tube, exserted, with long filaments;
anthers laterally dehiscent or extrorse, with apical
Sapotaceae 415
tuft of hairs or glabrous. Staminodes 5, well- placentation axile; style short. Berry 1-seeded.
developed, fixed in the corolla sinuses. Disk Seed broadly ellipsoid, not or sometimes laterally
annular, stipitate or 0. Ovary (4)5(6)-locular, pla- compressed, testa smooth to slightly wrinkled,
centation axile; style exserted. Berry 1-seeded. shining; hilum adaxial, full-length, narrow or
Seed narrowly to broadly ellipsoid, not laterally broad. Embryo cotyledons plano-convex, radicle
compressed, testa smooth; hilum adaxial, full- slightly exserted; endosperm thin. Four species
length, broad, covering from to almost all the seed in southern Venezuela, the Guianas, Brazilian
surface, and then only a narrow abaxial strip Amazonia and Panama.
remaining free. Embryo with thick plano-convex
cotyledons and included radicle; endosperm 0.
42. Niemeyera F. Muell.
About 5 species in New Caledonia, Papua New
Guinea and the Solomon Islands. Niemeyera F. Muell., Fragm. 7: 114 (1870).
Trees or treelets, sometimes pachycaulous. Leaves

40. Sarcaulus Radlkofer
spirally arranged; venation eucamptodromous or
Sarcaulus Radlkofer, Sitzungsber. Math.-Phys. Cl. Konig!. brochidodromous, higher order venation oblique,
Bayer. Akad. Wiss. Miinchen 12: 310 (1882); Pennington, Fl. or descending from the margin and parallel to
Neotrop. 52: 232-239 (1990).
secondaries; stipules 0. Inflorescence fasciculate,
axillary or ramiflorous. Flowers bisexual. Sepals
Leaves spaced, distichous or less frequently weakly 5, free, imbricate or quincuncial. Corolla rotate or
spirally arranged, venation eucamptodromous or cyathiform with spreading lobes, tube nearly
brochidodromous. Stipules 0. Flowers unisexual always shorter than the lobes or equalling them;
(plant dioecious). Sepals usually 5. Corolla lobes (4)5-10, simple. Stamens (4)5-10, exserted,
globose, broadly cyathiform or subrotate, tube usually inserted at or near the top of the corolla
weakly to strongly carnose, equalling or slightly tube, rarely only halfway up the tube; filaments
exceeding the lobes; lobes usually 5, slightly imbri- well-developed, free, geniculate in bud; anthers
cate or subvalvate. Stamens usually 5, fixed at the versatile in open flower, hairy or glabrous. Sta-
top of the corolla tube, exserted; filaments short, minodes 0. Disk usually 0. Ovary (2)3-5-locular;
swollen; anthers strongly inflexed. Staminodes placentation axile; style exserted or not. Berry 1-2-
usually 5, thick, carnose, erect or incurved against seeded. Seed narrowly ellipsoid to globose, not lat-
the style. Disk 0. Ovary 2-5-locular. Berry erally compressed; hilum adaxial, broad, often
1-several-seeded. Seeds laterally compressed, testa covering up to or more of the seed surface. Embryo
smooth or wrinkled, shining; hilum adaxial, full- with plano-convex cotyledons; radicle included;
length, rather narrow; embryo with plano-convex, endosperm 0. About 20 species in New Caledonia
free cotyledons, radicle extending to the surface; and Australia.
endosperm 0. Five species in tropical South
America.
43. Pradosia Liais
41. Elaeoluma Baillon Pradosia Liais, Climat., Geol., Faune Bresil: 614 (1872);
Pennington, Fl. Neotrop. 52: 639-668 (1990).
Elaeoluma Baillon, Hist. Pl. 11: 293 (1891); Pennington, Fl.
Neotrop. 52: 240-247 (1990).
Leaves opposite or verticillate, less frequently spi-
rally arranged; venation usually eucamptodro-
Trees or shrubs. Leaves spirally arranged, usually mous, less frequenly brochidodromous, midrib
minutely punctate on the lower surface. Venation sunken on the upper surface, rarely flat or raised,
eucamptodromous or brochidodromous, higher secondaries often impressed on the upper surface;
order veins often obscure, forming a lax reticulum. intersecondaries usually 0, rarely well-developed;
Stipules 0. Inflorescence mostly axillary. Flowers tertiaries usually oblique or horizontal; minute
unisexual. Sepals (4)5, imbricate or quincuncial. paired scales (stipels) sometimes present on
Corolla broadly cyathiform to rotate, tube shorter petiole. Stipules 0. Inflorescences cauliflorous or
than the lobes or rarely equalling the lobes in ramiflorous, less frequently axillary. Flowers bisex-
female, lobes 5( 6), spreading, simple. Stamens 5( 6), ual. Sepals usually 5. Corolla rotate, tube nearly
fixed at the top of the corolla tube and exserted; always shorter than the widely spreading lobes,
filaments well-developed; anthers glabrous. Sta- corolla lobes 5. Stamens 5, fixed at top of corolla
minodes 0 (1-2). Disk 0. Ovary 2-3( -4)-locular, tube or on base of lobes, exserted; filaments long,
416 T.D. Pennington
geniculate below the apex, and strongly narrowed ing the lobes; lobes 5-10, widely spreading or
below insertion of anthers. Staminodes 0. Disk 0. reflexed, simple. Stamens (7-)10, 1-2 opposite
Ovary (4)5(6)-locular, style short. Fruit a drupe each corolla lobe, less frequently up to 25 and then
with thinly cartilaginous endocarp, often slightly 3-4 opposite each corolla lobe, exserted, fixed in a
asymmetric. Seed solitary, with smooth, shining single whorl in the upper half or at the top of the
testa and full-length adaxial hilum covering up to corolla tube; filaments well-developed, free or
one third of the seed surface; embryo with thinly sometimes partially united in pairs, geniculate in
plano-convex cotyledons, radicle often exserted or bud; anthers extrorse, glabrous or rarely hairy. Sta-
only extending to the surface; endosperm a thin minodes 0. Disk 0 (small annulus at ovary base in
sheath or 0. Twenty three species in South P. vieillardii). Ovary 5-8( -10 )-locular, placentation
America, with one extending into Panama and axile; style usually included. Fruit a 1-seeded
Costa Rica. berry. Seed broadly ellipsoid, not laterally com-
pressed, hilum adaxial, full-length, covering about
1/3 to 1/2 the surface, or almost all the surface,
44. Leptostylis Bentham
hilum area sometimes strongly rugose or rumi-
Leptostylis Bentham in Bentham & Hooker, Gen. Pl. 2: 659 nate. Embryo with plano-convex, smooth or rumi-
(1876). nate cotyledons, radicle extending to the surface;
endosperm 0. About 12 species confined to New
Shrubs or small trees. Leaves (sub)opposite; vena- Caledonia.
tion usually brochidodromous, less frequently
eucamptodromous; higher order venation reticu-
late. Stipules 0. Inflorescence fasciculate, axillary 46. Synsepalum (A. de Candolle) Daniell Fig. 123
or ramiflorous. Flowers bisexual. Sepals 4, free or Synsepalum (A. de Candolle) Daniell, Pharm. J. Trans. 11: 445
slightly united, strongly imbricate. Corolla short (1852).
to long-tubular, infundibuliform or rotate, tube Sideroxylon sect. Synsepalum A. de Candolle (1844).
Vincente/la Pierre (1891).
greatly exceeding, equalling or shorter than the
Pachystela Pierre ex Baillon (1891).
lobes; lobes 4-10, widely spreading or reflexed, Afrosersalisia A. Chevalier (1943).
simple. Stamens 4-10, exserted, inserted in the
upper half or at the apex of the corolla tube; fila- Shrubs or trees. Leaves spirally arranged. Venation
ments well-developed, free, geniculate in bud; nearly always eucamptodromous, rarely brochido-
anthers extrorse, usually glabrous, sometimes with dromous, higher order venation oblique or reticu-
a tuft of hairs at the apex. Staminodes 0. Disk 0. late; leaves not striate. Stipules + and often
Ovary 3-4(5)-locular, placentation axile; style conspicuous, or 0. Inflorescence axillary or rami-
exserted, often very long and filiform. Fruit florous, rarely cauliflorous. Flowers in fascicles,
(known in only L. jilipes and L. petiolata) a 1- bisexual. Sepals 5(6), free or partly united, quin-
seeded berry. Seed ellipsoid to narrowly ovate, not cuncial or rarely open. Corolla usually with short
laterally compressed; hilum adaxial, full-length, tube exceeded by the widely spreading lobes,
covering about to the seed surface. Embryo with rarely exceeding the lobes and then cyathiform or
plano-convex cotyledons, radicle included or tubular, lobes 5(6), usually widely spreading,
slightly exserted; endosperm 0. Eight species in simple; aestivation imbricate or less frequently
New Caledonia. induplicate valvate. Stamens 5(6), fixed at the top
of the corolla tube or rarely slightly inside the
45. Pycnandra Bentham throat, usually strongly exserted; filaments usually
well-developed, free; anthers usually extrorse,
Pycnandra Bentham in Bentham & Hooker, Gen. Pl. 2: 658 glabrous or less frequently hairy. Staminodes vari-
(1876).
able in size, glabrous, or 0. Disk 0. Ovary 5(-7)-
locular, placentation axile; style usually exserted.
Trees or treelets, often pachycaulous. Leaves spi-
Berry 1-seeded. Seed ellipsoid, not laterally com-
rally arranged or occasionally opposite; venation
pressed; testa smooth; hilum adaxial, broad,
usually eucamptodromous, less frequently brochi-
usually covering at least one third of the surface
dodromous; higher order venation oblique or
and sometimes nearly all of the surface. Embryo
reticulate. Stipules 0. Inflorescence axillary or ram-
with plano-convex cotyledons, radicle extending
iflorous. Flowers bisexual. Sepals 5(6), free, quin-
to the surface or slightly exserted; endosperm 0.
cuncial. Corolla tube cyathiform, usually shorter
Possibly 20 species in tropical Africa.
than or equalling the lobes, rarely slightly exceed-
Sapotaceae 417
+,often large and persistent. Inflorescence axillary,

ramiflorous or cauliflorous. Flowers in fascicles,
bisexual or unisexual. Sepals 5(6), free or slightly
united, quincuncial. Corolla shortly tubular or
cyathiform, the tube slightly shorter than,
equalling or slightly exceeding the lobes; lobes
5{-10), often widely spreading, simple, imbricate
or quicuncial. Stamens 5{ -10), fixed at the top of
the corolla tube, usually exserted; filaments
usually well-developed, and often partially or
completely fused into a short tube or thick fleshy
collar; anthers usually extrorse, glabrous; stamens
sometimes converted into petaloid staminodes in
female. Staminodes 0 or occasionally a single
whorl of small subulate structures. Disk 0. Ovary
(4)5(-10)-locular, placentation axile; style
included or slightly exserted. Berry 1-seeded. Seed
ellipsoid, not or only slightly laterally compressed;
testa smooth, often thin; hilum adaxial; variable in
width, from narrow to broad, and sometimes
extending to cover most of the seed surface.
Embryo with plano-convex cotyledons, radicle
extending to the surface; endosperm 0. Five to ten
poorly defined species in tropical Africa.
48. Xantolis Rafinesque

Xantolis Rafinesque, Sylva tellur. 36 (1838); van Royen, Blumea
8: 207 (1957); Aubreville, Fl. Cambodge, Laos et Vietnam 3.
Sapotacees: 74 (1963).
Trees or shrubs, often spiny. Stipules 0. Leaves

spirally arranged, sometimes fascicled on short
ti\
shoots. Venation eucamptodromous or brochido-
dromous. Inflorescence axillary or ramiflorous,
\I fasciculate, sometimes clustered along short leaf-
E
\
"~..---------:...~· I V!!JJJ H
less shoots. Flowers bisexual. Sepals 5, free, quin-
cuncial. Corolla with short tube nearly always
Fig. 123. Sapotaceae-Chrysophylleae. Synsepalum cerasi- exceeded by the spreading lobes; lobes 5, simple,
ferum . A Flowering branch. B Young flower. C Anthetic flower. imbricate; corolla tube nearly always hairy around
D Dissected calyx. E Dissected corolla showing stamens and the base of the filaments. Stamens 5, fixed at the
small staminodes. F Corolla segment with stamens. GOvary. top of the corolla tube, exserted or closely applied
H Fruit. I Seed. (Hemsley 1968)
to the corolla lobes; anthers large, connective pro-
duced at the apex, extrorse, glabrous. Staminodes
5, well-developed, often equalling the corolla
lobes, often fimbriate and aristate, often hairy.
47. Englerophytum Krause Disk 0. Ovary (4)5-locular, placentation axile; style
long-exserted. Berry 1(2)-seeded. Seed slightly to
Englerophytum Krause, Bot. Jahrb. Syst. 50, Suppl.: 343 (1914).
Bequaertiodendron De Wildeman (1919).
strongly laterally compressed; hilum adaxial,
narrow, equalling the length of the seed to only
Shrubs or trees. Leaves spirally arranged. Venation half as long. Embryo with thin, flat, foliaceous
brochidodromous, usually with a submarginal cotyledons and long exserted radicle; copious
vein, secondaries usually close and parallel, higher endosperm. About 14 species from southern India
order venation parallel to the secondaries, the to Vietnam and south China, one species in the
leaves appearing coarsely striate. Stipules usually Philippine Islands.
418 T.D. Pennington
49. Capurodendron Aubn!ville 50. Tsebona Capuron

Capurodendron Aubreville, Adansonia II, 2: 921 (1962); Fl. Tsebona Capuron, Adansonia II, 2: 122 (1962); Baehni,
Madag. 164. Sapotacees: 68 (1974). Boissiera 1: 105 (1965); Aubreville, Fl. Madag. 164,
Sapotacees: 123 (1974).
Trees or shrubs. Leaves spirally arranged. Venation
eucamptodromous or brochidodromous. Stipules Tree. Leaves spirally arranged. Venation eucamp-
usually +, caducous. Inflorescence axillary, fascic- todromous. Stipules +, well-developed, caducous.
ulate. Flowers bisexual. Sepals 5, free or slightly Flowers solitary, axillary, bisexual. Sepals 5, free,
fused, quincuncial. Corolla tube often slightly con- quincuncial, both margins of the outer 2 sepals
stricted at the apex, slightly shorter than, equalling and 1 margin of the third sepal in duplicate valvate,
or slightly longer than the lobes; lobes 5, spread- and completely enclosing the remaining sepals in
ing, simple, contorted. Stamens 5, fixed at the top bud. Corolla tube much shorter than the lobes,
of the corolla tube; anthers extrorse, usually lobes 5, slightly spreading, simple, contorted.
closely applied to the corolla lobes, hairy or Stamens in groups of 2-4 opposite each corolla
glabrous. Staminodes 5, well-developed, nearly lobe, fixed at the top of the corolla tube; filaments
always narrowly triangular and densely woolly, free, short, hairy; anthers about the length of the
closely applied to the style. Disk 0. Ovary 5(6)- corolla lobes, extrorse, hairy. Staminodes 5, much
locular, placentation axile; style exserted. Berry 1- shorter than the corolla lobes, doubly geniculate,
seeded. Seed globose to narrowly ellipsoid, not and pressed against the style, densely hairy. Disk
laterally compressed; testa smooth; hilum adaxial minute, free, annular, surrounding the base of the
(basi-ventral), occupying about half the length of ovary. Ovary 5-locular, placentation axile; style
the seed or full-length, broad, sometimes covering exserted. Berry 1-several-seeded. Seed broadly
about one third of the seed surface. Embryo with ovoid to globose, not laterally compressed; testa
plano-convex cotyledons, radicle extending to the smooth; hilum covering about 3/4 to 7/8 of the
surface; endosperm 0. Twenty three species con- seed surface. Embryo with plano-convex cotyle-
fined to Madagascar. dons and included radicle; endosperm 0. A single
species in Madagascar (T. macrantha Capuron).
V. TRIBE 0MPHALOCARPEAE Dubard ex 51. Magodendron Vink

Aubn!ville (1964).
Magodendron Vink, Nova Guinea II 8: 124 (1957); Baehni,
Boissiera 11: 116 1965.
Calyx a single whorl of 5 valvate or quincuncial
sepals; corolla lobes same number as sepals,
simple; corolla cyathiform or tubular, tube shorter Tree. Leaves spirally arranged. Venation eucamp-
than or exceeding the lobes; stamens in a single todromous. Stipules 0. Inflorescence fasciculate,
whorl in groups of 2-6 opposite each corolla lobe; cauliflorous. Flowers bisexual. Sepals 5, free, quin-
staminodes +; hilum adaxial. cuncial. Corolla broadly tubular, expanded above,
tube exceeding the lobes; lobes 5, spreading,
KEY TO THE GENERA OF 0MPHALOCARPEAE simple, quincuncial. Stamens 10 with 2 opposite
each corolla lobe and fixed at the top of the
1. Stipules +; anthers c. length of corolla lobes 50. Tsebona corolla tube, long-exserted; filaments free, well-
- Stipules 0; staminodes much shorter than corolla lobes 2 developed; anthers locellate, glabrous. Staminodes
2. Anthers locellate; disk patelliform; seed not laterally com-
pressed, without endosperm 51. Magodendron 5, alternating with the corolla lobes and fixed at the
- Anthers not locellate; disk 0; seed strongly laterally com- level of the stamens, strongly inflexed, margin
pressed, endospermous 3 fringed, glabrous. Disk patelliform, free, sur-
3. Corolla lobes slightly longer than tube; stamens fixed near rounding the base of the ovary, sparsely hairy.
top of corolla tube; filaments mostly free, glabrous; anthers
Ovary 5-locular, placentation axile; style included.
glabrous 52. Omphalocarpum
- Corolla lobes free almost to base, stamens fixed in lower Berry 1-seeded. Seed broadly ellipsoid, testa
half of corolla tube, almost completely fused in pairs or 3s; smooth, shining; hilum adaxial, full-length, cover-
filaments hairy, anthers hairy or glabrous ing about half the seed surface. Embryo with
53. Tridesmostemon plano-convex cotyledons; radicle slightly exserted;
endosperm 0. A single species in Papua New
Guinea.
Sapotaceae 419
52. Omphalocarpum Palisot de Beauvois sub globose, several seeded. Seed strongly laterally
compressed, testa smooth, shining; hilum adaxial,
Omphalocarpum Palisot de Beauvais ex Ventenat, Bull. Sci. Soc.
Philom. Paris 2: 146 (1800); Aubreville, Fl. Forest. Cote almost full-length, very narrow. Embryo with thin
d'Ivoire ed. 2, 3: 109 (1959), Fl. Gabon 1, Sapotacees: 75 foliaceous cotyledons and exserted radicle;
(1961), Fl. Cameroun 2, Sapotacees: 57 (1964). endosperm +. Two or three species in Central
Ituridendron De Wildeman (1926). Africa (Cameroon, Gabon, Zaire).
Trees. Leaves spirally arranged. Venation eucamp-
todromous. Stipules 0. Cauliflorous or ramiflor- Doubtful Genus
ous. Flowers in fascicles, bisexual or unisexual
(dioecious), often subtended by spirally arranged Boerlagella Cogniaux
bracts. Sepals 5, free, quincuncial. Corolla broadly
tubular or cyathiform, tube shorter than the lobes; Boerlagella Cogniaux in A. de Candolle & C. de Candolle,
Monogr. Phan. 7: 1173. (1891); H.J. Lam, Bull. Jard. Bot.
lobes 5-11, erect or slightly spreading, simple, Buitenzorg III, 7: 250 (1925).
imbricate or quincuncial. Stamens in groups of Boerlagia Pierre (1890).
3-6 opposite each corolla lobe and inserted at the
top of the corolla tube, exserted or not; filaments Known only from incomplete material of leaf, fruit
well-developed, exceeding the anthers, free or a and seed. It was placed in Planchonella by Dubard
few partially united near the base (rarely irregu- (1912: 61) on account of the narrow hilum and
larly united to halfway or above), glabrous; anthers exendospermous seed, and subsequent authors,
extrorse, glabrous, 0 in female flowers. Staminodes except Lam, have regarded it as doubtfully Sapota-
5-11, much shorter than the corolla lobes, usually ceous (Aubreville 1964a; Baehni 1938, 1965).
petaloid, fimbriate, glabrous, often inflexed and Lam (1925: 250) placed it (with Dubardella) in
covering the ovary. Disk 0. Ovary 5-30-locular the Boerlagellaceae, for reasons which are not
(number of loculi reduced in male flowers), pla- clear, although possibly on account of the cotyle-
centation axile; style usually included. Fruit dons which are described as involute or inrolled.
usually depressed globose, the outer pericarp As no new material has been forthcoming, it is
often woody, several to many-seeded. Seed still not possible to place this plant more accu-
strongly laterally compressed, testa smooth, rately. The leaf and seed structure appear to be
shining; hilum nearly full-length, very narrow. perfectly consistent with the Sapotaceae, as are the
Embryo with thin foliaceous cotyledons and malpighiaceous hairs at the base of the fruit. The
exserted radicle; endosperm copious. About six cotyledon condition, described by Lam, is anom-
species in West and Central Africa. alous and not recorded elsewhere in the family, but
I have not been able to confirm it, as the only avail-
53. Tridesmostemon Engler able seed seen by me is immature.
In the absence of any further material, I agree
Tridesmostemon Engler, Bot. Jahrb. Syst. 38: 99 (1905); De with Dubard and place it provisionally in Pouteria
Wildeman, Pl. Bequaert. 4: 144 (1926); Pellegrin, Bull. Soc.
Bot. France 85: 179 (1938); Aubreville, Fl. Gabon 1.
sect. Oligotheca.
Sapotacees: 82 (1961); Fl. Cameroun 2. Sapotacees: 66
(1964).
Trees. Stipules 0 (or caducous?). Leaves spirally
arranged. Venation eucamptodromous. Flowers Anderberg, A.A. et a!. 2002. See general references.
bisexual or rarely unisexual?, solitary or in few- Arends, J.C. 1976. Somatic chromosome numbers of some
African Sapotaceae. Acta Bot. Neerl. 25: 449-457.
flowered fascicles, axillary or in the axils of fallen Assem, J. van den. 1953. Revision of the Sapotaceae of the
leaves. Sepals 5, free, imbricate. Corolla cyathi- Malaysian area in a wider sense. 4. Ganua Pierre ex Dubard.
form, corolla lobes 5, simple, imbricate, spreading Blumea 7: 364-400.
or reflexed at the tip, free almost to the base. Aubreville, A. 1961a. Notes sur les Sapotacees de !'Afrique
Stamens 10-15, 2-3 opposite each corolla lobe, equatoriale. Notul. Syst. (Paris) 16: 223-279.
Aubreville, A. 1961b. Notes sur les Pouteriees africaines et sud
fixed to way up the corolla lobes, with prominent americaines. Adansonia II, 1: 6-38.
traces to the base; filaments of each pair or group Aubreville, A. 1962a. Notes sur des Pouteriees americaines.
of 3 fused almost to the apex, hairy; anthers Adansonia II, 1: 150-191.
extrorse, hairy or glabrous. Staminodes 5, usually Aubreville, A. 1962b. Notes sur les Sapotacees de Ia Nouvelle
3-dentate at apex, hairy. Disk 0. Ovary 10-locular, Caledonie. Adansonia II, 2: 172-199.
Aubreville, A. 1963a. Notes sur les Sapotacees. Adansonia II, 3:
placentation axile; style slightly exserted. Fruit 19-42.
420 T.D. Pennington
Aubreville, A. 1963b. Flore du Cambodge, du Laos et du Gilly, C.L. 1943. Studies in the Sapotaceae 2. The Sapodilla-
Vietnam, Sapotacees, 1-105. Nispero complex. Trop. Woods 73: 1-22.
Aubreville, A. 1964a. Sapotacees. Adansonia, Memoire no. 1: Govaerts, R., Frodin, D. G., Pennington, T.D. 2001. World check-
1-157. list and bibliography of Sapotaceae. Kew: Royal Botanic
Aubreville,A.1964b. Notes sur des Sapotacees 3.1. Rehabilita- Gardens, Kew.
tion des genres americains Raga/a Pierre et Prieurella Hall, J.B. 1996. Vitellaria paradoxa: a monograph. Bangor:
Pierre. Adansonia II, 4: 367-391. University of Wales, 125 p.
Aubreville, A. 1964c. Systeme de classification des Sapotacees. Halle, F., Oldeman, R.A.A. 1970. Essai sur !'architecture et la
Adansonia II, 4: 38-42. dynamique de croissance des arbres tropicaux. Paris:
Aubreville, A. 1965. Notes sur des Sapotacees australiennes. Masson.
Adansonia II, 5: 21-26. Harley, M.M. 1986a. Distinguishing pollen characters for the
Aubreville, A. 1967. Flore de Ia Nouvelle Caledonie et Depen- Sapotaceae. Can. J. Bot. 64: 3091-3100.
dences. 1. Sapotaceae 1-168. Harley, M.M. 1986b. The nature of the endoaperture in the
Aubreville, A. 1971. Essais de geophyletique des Sapotacees 2. pollen of the Sapotaceae. In: Blackmore, S., Ferguson, I.K.
Adansonia II, 11: 425-436. (eds.) Pollen and spores: form and function. London:
Aubreville, A., Pellegrin, F. 1934. De quelques Sapotacees Academic Press. Linn. Soc. Symp. Ser. 12:417-419.
de Ia Cote d'Ivoire. Bull. Soc. Bot. France 81: 792- Harley, M.M. 1990a. Pollen morphology of neotropical
800. Sapotaceae. In: Pennington, T.D. 1990: A monograph of
Baehni, C. 1938. Memoires sur les Sapotacees. 1. Systeme de neotropical Sapotaceae. Flora Neotrop. 52: 11-29, 710-
classification. Candollea 7: 394-507. 741.
Baehni, C. 1942. Memoires sur les Sapotacees. 2. Le genre Harley, M.M.1990b. The pollen morphology of the Sapotaceae.
Pouteria. Candollea 9: 147-475. Kew Bull. 46: 379-491.
Baehni, C. 1964. Genres nouveaux de Sapotacees. Arch. Sci. 17: Harley, M.M. 1991. Pollen morphology of the Sapotaceae. In:
77-79. Pennington, T.D. 1990: The genera of Sapotaceae. Kew: Royal
Baehni, C. 1965. Memoire sur les Sapotacees. 3. Inventaire des Botanic Gardens Kew, New York: New York Botanical
genres. Boissiera 11: 1-261. Garden, pp. 23-50.
Bokdam, J. 1977. Seedling morphology of some African Harley, M.M., Kurmann, M.H., Ferguson, I.K. 1991. Systematic
Sapotaceae and its taxonomical significance. Meded. Land- implications of comparative morphology in selected fossil
bouwhogesch. Wageningen 77/20: 1-84. and extant pollen from the Palmae and the Sapotaceae.
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Ocean islands, with special reference to Pteropus. Mammalia patterns of diversification. Oxford: Clarendon Press, pp.
45: 205-238. 225-238.
Cleghorn, M.L. 1922. Observations on the bat flowers of the Hartog, M.M. 1878. On the floral structure and affinities of
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II, 18: 571-576. Hartog, M.M. 1879. Notes on Sapotaceae II. J. Bot. 17: 356-359.
Corner, E.J.H. 1976. See general references. Hegnauer, R. 1973, 1990. See general references.
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Bumelia. J. Arnold Arbor. 26: 435-471. ed., vol. II: 16-30.
Cronquist, A. 1945b. Studies in the Sapotaceae 4. The North Heine, H., Hemsley, J.H. 1960. Notes on African Sapotaceae II.
American species of Manilkara. Bull. Torrey Bot. Club 72: The genus Bequaertiodendron De Wild. Kew Bull. 14:
550-562. 304-309.
Cronquist, A. 1946. Studies in the Sapotaceae 2. Survey of the Hemsley, J.H. 1966. Notes on African Sapotaceae. Kew Bull. 20:
North American Genera. Lloydia 9: 241-292. 461-510.
Cronquist, A. 1946a. Studies in the Sapotaceae 5. The South Hemsley, J.H.1968. Sapotaceae. In: Flora of tropical East Africa.
American species of Chrysophyllum. Bull. Torrey Bot. Club London: Crown Agents.
73: 286-311. Hermann-Erlee, M.P.M., Royen, P. van 1957. Revision of the
Cronquist, A. 1946b. Studies in Sapotaceae 6. Misc. Notes Bull. Sapotaceae of the Malaysian area in a wider sense. 9.
Torrey Bot. Club. 73: 465-471. Pouteria Aublet. Blumea 8: 452-509.
Dubard, M. 1908. Les Sapotacees du groupe des Illipees. Rev. Hickey, L.J. 1973. Classification of the architecture of dicotyle-
Gen. Bot. 20: 193-206. donous leaves. Am. J. Bot. 60: 17-33.
Dubard, M. 1909. Les Sapotacees de groupe des Isonandrees. Johnson, M.A.T. 1991. Cytology. In: Pennington, T.D. The
Rev. Gen. Bot. 21: 392-398. genera of Sapotaceae, chap. 2. Kew: Royal Botanic Gardens
Dubard, M. 1912. Les Sapotacees de groupe des Sideroxylinees. Kew, New York: New York Botanical Garden, pp. 15-
Ann. Inst. Bot.-Geol. Colon. Marseille II, 10: 1-98. 22.
Dubard, M.1915. Les Sapotacees du groupe des Sideroxylinees- Johri, B.D. et al. 1992. See general references.
Mimusopees. Ann. Inst. Bot. Geol. Colon. Marseille III, 3: Kukachka, B.F. 1978. Wood anatomy of the neotropical
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Engler, A. 1904. Monographien afrikanischer Pflanzen- U.S.D.A. Madison. Res. Pap. 327: 1-7.
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Engelmann. Sapotaceae. 8. Diploon. Forest Products Research Labora-
Eyma, P.J. 1936. Notes on Guiana Sapotaceae. Recueil Trav. Bot. tory, U.S.D.A. Madison. Res. Pap. 349: 1-4.
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Botanic Gardens Kew, New York: New York Botanical
Garden.
422 K. Kubitzki
Sarraceniaceae
K. KUBITZKI
Sarraceniaceae Dumort., Anal. Fam. Pl.: 53 (1829), nom. cons. A distinctive family of three genera and ca. 15
species from North and northern South America,
Perennial, rhizomatous, mostly acaulescent, car- all occurring in nutrient-poor habitats.
nivorous herbs. Leaves alternate, shortly petio-
late, exstipulate, borne in a rosette or (some VEGETATIVE STRUCTURES. Sarraceniaceae are
Heliamphora) on an upright, sometimes branching predominantly rhizomatous herbs with a rosette
stem, all or some highly modified and transformed of pitcher-like leaves. Only some species of
into more or less elongate, ascidiate, often pitcher- Heliamphora, such as H. nutans and H. minor, have
like traps ("amphores") partly filled with digestive erect stems and can attain a dendroid habit.
liquid; petiole short and passing into the ascidiate The comparative morphology of the highly
portion of the leaf which, on the ventral side, bears modified ascidiate leaves has been dealt with by
a double or simple ridge or laminar wing and, on numerous authors, among them Troll (1939),
the apex of the dorsal side, a flattened, small or Arber (1941) and Markgraf (1955). The pitchers
large, often hood-like appendix; the opening of the are best interpreted in terms of Troll's (1932)
trap provided with a more or less distinctive collar; peltation theory, which implies unifaciality and
the outer and inner side of the pitcher around peltation including a "Querzone", although some
the opening provided with specialized glands and difficulties are inherent in this explanation (see
retrorse hairs that serve to entrap insects and other Juniper et al. 1989: 57 seq.).
small animals; scale-like or sword-like leaves The pitcher rosettes of Heliamphora and Sar-
sometimes produced late in the season. Flowers racenia form ± symmetrical circles, facing inwards
perfect, actinomorphic, nodding, solitary on a with their openings, whereas the openings of Dar-
scape or (Heliamphora) in few-flowered, some- lingtonia face outwards as a result of a tubular
times axillary racemoids; prophylls present twist of about 180°.
(Heliamphora); perianth (3)4-6-merous, uniseri- As in other carnivorous families, the pitcher
ate, petaloid (Heliamphora), or biseriate of persist- traps of Sarraceniaceae are covered with numer-
ent sepals and showy, caducous petals; all perianth ous nectarial glands that attract insects; these
members distinct, imbricate; stamens 10-20 or glands are densely packed around the entrance of
numerous, in Sarracenia usually arising from 10 the pitchers. Downwards-directed trichomes of
primordia; filaments short; anthers basifixed or varying length are another important part of the
(Sarracenia) versatile, introrse, dehiscing with trap mechanism, and Darlingtonia shares with
longitudinal slits or (Heliamphora) short slits the unrelated Nepenthes and Brocchinia the use of
on caudal anther appendages; gynoecium of 5 or detachable wax platelets to prevent arthropod
(Heliamphora) 3 united carpels; ovary superior, escape.
5(3)-locular, but the partition of the upper part Heliamphora has the simplest pitcher structure
often incomplete and placentation therefore axile of all Sarraceniaceae (Arber 1941), possessing a
below and parietal above; style slightly 3-lobed and poorly developed lid and lacking a rolled collar on
more or less truncate at the apex (Heliamphora), the rim, whereas the ventral keel (interpreted as
with 5 short style branches (Darlingtonia), or the leaf margins) is a double structure (Fig. 124A).
apically expanded and peltate, umbrella-shaped, In Darlingtonia, the pitcher consists of a broadly
with small stigmas under the tip of each of the 5 winged, bifacial sheathing base, a tubular region
lobes (Sarracenia); ovules numerous, anatropous, provided with a simple keel, and a strongly curved
unitegmic or (Darlingtonia?) bitegmic, tenuinucel- hood, which closes the top of the pitcher. The
late. Fruit a loculicidal capsule with numerous tissue of the hood becomes transparent and forms
seeds; seeds small, often winged, with exotestal a "window". Access for insects is possible only
seed coat, copious endosperm rich in oil and from underneath and leads over a peristome rim.
protein, and small linear embryo. Additionally, a fishtail-shaped appendage covered
Sarraceniaceae 423
with nectaries hangs down from the entrance and by bumble-bees (Renner 1989 and literature cited
is obviously homologous with the lid or flap of the therein). Darlingtonia does not seem to produce
other genera. Sarracenia also has a single keel, a nectar, and pollinators are unknown. Heliamphora
peristome rim and a lid that usually does not flowers are scentless and produce no nectar; in the
occlude the mouth of the pitcher. pendent flowers the short slits of the caudal
One could speculate that the flaps or hoods of appendages of the reflexed anthers face downward
the pitchers have the function of preventing rain- and make the anthers technically poricidal;
water from entering into the pitcher and diluting H. tatei was observed to be buzz-pollinated by
the digestive fluid. However, the Heliamphora bumble-bees, Xylocopa and Eulaema (Renner
species, which are most ineffective in this respect, 1989). Since buzz-pollinated flowers are frequent
grow in regions where heavy rains fall all over in the montane scrub communities of the Roraima
the year. Various insects dwell in the pitchers of formation, including several genera of Melastom-
Sarracenia (Uphof 1936), and the larvae of the ataceae and Rapateaceae, Heliamphora obviously
tropical mosquito Wyeomyia even overwinter in shares the common pollinator-pool of these
them (Wood 1960). communities.
Since the axial cambium in Sarraceniaceae is
active only for a short period of time, a limited SEED. The seed coat is formed by the outer epi-
amount of secondary xylem is present in the dermis of the integument, which has thickened
rhizomes. It has a number of primitive features radial and inner walls; the rest of the tissue is
including solitary vessels with scalarifom perfora- crushed. The endosperm is cellular and oily, the
tion plates and long and oblique end walls, tra- embryo is usually minute (Netolitzky 1926).
cheids with scalariform pitting, and diffuse axial
parenchyma. Vessel elements in Heliamphora have PHYTOCHEMISTRY. Chemosystematically signifi-
the greatest number of allegedly primitive traits cant is the occurrence of secoiridoids of route I of
(DeBuhr 1977). Jensen (1991). Sarracenin, an enol diacetal moo-
terpene, seems to be a compound unique to
EMBRYOLOGY. Pollen grains are 2-celled when Sarraceniaceae and has been recorded from five
shed. Ovules are anatropus, unitegmic (bitegmic species of Sarracenia and two of Heliamphora. For
in Darlingtonia?), and tenuinucellar. The arches- further compounds see Hegnauer (1990).
pore functions directly as the megaspore mother
cell; the chalaza! megaspore of the tetrad develops AFFINITIES. On the basis of a broad comparison
into a Polygonum-type embryo sac. An endothe- of the floral morphology and embryology, DeBuhr
lium encloses the embryo sac. Endosperm forma- (1975) refuted the former hypothesis of relation-
tion is cellular (Davis 1966). ship between Sarraceniaceae and other insec-
tivorous families such as Droseraceae and
KARYOLOGY AND HYBRIDISATION. Chromosome Nepenthaceae. Instead, he noted a significant
numbers are n = 13 (Sarracenia), 15 (Darlingto- agreement in these characters with Actinidiaceae,
nia), and 21 (Heliamphora); polyploidy has not Marcgraviaceae and Theaceae. Analysis of nuclear
been observed. All Sarracenia species are inter- and plastid DNA sequences (Bayer et al. 1996)
fertil and many natural hybrids are known from revealed Sarraceniaceae as a monophylum sister
the wild, yet the species remain morphologically to Roridula; Darlingtonia appears sister to
distinct (Bell1952). Sarracenia + Heliamphora. Other analyses
(Albach et al. 2001) support the association of Sar-
POLLEN MORPHOLOGY. Pollen is oblate to sub- raceniaceae and Roridulaceae, with Actinidiaceae
spherical and 3-9-colporate, in Heliamphora (3)4- the next closest family, and Cyrillaceae and
5-colporate and verruculate, in Darlingtonia Clethraceae more distantly related. These families
4-6-colporate and minutely scrobiculate, and in share many morphological characters.
Sarracenia 6-9-colporate and minutely scrobicu- Up to the present, on account of its relatively
late (Thanikaimoni and Vasanthy 1972). unspecialised pitchers and the racemoid in-
florescence, Heliamphora had usually been con-
POLLINATION. In Sarracenia, nectar is produced sidered the most basal genus of Sarraceniaceae.
at the base of the style; pollen is shed from the ver- A comparison with the sister family Roridu-
satile anthers and falls onto the stylar disk of the laceae shows that the racemoids but not the
pendent flowers; in S. rubriflora it sticks together uniseriate perianth of Heliamphora may be
in little pellets (Burr 1979); pollination takes place plesiomorphic.
424 K. Kubitzki
DISTRIBUTION AND HABITATS. The six species

of Heliamphora are restricted to the Roraima
sandstone formation of the Guayana Highlands in
northern South America, where they grow in open,
marshy or wet savannahs and altitudinal heath
formations. The monotypic Darlingtonia grows in
bogs and wet meadows in the western United
States (California and Oregon). Eight of the nine
species of Sarracenia grow in flatwoods, open bogs
and savannahs of the south-eastern United States;
S. purpurea extends north to Canada and west to
British Columbia.
ECONOMIC IMPORTANCE AND CONSERVATION.

Because of their beauty and amazing insectivorous
lifestyle, all members of the family have attracted
the interest of gardeners, traders and the public
(see Simpson 1994). Whereas the habitats of
Heliamphora are of difficult access, Sarracenia and
Darlingtonia are considered under such threat in
the wild that all taxa of these genera are included
in CITES. Apart from efforts of in-situ conserva-
tion of the nutrient-deficient wetlands, the cre-
ation of collections in Botanic Gardens and public
collections is an additional safeguard against loss
in the wild.
KEY TO THE GENERA

1. Flowers in a racemoid; perianth simple; ovary 3(4)-locular
1. Heliamphora
- Flowers solitary on a scape; perianth of calyx and corolla;
ovary 5-locular 2
2. Scape aphyllous; stamens numerous; style expanded into
broad umbraculiform disk 3. Sarracenia Fig. 124. Sarraceniaceae. Heliamphora tatei var. neblinae. A
- Scape bracteose; stamens 15; style with 5 horizontally Two amphores. BTrichomes above the "waist". C Distal part
patent style branches 2. Darlingtonia of racemoid. D Flower, one tepa! removed. E Flower with
perianth removed, anthers still erect. F Same, later stage with
anthers inverted. G Young stamens. H Inverted stamen. I
1. Heliamphora Benth. Fig. 124 Inverted stamens and pistil, with stigma enlarged. J Pistil. K
Same, transverse section. L Placenta with ovules. (Maguire
Heliamphora Benth., Trans. Linn. Soc. 18: 432 (1840); Steyer-
1978)
mark, Ann. Missouri Bot. Gard. 71: 302-312 (1984), rev.
Pitcher with a double keel. Tepals petaloid; Pitchers with simple keel, twisted for about 180°.
stamens 10-20; anthers basifixed, at anthesis Anthers basifixed, erect, dehiscing longitudinally;
reflexed, dehiscing through caudal appendages; ovary 5-locular; style with 5 stylodia. Seeds oval,
ovary 3(4}-locular; style short, stigma truncately basally tailed; embryo relatively large, embedded
3(4}-lobed; seed compressed, irregularly scario- in endosperm. 2n = 30. One sp., D. californica
fimbriately winged; embryo small, embedded in Torrey, N California, Oregon.
copious endosperm. 2n = 42. Five spp., partly
polymorphic, at middle and upper altitudes of the
Guayana Highland in Venezuela and northern 3. Sarracenia L.
Brazil. Sarracenia L., Sp. Pl.1: 510 (1753); McDaniel, Bull. Timbers Res.
Stat. 9: 1-36 (1971), rev.
2. Darlingtonia Torrey Pitchers with simple keel. Sepals 5, with 3

Darlingtonia Torrey, Smithson. Contrib. 6: 4, t. 12 (1854). appressed bracts; petals 5; stamens 70-80, arising
Sarraceniaceae 425
in 10 groups; anthers dorsifixed, not reflexed; style DeBuhr, L.E. 1977. Wood anatomy of the Sarraceniaceae: eco-
expanded into broad disk, with a small stigma logical and evolutionary implications. Plant Syst. Evol. 128:
159-169.
under each of the notched lobes. Seeds clavate to Hegnauer, R. 1990. Chemotaxonomie der Pflanzen, vol. 9.
obovate, unilaterally inconspicuously winged or Basel: Birkhaeuser.
ridged; embryo small, embedded in copious Jensen, S.R., Nielsen, B.J., Dahlgren, R. 1975. Iridoid com-
endosperm. 2n = 26. Two sections: sect. Erectae pounds, their occurrence and systematic importance in the
Uphof with upright, tubular or trumpet-shaped angiosperms. Bot. Notiser 128: 148-180.
Maguire, B. 1978. Sarraceniaceae. In: The Botany of the
leaves, seven or eight spp. in the SE United States; Guayana Highland - Part X. Mem. New York Bot. Gard. 29:
and sect. Sarracenia, with highly modified, 36-62.
decumbent pitchers and maroon petals, two spp., Markgraf, F. 1955. Ober Laubblatt-Homologien and ver-
North America from Labrador to Florida. wandtschaftliche Zusammenhange bei Sarraceniales. Planta
46: 414-446.
Newman, T., Ibrahim, S., Wheeler, J.W., McLaughlin, W.B.,
Petersen, R.L., Duffield, R.M. 2000. Identification of sar-
Selected Bibliography racenin in four species of Sarracenia (Sarraceniaceae).
Biochem. Syst. Ecol. 28: 193-195.
Albach, D., Soltis, P.S., Soltis, D.E., Olmstead, R.G. 2001. Phylo- Renner, S.S. 1989. Floral biological observations on
genetic analysis of asterids based on sequences of four Heliamphora tatei (Sarraceniaceae) and other plants from
genes. Ann. Missouri Bot. Gard. 88: 163-212. Cerro de la Neblina in Venezuela. Plant Syst. Evol. 163:
Arber, A. 1941. On the morphology of the pitcher-leaves in 21-29.
Heliamphora, Sarracenia, Darlingtonia, Cephalotus, and Shreve, F. 1906. The development and anatomy of Sarracenia
Nepenthes. Ann. Bot. II, 5: 563-578. purpurea. Bot. Gaz. 42: 107-126.
Bayer, R.J., Hufford, L., Soltis, D.E. 1996. Phylogenetic relation- Simpson, R.B. 1994. Pitchers in trade: a conservation review of
ships in Sarraceniaceae based on rbcL and ITS sequences. the carnivorous genera of Sarracenia, Darlingtonia, and
Syst. Bot. 21: 121-134. Heliamphora. Kew: Royal Botanic Gardens Kew.
Bell, C.R. 1949. A cytotaxonomic study of the Sarraceniaceae Thanikaimoni, G., Vasanthy, G. 1972. Sarraceniaceae: Palynol-
of North America. J. Elisha Mitchell Sci. Soc. 65: 137-166. ogy and systematics. Pollen Spores 14: 143-155.
Bell, C.R. 1952. Natural hybrids in the genus Sarracenia. I. Uphof, J.C.Th. 1936. Sarraceniaceae. In: Engler, A., Harms, H.
History, distribution, and taxonomy. J. Elisha Mitchell Sci. (eds.) Die natiirlichen Pflanzenfamilien, 2nd edn., 17b.
Soc. 68: 55-80. Leipzig: W. Engelmann, pp. 704-727.
Davis, G.L. 1966. Systematic embryology of the angiosperms. Wood, C.R. 1960. The genera of Sarraceniaceae and Droser-
New York: Wiley. aceae in the southeastern United States. J. Arnold Arbor. 41:
DeBuhr, L.E. 1975. Phylogenetic relationships of the Sarrace- 152-163.
niaceae. Taxon 24: 297-306.
426 0. Appel
Scytopetalaceae
O.APPEL
Scytopetalaceae Engl. in Engler & Prantl, Nat. Pflanzenfam. The leaves of the Scytopetalaceae are simple,
Nachtr. I: 242-245 (1897); nom. cons. estipulate or rarely stipulate (Asteranthos) and on
side branches usually arranged distichously. The
Evergreen trees or shrubs; young stems with cor- venation is generally brochidodromous. Usually,
tical bundles; indumentum, if present, of uniseri- the leaf-margin is entire and often undulate, but
ate hairs. Leaves alternate, simple, entire or serrate, dentate in Brazzeia soyauxii and Pierrina zenkeri.
papery to coriaceous, minutely stipulate. Inflores- Some species are characterized by slanting leaf
cences axillary, often ramiflorous or cauliflorous, blades. Stomata are anisocytic. They are present
panicles, thyrsoids, botryoids or racemes, rarely on both leaf surfaces in Oubanguia and Scyto-
reduced to single flowers (Asteranthos). Flowers petalum, but restricted to the lower surface in
regular, hypogynous or rarely perigynous; sepals the remaining genera. In species with thin leaves,
united; calyx cupular, thick, leathery, persistent on the palisade layer is lacking. The leaves of all Scy-
ripe fruit, occasionally accrescent; petals lacking; topetalaceae are distinguished by the occurrence
staminodes 6-28, completely fused, forming of spicular cells. In Oubanguia and Scytopetalum
a showy corolla-like structure; pseudocorolla calcium oxalate crystals were observed in tissues
glabrous or rarely pubescent, usually very thick sheathing the veins. Van Tieghem (1905), Metcalfe
and leathery (but thin in Asteranthos); stamens and Chalk (1950), and Amaral (1991) described
numerous; filament bases united, adnate to these crystalliferous cells as "cristarque cells".
pseudocorolla; anthers basifixed, tetrasporangiate; Metcalfe and Chalk (1950), Carlquist (1988), and
anther dehiscence lengthwise, complete or apical de Zeeuw (1990) studied the wood anatomy of
and pore-like; ovary syncarpous, 3-8-locular; pla- several members of the family. Important features
centation axile; style solitary; stigma undifferenti- are scalariform perforation plates, alternate
ated or lobed; ovules bitegmic, anatropous, medium-sized intervascular pits, multi- and
pendulous, 2, 4, or many in each locule. Fruits 1(2)- uniseriate rays, and apotracheal parenchyma in
or many-seeded, drupes, loculicidally dehiscent numerous uniseriate bands. The phloem is strati-
capsules, or leathery and berry-like. Seeds large, fied in lignified and unlignified portions. Cork
ca. 1 em long; testa poorly differentiated; develops surficially.
endosperm horny, often markedly ruminate;
embryo well differentiated; cotyledons large, flat, INFLORESCENCE MORPHOLOGY. The inflores-
cordate, rarely reduced (Asteranthos). cences of the Scytopetalaceae are always axillary
Six genera and about 21 species in humid trop- in position. The basic inflorescence type is a
ical West Africa and South America. panicle. All other types found in the family can be
interpreted as derived from panicles through
VEGETATIVE STRUCTURES. The shoot system of reductions and structural changes (Appel 1996).
Scytopetalaceae proliferates by sympodial branch- Thyrsoids are most common within the family
ing. At the base of each module usually the scars (Fig. 125A) but botryoids, racemes and single
of several scale leaves can be recognized, which flowers also occur. In Scytopetaloideae the inflo-
in Rhaptopetaloideae may subtend inflorescences. rescences appear in the leaf axils of the youngest
Young shoots are often angular to broadly winged. branches, whereas in Rhaptopetaloideae they have
The position of the wings is correlated with the shifted to a ramiflorous or cauliflorous position.
path of a peculiar system of cortical bundles,
which are borne at one node and usually enter the FLOWER STRUCTURE. In all members of Scyto-
leaf of the node after next (van Tieghem 1905). The petalaceae the sepals are thick, leathery, and
primary cortex, phloem and medullary rays are completely fused. Early in ontogeny the cupular
remarkable for the abundance of octaedric calyx opens rhexigenously, thus often having
calcium oxalate crystals. irregular incisions or dentations that do not cor-
Scytopetalaceae 427
respond to the number of sepals. In Asteranthos FRUIT AND SEED. In all genera except Oubanguia
the calyx is strongly accrescent and serves as a the fruits appear to be indehiscent. The mesocarp
floating device for its hydrochorous fruits (Ducke is always woody, en do carp and exocarp are usually
1948). In Oubanguia the abaxial surface of the leathery but in Scytopetalum the exocarp is fleshy.
calyx is densely covered with uniseriate hairs. The seeds are glabrous (Scytopetaloideae) or
The flowers are here interpreted as apetalous, embedded into a shiny, white haircoat oflong uni-
but possessing a showy corona or corolla-like cellular hairs formed by epidermal cells along the
structure of presumably staminodial origin. The testal bundles (Rhaptopetaloideae). The testa is
number of staminodes varies widely from 6 poorly differentiated and lacks any sclerotic tissue.
(Oubanguia) to 16 (Scytopetalum) or even 24-28 The abundant endosperm has strongly thickened
(Asteranthos). They are always completely fused, cell walls that contain amyloid, a hemicellulose, as
forming a pseudocorolla that bears several struc- main storage compound. Starch and oil are of
tural differences: usually the pseudocorolla has a minor importance. In all genera except Oubanguia
thick leathery texture and opens rhexigenously the endosperm is ruminate (Fig. 125E, F). Both
at anthesis, whereas in Asteranthos it is thin subfamilies are characterized by distinct rumina-
and pleated in bud (Fig. 125B), unfolding like an tion forms, which apparently evolved independ-
umbrella at anthesis. In Oubanguia and Scyto- ently (Appel 1996). In Rhaptopetaloideae the
petalum the pseudocorolla splits into lobes that position of the endosperm foldings is correlated
presumably correspond to single staminodes, with the path of the testal bundles, whereas in Scy-
whereas in Rhaptopetalum, Pierrina, and Brazzeia topetaloideae there is no such correlation (Appel
the number of pseudocorolla lobes in opened 1996).
flowers does not reflect the number of component
staminodes. In Scytopetalum and Oubanguia RELATIONSHIPS WITHIN THE FAMILY. Within the
the pseudocorolla bears apical outgrowths that Scytopetalaceae two clearly delimitated subfami-
project into the bud, filling its apex like a plug. In lies can be distinguished (for distinguishing char-
Asteranthos the margin of the pseudocorolla is acters see key). The neotropical Asteranthos is still
ciliate and curved inward in bud. The stamens are extremely similar to Scytopetalum and Ouban-
numerous and arise in centrifugal succession from guia, although it has apparently been separated
a ring-shaped meristematic wall. The gynoecium from its African stock long ago.
is syncarpous with axile placentation. Usually, the
ovaries are superior, but semi-inferior in some AFFINITIES. Traditionally the Scytopetalaceae
species of Rhaptopetalum and in Asteranthos. The were placed in the Malvales. This position was sup-
stigma is entire and somewhat papillose in Rhap- ported mainly by the stratified phloem common
topetalum, but lobed in the other genera. to both groups, and by the embryological charac-
ters provided byVijayaraghavan and Dhar (1976).
EMBRYOLOGY. Vijayaraghavan and Dhar (1976) In more recent publications, generally a position
and Tsou (1994) studied the embryology of near Ochnaceae and Quiinaceae in a broadly
Scytopetalum tieghemii and Asteranthos brasilien- defined order Theales or in Ochnales was sug-
sis. Pollen grains are shed at the 2-celled stage. gested (see Amaral 1991). Carlquist (1988),
Ovules are anatropous, bitegmic and tenuinucel- however, revealed striking similarities in wood
late. The inner integument is longer than the characters between Scytopetalaceae and Lecythi-
outer and forms the mycropyle. The development daceae. Asteranthos, which is now included in the
of the embryo sac conforms to the Polygonum Scytopetalaceae (see Tsou 1994), formerly has
type. often been placed in Lecythidaceae or (together
with Napoleonaea) at least close to this family.
POLLEN MORPHOLOGY. The pollen of several Affinities between Scytopetalaceae, Lecythidaceae,
members of the family was studied by Erdtman Foetidiaceae and Napoleonaeaceae are supported
(1952), Muller (1972) and Tsou (1994). The grains not only by wood anatomical features, such as the
are monads of medium size (20-30 flm), suboblate occurrence of cortical bundles, stratified phloem,
to spheroidal, tricolpate (Scytopetaloideae) or tri- and crystalliferous strands in secondary xylem
colporoidate (Rhaptopetaloideae) and circum- and primary cortex (Lignier 1890; de Zeeuw 1990),
aperturate to angulaperturate. The sexine but also with regard to embryology and palynol-
sculpture is microreticulate with duplibaculate ogy (Erdtman 1952; Muller 1972; Vijayaraghavan
muri and granulate lumina. The colpus mem- and Dhar 1976; Tsou 1989, 1994). Moreover, a
branes are granulate. systematic position of Scytopetalaceae near
428 O.Appel
Lecythidaceae, Foetidiaceae and Napoleonaeaceae

is further substantiated by rbcL sequence data.
According to Morton et al. (1997), Lecythidaceae
s.l. would form a monophylum if Scytopetalaceae
were included. This inclusion was· formally sub-
stantiated by Morton et al. (1998). However, since
Scytopetalaceae form a readily separated and well-
established family, their inclusion into the large
and highly diversified Lecythidaceae is not fol-
lowed here.
KEY TO THE GENERA
1. Inflorescences (or single flowers) inserted in leaf axils of

youngest branches; number of staminodes recognizable in
bud, their apices curved inwards in bud; anther dehiscence
complete; ovules 2 or 4 in each locule; fruits 1(2)-seeded;
seeds glabrous. Scytopetaloideae 2
- Inflorescences in ramiflorous or cauliflorous position;
number of staminodes not recognizable in bud, their
apices not curved inwards; anther dehiscence apical, pore-
like; ovules usually many in each locule; fruits many-
seeded; seeds pubescent. Rhaptopetaloideae 4
2. Flowers solitary; calyx accrescent; pseudocorolla more
than 20-partite, thin, pleated in bud, unfolding like an
umbrella at anthesis; ovules usually 4 in each locule
1. Asteranthos
- Flowers in few- to many-flowered inflorescences; pseudo-
corolla up to 16-partite, thick, leathery, opening rhexige-
nously; ovules usually 2 in each locule 3
3. Inflorescences determinate, with pubescent axes; carpels
3-5; fruits capsular; endosperm not ruminate
2. Oubanguia
- Inflorescences indeterminate, with glabrous axes; carpels
(6)7-8; fruits drupaceous; endosperm ruminate Fig.125. Scytopetalaceae. A Oubangia africana, inflorescence;
3. Scytopetalum note basal articulations of pedicels. B, CAsteranthos brasilien-
4. Flower stalks with distinct abscission zones; filaments sis. B Single axillary flower. C Vertical section of flower. D
much shorter than anthers; stigma simple; seeds not flat- Rhaptopetalum sp., vertical section of flower. E, F Scytopetalum
tened, without a circular notch 4. Rhaptopetalum tieghemii, seed. E Surface view. F Vertical section. (Appel
- Flower stalks without distinct abscission zones; filaments 1996)
longer than anthers; stigma with as many lobes as carpels;
seeds flattened, with a circular notch 5
5. Older inflorescences forming dense clusters; fruits
globose, rounded at apex 5. Brazzeia Shrubs or small trees; leaf-margin entire; flowers
- Inflorescences never forming dense clusters; fruits solitary in leaf axils; calyx accrescent; staminodes
fusiform, acute at apex 6. Pierrina 24-28, glabrous; pseudocorolla thin, pleated in
bud, at anthesis unfolding like an umbrella; ovary
semi-inferior; carpels 5-8; ovules 4in each locule;
I. SUBFAM. SCYTOPETALOIDEAE 0. Appel (1996). endosperm ruminate. n = 21. One sp.,A. brasilien-
sis Desf., N tropical South America.
Inflorescences or single flowers on youngest
branches. Staminodes 6- 28, their number dis-
2. Oubanguia Baill. Fig. 125A
tinguishable in bud, apically curved inwards.
Stamens 100-240; filaments more than 3 times Oubanguia Bail!., Bull. Soc. Linn. Paris 2: 869 (1890).
longer than anthers; anther dehiscence complete,
by longitudinal slits. Fruits usually 1-seeded. Seeds Trees or shrubs; leaf-margin entire or serrate;
glabrous. Pollen grains tricolpate. inflorescences determinate, with pubescent axes;
calyx not accrescent; staminodes 6- 12, pubescent;
pseudocorolla thick leathery, rhexigenously
1. Asteranthos Desf. Fig. 125B, C
opening; ovary superior; carpels 3-5; ovules 2 in
Asteranthos Desf., Mem. Mus. Hist. Nat. 6: 9 (1820). each locule; fruits loculizidally dehiscent capsules.
Scytopetalaceae 429
Endosperm non-ruminate. About 3 spp., tropical Trees or shrubs; leaf-margin entire or serrate;
West Africa. inflorescences usually in dense clusters. Petioles
without abscission zones; filaments longer than
anthers; carpels (5)6(7); stigma lobed; fruits
3. Scytopetalum Pierre ex Engl. Fig. 125E, F
globose; testal bundle unbranched. About 3 spp.,
Scytopetalum Pierre ex Engl. in Engler & Prantl, Nat. Pflanzen- tropical West Africa.
fam. Nachtr. I: 244 (1897).
Trees or shrubs; leaf-margin entire; inflorescences Selected Bibliography

indeterminate, with glabrous axes; petioles with a
basal abscission zone; calyx not accrescent; sta- Amaral, M.C.E. 1991. Phylogenetische Systematik der
minodes 12-16, glabrous; pseudocorolla thick, Ochnaceae. Bot. Jahrb. Syst. 113: 105-196.
leathery, opening rhexigenously; ovary superior; Appel, 0. 1996. Morphology and systematics of the Scytopeta-
laceae. Bot. J. Linn. Soc. 121: 207-227.
carpels (6)7-8, ovules 2 in each locule; fruits
Carlquist, S. 1988. Wood anatomy of Scytopetalaceae. Aliso 12:
drupaceous; endosperm ruminate. About 3 spp., 63-76.
tropical West Africa. Dehay, C. 1955. Caracteres de Ia feuille chez les Scytopeta-
lacees. Bull. Soc. Bot. Nord France 8: 76-81.
Ducke, J.A. 1948. Arvores Amaz6nicas e sua propagacil.o. Bol.
Mus. Paraense Hist. Nat. 10: 81-92.
II. SUBFAM. RHAPTOPETALOIDEAE
Engler, A. 1897. Scytopetalaceae. In: Engler & Prantl, Nat.
0. Appel (1996). Pflanzenfam. Nachtr. 1. Leipzig: Engelmann, pp. 242-245.
Engler, A. 1909. Scytopetalaceae africanae II. Bot. Jahrb. Syst.
Inflorescences ramiflorous or cauliflorous; stamin- 43: 373-377.
odes completely united into a thick, leathery Erdtman, G. 1952. See general references.
Knuth, R. 1934. tiber die Gattung Asteranthos. Notizbl. Bot.
pseudocorolla, their number not distinguishable Gart. Berlin-Dahlem 11: 1034-1036.
in bud, apically not curved inwards; stamens Knuth, R. 1939. Asteranthaceae. In: Engler, Pflanzenreich IV,
60-100; anthers dehiscent by short, pore-like lon- 219b. Leipzig: W. Engelmann.
gitudinal slits; ovules many in each locule; fruits Kowal, R.R. 1989. Chromosome numbers of Asteranthos and
many-seeded; seeds pubescent along testal the putatively related Lecythidaceae. Brittonia 41: 131-
135.
bundles; pollen grains tricolporoidate. Kravcoca, T.I. 1991. Scytopetalaceae. In: Takhtajan, A. (ed.)
Anatomia seminum comparativa. Vol. 3. Caryophyllidae -
Dilleniidae. Leningrad: Nauka, pp. 224-227.
4. Rhaptopetalum Oliv. Fig. 125D Letouzey, R. 1961. Notes sur les Scytopetalacees (Revision des
Scytopetalacees de l'herbier de Paris). Adansonia II, 1:
Rhaptopetalum Oliv., J. Linn. Soc. 8: 159 (1865).
106-142.
Letouzey, R. 1977. Nouvelle especes de Rhaptopetalum O!iv.
Trees or shrubs; leaf-margin entire; inflorescences (Scytopetalacees) du Cameroun et du Gabon. Adansonia II,
not in dense clusters; petioles with distinct abscis- 17: 129-138.
sion zones; filaments much shorter than anthers; Letouzey, R. 1978a. Scytopetalacees. In: Aubreville, A., Leroy,
J.-F. (eds.) Flore du Cameroun, vol. 20, pp. 139-193.
carpels 3-5; stigma simple, papillose; fruits Letouzey, R. 1978b. Scytopetalacees. In: Aubreville, A., Leroy,
globose; testal bundle branched. About 10 spp., J.-F. (eds.) Flore du Gabon 24, pp. 139-193.
tropical West Africa. Lignier, 0. 1890. Recherches sur l'anatomie des organes vege-
tatifs des Lecythidees, des Napo!eonees et des Barring-
toniees. Bull. Sci. France Belgique 21: 291-420.
5. Pierrina Engl. Metcalfe, C.R., Chalk, L. 1950. See general references.
Morton, C.M., Mori, S.A., Prance, G. T., Karol, K.G., Chase, M.C.
Pierrina Engl., Bot. Jahrb. Syst. 43: 374 (1909). 1997. Phylogenetic relationships of Lecythidaceae: a cladis-
tic analysis using rbcL sequence and morphological data.
Shrubs; leaf-margin entire or serrate; inflores- Am. J. Bot. 84: 530-540.
cences not in dense clusters; petioles without Morton, C.M., Prance, G.T., Mori, S.A., Thorburn, L.G. 1998.
Recircumscription of Lecythidaceae. Taxon 47: 817-
abscission zones; filaments longer than anthers; 827.
carpels (3)4(5); stigma lobed; fruits fusiform, Muller, J. 1972. Pollen morphological evidence for subdivision
acute; testal bundle unbranched. One sp.,P. zenkeri and affinities of Lecythidaceae. Blumea 20: 350-355.
Engl., humid tropical West Africa. Tsou, C.-H. 1989. The floral morphology and embryology of
Asteranthos and its systematic consideration. Am. J. Bot. 76
suppl.: 275-276.
6. Brazzeia Baill. Tsou, C.-H. 1994. The embryology, reproductive morphology
and systematics of Lecythidaceae. Mem. New York Bot.
Brazzeia Baill., Bull. Soc. Linn. Paris 1: 609 (1886). Gard. 71: 1-112.
430 O.Appel
van Tieghem, P. 1905. Surles Rhaptopetalacees. Ann. Sci. Nat., Zeeuw, C.H. de 1990. Secondary xylem of Neotropical Lecythi-
Bot. IX, 1: 321-388. daceae. In: Mori, S.A., Prance, G.T., Lecythidaceae- Part II,
Vijayaraghavan, M.R., Dhar, U. 1976. Scytopetalum tieghemii- chap. II. The zygomorphic-fiowered New World genera. Fl.
embryologically unexplored taxon and affinities of the Neotrop. Monogr. 21 (II), pp. 4-59.
family Scytopetalaceae. Phytomorphology 26: 16-22.
Sladeniaceae 431
Sladeniaceae
P.F. STEVENS and A.L. WEITZMAN
Sladeniaceae (Gilg & Werdermann) Airy Shaw, Kew Bull. 18: nodes are unilacunar and the phellogen is deep-
267 (1964). seated.
Evergreen trees; hairs unicellular. Leaves spiral or INFLORESCENCE AND FLOWER STRUCTURE. The
distichous, margins toothed or entire, secondary inflorescences are dichasia, simple in the case of
veins pinnate, stipules none. Inflorescences axil- Sladenia (Fig. 126A). Those of Ficalhoa are often
lary, dichasial. Flowers small, 5(6)-merous; sepals paired, originating in the axils of the prophylls
and petals imbricate; sepals free; petals free or of the axillary bud. The flowers are sessile (apart
basally connate; stamens (8-) 10-15, free or adnate from the ultimate flowers of S. integrifolia), with 2
to corolla, anthers basifixed, dehiscing by apical prophylls, small, usually 5-merous and apparently
pores or slits; nectary none; gynoecium 3- or 5- perfect. The calyx is quincuncial and the corolla is
carpellate; ovules axile, 2 or many per carpel; style imbricate (Sladenia) or quincuncial (Ficalhoa);
short, with sometimes very short style branches; the sepals are free. The corolla of Sladenia is
fruit schizocarpic, endocarp crustaceous, or a connate basally, that of Ficalhoa may be very
loculicidal capsule with persistent columella, calyx shortly connate (Shui et al. 2002). The stamens
persistent; seeds winged or not, testa crustose; of Sladenia are borne in a single whorl, and
embryo straight, endosperm copious; n = 24. the swollen filaments are tightly pressed to one
Two monotypic genera, one from East Africa, the another, but apparently free. The thin filaments of
other from SE Asia. Ficalhoa are adnate to the corolla. Sladenia is tri-
carpellate and there are two apical epitropous
VEGETATIVE STRUCTURES. All hairs are unicellu- ovules per carpel. Ficalhoa has five carpels oppo-
lar. Those of Sladenia are very short, and the plant site the petals; the placentas are axile and swollen
may even be glabrous, while those of Ficalhoa may and bear numerous ovules. Sladenia has a rather
be over 1 mm long. Petiole bundles are arcuate, but stout or almost non-existent style continuous with
associated fibers are lacking in Ficalhoa; there the ovary (Fig. 126E); the short style of Ficalhoa is
are also wing bundles in Sladenia. Ficalhoa con- impressed.
tains copious white "latex" (e.g., Verdcourt 1962);
mucilaginous substances in the epidermis are POLLEN MORPHOLOGY. The pollen grains of
reported from Sladenia; there is no hypodermis. Sladenia are tricolpate, broadly ellipsoid and ca.
Stomata are anomocytic. 15 f.lm across. The colpi lack internal thickenings
Growth rings are faint. The vessels are diffuse- and a well-defined pore, although it more or less
porous, and are in radial multiples (Sladenia) or bulges in the center. The exine is moderately thick,
not, with scalariform perforation plates that have smooth or faintly and finely granular; the intine is
(11-)16-17(-30) bars. Intervessel pits are scalari- thin, but slightly thickened beneath the colpi
form (Ficalhoa only) and opposite to alternate. (Wodehouse in Kobuski 1951).
Fiber tracheids are thick- to thin-walled and
have distinctly bordered pits. Parenchyma is FRUIT AND SEEDS. It seems that no mature fruits
diffuse-in-aggregates, vasicentric, or scanty para- of Sladenia have been studied so far (Gilg and
tracheal. Rays are 1-3(4) cells across (Sladenia) or Werdermann 1925; Kobuski 1951; Airy Shaw 1964;
of two distinct sizes, to 1.4mm high, 1-2-seriate pers. obs. S. Dressler). They were described as
and (Ficalhoa) (3- )5-7-seriate, heterogeneous, indehiscent but, since the carpels split easily along
with 1-6(-10) rows of square to upright marginal the septae, they are most probably schizocarpic
cells. There are chambered crystals in strands or and break into 3 mericarps. The exocarp is papery
axial parenchyma cells (Sladenia), or crystals are and the endocarp crustaceous. The small long-
absent (Ficalhoa) (Liang and Baas 1990, 1991). ellipsoid seeds have a ventral raphe and a thin, fine
Sclereids are absent; druses are widespread. The reticulate testa. They have been described as
432 P.F. Stevens and A.L. Weitzman
winged (Keng 1962; Shui et al. 2002) but this seems

to be a drying artifact. When fresh they are swollen
and apparently triangular (Kobuski 1951). Testal
d
cells are polygonal in surface view and probably
little thickened. Ficalhoa has smallloculicidal cap-
sules with several seeds on swollen, spongy pla-
centae. The seed is tiny, compressed ovoid with an
obscure circumferential wing which suggests ~
anemochory (pers. obs. S. Dressler). The walls of i
the testa are unthickened, and the embryo is terete,
with short cotyledons, and surrounded by copious G
endosperm. ~
A
AFFINITIES. Sladeniaceae are a very poorly known B \;J)I F

family. Sladenia has often been included Fig. 126. Sladeniaceae. Sladenia ce/astrifolia. A Floriferous
in Theaceae, either as a separate subfamily branch. B Flower bud. C Flower, vertical section. D Stamens.
(Takhtajan 1997) or as a tribe ofTernstroemioideae E Ovary. F Same in transverse section. G Ovule. (Gilg and
(Keng 1962); Airy Shaw (1964) described it as a Werdermann 1925)
family distinct from Actinidiaceae, in which it had
been included; others have placed it in Dilleniaceae
or Linaceae (see Liang and Baas 1990 for a
summary). Ficalhoa was originally placed in Eri-
caceae but it, too, has been placed in Actinidiaceae 1. Sladenia Kurz Fig. 126
(and also Sapotaceae) and is now often included in Sladenia Kurz, J. Bot. ll: 194 (1873); Sprague, Hook. Icon. 51:
Theaceae (Liang and Baas 1991; these authors t. 3026 (1915); Shui et al., Novon 12: 539- 542 (2002).
favored an association with Ternstroemioideae).
Sladenia was sister to Ternstroemiaceae in rbcL Plant glabrous or with short hairs; leaves spiral,
studies (Savolainen, Fay et al. 2000), although its serrate or entire; inflorescences simply cymose;
DNA was rather degraded. A more comprehensive calyx and corolla free, similar in size, different in
analysis has confirmed this position (support mod- texture; stamens 8-13, filaments swollen, anthers
erate), in addition placing Ficalhoa as its sister sagittate, pilose, porose; gynoecium tricarpellate,
group (support also moderate: Anderberg et al. placentation apical-axile, ovules pendulous, 2per
2002). The two share a number of similarities, most carpel, style with short branches; fruit schizo-
conspicuously their deep-seated, non-surficial carpic, sepals and style persistent, endocarp crus-
phellogen (polarity uncertain), their cymose inflo- taceous; testa crustose, endosperm none; 2n = 48.
rescences with small flowers (possible apomor- Two species, SE Asia.
phies), and their straight embryos (a probable
plesiomorphy). Their recognition as afamily sepa-
rate from Ternstroemiaceae is justified. 2. Ficalhoa Hiern
Despite their evident similarities, Sladenia, from Fica/hoa Hiern, J. Bot. 36: 329 (1898); Robson, Fl. Zambesiaca
the South East Asian mainland, and Ficalhoa, from 1: 405- 406 (1961), tab. 79.
tropical East Africa, show substantial differences
in flower, fruit and wood anatomy. Fossil wood Plant often with long, unicellular hairs; leaves dis-
named Sladenioxy lum africanum has been found tichous, serrate; inflorescences cymose, with two
in Albian-Cenomanian deposits from the north- main spreading branches at base; corolla connate;
ern Sudan (Giraud et al.1992); it is remarkably like stamens 15, in antesepalous groups of 3 and
the wood of extant Sladenia. adnate to corolla, connective much swollen at base
of anthers, anthers transversely dehiscent apically;
KEY TO THE GENERA gynoecium 5-carpellate, placentation axile, ovules
many per carpel, style impressed, style branches
1. Plant glabrous or with short hairs; petals free or almost so;
stamens forming a w horl, filaments swollen, anthers sagit- spreading, recurved, stigmas punctate; fruit a
tate, porose; fruit few-seeded 1. Sladenia loculicidal capsule with a central columella, calyx
- Plant often with hairs to l mm long; p etals basally connate; persistent; seeds minute. One species, F. laurifolia
stamens in groups of three, filaments linear, anthers linear, Hiern, a prominent tree of montane rain forest on
dehiscing byterminal transverse slits; fruit many-seeded the east African mountains.
2. Ficalhoa
Sladeniaceae 433
Selected Bibliography Li, L., Liang, H.X., Peng, H. 2003. Karyotype of Sladenia and its
systematic insights. Acta Bot. Yunn. 25: ~21-326. (In
Chinese, with English summary). Sladema for from
Airy-Shaw, H.K. 1964. Diagnoses of new families, new names, Theaceae, not close to Ternstroemiaceae.
etc. for the seventh edition of Willis's dictionary. Kew Bull. Liang, D., Baas, P. 1990. Wood anatomy of trees and shrubs
18: 249-273. from China II. Theaceae. I.A.W.A. Bull. II, 11: 337-378.
Anderberg, A.A. et al. 2002. See general references. Liang, D., Baas, P. 1991. The wood anatomy of the Theaceae.
Gilg, E., Werdermann, E. 1925. Actinidiaceae. In: Engler & I.A.W.A. Bull. II, 12: 333-353.
Prantl, Nat. Pflanzenfam. ed. 2, 21. Leipzig: W. Engelmann, Savolainen, V., Fay, M.F. et al. 2000. See general references.
pp. 36-47. Shui, Y.-M., Zhang, G.-J., Zhou, Z.-K., Mo, M.-Z. 2002. Sladenia
Giraud, B., Bussert, R., Schrank, E. 1992. A new Theacean wood integrifolia (Sladeniaceae), a new species from China.
from the Cretaceous of northern Sudan. Rev. Palaeobot. Novon 12: 539-542.
Palynol. 75: 289-299. Takhtajan,A.L.1997. See general references.
Hiern, W.P. 1898. A new genus of Ericaceae from Angola. J. Bot. Verdcourt, B. 1962. Flora of Tropical East Africa. Theaceae.
36:329-330,pl. London: Crown Agents.
Keng, H. 1962. Comparative morphological studies in the Wei, Z.-X., Li, D.-Z., Fan, X.-K., Zhang, X.-L. 1999. Pollen ultra-
Theaceae. Univ. Calif. Publ. Bot. 33: 269-384. structure of Pentaphylacaceae and their relationships to
Kobuski, C.E. 1951. Studies in the Theaceae. XXIV. The genus the family Theaceae. Acta Bot. Yunn. 21: 202-206, 3 pl. (in
Sladenia. J. Arnold Arbor. 32: 402-408, pl. 1. Chinese, with English summary).
Li, L. 2001. Chromosome number of Sladenia celastrifolia.Acta
Bot. Yunn. 23: 223-224.
434 P.W. Fritsch
Styracaceae
P.W. FRITSCH
Styracaceae Dumort., Anal. Fam. Pl.: 28, 29 (1829), nom. cons. underground runners, forming open thickets. In
Alniphyllum, Bruinsmia, Huodendron, and Styrax
Evergreen or deciduous trees or shrubs. Leaves the buds are naked, i.e., the outer leaves that con-
alternate, simple, pinnately nerved, petiolate, stitute the buds on twigs of the dormant season
exstipulate. Plants with stellate or peltate tri- typically expand fully during the expansion of the
chomes. Inflorescences terminal and/or axillary new shoot. The remaining genera possess at least
cymes, racemes, or panicles, or sometimes 1-2- two bud scales that abscise quickly after shoot
flowered, usually bracteolate. Flowers bisexual or expansion. In several of the latter genera, a± con-
rarely female in gynodioecious species, actin- tinuous series exists from indurate, non-expansive
omorphic, with hypanthium adnate to ovary wall bud scales proximally through green, leafy bracts
at various levels, usually pendent; calyx synsepa- and finally fully formed leaves distally. Thorns are
lous, truncate or 4-5( -9)-toothed, teeth valvate or present in the species of Sinojackia. Several species
open in bud; corolla sympetalous, (4)5( -8)-lobed of Styrax in Brazil develop xylopodia, which have
or -parted, imbricate or subinduplicate-valvate in a strong capacity to form new sprouts after fire.
bud, generally campanulate to open with the lobes Phyllotaxis is spiral. In Bruinsmia and most trop-
spreading to reflexed; stamens usually twice, rarely ical species of Styrax, the leaves are persistent,
up to four times or equal the number of corolla whereas in the rest of the genera they are decidu-
lobes, uniseriate, replaced by 5 staminodes in ous (the condition of Parastyrax is unknown).
female flowers; filaments usually flattened at least Leaves are usually chartaceous (Fig. 127 A), but are
at the base, adnate to the corolla, sometimes membranous or coriaceous in some species of
forming a tube distally; anthers basifixed, introrse, Styrax. They can be entire or serrate; serrations are
tetrasporangiate, longitudinally dehiscent, oblong capped by a gland-like structure. Stellate hairs of
to linear, the connective roughly equal to or various shapes or sometimes radiate to peltate
surpassing the thecae; ovary 2-4(5)-carpellate, scales are always present (Fig. 127B, C); rust-
2-4(5)-septate at the base but usually 1-locular colored to black glands are scattered over the peti-
through the distal attenuation of the septa, partly oles, inflorescence branches, pedicels, and calyces
to completely inferior, with essentially axile plain many species of Styrax.
centation; style filiform, usually hollow; stigma
terminal, truncate or minutely lobed; ovules VEGETATIVE ANATOMY. Wood anatomical char-
(1- )4-9( -ca. 30) per carpel, anatropous, unitegmic acters occurring in most or all genera include
or bitegmic, tenuinucellate. Fruits mostly dry and growth rings, diffuse porosity, combinations of
usually capsular with loculicidal dehiscence, or both solitaries and pore multiples, scalariform
indehiscent and sometimes samaroid, or drupa- perforation plates, opposite to alternate inter-
ceous, with persistent calyx. Seeds 1-4( -ca. 50), ± vessel pitting, imperforate tracheary elements
globose to fusiform, rarely winged; testa brown, with indistinctly bordered pits, both uniseriate
thin to indurate; cotyledons flattened or nearly and multiseriate heterocellular rays, and scanty
terete; endosperm copious, oily. axial parenchyma distributed as a combination of
A family of 11 genera and about 160 spp., diffuse and diffuse-in-aggregates. Prismatic crys-
tropical and warm-temperate regions of North tals are present in some genera, and silica is
and South America, S Europe, E and SE Asia, and present in many species of Styrax series Valvatae.
Malesia. Pore outline is generally a combination of angular
and circular. Imperforate tracheary elements
VEGETATIVE MoRPHOLOGY. Members of Styra- range from fiber-tracheids with distinctly bor-
caceae are typically medium-sized trees up to dered pits to elements having small, indistinctly
20m tall; occasionally they reach 30m or are bordered pits that approach the simple condition.
shrubs 1-4m tall. Styrax grandifolius produces The species of Styrax series Styrax have a number
Styracaceae 435
of characters often associated with dry climates, trace or one-trace structures, whereas the petals
e.g., simple perforation plates in the wider, early- are always one-trace.
wood vessel elements or plates with a reduced The androecium is seemingly 1-whorled (Fig.
number of bars, increased pore frequency, and 127N), but in some genera the stamens that alter-
decreased vessel element length (Metcalfe and nate with the petals are longer, with their anthers
Chalk 1950; Dickison and Phend 1985). nearer the style. This arrangement led van Steenis
Anatomical characters that are found in leaves {1932) to hypothesize an obdiplostemonous
of nearly all genera include bifacial mesophyll; ancestor for the family. This hypothesis is sup-
anomocytic stomata confined to the abaxial ported by the lower level of origin of the vascular
surface; crystals in the form of druses or prisms or traces to the stamens opposite the petals relative
both; stellate, peltate, or simple-cylindrical tri- to those that alternate with the petals in some
chomes; sheathing and supporting elements asso- flowers, as well as the fact that petal and stamen
ciated with the venation; and pinnate venation traces are more commonly fused than are sepal
accompanied by camptodromous secondary vena- and stamen traces (Dickison 1993). The stamens
tion. Almost all variation observed in the leaf that alternate with the petals develop first in
structure of the ten smaller genera of Styracaceae Styrax and Halesia (Payer 1966), from which it
is found within Styrax. Petiolar vasculature, ex- can be deduced - if the hypothesis of an
cept for Parastyrax, consists of an arc, an arc obdiplostemonous ancestor for the family is
with invaginated ends, or a medullated cylinder correct - that stamen development in Styracaceae
(Schadel and Dickison 1979). is centrifugal.
In many Neotropical species of Styrax, a mass of
FLORAL MORPHOLOGY AND ANATOMY. Flowers of trichomes occurs on the ventral portion of the
Styracaceae are perfect except in females of the stamen filament (Fig. 127P). The connective is ±
gynodioecious species. Female flowers have sta- linear in all genera (Fig. 1270). The anther
minodes with nonfunctional pollen sacs. At least connectives are characterized by conspicuously
some parts of the flower usually have stellate or enlarged, thin-walled epidermal cells on both the
peltate trichomes (Fig. 127E, L, R). The pedicel is ventral and dorsal surfaces that may aid in anther
articulated apically in all genera except Huoden- dehiscence through changes in turgor. At maturity
dron and Styrax, in which pedicel articulation is the microsporangium wall has an endothecium
absent. (consisting of a single layer), except in Huoden-
The degree of connation and adnation of parts dron and some species of Styrax. The stamens are
is highly variable. All Styracaceae have a hypan- always one-trace structures.
thium that is adnate to the ovary wall from just Ovaries are incompletely septate in all genera
above the base of the ovary to nearly the complete except Parastyrax, which has completely septate
length of the ovary. The hypanthium and free ovaries. In most of the incompletely septate
portion of the calyx appear as a continuous, gen- members, the septa are united along a central axis
erally cupuliform or funnelform structure that is until a point± midway distally, where the axis ends
thickened below and typically gradually dimin- and the septa begin to diverge. The edges of the
ishes in thickness distally. Above the hypanthium, septa recede distally toward the ovary wall and
sepals range from distinct to completely connate, continue uninterrupted into the hollow style in the
petals and stamens range from nearly distinct form of ridges. This results in a single, continuous
(coherent at base) to connate most of their length, chamber within the ovary, partially compartmen-
and carpels are nearly always completely connate; talized proximally, that is also continuous with the
stylodia are usually distally distinct in Huoden- ridged and angled stylar canal (Fig. 127E-G, R).
dron). Distal to the hypanthium, the calyx is always Placentation is axile in all genera except some of
free from the corolla and androecium (Fig. 127E, the gynodioecious species of Styrax, in which pla-
R), whereas the stamens are slightly to nearly com- centation is near-basal. In all genera except Styrax
pletely adnate to the petals (Fig. 127N). Connation placentas are divided distally into two lobes by
of either the petals or the stamens or both may ventral carpellary sutures that meet at the apex of
continue for a variable distance distal to the point the central axis. In all species of Styrax except
of corolla and androecium divergence. some of the gynodioecious species, placental out-
The perianth is always differentiated into a growths (obturators) occur between the ovules
calyx and a usually white, sometimes rose-pink (Fig. 127E). The style always appears as an attenu-
or yellow, corolla. Corolla thickness ranges from ation of the ovary (Fig. 127D, E, R). The stigma may
membranous to ± fleshy. Sepals are either three- be flat and not wider than the style, or lobed to
436 P.W. Fritsch
Fig. 127. Styracaceae. A, B Styrax leprosus. A Flowering shoot. nearly capitate and wider than the style (Fig. 127H,
B Close-up of abaxial leaf surface. C-K Styrax redivivus. C R). The orientation of the ovules is usually
Leaf, abaxial view and close-up of abaxial leaf surface. D Pistil.
E Ovary with perianth and androecium, vertical section. F apotropous, but is modified in several genera such
Ovary in transverse section, lower part. G Ovary in transverse that the upper ovules are apotropous and the
section, upper part. H Stigma. I Fruit. J Seed in lateral view. lower are epitropous. Several genera possess intra-
K Seed in polar (hilum end) view. L-M Styrax ramirezii. L ovarian trichomes. Gynoecial vascular features
Flower. M Fruit. N-P Styrax pallidus. N Corolla and androe-
cium opened out. 0 Upper portion of stamen in ventral and
include three-trace carpels; numerous carpellary
lateral view. P Lower free portion of stamen filament show- wall bundles most commonly arising on the inner
ing ventral mass of trichomes. Q- T Alniphyllum fortunei. Q and lateral edges of the hypanthial wall bundles; a
Flower. R Ovary with perianth and androecium, vertical single dorsal bundle per carpel; and ovules vascu-
section, and style in lateral view. S Capsular fruit. T Seed. U larized by a single vein that branches from a pla-
Huodendron biaristatum seed. V Melliodendron xylocarpum
fruit (distal end at bottom). (Orig.)
cental bundle derived from the ventral ovarian
venation (Dickison 1993).
Styracaceae 437
EMBRYOLOGY. Embryological characters have been described as having a mix of floral features
been studied in Alniphyllum, Halesia, Pterostyrax, characteristic of bee, butterfly, moth, and perhaps
and Styrax (Manshard 1936; Copeland 1938; hummingbird syndromes (Sugden 1986). Both
Veillet-Bartoszewska 1960; Yamazaki 1970; nectar and pollen serve as floral rewards for polli-
Yakovlev 1983; Johri et al. 1992). The microspore nators. The stellate hairs present on the exterior
mother cells undergo meiotic divisions, cytokine- surface of the corolla in most species of Styra-
sis is simultaneous, and tetrads are tetrahedral caceae have been suggested as an adaptation for
and isobilateral. Ovules of Styrax are bitegmic, supporting large pollinators, which use them as
whereas those of the other genera are unitegmic. "toe holds" to gather nectar and pollen (Sugden
An integumentary tapetum is missing in most 1986). Both Halesia and Styrax flowers are sweetly
Styracaceae. The archesporia! cell functions as the fragrant (Perkins 1907; Copeland 1938).
megaspore mother cell, undergoes meiotic divi-
sions, produces a linear or T-shaped tetrad, REPRODUCTIVE SYSTEMS. Nearly all species have
and the chalazal megaspore develops into a exclusively hermaphroditic flowers. Six species of
Polygonum-type embryo sac. The polar nuclei fuse Styrax are known to have hermaphroditic and
upon fertilization, and antipodals are ephemeral. female flowers on separate plants (WallnOfer 1997;
Numerous large, unstained starch grains oc- Fritsch 1999). These species are assumed to be
cur in the megagametophyte of some genera functionally gynodioecious, although this has
(Dickison 1993). Endosperm development is cel- not been confirmed with breeding experiments
lular. Embryo development is of the Polygonad (Fritsch 1999). Reports of polygamo-dioecy in
type-Solanad variation and the embryo is straight Bruinsmia are unfounded (van Steenis 1932).
or slightly curved (Johri et al. 1992). Observations of andromonoecy in Halesia are
POLLEN MORPHOLOGY. Pollen morphology of suspected to have been based on immature her-
the family is fairly uniform. Grains of nearly all maphroditic flowers (Wood and Channel11960).
genera are solitary, radially symmetrical, tricol- Predominant to obligate xenogamy is documented
porate with lalongate endoapertures each with for the several species of Styrax examined for
an equatorial bridge, and a tectate-perforate- breeding system type (Sugden 1986; Saraiva et al.
columellate exine. Mean grain size ranges from 1988; Tamura and Hiura 1998).
15.3 x 21.4Jlm to 43.3 x 37.7Jlm, with an overall
mean of 32.7 x 37.7Jlm. The exine is stratified into FRUIT AND SEED. Styracaceae possess a wide
ektexine and endexine, with the ektexine com- array of fruit types, including loculicidal capsules
posed of a distinct tectum, columellae, and foot (Alniphyllum, Huodendron, Styrax; Fig. 1271, S),
layer. Outer tectal surfaces are smooth, granulose, drupes (Parastyrax, Styrax; Fig. 127M), samaras
microspinulose, beaded, or banded (Morton and (Halesia, Pterostyrax), baccoid fruits (Bruinsmia),
Dickison 1992). and nut-like (i.e., dry, indehiscent, one- to several-
seeded) fruits (Changiostyrax, Pterostyrax,
KARYOLOGY. The basic chromosome humber of Melliodendron, Rehderodendron, Sinojackia,
Styrax is x = 8. In Styrax two polyploid species are Styrax; Fig. 127V). The fruits of Styrax series Val-
known (Gonsoulin 1974). Halesia, Sino-jackia and vatae possess a dark purple, often sweet-tasting
Pterostyrax, the only other genera of Styracaceae drupe (Fig. 127M), whereas those of the other
sampled for chromosome number, have n = three series of the genus are almost always dry
12 (Manshard 1936; Lewis et al. 1962, Hsu et al. (fleshy in one species) and either indehiscent or 3-
1994). valvate-dehiscent (Fig. 1271). In all genera the
calyx and hypanthium are persistent in fruit (Fig.
POLLINATION. Species of the family are primarily 127M, S, V), and in seven genera the hypanthial
bee-pollinated. Halesia and Styrax species are gen- tissue greatly elongates during fruit development,
erally most frequently pollinated by bumble bees eventually constituting most of the fruit wall
and honey bees (Chester 1966; Gonsoulin 1974; (Fig. 127V). The style base also elongates and
Sugden 1986). Other pollinators reported for thickens in these genera to form a narrow to broad
Styrax species are papilionoid butterflies, syrphid rostrum in the mature fruit (Fig. 127W). The
flies, sphingid moths, wasps of various families, mesocarp can be fleshy (Parastyrax, some Styrax),
and a wide variety of bees (e.g., carpenter spongy (Sinojackia), woody (Melliodendron),
bees, halictids, anthophorids; Copeland 1938; or not differentiated from the remaining pericarp
Gonsoulin 1974; Sugden 1986; Saraiva et al. 1988; (Huodendron, Pterostyrax). The endocarp is thick,
Tamura and Hiura 1998). Styrax redivivus has indurate, and ribbed in many genera.
438 P.W. Fritsch
There are usually one to four seeds per fruit; in and Dickison 1992), whereas floral morphology
Alniphyllum, Bruinsmia, and Huodendron there suggests a close relationship with members of Eri-
are many. The seed coat is indurate and thick in cales sensu Cronquist (1981). Data from rbcL and
Styrax (Fig. 127J, K), thin in all others (Fig. 127T, atpB chloroplast DNA sequences, as well as 18 S
U). Papillae or hairs cover the seeds of some ribosomal DNA sequences (Soltis et al. 1997),
species of Styrax. Seed shape can be ± globose strongly support the placement of the family
or ellipsoid (Styrax; Fig. 127J, K), irregularly within a clade comprising the families of Ericales
angular (Bruinsmia), flattened-alate or -auriculate s.l. (APG 1998), which includes the Ebenales and
(Alniphyllum, Huodendron; Fig. 127U, V), or fusi- many members of Cronquist's Theales (Morton et
form to narrowly cylindrical (other genera). In al. 1996; Savolainen, Fay et al. 2000), although
Halesia and Styrax, the testa is at first thick, with unequivocal morphological synapomorphies for
postchalazal vascular bundles, then crushed. this clade have not been identified. Close relation-
The endosperm is cellular, oily, and copious ships between Styracaceae, Diapensiaceae, Actini-
(Netolitzky 1926; Corner 1976). diaceae, Clethraceae, Ericaceae, and Cyrillaceae
have been suggested by different phylogenetic
DISPERSAL. Little data exist on the dispersal studies based on DNA sequences, but statistical
mechanisms of Styracaceae. Fruits of Halesia are support for these relationships was weak and con-
dispersed by wind, water, and possibly wildlife tradictory. The five-gene analysis of Anderberg et
(Ridley 1930; U.S.D.A. Forest Service 1948). Fruits al. (2002) has now revealed a strongly supported
of Styrax obassia are dispersed by ground mam- relationship between Styracaceae and Diapensi-
mals and birds (Kato and Hiura 1999). After the aceae, which is also reflected in embryological
fruit wall has become detached, the seeds of the traits.
riparian species S. faberi remain attached to
the infructescence. The seeds, which would other- RELATIONSHIPS WITHIN THE FAMILY. Infra-
wise sink, can thus be transported down-river familial relationships have recently been investi-
by the floating infructescence. The seeds of S. gated with morphological and molecular data
americanus reportedly have been found attached (Fritsch et al. 2001). A combined analysis of three
to the feet of waterfowl (Ridley 1930), but this is DNA sequence datasets and a morphological
probably not a primary means of dispersal of the dataset supports Styracaceae as monophyletic
species because the seed is smooth and ellipsoid. with Styrax and Huodendron in one clade, and the
rest of the genera in another. Alniphyllum and
PHYTOCHEMISTRY. Leaf extracts of Ha[esia and Bruinsmia form a clade that is sister to the remain-
Pterostyrax contain quercetin, kaempferol, tan- ing genera. Halesia macgregori (China) groups
nins, a small amount of leucoanthocyanin, with Rehderodendron (China) rather than to the
saponins, and caffeic, p-coumaric and ferulic acids. other species of Halesia (United States). Unam-
The benzofuran egonol and its glycosides occur biguously placed synapomorphies supporting
in the seed oil of a number of species of Styrax. the monophyly of the Styracaceae are stellate
Fruits of Styrax contain significant amounts of trichomes, linear anther connectives of equal
jegosaponin, a potent defense chemical. Various width throughout, and apotropous ovules (modi-
species of Styrax also contain styracitol, ~-phenyl fied within the family to mixed apotropous/
ethyl alcohol, and coniferin (Hegnauer 1962; epitropous). The genus Lissocarpa was originally
Gibbs 1974). included in Styracaceae (Bentham and Hooker
In many species of Styrax a balsamic resin 1873) but is now commonly placed within the
(benzoin, gum benjamin) exudes from the bark monogeneric family Lissocarpaceae. Lissocarpa
and wood tissues following injury to the cambium. lacks the stellate or peltate trichomes that charac-
This resin consists chiefly of coniferyl cinnamate, terize the Styracaceae.
cinnamyl cinnamate (styracin), and coniferyl
benzoate associated with cinnamic and benzoic DISTRIBUTION AND HABITATS. In the Americas,
acids; minor components are fragrant benzal- the family occurs from the United States to north-
dehyde, vanillin, and styrene (Hegnauer 1962; western Argentina and Uruguay, including the
Langenheim 2003). Antilles. In Asia, the family ranges from Japan and
Korea, south to Nepal, eastern India, Myanmar,
AFFINITIES. Traditionally Styracaceae have been southeast Asia, Indonesia, and Papua New Guinea.
placed in Ebenales (Dahlgren 1980; Cronquist One species of Styrax occurs in the Mediterranean
1981; Thorne 1992; Takhtajan 1997). Pollen region, and another extends into several island
morphology suggests a link to Theaceae (Morton chains of the South Pacific. The distribution of
Styracaceae 439
Styrax essentially mirrors that of the family, and it tures, such as stellate or scale-like trichomes, can
is the only genus occurring in the Neotropics. be observed; hence, the fossil occurrence of the
Halesia is disjunct between the eastern United family in the New World is unproven.
States and China (but see above). The rest of the The results of phylogenetic analysis of the family
genera are distributed among Asia and Malesia. (Fritsch 2001; Fritsch et al. 2001) are consistent
Changiostyrax, Melliodendron, and Sinojackia are with the fossil record in suggesting an Eurasian
endemic to mainland China. Most species are origin of the family. During the Eocene, Styracaeae
understorey trees of humid warm-temperate and must have crossed the North Atlantic land bridge
tropical lowland or montane forests from 100 to and later.
3000 m elevation. Two species of Styrax occur
under summer-dry climates in California and the EcoNOMIC IMPORTANCE. Several Asian species of
Mediterranean region; several others occur in the Styrax (S. benzoin, S. paralleloneurus, S. tonkinen-
cerrado of southern Brazil and Andean subparamo sis) are commercial sources of the balsamic resin
vegetation. The several intercontinental disjunc- benzoin, used in the pharmaceutical, confection,
tions among species of Styrax (Fritsch 1999) and and fragrance industries (Duke 1985). The resin of
several Neotropical species of Styrax is used med-
Halesia, as well as the occurrence of Tertiary fossils
in areas of the northern hemisphere in which the icinally and for incense on a local scale (Standley
family is not known to occur today, suggest that this1924; Schultes and Raffauf 1990). Resin production
family once had a more continuous and wider dis- in the Mediterranean species S. officinalis has been
tribution in North America and Eurasia. widely reported anecdotally, but bark-incision
experiments have failed to induce resin formation
PARASITES. Some species of Styrax (series Cyrta in this species (Zeybek 1970). This discrepancy
in part and series Benzoin) serve as the primary may be a result of nomenclatural confusion with
host for species of gall-forming aphids constitut- the resin produced by Liquidambar. Alternatively,
ing the family Hormaphididae (tribe Cerataphi- genetically based or environmentally controlled
dini). Each cerataphidine species produces galls variation in resin production may exist among
and reproduces sexually on only one species of individuals of S. officinalis (Langenheim 2003).
Styrax. The galls are formed on either vegetative
or floral meristems and come in a wide variety of KEY TO THE GENERA
shapes, including globose, spiral, coralliniform,
1. Fertile shoots produced both laterally and (on at least
alcicorniform, and tubular (Docters van Leeuwen some twigs) terminally on twig of the previous growth
1922; Docters van Leeuwen-Reijnvaan and Docters period; ovules not arranged in 2 longitudinal rows in each
van Leeuwen 1926). Although cerataphidines are carpel 2
distributed pantropically, the galls are found only - Fertile shoots produced only laterally on twig of the pre-
vious growth period; ovules arranged in 2 longitudinal
on Asian species of Styrax. Phylogenetic patterns rows in each carpel 4
provide evidence for host-switching between 2. Stamen connectives 2-3-toothed at apex; stylodia usually
members of series Benzoin and series Cyrta (Stern distinct distally; fruiting calyx epicarpous 1/3 to 2/3 the
1994; Fritsch 1999). length of the fruit 2. Huodendron
- Stamen connectives not toothed at apex; style completely
unbranched; fruiting calyx epicarpous only slightly
PALEOBOTANY AND DISTRIBUTIONAL HISTORY. beyond the fruit base 3
The fossil record of Styracaceae has been reviewed 3. Pedicels articulated; seed to carpel ratio >2; seed coat thin,
in Fritsch et al. (2001}. Eocene and Miocene fossil fragile 3. Bruinsmia
fruits of Rehderodendron are known from France - Pedicels not articulated; seed to carpel ratio ,; 1; seed coat
thick, indurate 1. Styrax
(Mai 1970; Vaudois-Mit* 1983). Fruits of Halesia
4. Fruit capsular; fruiting calyx epicarpous only slightly
are well represented by fossil endocarps in the beyond the fruit base; seed to carpel ratio >2; seeds
European Tertiary (Kirchheimer 1957; Manchester winged 4. Alniphyllum
1999). Fossil fruits of Melliodendron and - Fruit indehiscent; fruiting calyx epicarpous the complete
Pterostyrax are known from the late Tertiary of length of the fruit (excluding rostrum); fruit indehiscent;
seed to carpel ratio !S;2; seeds unwinged 5
Japan (Miki 1941, 1963, 1968). Styrax fossil seeds 6
5. Corolla 4-lobed
are reported for various European localities - Corolla 5-lobed 7
ranging from late Eocene to upper Pliocene 6. Calyx 4-toothed; fruit winged, rostrum <1.5cm long
(Kirchheimer 1957) and for Japan from late 9.Halesia
Miocene through late Pliocene (Miki 1941). - Calyx truncate; fruit not winged, rostrum 4-5.5 em long
5. Changiostyrax
Reports of various genera of Styracaceae in the 7. Thorns present on trunk; stamen filament trichomes
fossil record based on leaf impressions cannot be glossy, arms cylindrical; ovules all apotropous
considered reliable unless detailed anatomical fea- 6. Sinojackia
440 P.W. Fritsch
- Thorns absent; stamen filament trichomes dull, arms flat- 3. Bruinsmia Boerl. & Koord.
tened; upper 2 ovules of each carpel apotropous, lower 2
ovules epitropous 8 Bruinsmia Boerl. & Koord., Natuurk. Tijdschr. Ned.-Indie
8. Ovary 3-locular; fruit drupaceous; exocarp not ribbed 53(1): 68 (1893).
7. Parastyrax
- Ovary 1-locular through distal attenuation of the septa; Bud scales absent; fertile shoots produced both
fruit dry; exocarp ribbed (and sometimes winged) 9
9. Inflorescences ca. 30-70-flowered, branches one-sided; laterally and (on at least some twigs) terminally
corolla lobes connate (coherent) at the base only on twig of the previous growth period; pedicels
8. Pterostyrax articulated; calyx truncate or slightly 5-toothed;
- Inflorescences 1-10-flowered, branches not one-sided; stamen filament trichomes glossy, arms cylindri-
corolla lobes connate distinctly beyond the base 10
10. Pedicels borne on inflorescence axis; calyx toothed; meso-
cal; ovules many per carpel, irregularly arranged;
carp indistinct from endocarp, internally lacunate fruit baccoid; fruiting calyx epicarpous only
10. Rehderodendron slightly beyond the fruit base; seeds numerous.
- Pedicels borne directly on twig; calyx truncate; mesocarp Two spp., India, China, Malaysia, Papua New
distinct from endocarp, both solid 11. Melliodendron Guinea.
GENERA OF STYRACACEAE
4. Alniphyllum Matsum. Fig. 127Q-T
1. Styrax L. Fig. 127 A-P Alniphyllum Matsum., Bot. Mag. (Tokyo) 15: 67 (1901).
Styrax L., Sp. Pl. 1: 444 (1753); Perkins, Pflanzenreich IV, 241:
1-111 ( 1907), rev.; van Steenis, Bull. Jard. Bot. Buitenzorg III, Bud scales absent; fertile shoots produced only
12: 212-272 (1932), Malesian spp.; Hwang, Fl. Rei. Pop. Sin. laterally on twig of the previous growth period;
60(2): 77-150 (1987), Chinese spp.; Svengsuksa & Vidal, Fl. pedicels articulated; calyx 5-toothed; stamen fila-
Cam. Laos Viet. 26: 145-195 (1992), Indochinese spp.;
Fritsch, Ann. Missouri Bot. Gard. 84: 705-761 (1997),
ment trichomes glossy, arms flattened; ovules 8-10
Mesoamerican spp.; Wallnofer, Ann. Naturhist. Mus. Wien per carpel, in 2 longitudinal rows, all apotropous;
99B: 681-720 (1997), 5-anthered spp.. fruit a 5-locular capsule; fruiting calyx epicarpous
Pamphilia Mart. ex A.DC. (1844). Styrax sectio Pamphilia only slightly beyond the fruit base; seeds
(Mart. ex A.DC.) B. Walln. numerous, winged. Three spp., India, China, Laos,
Myanmar, Vietnam.
Bud scales absent; fertile shoots produced both lat-
erally and (on at least some twigs) terminally
on twig of the previous growth period; pedicels not 5. Changiostyrax C.T. Chen
articulated; stamens attached high on corolla tube; Changiostyrax C.T. Chen, Guihaia 15: 291 (1995).
stamen filament trichomes glossy, arms cylindri-
cal; ovules 1-many per carpel; placental obturators Bud scales present; pedicels articulated; calyx
usually present; mesocarp present; seeds 1-3; seed truncate; corolla 4-lobed; ovules 8 per carpel, in 2
coat thick, indurate. n = 8, 16, >20. About 130 spp., longitudinal rows, all apotropous; fruit dry, inde-
E and SE Asia, Malesia, North and South America, hiscent; ectocarp 8-ribbed; mesocarp corky; fruit-
Mediterranean. Fritsch (1999) has divided the ing calyx epicarpous the complete length of the
genus into sect. Styrax, subdivided into series fruit (excluding the 4-5.5-cm-long rostrum). A
Styrax and Cyrta (Lour.) P.W. Fritsch; and sect. Val- single species, C. dolichocarpus (C.J. Qi) C.T. Chen,
vatae Giirke, subdivided into series Valvatae SE China.
Perkins and Benzoin P.W. Fritsch.
6. Sinojackia Hu
2. Huodendron Rehder Fig. 127U
Sinojackia Hu, Contrib. Biol. Lab. Chin. Assoc. Adv. Sci., Sect.
Huodendron Rehder, J. Arnold Arbor. 16: 341 (1935). Bot. 4(1): 1 (1928).
Bud scales absent; fertile shoots produced both Thorns present on trunk; bud scales present; calyx
laterally and (on at least some twigs) terminally on 5-toothed; pedicels articulated; stamen filament
twig of the previous growth period; pedicels not trichomes glossy, arms cylindrical; ovules 6-8 per
articulated; calyx 5-toothed; stamen filament tri- locule, in 2longitudinal rows, all apotropous; fruit
chomes glossy, arms cylindrical; stamen connec- indehiscent; exocarp distinctly lenticellate, not
tives 2-3-toothed at apex; ovules many per carpel, ribbed; mesocarp spongy; fruiting calyx epicar-
irregularly arranged; fruit a 3-locular capsule; pous the complete length of the fruit (excluding
mesocarp absent; seeds numerous. Four spp., the broad rostrum). n = 12. Five spp., China.
China, Myanmar, Thailand, Vietnam.
Styracaceae 441
7. Parastyrax W.W. Sm. 11. Melliodendron Hand.-Mazz. Fig. 127V

Parastyrax W.W. Sm., Notes Roy. Bot. Gard. Edinburgh 12: 231 Melliodendron Hand.-Mazz., Anz. Akad. Wiss. Wien.,
(1920). Math.-Naturwiss. Kl. 59: 109 (1922).
Pedicels articulated; calyx truncate; stamen fila- Bud scales present; flowers 1-2 per leaf axil; calyx
ment trichomes dull, arms flattened; ovary 3- truncate; pedicels articulated; ovules 4 per carpel,
locular throughout; style solid; ovules 4 per carpel, the upper 2 of each carpel apotropous, lower 2
the upper 2 of each carpel apotropous, lower 2 epitropous; fruit dry, indehiscent, turbinate; meso-
epitropous; fruit drupaceous; exocarp distinctly carp distinct from endocarp, both solid; fruiting
lenticellate, not ribbed; fruiting calyx epicarpous calyx epicarpous the complete length of the fruit
the complete length of the fruit (excluding the (excluding the broad rostrum). A single species,
short rostrum). Two spp., China, Myanmar. S China.
8. Pterostyrax Sieb. & Zucc.

Pterostyrax Sieb. & Zucc., Fl. Jap. 1: 94 (1839).
Bud scales present; inflorescences ca. 30-70- APG (Angiosperm Phylogeny Group) 1998. See general
flowered, branches one-sided; pedicels articulated; references.
Bentham, G., Hooker, J.D. 1873. Genera plantarum. Vol. 2.
stamen filament trichomes dull, arms flattened; London: Reeve.
ovules 4 per carpel, the upper 2 of each carpel Chester, E. 1966. A biosystematic study of the genus Halesia,
apotropous, lower 2 epitropous; fruit dry, indehis- Ellis (Styracaceae). Ph.D. dissertation. Ann Arbor, MI: The
cent; exocarp 10-ribbed or 5-winged; mesocarp University of Tennessee, University Microfilms.
Copeland, H.F. 1938. The Styrax of northern California and the
absent; fruiting calyx epicarpous the complete
relationships of the Styracaceae. Am. J. Bot. 25: 771-780.
length of the fruit (excluding the rostrum). n = 12. Corner, E.J.H. 1976. See general references.
Four spp., China, Japan, Myanmar. Cronquist, A. 1981. See general references.
Dahlgren, R. T.M. 1980. A revised system of classification of the
angiosperms. Bot. J. Linn. Soc. 80: 91-124.
9. Halesia J. Ellis ex L. De Candolle,A.L.P.P.1844. Styraceae. In: Prodromus Systema-
tis Naturalis Regni Vegetabilis. Vol. 8. Paris: Treuttel &
Halesia J. Ellis ex L., Syst. Nat. ed. 10, 1044 (1759); Fritsch Wurtz, pp. 244-272.
& Lucas, Syst. Bot. 25: 197-210 (2000), part rev. of North Dickison, W.C. 1993. Floral anatomy of the Styracaceae, includ-
American spp. ing observations on intra-ovarian trichomes. Bot. J. Linn.
Soc. 112: 223-255.
Bud scales present; calyx 4-toothed; pedicels artic- Dickison, W.C., Phend, K.D. 1985. Wood anatomy of the Styra-
ulated; corolla 4-lobed; stamen filament trichomes caceae: evolutionary and ecological considerations. I. A. W.A.
Bull. N.S. 6: 3-22.
dull, arms flattened; ovules 4 per carpel, the upper Docters van Leeuwen, W.M.1922. Uber einige von Aphiden an
2 of each carpel apotropous, lower 2 epitropous; Styrax-Arten gebildete Gallen. Bull. Jard. Bot. Buitenzorg,
fruit dry, indehiscent, 2-4-winged; fruiting calyx ser. 3 4: 147-162.
epicarpous the complete length of the fruit Docters van Leeuwen-Reijnvaan, J., Docters van Leeuwen,
(excluding the <1.5-cm-long rostrum). n = 12. W.M. 1926. The zoocecidia of the Netherlands East Indies.
Batavia: Drukkerij de Unie.
Three spp., eastern U.S.A., China. Duke, J.A. 1985. CRC handbook of medicinal herbs. Boca
Raton, FL: CRC Press.
Fritsch, P.W. 1996a. Population structuring and patterns of
10. Rehderodendron Hu morphological variation in California Styrax (Styracaceae).
Aliso 14: 205-218.
Rehderodendron Hu, Sinensia 2: 109 (1932).
Fritsch, P.W. 1996b. Isozyme analysis of intercontinental dis-
juncts within Styrax (Styracaceae): implications for the
Bud scales present; inflorescences 3-10-flowered; Madrean-Tethyan hypothesis. Am. J. Bot. 83: 342-355.
calyx 5-toothed; pedicels articulated; ovules 4 per Fritsch, P. W. 1997. A revision of Styrax (Styracaceae) for
carpel, the upper 2 of each carpel apotropous, western Texas, Mexico, and Mesoamerica. Ann. Missouri
Bot. Gard. 84: 705-761.
lower 2 epitropous; fruit dry, indehiscent, broadly Fritsch, P.W. 1999. Phylogeny of Styrax based on morphologi-
cylindrical; exocarp ca. 10-ribbed; mesocarp cal characters, with implications for biogeography and
indistinct from endocarp, internally lacunate; infrageneric classification. Syst. Bot. 24: 356-378.
fruiting calyx epicarpous the complete length of Fritsch, P.W. 2001. Phylogeny and biogeography of the flower-
the fruit (excluding the short rostrum). Five spp., ing plant genus Styrax (Styracaeae) based on chloroplast
DNA restriction sites and DNA sequences of the internal
China, Myanmar, Vietnam. transcribed spacer region. Molec. Phylog. Evol. 19: 387-408.
442 P. W. Fritsch
Fritsch, P. W. In press. Styracaceae. In: Flora of North America. nales based upon rbcL sequence data. Syst. Bot. 21: 567-586.
New York: Oxford University Press. Netolitzky, F. 1926. Anatomie der Angiospermen-Samen. In:
Fritsch, P.W., Lucas, S.D. 2000. Clinal variation in the Halesia Linsbauer, K. (ed.) Handbuch der Pflanzenanatomie. Vol.10,
carolina complex (Styracaceae). Syst. Bot. 25: 197-210. 4. Berlin: Borntrager.
Fritsch, P.W., Morton, C.M., Chen, T., Meldrum, C. 2001. Phy- Payer, J.-B. 1966. Traite d'organogenie comparee de Ia fleur.
logeny and biogeography of the Styracaceae. Int. J. Pl. Sci. New York: J. Cramer.
162 (6 Suppl.): S95-S116. Perkins, J. 1907. Styracaceae. In: Engler, A. (ed.) Pflanzenreich
Gibbs, R.D. 1974. Chemotaxonomy of the flowering plants, IV. Vol. 241 (Heft 30). Leipzig: W. Engelmann, pp. 1-111.
4 vols. Montreal: MeGill-Queen's University Press. Perkins, J. 1928. Ubersicht tiber die Gattungen der Styracaceae.
Gonsoulin, G.J. 1974. A revision of Styrax (Styracaceae) in Leipzig: W. Engelmann.
North America, Central America, and the Caribbean. Sida Ridley, H.N. 1930. The dispersal of plants throughout the
5: 191-258. world. Ashford, Kent: L. Reeve.
Giirke, M.1891. Styracaceae. In: Engler, A., Prantl, K. Die natiir- Saraiva, L., Cesar, 0., Monteiro, R. 1988. Biologia da poliniza-
lichen Pflanzenfamilien, IV, 1. Leipzig: W. Engelmann, <;iio e sistema de reprodu<;iio de Styrax camporum Pohl e
pp. 172-180. S. ferrugineus Nees et Mart. (Styracaceae). Revista Brasil.
Hegnauer, R. 1962. See general references. Bot. 11: 71-80.
Hsu, P.S., Weng, R.F., Kurita, S. 1994. New chromosome counts Savolainen, V., Fay, M.F. et al. 2000. See general references.
of some dicots in the Sino-Japanese region and their sys- Schadel, W.E., Dickison, W.C. 1979. Leaf anatomy and venation
tematic and evolutionary significance. Acta Phytotax. Sin. patterns of the Styracaceae. J. Arnold Arbor. 60: 8-37.
32: 411-418. Schultes, R.E., Raffauf, R.F. 1990. The healing forest. Portland,
Hwang, S.-M. 1987. Styracaceae. In: Flora Reipublicae Popu- OR: Dioscorides Press.
laris Sinicae. Vol. 60(2). Beijing: Science Press, pp. 77-150. Soltis, D.E. et al. 1997. See general references.
Hwang, S.-M., Grimes, J. 1996. Styracaceae. In: Wu, Z.-Y., Spongberg, S.A. 1976. Styracaceae hardy in temperate North
Raven, P.H. (eds.) Flora of China. Vol. 15. Beijing: Science America. J. Arnold. Arbor. 57: 54-73.
Press, pp. 253-271. Standley, P.C. 1924. Trees and shrubs of Mexico. Contrib. U.S.
Johri, B.M. et al. 1992. See general references. Natl. Herb. 23: 1-1721.
Kato, E., Hiura, T. 1999. Fruit set in Styrax obassia (Styra- Stern, D.L. 1994. Phylogenetic evidence that aphids, rather than
caceae ): the effect of light availability, display size, and local plants, determine gall morphology. Proc. Roy. Soc. Lond. Ser.
floral density. Am. J. Bot. 86: 495-501. B, Biol. Sci. 256: 203-209.
Kirchheimer, F. 1957. Die Laubgewachse der Braunkohlenzeit. Sugden, E.A. 1986. Anthecology and pollinator efficacy of
Halle (Saale): VEB Wilhelm Knapp. Styrax officinale subsp. redivivum (Styracaceae). Am. J. Bot.
Langenheim, J. 2003. Plant resins: chemistry, evolution, 73: 919-930.
ecology, and ethnobotany. Portland: Timber Press. Svengsuksa, B.K., Vidal, J.E. 1992. Styracacees. In: Morat, P.
Lewis, W.H., Stripling, H.L., Ross, R.G. 1962. Chromosome (ed.) Flore du Cambodge du Laos et du Vietnam. Vol. 26.
number reports LXIX. Taxon 29: 728. Paris: Museum National d'Histoire Naturelle, pp. 145-195.
Magellan, S., Crane, P.R., Herendeen, P.S. 1999. Phylogenetic Takhtajan, A. 1997. See general references.
pattern, diversity, and diversification of eudicots. Ann. Mis- Tamura, S., Hiura, T. 1998. Proximate factors affecting fruit set
souri Bot. Gard. 86: 297-372. and seed mass of Styrax obassia in a masting year. Eco-
Mai, D.H. 1970. Subtropische Elemente im europaischen science 5: 100-107.
Tertiar I. Palaont. Abh. B 3: 441-503. Thorne, R.F. 1992. Classification and geography of the flower-
Mai, D. H. 1988. Uber antillanische Styracaceae. Feddes Repert. ing plants. Bot. Rev. 58: 225-348.
99: 173-181. U.S.D.A. Forest Service. 1948. Woody-plant seed manual.
Manchester, S.R. 1999. Biogeographical relationships of North Washinton, D.C.: U.S. Government Printing Office.
American Tertiary floras. Ann. Missouri Bot. Gard. 86: van Steenis, C.G.G.J. 1932. The Styracaceae of Netherlands
472-522. India. Bull. Jard. Bot. Buitenzorg III, 12: 212-272.
Manshard, E. 1936. Embryologische Untersuchungen an Styrax Vaudois-Mieja, N. 1983. Extention palaeogeographique en
obassia Sieb. et Zucc. Planta 25: 264-383. Europe de l'actuel genre Rehderodendron Hu (Styracaceae).
Metcalfe, R.C., Chalk, L. 1950. Anatomy of dicotyledons. C. R. Acad. Sci. Paris II, 296: 125-130.
Oxford: Clarendon Press. Veillet-Bartoszewska, M. 1960. Embryogenie des Styracacees.
Miers, J. 1859. On the natural order Styraceae, as distinguished Developpement de 1' embryon chez le Styrax officinalis L. C.
from the Symplocaceae. Ann. Mag. Nat. Hist. III, 3: 394-404. R. Acad. Sci. Paris II, 250: 905-907.
Miki, S. 1941. On the change of flora in Eastern Asia since Ter- Wallnofer, B. 1997. A revision of Styrax L. section Pamphilia
tiary period. I. The clay of lignite beds flora in Japan with (Mart. ex A. DC.) B. Walln. (Styracaceae). Ann. Naturhist.
special reference to the Pinus trifolia beds in Central Hondo. Mus. Wien 99B: 681-720.
Japan. J. Bot. 11: 237-304. Wood, C.E., Channell, R.B. 1960. The genera of the Ebenales in
Miki, S. 1963. Further study on plant remains in Pinus trifoli- the southeastern United States. J. Arnold Arbor. 41: 1-35.
ata beds, Central Hondo, Japan. Special issue, Chigaku- Yakovlev, M.S. (ed.) 1983. Comparative embryology of
kenkyu 80-93. flowering plants, Phytolaccaceae-Thymelaeaceae.
Miki, S. 1968. Paleodavidia, synonym of Melliodendron and Leningrad: Nauka (in Russian).
fossil remains in Japan. Bull Mukogawa Women Univ. 16: Yamazaki, T. 1970. Embryological studies in Ebenales 1. Styra-
287-291. caceae. J. Japan. Bot. 45: 267-273.
Morton, C.M., Dickison, W.C. 1992. Comparative pollen mor- Zeybek, N. 1970. Liefert Styrax officinalis L. ein Harz? Ber.
phology of the Styracaceae. Grana 31: 1-15. Schweiz. Bot. Ges. 80: 189-193.
Morton, C.M., Chase, M.W., Kron, K.A., Swensen, S.M. 1996. A
molecular evaluation of the monophyly of the order Ebe-
Symplocaceae 443
Symplocaceae
H.P. NooTEBOOM
small scales. The hairs are simple and 1-celled but

Symplocaceae Desf., Mem. Mus. Hist. Nat. Paris 6: 9 (1820),
nom. cons. have transverse septa; rarely they are bulbous-
based. The indumentum varies greatly and some-
Evergreen shrubs or trees (but S. paniculata is times is important for the discrimination of the
deciduous). Leaves spirally or distichously species but often it is of no systematic value at all.
arranged, exstipulate, simple, petiolate; leaf Sometimes the absence of hairs is taxonomically
margin blade dentate, glandular dentate, or entire, important.
midvein adaxially impressed (rarely flat or pro- The leaves are mostly spirally or distichously
minent). Inflorescence a thyrse or more often arranged, rarely they are pseudoverticillate. In
reduced or compound to a spike, raceme, panicle, general, phyllotaxis is constant for a species but,
glomerule, or a solitary flower. Flowers actin- in some species with distichous leaves, the 'leaders'
omorphic, bisexual, rarely (unisexual), in spikes, often have spirally arranged leaves. The shape, size
racemes, panicles, or glomerules (solitary), sup- and venation of the leaves often yield good specific
ported by 1 bract and 2 prophylls (exceptionally characters but in some species vary so widely that
bractless or with several bracts in the axil of a species cannot be distinguished vegetatively.
leaf); calyx lobes (3- )5, valvate or imbricate, per- With the exception of a few species, in the dry
sistent; corolla of (3- )5( -11) imbricate lobes, white state the midrib is sulcate. The number of lateral
or rarely yellow, gamopetalous but divided nearly nerves varies greatly. The reticulation of the veins
to base (or sometimes to middle in sub g. Symplo- is often an important character. In some species
cos); stamens many (4 or 5), rarely few, adnate the secondary veins are transverse to the nerves.
to base of the corolla tube, monadelphous fila- The tertiary and the quaternary veins can be
ments all united in subg. Symplocos, and all united prominent or obscure, the reticulation dense or
or monadelphous or pentadelphous, the same or lax, and sometimes a fine reticulation is visible
in 5 antesepalous bundles in subg. Hopea; anthers only in translucent light. Usually details of the
subglobose, 2-locular, opening with longitudinal venation are only distinct on the undersurface.
slits; ovary inferior or half-inferior, (2)3( -5)- The margin of the leaves is often provided with
celled, usually with an apical 5-glandular, annular, vesicular or tooth-like glands. The latter are
cylindrical, or 5-lobed disk; style filiform to thick; usually lost during the expansion of the leaves.
stigma small, capitate or 2-5-lobed; ovules 2-4 per The length of the petiole is very variable in some
cell. Fruit a drupe; seeds with copious endosperm; species, but rather constant and providing a good
embryo straight or curved; cotyledons very short. character in others. In cross-section the petiole is
A monogeneric family of ca. 3200 species, widely usually convex beneath and grooved or flat above.
distributed in tropical and subtropical Asia, In some species it has decurrent edges and may
Australia, and America, with few species extend- even appear winged by the decurrent leafblade.
ing to temperate Asia and North America.
VEGETATIVE ANATOMY. The leaves of Symplocos
VEGETATIVE MoRPHOLOGY. Symplocaceae are are bifacial. The cuticle is smooth, striated or
shrubs or trees, varying, in some species, from ridged. The lower epidermis is generally smooth
small shrubs to moderately large trees. The twigs but sometimes papillate. The thickness of the
are slender to rather thick and in some species are leaves is very variable for the genus as a whole, but
covered with pulvinate leaf scars. When the leaves relatively constant for the species. The size of the
are distichously arranged, the twigs are often epidermal cells varies considerably from species to
zigzag. In most species growth is discontinuous species; their radial walls are straight or undulate.
and the twigs are terminated by a bud protected A hypodermal layer is present in several species,
by large, leathery scales. In the species with con- but is sometimes not continuous. In one and the
tinuous growth, the buds are naked or covered by same species, the number of layers of palisade
444 H.P. Nooteboom
parenchyma can vary from 1 to 3, but mostly bordered to almost simple, scalariform, transi-
this character is rather constant. The spongy tional, opposite or even tending to alternate. In
parenchyma is very compact to very loose; in very oblique end walls, perforations are scal-
some species it contains sclereids. The midrib ariform with 9-59 bars. Spiral thickenings are
encloses one large vascular bundle, usually en- absent or present in tails only, or are well devel-
veloped by a sclerenchymatic sheath which can oped. Tyloses or warty layers are noted in few spec-
be continuous or not. Above the vascular bundle, imens only. The ground tissue is composed of long
usually several layers of parenchymatic or col- fibre-tracheids (1170-2920f.lm). The pits are con-
lenchymatic tissue can be found. Beneath the vas- spicuously bordered (4-8 f.lm in diameter) on both
cular bundle, mostly a thick sclerenchymatic cap radial and tangential walls, with slit-like, included
is present. In all but three species, the midrib is or extending apertures. Spiral or annular thicken-
grooved in the dried state. ings are absent, restricted to fibre tips, or well
The leaf surface is totally glabrous in some developed. Axial parenchyma is scanty paratra-
species, in other species it possesses a more or cheal and apotracheally diffuse, very sparse to
less dense indumentum, especially underneath on fairly abundant, in strands of 4-8 cells. Rays are
the nerves. The hairs are nearly always septate; of the Kribs' type heterogeneous I, sometimes
they are appressed to erect and sometimes ligni- tending to heterogeneous II. Crystals are mostly
fied and heavily cutinised, especially in the basal absent; if present they are irregularly rhomboidal
part. or fragmented and form crystalline masses mostly
The stomata are of the paracytic type. In most deposited in ordinary ray cells, rarely in cham-
(or all?) species there are two kinds of stomata: bered ray cells or axial parenchyma. Silica bodies
besides the "normal" ones, a few, very large, so- are absent. Pith flecks are present in some speci-
called water stomata occur. Within certain limits, mens only (Metcalfe and Chalk 1950; van den
the size of the stomata is constant for a species. Oever et al. 1981).
Clustered calcium Ca-oxalate crystals probably A similar wood structure anatomy is found in
occur in all species and can be found in most parts quite a few other families, such as Theaceae,
of the leaves. Tannin-like compounds, probably Cornaceae, Nyssaceae, Saxifragaceae, Clethraceae,
condensed proleucoanthocyanins and catechins Cyrillaceae, and Hamamelidaceae (e.g. Janssonius
(so-called Inklusen) are common in the epidermal 1920), representing the Comus type of Huber
cells, sometimes giving them a very characteristic, (1963).
brown colour. The leaves of many species contain The results of a fairly large sample of Symplocos
a high amount of aluminium compounds; these species and specimens (van den Oever et al. 1981)
are often visible as dark bodies in the leaf tissues confirm the general validity of most of the altitu-
(cf. fat-like bodies of Metcalfe and Chalk 1950). dinal and latitudinal trends in wood anatomical
Leaf anatomy is rather constant for some species characters reported by Baas ( 1973) for the genus
and highly variable for others, probably often due !lex.
to altitudinal variation. For further information on
leaf anatomy, see Loesener (1896), Cador (1900), INFLORESCENCE AND FLOWER STRUCTURE. The
Wehnert (1906), Neger (1923), and Metcalfe and inflorescences are inserted in the axils of the upper
Chalk (1950). leaves, or on the wood beneath them; rarely, they
The wood is light and soft, white to yellowish, are terminal. Racemose in appearance, they are
sometimes with a pinkish tinge. Growth rings, often true racemes or spikes, but with a tendency
when present, are usually visible to the naked eye, to branch. Sometimes the inflorescences are true
as are the vessels and broad rays. Microscopically, panicles, or condensed to fascicles or very short
the wood is diffuse-porous, very rarely tending to clusters of flowers, or even reduced to one flower.
semi-ring-porous. The frequency of vessels is Morphologically, they are probably thyrses or
12-271/mm2; typically they are solitary but derived there from.
sometimes are arranged in radial pairs or more Principally one bract and two prophylls are
rarely in multiples of up to 4. Vessels are angular present below each flower, the latter directly
to oval in transverse section and the average vessel beneath the flower (Fig. 128B). Occasionally, a
member length is 772-1836 f.lm. Inter-vessel pits pedicel is present between the bract and the pro-
are scalariform, transitional or opposite with a phylls. The bracts and/or prophylls are persistent
tangential diameter of 4-15 f.lm, or in scalariform or caducous. Sometimes the presence of additional
pits even wider, with slit-like, included apertures. bracts (metaxyphylls) are indicatives of an origin
Vessel-ray and vessel-parenchyma pits are half- from a more elaborate inflorescence. Sometimes
Symplocaceae 445
the bracts and prophylls bear vesicular glands on controversial character by Nooteboom (1975)
the margin. showed that the ovules are attached on the indu-
The flowers are usually bisexual, but rarely func- plicate part of the carpels, close to the centre. The
tionally unisexual flowers occur in taxa that are septa are not connate to each other at the centre in
either dioecious or polygamous. In male flowers the upper part of the ovary (see also the develop-
the style is small without a stigma; in female mental study of Symplocos paniculata flowers by
flowers the number of stamens, the anthers of Caris et al. 2001).
which are sterile, is reduced, sometimes even to
less than ten. EMBRYOLOGY. The ovule is anatropous, uniteg-
The ovary is inferior and the calyx tube is adnate mic, and tenuinucellar; the archesporia! cell fun-
to it; the free part of the calyx is called the calyx ctions as the megaspore mother cell; the chalaza!
limb, consisting of (3-)5 free or basely connate or subchalazal megaspore develops into a
lobes. The calyx lobes are normally ± equal, but Polygonum-type embryo sac (Davis 1966; Corner
in several species the calyx splits after anthesis, 1976).
thus becoming symmetrically cleft. The incision
between the calyx lobes sometimes becomes POLLEN MORPHOLOGY (by R.W.E.J.M. VAN DER
longer with age. The presence or absence of an HAM). Pollen grains of Symplocos are 3(2 or 4)-
indumentum on the calyx, and the shape and the aperturate monads of 20-70 (av. 30) J..tm equatorial
length of the calyx lobes sometimes offer useful diameter. In polar view the grains are circular, with
characters. more or less protruding apertures, to angular; in
In subg. Symplocos the petals are connate into a equatorial view they are oblate to oblate spher-
tube which often can be seen only from the inside oidal. The colpate ectoapertures are usually
because the overlapping margins are free; in subg. (very) short and often inconspicuous in light
Hopea the petals are connate only at their very microscopy, unless a thickened margo is present.
base. Mostly the petals are glabrous or provided The endoapertures are mostly circular to lalongate
with some minute hairs towards the outer base. pores, sometimes lalongate colpi, provided with
Rarely they are covered with a dense indumentum, an irregularly delimited and mostly fragmented
at least in bud. For most species the corolla does endoannulus or polar costae. The total thickness
not offer good specific characters. of the exine is 1-4J..tm. The nexine is usually
In sub g. Hopea the stamen filaments are connate thicker than the sexine, and sometimes shows
only towards the base for at most 2 mm, and form endocracks, especially around the apertures. A
5 antepetalous-alternipetalous bundles or are very thin, fragmented endexine is probably
evenly arranged around the gynoecium. In subg. present (Barth 1982; Nagamasu 1989). The infrate-
Symplocos the stamen filaments are fused to form ctum is distinct (columellate) to thin and indis-
a long tube. The number of the filaments and the tinct. The tectum is finely punctate, perforate to ±
relative length of their free part provide good reticulate, and psilate or provided with supratectal
characters. In some species the stamens are hairy. elements (scabrate, rugulate, areolate, verrucate,
The upper part of the ovary is covered by an echinate). Irregularly distributed globules of 2-
often stellate disk which bears 5 glands. In some 6J..tm have been interpreted as Ubisch bodies.
species the shape of this part of the ovary is vari- On the basis of the presence/absence of grooves
able. The disk can be globose, low-cylindrical, or around the endoannulus (or along the equatorial
fiat and then inconspicuous. It is often covered by sides), Van der Meijden (1970) distinguished
an indumentum. two main pollen types, which correspond with the
The style is of the same shape in the whole two subgenera Symplocos and Hopea delimited by
genus; its indumentum is often coincident with Nooteboom (1975). Other attempts at relating
that of the disk. In male flowers of dioecious or pollen morphology and infrageneric classification
polygamous species, the peltate or punctiform of Symplocos have been undertaken by Gupta and
stigma is absent. Sharma (1977) and Barth (1979).
The ovary is mostly rather constantly 2-, 3-, 4-,
or 5-celled; the number of ovules being 2-4, but KARYOLOGY. The base number seems to be x = 11;
usually 4 in each locule. Placentation is mostly in subgen. Hopea n = 11 is most frequent (see tab-
considered axile but, according to Chirtoiii (1918) ulation by Nooteboom 1975); the very few records
and Nakai (1924), for Asiatic species placentation of 2n = 23 and 24 may be due to B-chromosomes
is parietal. This was denied by Randel-Mazzetti (Mehra and Gill1968). The chromosome number
and Peter-Stibal (1943). Re-examination of this of Symplocos pendula (the only species of sub g.
446 H.P. Nooteboom
Symplocos studied so far) was determined as thocyanins seem to be rather common. The record
approximately 2n = 90 and is probably octoploid, of the iridoid compound cornin is remarkable
based on x = 11 (Nooteboom 1975). and agrees with a position within the Ericales. The
Karyotypes of all species of sub g. Hope a are very leaves of subg. Hopea contain much alumin-
similar. This is consistent with the assumption that ium and turn yellow at drying when the alumin-
hybridisation between them is possible. This is ium forms complexes with the leaf flavonoids
also indicated by the frequency of intermediate or (Chenery 1948a; Nooteboom 1975). In subg.
transitional specimens found in the herbarium. Symplocos no accumulation of aluminium has
There are also certain hermaphroditic taxa which been observed, and their leaves never turn
always show a high proportion of sterile pollen, yellow.
which may be due to their hybrid origin. Already Radlkofer (1904) reported that the ash
of the leaves contain ca. 50% aluminium oxide. He
FRUIT AND SEED. The fruit is a drupe with a also described the so-called Tonerdekorper in the
fleshy, corky or woody mesocarp and a very hard leaves of Symplocos. These are masses of colour-
stone (endo carp). Shape and size of the fruit less material filling often large parts of the cells,
provide good characters, particularly for groups predominantly in the palisade parenchyma.
of related species. The shape can be globose, Radlkofer stated that most of the aluminium of
ellipsoid, ovoid, obovoid, ampulliform, spindle- the leaves is accumulated in these masses. Neger
shaped, and cylindrical. The shape of the stone ( 1923) cultivated individuals of S. lucida S. & Z.
may differ from the shape of the fruit, and ovoid (= S. japonica DC) in solutions containing differ-
fruits sometimes contain ampulliform stones. ent amounts of aluminium salts. He observed that
The seeds are exotegmic-ericoid (Fig. 2D). The plants grew best on a solution of 1% aluminium,
seed coat is very thin. The embryo can be straight an amount that is toxic for other xerophyllous
or curved to U-shaped, or curved twice in two plants.
planes. Almost always the shape of the embryo is
similar to that of the seed. Globose and ampulli- AFFINITIES. The taxonomic history of the
form fruits are mostly correlated with curved family has been dealt with by Nooteboom (1975).
embryos; cylindrical fruits always have straight Following Jussieu (1789), most later authors
embryos. Ovoid and ellipsoid fruits may contain placed Symplocaceae in the same alliance with
straight or curved seeds and embryos; in some Ebenaceae, Sapotaceae, and Styracaceae, a posi-
species there are even intergradations between tion that was maintained. All these taxa were
them. The endosperm often contains starch as well part of the Ebenales of most later authors, up to
as fatty oil and probably aleuron (pers. obs.). SeedsTakhtajan (1997).
of S. paniculata are reported to be starch-free Formerly Symplocos was frequently included in
and to contain 21.2% proteins and 51.7% fatty oil. Styracaceae but, besides the well-known morpho-
Palmitinic, oleic and linolic acids were found to be logical and anatomical differences exhibited
the main fatty acids in samples of Symplocos seeds with this family, Kirchheimer (1949) noted that in
investigated to date. Symplocos fruits each locule possesses an apical
Germination of S. paniculata is phanerocotylar, pore which is absent in the fruits of Styracaceae;
and has been described by Lubbock (1892). according to him this difference goes back to
Eocene times at least.
DISPERSAL AND REPRODUCTION. Birds are prob- In the ordinal classification of APG (1998),
ably the main dispersal agents. Seedlings are, Symplocaceae were assigned to a broadly con-
however, rarely encountered in the forest (pers. strued order Ericales, comprising some 24 families
obs.). In S. cochinchinensis var. laurina on the Doi including those mentioned above. This affiliation
Inthanon (Thailand), it was observed that most, if has been confirmed by several recent molecular
not all, young trees sprouted from fallen branches. studies, but none of them, including the five-
One fallen treelet gave rise to more than ten new gene analysis by Anderberg et al. (2002), has been
trees. This probably reflects a high degree of seed able to determine a precise position of the family
sterility (Nooteboom 1975). within this clade. When comparing the broader
based, recent molecular studies (as available at the
PHYTOCHEMISTRY. The reader is referred to time of writing, Nov. 2000), Symplocaceae invari-
Hegnauer (1973, 1990). Phenolic compounds, ably form part of this Ericales clade or grade, but
including gallic and ellagic acids and the the closest relatives vary considerably and in-
hydrolysable tannins based on them, and leucoan- clude families such as Lecythidaceae, Sapotaceae,
Symplocaceae 447
Styracaceae, Theaceae, Ternstroemiaceae, and Only the Japanese Pliocene fossils of S. paniculata,
Diapensiaceae. S. lancifolia and S. sumuntia have curved seeds. Of
the extant Old World species, ca. 20% possess 1- or
DISTRIBUTION AND HABITATS. The large genus of 2-curved seeds, among them S. cochinchinensis.
nearly 300 species is distributed in the eastern This species not only occupies nearly the whole
parts of the Old World from the Deccan Peninsula range of Symplocos in the Old World, but also has
to northern China/Japan, extending beyond the many infraspecific taxa and in some places domi-
Tropic of Cancer, in Australia reaching as far as nates the vegetation. Taken together, the species
New South Wales and Lord Howe Island, and in the with curved embryos represent the vast majority
Pacific as far as Fiji. In the New World Symplocos of individual plants of the genus in SE Asia. As
is found from the south-eastern United States to far as is known from the fossil record, no relatives
southern Brazil. of this species with curved seeds ever reached
Subg. Symplocos is represented in Asia only Europe.
by two species, while the New World harbours Several fossil Symplocos endocarps are so
approximately 130 species. The range of this sub- similar to endocarps of Recent species that it is
genus is almost entirely tropical. Subg. Hopea has almost certain that, from the Lower Tertiary up to
ca. 110 species in Asia, one species extending north Recent times, no major evolutionary changes have
to Korea, and ca. 50 in the New World. occurred. Some fossil species, particularly from
Symplocos grows in warm-temperate and tropi- Japan, such as S. paniculata, S. lancifolia, and S.
cal zones, preferably in moist to wet mixed, mostly sumuntia, are hardly distinguishable from Recent
evergreen rainforest. The genus is most abundant species.
in mountain forests. On Mt. Kinabalu and the New Hardly any other genus of dicotyledons is
Guinean mountains, it ascends up to an altitude of known to have occurred in such local abundance
4000 m, where it is represented by microphyllous and diversity as Symplocos. Kirchheimer (1949)
dwarf shrubs. In the Andes it extends from 500 calculated the total mass of carpolithic coal in the
to 4000 m altitude but is most frequent from 2500 Vogelsberg mountains to be 3500m3 • In 125cm3 of
to 3500m (Stahl 1995). Symplocos avoids arid coal he found 99 Symplocos endocarps and 19
regions. fruits or seeds of other plants. The total number of
Symplocos endocarps was estimated to be more
DISTRIBUTIONAL HISTORY AND PALAEOBOTANY. than 2.5 milliards, mainly belonging to S. minutula
The oldest record of fossil Symplocos consists of (v. Sternb.) Kirchheimer, but also to S. lignitarum
pollen grains from the Maastrichtian of California (Quensted) Kirchheimer. According to Kirch-
(Muller 1981). According to Krutzsch (1989), heimer, this enormous mass of fruits must have
who has dealt with the distributional history of been produced in situ because the coal layers
Symplocos, by the Upper Eocene the genus had provided no evidence of the presence of running
acquired a coherent north-Tethyan distribution water which could have transported the endo-
area from western N America to E Asia. This belt carps. Kirchheimer estimated that the coal layer
disintegrated in the younger Tertiary, and in was deposited during a period of about 100 years,
Europe the genus became extinct at the beginning and that the surrounding plants would have fur-
of the Pleistocene. The expansion southward in nished at least 25 million endocarps a year. The
America and south-eastward in SE Asia occurred vegetation certainly was dominated by Symplocos
from the Pliocene onward, concomitantly with the trees, although fruits/seeds of Mastixia, Brasenia,
formation of secondary centres in America and SE Magnolia, and Vitis were found in the same beds.
Asia.
The oldest fossil fruits are from the Lower ECONOMIC IMPORTANCE. The reader is referred to
Eocene of C and W Europe. In some instances the Nooteboom (1991). The inner bark of S. cochinchi-
fruit stones of up to as much as four species were nensis ssp. cochinchinensis var. cochinchinensis
found together in European brown-coal beds. This and S. fasciculata was often used as a mordant in
makes it probable that in the Lower Tertiary Sym- the batik industry and, mixed with other plants,
plocos was already as diversified as it is today. as a dye. It gives a yellow colour by itself but is
About 25 species have been described from fossil more frequently used in the preparation of reds
fruits. About 22 of them, all occurring in Europe, derived from Morinda spp., Caesalpinia sappan,
possessed straight seeds; they belonged to subg. Butea spp., and other dye plants. Also the leaves are
Hopea. This is also the condition in all species of used as a yellow dye or mordant, as in S. cochinch-
subg. Symplocos, both in the Old and New World. inensis ssp. laurina var. laurina and S. Iucida. In
448 H.P. Nooteboom
A
c
Fig. 128. Symplocaceae. Symplocos ophirensis var. cumin- when drying, and straight embryo; about 130 spp.,
giana. A Flowering twig. B Flowers. C Fruit. D Fruit stone. E mostly New World; subg. Hopea, with petals con-
Same in transverse section. F Fruit, vertical section. me =
mesocarp; ec = endocarp; sc = seed cavity. G Seed. H Embryo.
nate only at the base, leaves usually becoming
(Nooteboom 1975) yellow when drying, and embryo straight or
curved; ca. 160 spp., Old and New World.
western Java the inner bark of S. odoratissima var.
odoratissima, known as "kulit seriawan", is exhib-
ited in every drugstore. It is pulped and rubbed on Selected Bibliography
the gums to cure thrush. An infusion of the leaves
of this variety is used for the same purpose. All Airy Shaw, H.K. 1966. A dictionary of the flowering plants and
these uses seem to be based on the astringent ferns, by J.C. Willis, ed. 7, 1214 pp.
properties of the plant tissue. Anderberg, A.A. et al. 2002. See general references.
Most species do not reach a volume adequate references.
for timber; usually the wood is reported as soft Baas, P. 1973. The wood anatomical range in Ilex
and light and used for light construction, but in (Aquifoliaccae) and its ecological and phylogenetical
Vietnam the wood of S. adenophylla var. adeno- significance. Blumea 21: 193-258.
phylia is reportedly very hard and good for col- Barth, O.M. 1979. Pollen morphology of Brazilian Sy mplocos.
Grana 18: 99-107.
umns. In S America the leaves of several species Barth, O.M. 1982. The sporoderm of Brazilian Symplocos
are used for making mate (Cador 1900). pollen types (Symplocaceae). Grana 21: 65-69.
A single genus: Borgmann, E. 1964. Anteil der Polyploiden in der Flora des
Bismarckgebirges von Ostneuguinea. Zeitschr. Bot. 52:
134.
Symplocos Jacq. Fig. 128 Brand, A. 1901. Symplocaceae. In: Engler, A. (ed.) Das
Pflanzenreich IV. 242. Leipzig: W. Engelmann.
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Leiden Bot. Ser. 1 (1 975), rev. Old World; Stahl, Fl. Ecuador Arten. Bot. Centralbl. 2184: 248- 251,345,369-371.
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49: 369-388 (1994), Bolivian spp., 51: 323-364 (1996), Vegetabilis 8. Paris: Treutel et WUrtz, 684 pp.
Colombian spp. Caris, P., Ronse Decraene, L.P., Smets, E., Clinckemaillie, D.
2001. The uncertain systematic position of Symplocos (Sym-
Description as for family. About 300 spp., sub- plocaceae): evidence from a floral ontogenetic study. Int. J.
Plant Sci. 163: 67-74.
tropical and tropical regions except Africa. Two
Chenery, E.M. 1948a. Aluminium in the plant world. Kew Bull.:
subgenera are distinguished by Nooteboom ( 1975): 173- 183.
subg. Symplocos, with petals connate at least Chenery, E.M. 1948b. Thioglycolic acid as an inhibitor for iron.
halfway up, leaves usually not becoming yellow Analyst: 501.
Symplocaceae 449
Chirtoiii, M. 1918. Lacistemacees et Symplocacees. Bull. Soc. Maguire, B., Huang, Y.C. 1978. Symplocaceae. In: Maguire, B.
Bot. Geneve XI, 10: 350-361. et al., The botany of the Guayana Highlands 10. Mem. New
Corner, E.J.H. 1976. See general references. York Bot. Gard. 29: 223-230.
Davis, G.L. 1966. See general references. Mehra, P.N., Bawa, K.S. 1969. Chromosomal evolution in trop-
Endlicher, S.L. 1839. Genera plantarum: 744. Vienna: Fr. Beck. ical hardwoods. Evolution 23: 466-481.
Erdtman, G. 1952. See general references. Mehra, P.N., Gill, B.S. 1968. IOPB chromosome number reports
Gupta, H.P., Sharma, C. 1977. Palynotaxonomy and phylogeny XIX. Taxon 17: 575.
of Indian Symplocaceae and Sapotaceae. Geophytology 7: Meijden, R. Vander, 1970. A survey of the pollen morphology
147-159. of the Indo-Pacific species of Symplocos (Symplocaceae).
Hallier, H. 1923. Beitrage zur Kenntnis der Linaceae. Beih. Bot. Pollen Spores 12: 513-551.
Centralbl. 39B b: 1-178. Metcalfe, C.R., Chalk, L. 1950. See general references.
Handel-Mazzetti, H., Peter-Stibal, E. 1943. Eine Revision der Muller, J. 1981. Fossil pollen records of extant angiosperms.
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Centralbl. 62B b: 1-42. Nagamasu, H. 1989. Pollen morphology of Japanese Symplocos
Hardin, J.W. 1966. An analysis of variation in Symplocos tinc- (Symplocaceae). Bot. Mag. Tokyo 102: 149-164.
toria. J. Elisha Mitchell Sci. Soc. 82: 6-12. Nakai, T.1924.Abstract from T. Nakai: trees and shrubs indige-
Hegnauer, R. 1973, 1990. See general references. nous in Japan proper, vol. 1 (1922), with additional remarks
Huang, T.C. 1967. Pollen grains of Formosan plants 2. on some species. Bot. Mag. Tokyo 38: 23-48.
Taiwania 13: 15-110. Neger, F.W. 1923. Neue Methoden und Ergebnisse der
Huang, T.C. 1972. Pollen flora of Taiwan. Taipei: National Mikrochemie der Pflanzen. Flora N.F. 16: 26-330.
Taiwan University. Nevling, L.I. Jr. 1969. The ecology of an elfin forest in Puerto
Huber, H. 1963. Die Verwandtschaftsverhaltnisse der Rico. 5. Chromosome numbers of some flowering plants.
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Hutchinson, J. 1959. The families of flowering plants, ed. 2, 1. Nooteboom, H.P. 1975. Revision of the Symplocaceae of the
Oxford: Clarendnon Press, 510 pp. Old World, New Caledonia excepted. Leiden Bot. Ser. 1.
Janssonius, H.H. 1920. Mikrographie des Holzes. 4. Leiden: Universitaire Pers.
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Keng, H. 1962. Comparative morphological studies in Oever, L. van den, Baas, P., Zandee, M. et al. 1981. Comparative
Theaceae. Univ. Calif. Publ. Bot. 33: 269-384. wood anatomy of Symplocos and latitude and altitude. IAWA
Kircheimer, F. 1949. Die Symplocaceae der erdgeschichtlichen Bull. n.s. 2: 3-24.
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Verbreitung des Aluminiums im Pflanzenreich. Sitz. Ber. Stahl, B. 1995. Diversity and distribution of Andean Symplo-
Akad. Wiss. Wien: 311-336. caceae. In: Churchill, S.P., Balslev, H., Forero, H., Luteyn,
Krutzsch, W. 1989. Paleogeography and historical plant geog- J.L. (eds.) Biodiversity and conservation of neotropical
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Liang, Y.H., Yu, C.H. 1985. Pollen morphology of Styracaceae Takhtajan, A.L. 1959. Die Evolution der Angiospermae. Jena: G.
and its taxonomic significance. Acta Phytotax. Sin. 23(2): Fischer.
81-90. Takhtajan, A.L. 1973. Evolution und Ausbreitung der
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Sapotaceae to Fabaceae. Pollen Spores 24: 331-368. Webb, L.J. 1954. Aluminium accumulation in the Australian
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Ber. Deutsch. Pharm. Ges. 6: 203. Wehnert, A. 1906. Anatomisch-systematische Untersuchungen
Lubbock, J. 1892. A contribution to our knowledge of seedl- der Gattung Symplocos. Thesis. Miinchen.
ings. London: 206-208.
450 A.L. Weitzman et a!.
Ternstroemiaceae
A.L. WEITZMAN, S. DRESSLER and P.F. STEVENS
The hairs are unicellular, although sometimes

Ternstroemiaceae Mirb. ex DC., Essai prop. med. pl., ed. 2: 203
(1816). aggregated into fascicles, as in some Freziera. In
Pentaphylacaceae Engler in Engler & Prantl, Nat. Pflanzenfam.,
leave_s of Adin~ndra the hair base may be large and
Nachtr. 1: 214 (1897), nom. cons.'
redd1sh, makmg the lower surface, at least dis-
tinctly punctate. However, not all hairs o~ the
E~e~green _trees or shrubs, accumulating alu-
leaf have such bases (e.g. A. clemensiae), and it is
mmmm; mdumentum of unicellular hairs.
unclear if the punctations on the leaves of all
Leaves distichous or spiral, conduplicate-involute
Ternstroemieae are comparable. Pentaphylax has
or supervolute, usually evergreen and coriaceous
perulate buds. Growth elsewhere in the family, as
margins entire or toothed, a small, deciduous set~
m Ternstroemia and Cleyera, is also clearly rhyth-
near the apex of each tooth; stipules absent.
mic, and the first leaves of the flush are small and
Plant dioecious or flowers hermaphroditic. Inflo-
fall off early. In many other genera, however, it is
rescences axillary, rarely pseudo-terminal, flowers
not easy to distinguish flushes of growth, at least
solitary or fasciculate, rarely in short racemes of
on herbarium material. The leaves are distichous
up to 15 flowers, pedicellate, regular, 5-merous,
to spiral, this varying infragenerically in Eurya
usually <2 em in diameter, pedicel with more or
(Barker 1982). In Ternstroemia and Anneslea the
less apical prophylls; sepals quincuncial, connate
expanded leaves are congested at the end of
or distinct, persistent in fruit, usually thick, con-
each innovation. The lamina margin is toothed to
cave, subequal, rarely unequal; petals distinct
se_rr~te, but t~is varies within genera, and perhaps
or connate basally, quincuncial, thick or mem-
w1thm a spec1es (Cleyera japonica). The teeth are
branous; stamens 5-many, free, connate and/or
glandular, black to red in colour and tend to be
adnate t? corolla basally, filaments equal or un-
deciduous; they are sometimes described as setae
equal, w1de and flat or thickened, often narrow-
(Hickey 1979). They are borne near the apex of the
ing and incurved apically, shorter than to 4x as
teeth. Interestingly, although the leaf blades of
long as the anthers; anthers dithecal basifixed
Ternstroemia are entire to crenulate, seedlings of
introrse, oblong, opening by longitudinal slits:
T. elongata are drawn with strongly serrate leaves
rarely by flaps or pore-like slits, connective often
(d~ Vogel 1980). Venation is commonly bro-
elongated; carpels (2)3-5(6), ovary superior,
chldodromous, sometimes reticulodromous. The
rar~ly inferior, placentation axile, rarely apical or
leaves are usually petiolate, but frequently only
panetal; ovules anatropous to campylotropous, (1,
very. shortly so or even sessile in genera such as
2)-many per carpel; style simple or with apical
Frezzera and Archboldiodendron. Colleters are
branches, or. stylodia separate; stigmas usually
known in some genera, e.g. Freziera.
separate. Frmt a berry, or rarely irregularly dehis-
. Leaf sclerification can be extensive (Keng 1962).
cent, or a loculicidal capsule; seeds 1-numerous
Smgl_e cells or cluste:s of palisade cells may be
re~ifor_m, small, reticulate or areolate, or larger: sclenfied, and may m turn be associated with
elhpsmd, ± smooth; endosperm slight to copious;
sclerified hypodermal cells. In addition, single or
embryo hippocrepiform to ± straight.
clustered, more or less isodiametric, angular to
A family of about 12 genera and about 340
shortly-branched sclereids are usually present in
species; worldwide in tropical and subtropical
the spongy parenchyma; they are absent in
regions, mostly in Asia and the neotropics, few in
Balthasaria and Pentaphylax. Both palisade scle-
Africa.
reids and isodiametric sclereids may be densely
clustered around veins or scattered evenly
VEGETATIVE MORPHOLOGY. Ternstroemiaceae
throughout the leaf and, in the dried leaf, they are
are evergreen shrubs to usually rather small trees.
often visible as minute "papillae". Ternstroemieae
have much-branched sclereids, whilst those of
1 See nomenclatural note below under "Affinities". Freziereae are less branched to rounded (excep-
Ternstroemiaceae 451
tions occur in Adinandra and Cleyera). On the the Ternstroemieae there is only a single flower
upper surface of the lamina there may be one or per axil of a reduced, non-photosynthetic leaf (but
two hypodermal layers of almost isodiametric Anneslea donnaiensis may have flowers in the axils
cells. Mucilage has been reported in both upper of photosynthetic leaves). In Pentaphylax such
and lower epidermes (Keng 1962). The palisade flowers are borne at the beginning of the innova-
tissue consists of a variable number of layers. tion (Fig. 129A), and ordinary expanded leaves
Stomata are present only on the lower surface may never be produced later, so the inflorescence
and are usually anomocytic (Keng 1962). In some appears to be a raceme. In Freziereae flowers are
genera they tend to be cyclocytic (Kva ek and either single or fasciculate (reduced axillary
Walther 1984), and in Freziera there may be a con- shoots) in the axils of expanded and sometimes
siderable difference in size between subsidiary also reduced leaves (Fig. 131A). The flowers may
cells and epidermal cells (Weitzman 1987b); also be single in the axils of leaves of more or less
Pentaphylax has paracytic stomata. In general, developed shoots arising from the current innova-
Ternstroemieae and Freziereae show little epider- tion; almost all this variation can be seen within
mal variation (Kva ek and Walther 1984). Calcium a single individual of Cleyera japonica (see
oxalate druses are found throughout the plant, but also Keng 1962). The pedicels have two more or
are scanty in Pentaphylax. less apical prophylls; these may be persistent or
Nodes in most Ternstroemiaceae are unilacunar caducous.
with one trace (Beauvisage 1920; Keng 1962; Flowers are either hermaphroditic or unisexual,
Schofield 1968; Weitzman 1987b). The leaf trace although this latter condition may be difficult to
leaves the stele as a flattened arc; lateral leaf observe, as in Freziera (see below). The flowers are
bundles diverge from the ends of the trace, and pentamerous and radially symmetrical. Sepals and
there are from one to five bundles in the petiole. petals are opposite each other in Ternstroemieae.
However, in Archboldiodendron and a few species The sepals are quincuncial, persistent, and usually
of Freziera, the ends of the arc curve inwards adax- free, although in Balthasaria Verdcourt (1962)
ially; lateral leaf bundles diverge from the shoul- draws three sepals as being completely outside the
ders of the main bundle. Freziera, the only genus others; there is no overlap of any of the sepals in
studied in detail, shows further variation in petiole what are depicted as two whorls. In most taxa
anatomy, and trilacunar nodes also occur in the there is a small apical seta on the sepals similar to
genus (Weitzman 1987b). those found on the leaf margins (see above), and
these have been observed in Pentaphylax as well;
WooD ANATOMY. Growth rings are usually setae may also occur on the margins of the sepals.
indistinct. Vessels are usually solitary and vessel The corolla is quincuncial, and is more or less
elements are 910-2340j..tm long (measurements campanulate and fleshy; there may be some scle-
are species means), 30-95( -62) jlm in tangential reids. In Freziera (and some other taxa, as in Eurya
diameter, sometimes with spiral thickening, and sandwicensis) it is urceolate and very highly scle-
with scalariform to opposite intervessel pits. rified (Weitzman 1987b ). The petals vary from free
Balthasaria and some species of Eurya have par- to more or less fused. Archboldiodendron alone
ticularly wide vessels. Perforation plates are scalar- has a biseriate corolla usually with ten petals (Fig.
iform, with (10-)20-55(-108) bars. Vessel/ray pits 130), although there may be as few as six, members
are half to fully bordered (note that the pits in of the inner series being absent (Kobuski 1940).
Theaceae lack a border or have only reduced Stamens are usually indefinite in number,
borders, Liang and Baas 1990). Fibre tracheids although in Pentaphylax and some Eurya there
are 1440-2930 jlm long. Parenchyma is apotra- are only five, alternating with the petals. The fila-
cheal diffuse to paratracheal. Prismatic crystals ments are commonly slightly connate at the base
are occasional. Rays are 1-10 cells across, and are and slightly adnate to the petals; those of Penta-
often of two different size classes; they are Kribs phylax are very broad and closely adpressed to the
types heterogeneous I-Ill. The rays in Adinandra corolla, forming a tube. The stamens are whorled,
and especially Cleyera are narrow, usually one cell although this can be difficult to see when they are
across (information is taken from Keng 1962; connate, as in Adinandra (Barker 1980). In Visnea
Baretta-Kuipers 1976; Carlquist 1984; Liang and there are antepetalous stamen pairs (Payer 1857;
Baas 1990, 1991). Corner 1946; Ronse Decraene and Smets 1996).
Although the family is sometimes characterised as
INFLORESCENCE AND FLOWER STRUCTURE. Inflo- having filaments at most only slightly longer than
rescences are always axillary. In Pentaphylax and the anthers (Judd in Schoenenberger and Friis
2001), in Freziereae such as Cleyera they are ap- (Corner 1976) and Pentaphylax are apotropous,
preciably longer (see also de Wit 1947). The those of Symplocarpon epitropous. Orientation
anthers are quite often hairy, and the connective is in Eurya, Freziera and Cleyera is variable.
often more or less produced, and it is especially The micropyle is endostoma! and the embryo sac
conspicuous in Anneslea. Anthers of Pentaphylax is eight-celled, its formation being of the Poly-
open by flaps near the apex of the thecae, not gonum type.
simply by pores (Cronquist 1981; Takhtajan 1997;
cf., amongst others, Gardner 1849; van Steenis POLLEN MORPHOLOGY. Pollen grains are spher-
1955). oidal to prolate, tricolporate and tectate; exine
The gynoecium is generally superior, although sculpturing is inconspicuous, varying from psilate
inferior in Anneslea and Symplococarpon, and to scab rate, rugulate, foveolate or rugose (Erdtman
apparently inferior in Visnea, partly because the 1952; Keng 1962). The exine of Euryodendron is
connate sepals closely invest the ovary in fruit. indistinctly rugose and sparsely and minutely
Carpel number is variable, although commonly perforate at the polar regions (Ying et al. 1993).
three (De Wit 1947), and placentation varies Weitzman (1987b) found Freziera pollen to be
from axile (the usual condition) to parietal, as in very small (9-20 !liD, usually about 10 !liD long);
Cleyera. Ovule number varies considerably; when that of Balthasaria is rather large (up to 28.51..lm
there are only a few, they are apical. There is either long). The maximum reported is an equatorial
a style with a stigma with as many radii as carpels, diameter of 37.81..lm in Ternstroemia mokof (Lee
as in Pentaphylax, or there may be as many style 1987). Pentaphylax pollen is 14-16.51..lm and
branches or stylodia as carpels. Infrageneric vari- smooth, with thin tectum, poorly developed col-
ation in this feature is common. The stigma is umellae and a thick endexine (Wei et al. 1999).
papillate.
KARYOLOGY. Haploid numbers in Ternstroemia
FLORAL ANATOMY. There are sclereids in the are 20 or 25, in Freziereae they are 12-13?, 15, 18,
sepals, petals and ovary. Crystal druses in anthers 21 (the commonest number), 22, 23, etc., suggest-
are reported (Tsou 1995). In Cleyera, sepals, petals ing polyploidy. However, the family is poorly
and stamens all have single traces. A single dorsal known cytologically.
trace and two ventral traces supply each carpel, the
two ventral traces proceeding up the hollow style FRUIT AND SEED. Pentaphylax has loculicidal
(the style is also hollow in Adinandra and Penta- capsules and apparently wind-dispersed seeds.
phylax). Symplocarpon and Anneslea, with inferior Ternstroemia, and probably Anneslea, have irregu-
ovaries, as well as Visnea, have recurrent vascular larly dehiscent fruits whose rupture may be caused
bundles from which the individual carpel traces by the expansion of the ripening seeds; when
diverge. This suggests that the ovary is immersed open, the seeds dangle and "the brilliant sarcotesta
in receptacular or axial tissue (Keng 1962). Devel- [is displayed] to birds" (Barker 1980: 23). The fruit
opment of the androecium appears to be centrifu- of Freziereae is usually a berry. A few species of
gal in Adinandra (Corner 1976). Freziera have drupes, with as many few-seeded
pyrenes as there are carpels, while in Visnea the
EMBRYOLOGY. Detailed anther and ovule devel- dry, indehiscent fruit is enclosed by the accrescent,
opment is known from only eight species of Adi- more or less fleshy calyx. The fruits of Freziereae
nandra, Cleyera and Eurya (Tsou 1995), although are probably eaten by birds or sometimes bats, as
Corner (1976) includes some information on in Adinandra (Ridley 1930, cit. in Barker 1980).
ovule morphology. Anther wall development is Seed size and number varies considerably. The
dicotyledonous, and the anther epidermis is hea- fruits of Symplococarpon have a single seed ca.
vily tanniniferous. Tapetal cells are 2-(Eurya) 5 mm long, while fruits of Balthasaria have hun-
or 4-nucleate (Cleyera, Adinandra). Microspore dreds of seeds ca. 1 mm long. Ternstroemia and
initiation is simultaneous, and tetrahedral tetrads Anneslea have fruits with rather few and large
are usually produced. Ovules are amphitropous or seeds (2-)5-10 mm long.
campylotropous, and the ovule epidermis is more Testa structure is variable. The seeds of
or less tanniniferous. Ovules are tenuinucellate Ternstroemieae dry reddish and are reported to be
and bitegmic, the integuments being three cells arillate (e.g. Keng 1962). However, in both
thick. However, there is a suggestion that Penta- Anneslea and Ternstroemia there is a sarcotesta,
phylax is crassinucellate (Mauritzon 1936). The and both have stout, multicellular, unthickened
apical ovules of Visnea, the Ternstroemieae hairs. Anneslea has a sarcotesta, then a layer of
thin-walled crystalliferous cells, and then a scle- know, dioecious, androdioecious, or perhaps,
rotic meso testa to 10 cells thick; other testal and hermaphroditic. In most instances, three different
tegmic cells are thin-walled. In T. lowii, the only specimens are necessary for a complete under-
species for which there is detailed information, standing of a single taxon" (see also Kobuski e.g.
there are pockets of"watery tissue with large cells" 1942a, 1942b, 1943, 1963). Eurya is dioecious.
(Corner 1976: 506) on either side of the seed. The Freziera appears to be gynodioecious; however, all
epidermal cells are small, the crystalline hypoder- species for which there are sufficient data are func-
mal cells are thickened, and again there is a tionally dioecious (Weitzman 1987a, 1987b).
well-developed sclerotic mesotesta. The testa of The gynoecium in staminate flowers of Tern-
Freziereae and Pentaphylaceae is reticulate and stroemia may be absent or represented by a with-
black to brown in colour. In at least some ered and clearly non-functional ovary with styles
Freziereae, the seeds are embedded in a fleshy (and rarely stigmas). Although the gynoecium in
placenta. Cleyera japonica has much enlarged staminate flowers of Freziera appears to be func-
exotestal cells, but C. theoides has very enlarged tional, such flowers nearly always fall off soon after
and thickened mesotestal cells with mucilaginous anthesis. The anthers of staminate flowers may
walls (Keng 1962). Other Freziereae have a thick- open in bud (Weitzman 1987b). In both genera,
walled mesotesta which is one (Adinandra), two staminodia in carpellate flowers look very much
(Eurya) or 4-5 (Visnea) cells thick, the cells con- like filaments without anthers. Staminate flowers
taining crystals; the last two genera have enlarged of Eurya lack or have only a very rudimentary
exotestal cells with more or less massive, U-shaped gynoecium, while carpellate flowers usually lack
thickenings, while the exotesta of Adinandra is not staminodes.
notably thickened (Corner 1976). The inner walls The corolla is greenish to yellowish and the
of the exotestal cells of Pentaphylax are somewhat flowers are fairly small, usually less than 1.5 em
thickened, but underlying cells are thin-walled. across, but substantially larger in Ternstroemieae
The embryo occupies the full length of the seed and especially in Balthasaria, where they are up to
cavity, or almost so; although Takhtajan (1997) 5 em long, yellow-orange in colour, and perhaps
described the embryo of Visnea as minute, this even pollinated by birds. Generally, pollination is
is belied by the illustration in Corner (1976). The probably by bees or other insects, but little is known
embryo is curved back on itself and hippocrepi- in detail. Vibrational ("buzz") pollination has been
form in Ternstroemieae and Pentaphylaceae. In reported in two species of Ternstroemia (Bittrich et
Freziereae like Freziera, it is slightly curved, al- al. 1993). Although the family is supposed to lack
though it is hippocrepiform in Freziera dudleyi. nectaries, there is a presumably nectariferous ring
Although the seed is clearly U-shaped in Arch- around the base of the ovary in Pentaphylax,
boldiodendron and Eurya, the embryo itself devel- flowers of Cleyera japonica secrete substantial
ops in only one arm of the "U", and hence is only amounts of nectar within the staminal ring, and
slightly curved (Barker 1980), while in Balthasaria other species apparently have a nectary at the base
the seed and embryo are strongly curved, but of the ovary, while Symplococarpon is described as
are not as narrowly U-shaped as is common having a disc on top of the inferior ovary. In genera
elsewhere. like Ternstroemia and Adinandra, the stamens
Germination of Cleyera is epigeal, but the radicle closely surround the ovary, and one may suppose
soon stops growing; Adinandra also has epigeal that any nectar would be hard to obtain.
germination, but there the behaviour of the radicle
was not mentioned (Keng 1962). Ternstroemia has PHYTOCHEMISTRY. Common proanthocyanini-
epigeal germination and a well-developed radicle dins, flavonols and ellagic acid are widespread,
(de Vogel1980); in T. bancana the cotyledons seem myricetin is rare; saponins are present, and large
to be short-lived (Corner 1976). amounts of triterpenic acids (betulinic acid) are
accumulated in the bark of Eurya and Ternstroemia
REPRODUCTIVE SYSTEMS AND POLLINATION. spp. (the latter compounds are not known from
Ternstroemia is reported to be basically dioecious. Theaceae). Accumulation of aluminium and fluo-
However, the situation needs more study. Barker ride is general (Hegnauer 1973, 1990).
(1980: 14) qualified his description of its breed-
ing system "?rarely monoecious, sometimes SUBDIVISION AND RELATIONSHIPS WITHIN THE
andro-dioecious or andro-monoecious ... :'while FAMILY. Ternstroemiaceae as circumscribed
Kobuski (1961: 263) noted of the Philippine here are divided into three tribes, Pentaphylaceae,
species "the genus in the Philippines is, as far as I Ternstroemieae and Freziereae.
454 A.L. Weitzman et al.
Pentaphylaceae are the only Ternstroemiaceae AFFINITIES. For details of the relationships of
to have regularly capsular fruits; these are loculi- Theaceae s.l., in which Ternstroemiaceae have
cidal and probably plesiomorphic, as is the ap- often been included, see the account of the former.
parent absence of sclereids. The inflorescence of Molecular data now place Theaceae s.l. in the
Pentaphylax is basically the same as that of asterids-Ericales (e.g. Morton et al. 1996). What
Ternstroemieae; the cymose inflorescence of then are the proper limits of Theaceae?
Sladeniaceae has little in common with either. The Within the old Theaceae, Ternstroemia and its
curved embryo of Pentaphylax also agrees with its relatives, whether or not including Sladenia, have
position in Ternstroemiaceae. long been considered separable from Camellia and
Ternstroemieae and Freziereae are linked by its relatives, at least at some level. Pentaphylax has
their foliar sclereids. Ternstroemieae are morpho- often not been considered part of Theaceae s.l.
logically very distinct in their pseudoverticillate (Mattfeld 1942; Keng 1962; Liang and Baas 1990,
leaves, short filaments, sepals opposite the petals, 1991; Takhtajan 1997; see van Steenis 1955 for a
fruit type, etc. Anneslea is very like Ternstroemia, summary). Its anthers are superficially like those
and it can be placed in a separate tribe only if of Diapensiaceae, placed in Theales (Takhtajan
the significance of its inferior ovary is over- 1997); Pentaphylax and Theaceae are superficially
emphasized. similar, and both Theaceae and Diapensiaceae
Freziereae are distinct enough because of their are included in Ericales here. The seed is Ericalean
often short sclereids and baccate fruit, but generic (Huber 1991), although the elaborated mesotesta
limits in the tribe are very difficult. Bentham is unlike that of Ericaceae, etc. There have been
(1861: 55) noted that the then recognized genera suggestions that Pentaphylacaceae go with
of Ternstroemioideae were "so closely connected Balsaminaceae, etc., also in Ericales (Nandi et al.
with each other that their distinct separation by 1998). Wei et al. (1999) compared the pollen of
positive characters is very difficult:' Several genera Pentaphylax with that of Clematoclethra (Actini-
have usually been considered to be separate for diaceae) - again, Ericales. Although Pentaphylax
150 years, yet as genera they are often distin- was associated with Cardiopteridaceae and Gono-
guishable only by those who are very familiar with caryum in Savolainen, Fay et al. (2000), the latter
the individual species. The tribe has often been two are strongly associated with Aquifoliales in
divided into two groups centred on Adinandra and other analyses (Soltis et al. 2000; Karehed 2001).
Eurya. The character most frequently used to sep- Recent studies confirm that Theaceae s.l. can
arate them has been stylar fusion, although from easily be separated into two morphologically quite
the descriptions it can be seen that it commonly distinct groups, Theaceae s.str. and Ternstroemi-
varies even within genera. Archboldiodendron, aceae plus Pentaphylax and Sladeniaceae. There is
which has separate stylodia, was segregated from no evidence that Theaceae s.l. are monophyletic,
Adinandra which has a single style (Kobuski but both of these groups are monophyletic (Prince
1940), yet no mention was made of other members and Parks 2001; Anderberg et al. 2002), forming
of the tribe like Eurya which have separate stylo- two smallish clades in a major polytomy in
dia. On the other hand, Freziera and Cleyera Ericales.
were segregated from Eurya largely on the basis What are the limits of Ternstroemiaceae?
of their single style! Symplococarpon has free sty- Sladeniaceae (Sladenia and Ficalhoa) are a mono-
lodia and was segregated from Cleyera, which has phyletic group that differs from Ternstroemiaceae
a single style - but the former also has an inferior + Pentaphylax in such features as stem anatomy
ovary. (deep-seated versus surficial phellogen), inflores-
Some genera of the Freziereae may not be mono- cence type (cymose versus racemose inflorescence
phyletic, but our understanding of the basic mor- or single axillary flowers; see above) and embryo
phological variation of the group is poor, type (straight versus U-shaped). These and other
Balthasaria and Euryodendron being particularly differences allow the two to be readily distin-
little known. Archboldiodendron, Visnea and guished. Pentaphylax and the other Ternstroemi-
Symplococarpon are small genera, but are highly aceae have similar anatomy, inflorescence and
derived and consequently difficult to place. seeds. The only problem in combining the two is
There may be relationships between Adinandra nomenclatural; the name Pentaphylacaceae is
and Asian species of Cleyera, between Neotropical conserved. Conservation of Ternstroemiaceae will
species of Cleyera and Euryodendron, and between be proposed. The name Pentaphylaceae used
Freziera and Ternstroemiopsis (here treated as a by Hallier (1923: pp. 133, 176) is invalid, lacking a
Hawaiian species of Eurya). description or reference to one.
DISTRIBUTION AND HABITATS. Ternstroemiaceae KEY TO THE GENERA

prefer warm temperate to subtropical conditions,
1. Stamens 5; fruit a loculicidally dehiscent capsule
being commoner in tropical colline and montane 1. Pentaphylax
habitats than in the lowlands. Some species of - Stamens {5-)15-many; fruit a berry or irregularly dehis-
Eurya at times almost dominate subalpine habitats cent, rarely leathery capsule 2
in New Guinea, although other species grow in the 2. Leaves pseudoverticillate; flowers solitary; seeds often 4
mm long, with reddish sarcotesta 3
lowlands. Species of Eurya and Freziera in partic- - Leaves scattered; flowers usually in axillary fascicles of 2 or
ular do well in disturbed habitats. more; seeds usually <4 mm long, with brown or black retic-
ulate testa 4
PALAEOBOTANY. Paradinandra was described 3. Flowers borne below the leaves; ovary superior; petals
(SchOnenberger and Friis 2001) from flowers from nearly free 2. Ternstroemia
- Flowers borne above the leaves; ovary inferior ; petals often
the Late Cretaceous of southern Sweden. Although connate in their basal half 3. Anneslea
they did not place it in a family, they compared it 4. Petals {6-9)10 6. Archboldiodendron
with Ternstroemiaceae, specifically Cleyera, but - Petals 5 5
their comparison was hindered by the prevailing 5. Corolla tubular, at least 3 em long; style longer than 2 em
belief that Ternstroemiaceae lack nectar and 12. Balthasaria
- Corolla urceolate, campanulate, rarely tubular, up to 1.5 em
have very short filaments, while Paradinandra had long; style shorter than 2 em long 6
relatively longer filaments and an intrastaminal 6. Ovary 2-locular, inferior; calyx on top of fruit
nectary (see above, both are features of Cleyera). 9.Symplococarpon
A perhaps more distinctive feature is its tricolpate - Ovary (2)3-5-locular, superior; fruit base± surrounded by
pollen, an unusual feature for Ternstroemiaceae. the calyx 7
7. Sepals ca. 1 em long, half connate (ovary appears to be infe-
A fossil flower and leaf fragments of Pentaphylax rior), enclosing the woody indehiscent fruit 11. Visnea
were described from Baltic amber (Conwentz - Sepals often shorter, free, persisting at the base of the
1886). Mai (1971) and Knobloch and Mai (1986) baccate fruit 8
described fossil fruits and seeds from the Late 8. Ovaries 2-3-locular; ovules 8-16; filaments more than
Cretaceous of central Europe as Allericarpus and twice as long as the anthers 5. Cleyera
- Ovaries 3-5-locular, ovules 20-100; filaments usually up to
Pentaphylax resp. Eurya, Visnea and Protovisnea, twice as long as anthers 9
although confirmation is needed; both Theaceae 9. Staminate flowers with clearly non-functional gynoecium;
and Ternstroemiaceae appear to have been com- carpellate flowers often lacking staminodia; styles
mon in the lauraceous forests that were then separate 7. Eurya
- All flowers with apparently functional gynoecia; carpellate
widespread (summary in Grote and Dilcher 1989). flowers (when recognisable) with staminodia; style single,
Cleyera seed is known from the Middle Tertiary divided or not 10
of Vermont (Tiffney 1994). Wijninga and Kuhry 10. Leaves spiral; flowers with style about the length of the
(1990) and Wijninga (1996) report seeds perhaps ovary, ca 3 mm long 10. Euryodendron
of Freziera from the Pliocene of Colombia near - Leaves distichous, if spiral, styles apically free 11
11. Corolla campanulate; style longer than ovary, to 1.5 em
Bogota. Ternstroemia-like leaves are widespread long; leaves entire or serrate 4. Adinandra
in the northern hemisphere, but such leaves (in - Corolla urceolate; style shorter than ovary, ca. 2 mm long;
the form genus Ternstroemites; also the form leaves serrate 8. Freziera
genera Ternstroemiacinium and Ternstroemioxy-
lon) resemble genera in both Theaceae and Basal condition for the family
Ternstroemiaceae (Grote and Dilcher 1989).
Growth rhythmic; hairs unicellular; leaves exstip-
EcoNOMIC IMPORTANCE. Species like Eurya ulate; flowers 5-merous, regular, ± campanulate;
japonica, Eurya chinensis, Ternstroemia japonica sepals 5, free, quincuncial; petals 5, connate
and Cleyera japonica are used as ornamentals. basally, quincuncial; placentation axile; embryo
long, U-shaped, endosperm+.
CONSPECTUS OF TERNSTROEMIACEAE
I. Tribe Pentaphylaceae
Genus 1 1. TRIBE PENTAPHYLACEAE P.F. Stevens &
II. Tribe Ternstroemieae A.L. Weitzman, trib. nov.
Genera 2-3
III. Tribe Freziereae Pentaphylacaceae Engler in Engler & Prantl, Nat. Pflanzenfam.,
Genera 4-12 Nachtr. 1: 214 {1897).
Leaves scattered; flowers single, in the axils of

reduced leaves; stamens 5, opposite the sepals.
456 A.L. Weitzman et al.
lt
Ternstroemieae + Freziereae
Stamens (5-)<8; fruit a berry; radicle longer than
cotyledons, cotyledons incumbent.
~ 2. TRIBE TERNSTROEMIEAE DC. (1822).
Leaves pseudoverticillate, often with black dots

on the lower surface; sclereids usually much
branched. Flowers single, in the axils of non-
expanded leaves, rather large (usually >1.5 em
across); sepals opposite petals, filaments shorter
than the anthers; ovules pendulous. Fruit irregu-
larly dehiscent; seeds usually >4 mm long, reddish-
brown; testa fleshy.
2. Ternstroemia Mutis ex L. f.
G Ternstroemia Mutis ex L. f., Suppl. Pl.: 39, 264 (1782), nom.
cons.; Kobuski, J. Arnold Arbor. 23: 298-343,464-478 (1942),
24: 60-76 (1943), 42: 81-86, 263-275, 426-429 (1961), 44:
421-435 (1963); Barker, Brunonia 1: 14-42 (1980); Hung T.
Chang & Kuan, Higher plants of China 4: 620-624 (2000).
Shrubs or trees. Leaf margin entire, rarely crenu-

late, dots on lower surface often lacking. Flowers
borne below expanded leaves, prophylls deciduous
or persistent; petals at most connate basally; sta-
mens many, 3-seriate; anthers retuse to apiculate,
base hardly raised; ovary 1-3-locular, or 4-6-
locular with false septae, 1-20 ovules per loculus,
Fig. 129. Ternstroemiaceae. Pentaphylax euryoides. A Flower- stylodia or style +. Endosperm sometimes sparse.
ing branch. B Flower. C Stamens. D Dehisced fruit. E Young Ca. 100 spp., Sri Lanka to SE and E Asia, tropical
fruit. F Pistil. G Seed. Drawn by R. van Crevel. (van Steenis and subtropical Americas (most species), Africa
1955)
(ca. 2 species).
Fruit a loculicidal capsule, the midribs of the
carpels forming long "teeth" between the valves, 3. Anneslea Wall.
columella persistent; cotyledons longer than Anneslea Wall., Pl. Asiat. Rar. 1: 5, pl. 5 (1829), nom. cons.;
radicle. Kobuski, J. Arnold Arbor. 33: 79-90 (1952).
Paranneslea Gagnep.
1. Pentaphylax Gardner & Champ. Fig. 129 Trees. Leaf margin crenulate-serrulate. Flowers
Pentaphylax Gardner & Champ., Hooker's J. Bot. Kew Gard. borne above expanded leaves; prophylls (sub)per-
Misc. 1: 244 (1849); Mattfeld in Engler & Prantl, Nat. sistent; petals connate half way; stamens many,
Pflanzenfam. ed. 2, 20b: 13-21 (1942); van Steenis, Fl. Males. 1-2-seriate; anther base cordate, apiculus very
I, 5: 121-124 (1955).
long; ovary 2-3-locular, inferior, 2-10 ovules per
loculus, style 2-5-fid at apex. Three spp., China, SE
Trees. Leaves spiral, entire. Flowers sub sessile; pro-
Asia to Sumatra.
phylls persistent; petals emarginate; stamens 5, fil-
aments very broad, anthers incurved, dehiscing
by two valves; ovary 5-locular, nectary basal, 2
3. TRIBE FREZIEREAE DC. (1822).
apical pendulous ovules per loculus, style+. Seed
reticulate. One sp., P. euryoides Gardner & Champ.,
Growth rarely obviously rhythmic; leaves scat-
China (Kwangtung and Hainan) to Sumatra,
tered, sclereids usually only slightly branched;
scattered.
flowers 1 or more together, at least sometimes in
the axils of expanded leaves; placentae median

(apical); seeds <4( -6) mm long, brown or black.
4. Adinandra Jack
Adinandra Jack, Malayan Misc. 2(7): 49 (1822); Kobuski, J.
Arnold Arbor. 28: 1-98 (1947); Hung T. Chang & Kuan,
Higher plants of China 4: 624-629 (2000).
Trees or shrubs. Leaves distichous, margins ser-

rate. Flowers 1-2 together; prophylls deciduous
to persistent; petals basally connate; stamens
15-many, I-S-seriate (in bundles}, more or less
connate, anthers apiculate, base ± rounded; ovary
(2}3(4}5-locular, many ovules per carpel; style
(stylodia) up to 15mm long. Embryo also J-
shaped. Ca. 80 spp.; E and SE Asia to Malesia.
5. Cleyera Thunb.
Cleyera Thunb., Nov. Gen. Pl. 3: 68 (1783), nom. cons., p.p.
emend. Sieb. & Zucc., Fl. Jap. 153, pl. 81 (1841); Kobuski,
J. Arnold Arbor. 18: 118-129 (1937), 22: 395-416 (1941),
Hung T. Chang & Kuan, Higher plants of China 4: 629- 633
tfJl
(2000).
Trees. Leaves distichous, margins entire to serrate.

Eu
Flowers hermaphroditic, 1-4 together; prophylls
deciduous; petals connate basally; stamens 25,
uniseriate; anthers opening by short gaping slits,
apiculate, base cuneate; ovary 2-3-locular, placen-
0
tation axile, but sometimes seemingly parietal by
long proliferating placentar strands, 5-l 0 ovules
Fig. 130. Ternstroemiaceae. Archboldiodendron calosericeum.
per loculus; style +, divided half way or not. Eight A Fruiting branch. B Flower. C Corolla spread open, with
spp., India to Japan, Mexico, Central America, stamens. D Immature fruit with persistent style and style
Caribbean. branches. E Seed, vertical section. Drawn by T. Iwagu. (Barker
1980)
6. Archboldiodendron Kobuski Fig. 130

Archboldiodendron Kobuski, J. Arnold Arbor. 21: 140 (1940);
Barker, Brunonia 3: 47-54 (1980).
Barker in van Royen, Alpine Fl. New Guinea 2: 1402-1454
(1982); Hung T. Chang & Kuan, Higher plants of China 4:
Trees. Leaves distichous, margins serrate, petioles 633- 655 (2000).
short. Flowers hermaphroditic, single; prophylls Ternstroemiopsis Urban
deciduous; petals (6-) 10, biseriate, basally con-
nate; stamens 30, uniseriate, anther apex acute, Trees or shrubs; leaves distichous, rarely spirally
base rounded; ovary 5( - 7}-locular, placentae T- arranged, margins serrate. Plant dioecious; flowers
shaped, many ovules per loculus, styles free, short. 1-8 together; prophylls persistent; petals connate
Embryo curved. One sp., A. calosericeum Kobuski, ca. 1/3-1/2, staminate flowers: stamens 5-25, unis-
montane New Guinea. eriate, anthers often locellate, apex apiculate to
obtuse, base rounded, rarely pistillode vestigial;
7. Eurya Thunb. pistillate flowers: staminodes rarely+, ovary 3( -5)-
locular, 4-many ovules per loculus, style bran-
Eurya Thunb., Nov. Gen. Pl. 67 (1783); Kobuski, J. Arnold ched. Embryo curved. 50- 100 spp., S China, SE
Arbor. 16: 347- 352 (1935), 25: 299- 359 (1938), 20: 361- 374
(1939); De Wit, Bull. Jard. Bot. Buit. III, 17: 329- 375 (1947);
Asia and nearby Pacific islands. The genus is badly
in need of revision.
9. Symplococarpon Airy-Shaw
Symplococarpon Airy-Shaw in Hooker's Icon. Pl. 34: pl. 3342
(1937); Kobuski, J. Arnold Arbor. 22: 188-196 (1941).
D
E
~ Trees. Leaves distichous, margins entire to
slightly serrate. Plant hermaphroditic; flowers
1-5 together; prophylls persistent; petals ± free;
stamens 25-40, uniseriate, filaments long, anthers
caudate apically, base cuneate; ovary 2-locular,
inferior, disk on top, 1(-2) pendulous ovules per
loculus, style free or stylodia +. Nine spp., Mexico,
Central America and NW South America.
10. Euryodendron Hung T. Chang

Euryodendron Hung T. Chang, Acta Sci. Nat. Univ. Sunyatseni
1963(4): 129 (1963); Ying et al., The endemic seed plants of
China: 693-695 (1993); Hung T. Chang & Kuan, Higher
plants of China 4: 633 (2000).
Trees. Leaves spiral, margins serrate. Flowers her-

maphroditic, 1-2 together, prophylls persistent;
petals ?free; stamens ca. 25, uniseriate, anthers
hairy, apex acute, base rounded; ovary 3-locular,
many ovules per loculus; style +. Embryo
unknown. One sp., E. excelsum Hung T. Chang,
Guangxi, China, low alt. The margins of the sepals
Fig. 131. Ternstroemiaceae. Freziera carinata. A Flowering are fimbriate.
branch. B Flower. C Petal of female flower, stamens adnate.
D Pistil of female flower. E Pistil and stamens of male flower.
F Fruit. G Seeds, side view and chalaza! view. Drawn by
11. Visnea L. f.
A.L. Weitzman. (Weitzman 1987a)
Visnea L. f., Suppl.: 36 (1782); Kobuski, J. Arnold Arbor. 33:
188-191 (1952).
8. Freziera Willd. Fig. 131 Small trees or shrubs. Leaves distichous, margins
Freziera Willd., Sp. Pl. 2(2): 1179 (1799) nom. cons.; Weitzman, serrate. Flowers hermaphroditic?, 1-3 together;
Systematics of Freziera Willd. (Theaceae), Ph.D. thesis, prophylls persistent; sepals half connate; petals
Harvard University (1987). connate basally; stamens 12( -21), uniseriate,
Patascoya Urb.
Killipiodendron Kobuski
anthers apiculate, base rounded; ovary (2-)3-
locular, 2-3 pendent ovules per loculus, stylodia
Small trees or shrubs. Leaves distichous, mar- free. Embryo J-shaped. One sp., V. mocanera L. f.,
gins serrate, petioles short. Plant functionally from cloud/laurel forests of the Canary Islands
dioecious; flowers 1-14 together; prophylls and Madeira.
usually persistent. Corolla urceolate, petals free or
slightly connate basally; staminate flowers: 12. Balthasaria Verde.
stamens (8- )15- 35(- 48), uniseriate, anthers apic-
Balthasaria Verde., Kew Bull. 23: 469 (1969).
ulate, ± rounded basally; pistillode +; carpellate Melchiora Kobuski, J. Arnold Arbor. 37: 154 (1956); Verde., Fl.
flowers: staminodes +; ovary (2-)3-5( -6)-locular, Trop. E. Afr. Theaceae: 3-6 {1962), non Melchioria Penz. &
(6-)many ovules per loculus, style short, with Sacc.
short lobes. Fruit a drupe. Embryo curved,
sometimes slightly, sometimes straight. 57 spp., S Large trees. Leaves spiral, margins serrate, petioles
Mexico and the West Indies through Central and short. Flowers hermaphroditic, single; prophylls
South America to S Bolivia, and E to the Guayana persistent; petals 3 em long, tubular, free; sta-
Highland; primarily montane. F. dudleyi Gentry mens 15-35, uniseriate, anthers long-apiculate,
has many characters anomalous in the genus. base rounded; ovary 4-5 locular, many ovules per
loculus, style >2cm long. Perhaps one sp., M. Karehed, J. 2001. Multiple origin of the tropical forest tree
schliebenii (Melchior) Verde., montane E Africa. family Icacinaceae. Am. J. Bot. 88: 2259-2274.
Keng, H. 1962. Comparative morphological studies in the
Theaceae. Univ. Calif. Pub!. Bot. 33: 269-384.
Knobloch, E., Mai, D.H. 1986. Monographie der Frtichte und
Selected Bibliography Samen in der Kreide von Mitteleuropa. Rozpr. Ustr. Ust.
Geol. 47: 1-219.
Kobuski, C.E. 1940. Studies in the Theaceae, V. The Theaceae
Anderberg A.A. et a!. 2002. See general references.
of New Guinea. J. Arnold Arbor. 21: 134-162.
Baretta-Kuipers, T. 1976. Comparative wood anatomy of
Kobuski, C.E. 1942a. Studies in Theaceae, XII. Notes on the
Bonnetiaceae, Theaceae and Guttiferae. Leiden Bot. Ser. 3:
South American species of Ternstroemia. J. Arnold Arbor.
76-101.
23: 298-343.
Bar~er, W.R.1980. Taxonomic revisions in Theaceae in Papua-
sia. I. Gordonia, Ternstroemia, Adinandra and Archboldio- Kobuski, C.E. 1942b. Studies in Theaceae, XIII. Notes on the
Mexican and Central American species of Ternstroemia.
dendron. Brunonia 3: 1-60.
Barker, W.R. 1982. Theaceae. In: van Royen, P., The Alpine J. Arnold Arbor. 23: 464-478.
Kobuski, C.E. 1943. Studies in Theaceae, XIV. Notes on the West
Flora of New Guinea. Vol. 3: Taxonomic part. Winteraceae
Indian species of Ternstroemia. J. Arnold Arbor. 24: 60-76.
to Polygonaceae. Vaduz: J. Cramer, pp. 1397-
Kobuski, C.E. 1961. Studies in Theaceae, XXXII. A review of the
1454.
genus Ternstroemia in the Philippine Islands. J. Arnold Arb.
Beauvisage, L. 1920. Contribution a I'etude anatomique de Ia
42: 263-275.
famille des Ternstroemiacees. Tours: Arrault.
Kobuski, C.E. 1963. Studies in Theaceae, XXXV. Two new
Bentham, G. 1861. Notes on Ternstroemiaceae. J. Proc. Linn.
species of Ternstroemia from the Lesser Antilles. J. Arnold
Soc. Bot. 5: 53-65.
Bittrich, V., Amaral, M.C.E., Melo, G.A.R. 1993. Pollination Arb. 44: 434-435.
Kva ek, Z., Walther, H. 1984. Nachweis tertiii.rer Theaceen
biology of Ternstroemia laevigata and T. dentata
Mitteleuropas nach blatt-epidermalen Untersuchungen.
(Theaceae). Pl. Syst. Evol. 185: 1-6.
Carlquist, S. 1984 (1985). Wood anatomy and relationships I. Teil - Epidermale Merkmalskomplexe rezenter Theaceae.
Feddes Repert. 95: 209-227.
of Pentaphylacaceae: significance of vessel features. Phyto-
Lee, S. 1987. A palynotaxonomic study on the Korean
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Conwentz, H. 1886. Die Angiospermen des Bernsteins. Danzig:
Liang, D., Baas, P. 1990. Wood anatomy of trees and shrubs
W. Engelmann.
from China II. Theaceae. I.A.W.A. Bull. II, 11: 337-378.
Corner, E.J.G.H. 1946. Centifugal stamens. J. Arnold Arbor. 27:
Liang, D., Baas, P. 1991. The wood anatomy of the Theaceae.
423-437.
I.A.W.A. Bull. II, 12: 333-353.
Corner, E.J.G.H. 1976. See general references.
Mai, D.H. 1971. Dber fossile Lauraceae und Theaceae in
Mitteleuropa. Feddes Repert. 82: 313-341.
De Vogel, E.F. 1980. Seedlings of dicotyledons. Wageningen:
Mattfeld, J. 1942. Pentaphylacaceae. In: Engler, A., Prantl, K.
Pudoc.
De Wit, H.C.D. 1947. A revision of the genus Eurya Thunb. Nat. Pfianzenfam., ed. 2, 20b. Leipzig: W. Engelmann, pp.
13-21.
(Theac.) in the Malay Archipelago (including New Guinea
and south of the Philippines). Bull. Jard. Bot. Buitenzorg, ser. Mauritzon, J. 1936. Zur Embryologie und systematischen
III, 17: 329-375. Abgrenzung der Reihen Terebinthales und Celastrales. Bot.
Not. (Lund) 1936: 161-212.
Melchior, H. 1925. Theaceae. In: Engler, A., Prantl, K. Nat.
Gardner, G.1849. Descriptions of some new genera and species
Pfianzenfam., ed. 2, 21. Leipzig: W. Engelmann, pp. 109-
of plants, collected in the island of Hong Kong by Capt. J. G.
Champion, 95'h Regt. Hooker's J. Bot. Kew Gard. Misc. 1: 154.
Metcalfe, C.R., Chalk, L. 1950. See general references.
240-246.
Morton, C.M. et a!. 1996. See general references.
Grote, P.J., Dilcher, D.L. 1989. Investigations of angio-
Nandi, O.I., Chase, M.W., Endress, P.K. 1998. A combined
sperms from the Eocene of North America: a new genus of
Theaceae based on fruit and seed remains. Bot. Gaz. 150: cladistic analysis of angiosperms using rbcL and non-
molecular data sets. Ann. Missouri Bot. Gard. 85: 137-212.
190-206.
Hallier, H.1923. Beitrii.ge zur Kenntnis der Linaceae (DC.1819) Payer, J.-B. 1857. Traite d'organogenie comparee de Ia fieur.
Dumort. Beih. Bot. Centralbl. 39(ii): 1-178. Paris: Victor Masson.
Hegnauer, R. 1973, 1990. See general references. Prince, L.M., Parks, C.M. 2001. Phylogenetic relationships of
Hickey, L.J. 1979. A revised classification of the architecture of Theaceae inferred from chloroplast DNA data. Am. J. Bot. 88:
2309-2320.
dicotyledonous leaves. In: Metcalfe, C.R., Chalk, L. (eds.)
Anatomy of the dicotyledons, ed. 2, vol. 1. Oxford: Oxford Ridley, H.N. 1930. The dispersal of plants throughout the
world. Ashford, Kent.
University Press, pp. 25-39.
Ronse Decraene, L.P., Smets, E.F.1996. The morphological vari-
Hitzemann, C. 1886. Beitrii.ge zur vergleichende Anatomie
der Ternstroemiaceen, Dilleniaceen, Dipterocarpaceen und ation and systematic value of stamen pairs in the Magnoli-
atae. Feddes Repert. 107: 1-17.
Chlaenaceen. Osterode: von Giebel & OehlschHigel.
Savolainen, V., Fay, M.P. et a!. 2000. See general references.
Huber, H. 1991. Angiospermen. Leitfaden durch die Ordnun-
gen und Familien der Bedecktsamer. Stuttgart: Gustav Schofield, E.K. 1968. Petiole anatomy of Guttiferae and related
families. Mem. New York Bot. Gard. 18: 1-55.
Fischer.
Schiinenberger, J., Friis, E.M. 2001. Fossil flowers of Ericalean
Hung T. Chang, Ling Lai Kuan, 2000. Theaceae. In: Fu, L., Chen,
T., Lang, K., Hong, T., Lin, Q. (eds.) Higher plants of China, affinity from the Late Cretaceous of Southern Sweden. Am.
vol. 4. Quingdao: Quingdao Publishing House, pp. 572-656 J. Bot. 88: 467-480.
Soltis, D.E. et a!. 2000. See general references.
(in Chinese).
Takhtajan,A.L.1997. See general references. relationships to the family Theaceae. Acta Bot. Yunn. 21:
Tiffney, B.H. 1994. Re-evaluation of the age of the Brandon 202-206, pl. 1-3 (in Chinese).
lignite (Vermont, USA) based on plant megafossils. Rev. Weitzman, A.L., 1987a. Taxonomic studies in Freziera
Palaeobot. Palynol. 82: 299-315. (Theaceae), with notes on reproductive biology. J. Arnold
Tsou, C.-H. 1995. Embryology of Theaceae - anther and ovule Arbor. 68: 323-334.
development of Adinandra, Cleyera and Eurya. J. Plant Res. Weitzman, A.L. 1987b. Systematics of Freziera Willd.
108: 77-86. (Theaceae). Ph.D. thesis, Department of Organismic and
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C.F.P. (ed.) Flora brasiliensis, vol. 12(1). Miinchen: Palaeobot. Palynol. 62: 249-290.
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structure of Pentaphylacaceae and Sladeniaceae and their
Tetrameristaceae 461
Tetrameristaceae
K. KUBITZKI
Tetrameristaceae Hutch., Fam. Fl. Pl. ed. 2: 277, fig. 140 (1959). Both the sepals and petals of Pentamerista are
quincuncially arranged. Maguire et al. (1972)
Trees or shrubs; nodes trilacunar. Leaves alter- suggest that in Tetramerista it is the fifth sepal and
nate, simple, entire, pinnately veined, petiolate or petal that have been reduced. A derived state for
sessile; leaf bases decurrent; stipules wanting. Tetramerista would correlate with the lack of
Inflorescences axillary, pedunculate, umbelliform scalariform perforations of vessel elements, in
or compactly corymbiform racemes; flowers 4- or contrast to the condition in Pentamerista.
5-merous throughout, small, with 2 persistent or Both genera have flask-shaped glands at the
caducous prophylls inserted directly below the basis of the sepals, which appear as pores or slits
sepals; sepals distinct, imbricate, with numerous on the inner surface and obviously function as
scattered pits near the middle of the upper sur- nectaries; similar structures are known from
face; petals distinct, imbricate, not much if at all Pelliciera.
longer than the sepals; filaments applanate, shortly
connate at base; anthers tetrasporangiate, introrse, EMBRYOLOGY, CHROMOSOMES, POLLINATION.
dehiscing by longitudinal slits; gynoecium syncar- Unknown.
pous, 4-5-carpellate; ovary 4-5-locular, capped by
a terminal style with a punctate or minutely lobed POLLEN MORPHOLOGY. Pollen is tricolporate,
stigma; ovules solitary in each locule, axile-basal, suboblate, and reticulate (Erdtman 1952; Maguire
anatropous, bitegmic, tenuinucellate. Fruit a 4- et al. 1972).
5-seeded coriaceous berry; seeds relatively large,
with copious endosperm surrounding a straight, AFFINITIES. Tetramerista has variously been
basal embryo; hypocotyl much longer than included in families such as Ochnaceae,
cotyledons. Marcgraviaceae or Theaceae and, after elevating
Two genera, the monotypic Pentamerista in Tetramerista to family rank, an affinity with
northern South America and Tetramerista (1 [3?] Pellicieraceae (Maguire et al. 1972) and Theaceae
spp.) in SE Asia-Malesia. (Takhtajan 1997) has been proposed. Both plastid
and nuclear gene sequences provide strong
MoRPHOLOGY AND ANATOMY. Young leaves are support for a clade comprising Balsaminaceae,
provided with marginal glands. In Tetramerista Marcgraviaceae, Pellicieraceae and Tetrameris-
the leaves are puncate with black glands below taceae (Savolainen, Fay et al. 2000; Soltis et al. 2000;
(Hutchinson 1959), and Pentamerista has pocket Anderberg 2002), with Pellicieraceae being closest
domatia along the median and lateral veins. The to Tetrameristaceae.
cortex of young stems contains numerous promi-
nent isolated masses of stone cells. Raphide cells KEY TO THE GENERA
are commonly found in the parenchyma and espe-
1. Flowers 5-merous; prophylls and sepals early caducous;
cially frequent in the palisade leaf tissue. Stomata seed obtusely triangular in cross section. N South
are mostly anomocytic. Sclereids are wanting. In America 1. Pentamerista
the wood, vessel elements have predominantly - Flowers 4-merous; prophylls and sepals persistent; seed
simple perforations; rays are 2-3 cells wide, hete- compressed, broadly oblong in cross section. SE
Asia/Malesia 2. Tetramerista
rocellular Kribs type I; axial parenchyma is apo-
tracheal diffuse to diffuse-in-aggregates (Maguire
et al. 1972). Both genera agree in all essential fea-
tures, which is confirmed by H.-G. Richter (pers.
comm.) who compared wood specimens of the two
genera available in the Bundesforschungsanstalt
fur Forst- und Holzwirtschaft, Hamburg.
462 K. Kubitzki
1. Pentamerista Maguire Fig. 132

Pentamerista Maguire, Mem. New York Bot. Gard. 23: 187
( 1972).
Shrub or small tree rarely up to 15m high. One

species, P. neotropica Maguire, lowland savannah
edges of Upper Orinoco-Casiquiare-drainage,
Venezuela, Colombia.
2. Tetramerista Miq.
Tetramerista Miq., Fl. Ind. Bat.: 534 (1860); Ic. Bog. 252 (4): tab.
83 (1901); Maguire, op.c.: 187; Sugan in Tree Fl. Saba! &
Sarawak 2: 379-382 (1996).
Tree up to 30m high; seeds ruminate (van Balgooy

1997). Malaya, Sumatra, Borneo. Probably only one
species, P. glabra Miq., in lowland marshes and
forests up to 1000m altitude.

Balgooy, M. van 1997. Malesian seed plants, vol. 1: spot char-
acters. Leyden: Rijksherbarium.
Hutchinson, J. 1959. The families of flowering plants, 2nd edn.,
vol. 1. Oxford: Clarendon Press.
Maguire, B., de Zeeuw, C., Huang, Y.-c. [sic], Clare, C. C. Jr. 1972.
Tetrameristaceae. In: The Botany of the Guayana Highland,
part IX. Mem. New York Bot. Gard. 23: 165-192.
Soltis, D.E. et al. 2000. See general references.
Takhtajan, A. 1997. See general references.
Fig. 132. Tetrameristaceae. Pentamerista neotropica. A Florif-

erous branch. B Flower with bract and prophylls. C Flower.
D Stamens with subsurface raphide bundles visible. E Fruit.
F Same, transverse section. (Maguire et al. 1972)
Theaceae 463
Theaceae
P.F. STEVENS, S. DRESSLER and A.L. WEITZMAN
Theaceae Mirb., Bull. Soc. Philom. 3: 381 (1813). VEGETATIVE ANATOMY. Druses or sometimes
single crystals of calcium oxalate may be found
Trees or shrubs, accumulating aluminium; indu- throughout the plant. The cork is usually deep-
mentum of unicellular hairs or 0. Leaves spiral or seated, being initiated near the pericycle, but it is
distichous, simple, usually evergreen, usually cori- subepidermal in Franklinia, Schima, and Gordonia
aceous, margins toothed, rarely entire, with a s. str. The pericyclic sheath is either collenchyma-
small, deciduous gland associated with each tooth; tous, lignified, or with groups of fibres. Nodes
stipules absent. Flowers hermaphroditic, solitary, are unilacunar and with a single trace, although
axillary, large and showy, prophylls two, or several cotyledons may have three or more traces. The
bracteoles intergrading with calyx and corolla; petiole bundle is more or less U- or V-shaped, but
calyx of five or more sepals, imbricate, connate or there may be additional smaller lateral bundles.
distinct basally, often persistent in fruit, usually Much-thickened sclereids are common in stem
thick, concave, equal to unequal; corolla of five, cortex, petiole and lamina, and they are usually
rarely numerous petals, distinct or connate basally, branched and with pointed, rarely rounded
imbricate; stamens 20+, free, rarely connate, often (Gordonia) ends; in some genera they are much-
adnate to base of corolla, anthers versatile, rarely elongated spicular cells which traverse the whole
basifixed, opening by longitudinal slits; pollen of the mesophyll. They are absent in Stewartia,
tricolporate, pseudopollen present; gynoecium except for the pedicels, and they are never found
syncarpous, (3- )5( -10)-carpellate, ovary superior, in seedlings (Beauvisage 1920; Keng 1962).
placentation mostly axile; 2-few ovules/carpel, There may be large and much-thickened fibres
ovules bitegmic, tenuinucellate, styles simple, in the phloem. The wood is diffuse porous or more
branched, or stylodia, stigmas usually lobed. Fruit or less ring porous. Vessels are small, usually
a loculicidal capsule, rarely irregularly dehiscent 25-100 )lm tangential diameter, usually solitary
or a drupe, columella persistent, rarely none; seeds and 16-400 per mm 2 • Spiral thickening is com-
few, sometimes winged; testa vascularised, more mon, and the perforation plates are very oblique
or less lignified; endosperm nuclear, usually slight; and with 7-42( -57) bars. Intervascular pitting is
embryo straight; germination epigeal, rarely scalariform to opposite. Vessel-ray pits have at
hypogeal; n = 15, 18. About 7 genera and 195-460 most narrow borders. Parenchyma is often scanty
species; most speciose in Southeast Asia, but also apotracheal, usually diffuse but sometimes in
Indo-Malesia, SE U.S.A., and the Caribbean and short tangential lines. Strands of crystalliferous
tropical America. cells occur in Schima and elsewhere. Wood fibres
are about 800-2000 )lm long and have thin to thick
CHARACTERS OF RARE OCCURRENCE. Gordonia walls. Rays are all about the same size and are
imbricata and a few other species have the con- 1-3(-8) cells wide and up to 2mm or more high.
nective produced at the anther apex. Hairy seeds Uniseriate rays are made up of tall to low upright
are known in Camellia sections Tuberculata and cells only. Multicellular rays are of Kribs types I
Pseudocamellia.Apterosperma has reniform seeds. and II, with narrow, procumbent cells in the body
of the ray and uniseriate margins of square to
VEGETATIVE MORPHOLOGY. The leaves are spiral upright cells (Metcalfe and Chalk 1952; Liang and
or distichous, involute or supervolute in bud, and Baas 1990, 1991).
evergreen, or sometimes deciduous, and the buds Hairs are unicellular, rarely fasciculate. Epider-
are usually perulate (but in Stewartia leaves may mal cells are mucilaginous or not, and there is
be deciduous or evergreen and buds perulate or sometimes a single-layered hypodermis. The
not). Venation is pinnate and more or less craspe- stomata are usually gordoniaceous, the guard cells
dromous. The leaf teeth are crowned by glandular, being surrounded by 2-4 narrow subsidiary cells,
deciduous tips, the Theoid leaf tooth. but Franklinia and many species of Pyrenaria have
0 Schima (Tsou 1998). In Camellia, by contrast, and

probably other genera as well, the stamens are
initiated centrifugally from a ring wall (a circular
primordium) (Ronse Decraene 1989), certainly a
derived state. The stamens are often shortly adnate
to the base of the corolla, and in some species of
Camellia the filaments are connate and form a
prominent tube in the centre of the flower. The
anthers are usually dorsifixed or versatile, the fila-
ments being relatively long and the anthers short
and usually without any prolongation of the con-
nective; basifixed anthers are known in Camellia,
etc. When the anthers are basifixed, the filament
may be narrowed and effectively articulated with
the anther, unlike the basifixed anthers (with short
filaments) of Ternstroemiaceae. Although the
ovary is superior, in Franklinia and Schima it is
partly enclosed by the fused bases of the stamens
plus petals, and so has been described as being
semi-inferior (Tsou 1997). The ovary is often hairy
Fig. 133. Theaceae. Floral diagram of Stewartia pseudocamel- and narrows more or less gradually into the style,
lia, with indication of sequence of inception of floral organs which sometimes is branched (Figs. 134C, 136D)
all of which originate in a consecutive sequence, following an or cleft into stylodia; there is commonly infra-
angle of 144°, except for the smaller angle between organs
10/ll (the transition from corolla to androecium); were this
generic variation in this feature. The stigmas are
divergence also 2/5, a superposed position of sepals and petals, more or less expanded or lobed. The carpels
as in Ternstroemiaceae-Ternstroemieae, would result. (Erbar are usually opposite the petals, although they are
1986) apparently opposite the sepals in Camellia. This
may be connected with the fact that sepals and
petals intergrade and so the arrangement of the
anomocytic stomata and Stewartia has paracytic petals differs from that of other taxa. The basic ori-
stomata. There is also extensive variation in cuticle entation in Camellia is the same as that in Gordo-
surface and details of stomatal morphology nia, where the calyx is clearly opposite the corolla
(Kvacek and Walther 1984a). There are 1-3 layers (Eichler 1875-1878). The young gynoecium shows
of palisade tissue and the fine venation is em- strongly ascidiate development and the degree of
bedded. In some species of Camellia there are congenital fusion is usually low, although high in
"cork-warts", rings of corky material surround- Franklinia and Schima.
ing shallow depressions, on the lower surface of
the lamina (Beauvisage 1920; Metcalfe and Chalk EMBRYOLOGY. Anther cells can have crystals
1952; Keng 1962). and/or tannin, or these may be absent. The con-
nective usually has stomata, but not in Camellia.
INFLORESCENCE AND FLOWER STRUCTURE. Anther wall formation is of the basic type. The
Flowers are large, solitary and axillary, although tapetum is glandular and microsporogenesis is
sometimes appearing terminal. In Camellia pique- simultaneous. The pollen is 2-celled. Pseudopollen
tiana several flowers occur in the axils of scale-like is produced from connective cells. It has large
bracts on a short axillary shoot with a terminal vacuoles and a marginally-situated nucleus when
bud. There has been much discussion over the mature; the wall itself has either rib-like thicken-
distinction between sepals, petals, and prophylls in ings (e.g. Camellia) or circular thickenings, and
Theaceae. In taxa such as species of Camellia, all pseudopollen with the latter thickenings can look
three grade insensibly into one another whereas in like pantoporate pollen grains (e.g. Schima: see
others, like Schima, prophylls, calyx and corolla Tsou 1997, for details).
are all clearly distinguished. There are 20 to well There are only a few apotropous or pleuro-
over 100 stamens. In Stewartia there are five ante- tropous ovules in each carpel and they are spread-
petalous primary primordia on which individual ing or more or less pendulous, except in Stewartia
stamens are initiated centrifugally (Fig. 133; Erbar where they are more or less basal. The ovules
1986); the same may be true of Franklinia and are anatropous (campylotropous in Schima),
Theaceae 465
bitegmic, and tenuinucellate. The micropyle is onal, rarely elongated. The mesotesta is lignified
endostoma!. The outer integument is 4-8 cells or contains sclereids; sometimes the cells are
thick and the inner integument is 4-7 cells thick; fibrous. The endotesta is lignified or not; the cells
in Schima the outer epidermis of the outer integu- are polygonal (Grote and Dilcher 1989, 1992).
ment proliferates in the antiraphal region, forming Mature seeds usually contain at most a little
a ridge of cells up to 12 cells high. In Franklinia oily endosperm (described as being abundant in
the raphe can be massive, to 30 cells thick. There Camellia: Sealy 1958) and a large, straight or
are vascular bundles in either the outer or the rarely curved embryo. The cotyledons are accum-
inner integument, or sometimes only at the base bent and are very large, being much longer than
of the inner integument. There is an integumen- the radicle. They are either fiat or longitudinally
tary tapetum and a hypostase. The archesporium folded (Pyrenaria). Germination is hypogeal
is 1-celled and the embryo sac is usually the (Camellia) or epigeal, the latter being com-
Polygonum type, but it is of the Allium type in monest, but in the former germination type the
Camellia. (General information is taken mainly cotyledons generally separate and may even be
from Yang and Ming [1995a: Pyrenaria, Tutcheria] photosynthetic.
and Tsou [1997: Camellia, Franklinia, Schima]).
The endosperm is nuclear. REPRODUCTIVE SYSTEMS, POLLINATION AND
DISPERSAL. Although most Theaceae have her-
POLLEN MORPHOLOGY. The pollen is tricol- maphroditic flowers, there are reports that Gordo-
porate, suboblate or oblate, and over 28J..Lm and nia (inc. Laplacea) may be dioecious or
sometimes over 50 J..Lm long. The exine is finely monoecious (e.g. Melchior 1925, but cf. Kobuski
granulate or finely reticulate; sexine and nexine 1949, 1950). Theaceae lack nectar, and pollen is
are indistinguishable. The endoaperture is rather presumably the reward for pollinators. The role of
large, lalongate, and indistinct (Keng 1962; Ying the pseudopollen in pollination is not well under-
et al. 1993). stood. Tsou (1997) found both pollen and
pseudopollen on three honey bees foraging in a
KARYOLOGY. Chromosome numbers in Camellia tea garden, although the bees had many fewer
species are most frequently 2n = 30, although some grains of pseudopollen than of pollen. Franklinia
species are polyploids; triploids are common in is highly unusual in that the showy and strongly
cultivars of Camellia japonica (Kondo 1977). The scented flowers appear in late summer, but the
other main number in the family is 2n = 36 (e.g. capsules do not dehisce and the seeds are not
Franklinia, Stewartia), again with possibly poly- ripe until the following summer. Dispersal of the
ploid series based on this. Information on chro- winged seeds is presumably largely by the wind.
mosome numbers is summarized by Tsou (1998). The wingless seeds of Franklinia and especially
Camellia are not likely to travel far. It is not known
FRUIT AND SEED. The fruit is usually a loculici- what eats the drupes of Pyrenaria.
dally dehiscent capsule with a persistent columella
(Fig. 135D), although the latter is at most poorly PHYTOCHEMISTRY. The reader is referred to
developed in Stewartia. The fruit of Franklinia Hegnauer (1973, 1990). Common polyphenols
also dehisces septicidally from the base up. The such as fiavonols, fiavones, and proanthocyanins
valves may soon fall off, or they may remain are widely distributed in Theaceae; the same is true
attached to the fruit. Some species of Camellia of ellagic acid; myricetin is rare in large amounts,
have irregularly-dehiscent capsules, while many and gallic acid and catechins (fiavan-3-ols and
species of Pyrenaria have a drupaceous fruit. fiavan-3,4-diols) are restricted to Camellia sect.
The calyx usually persists at the base of the fruit Thea (C. sinensis, C. taliensis and C. irrawadiensis ).
(Fig. 136B). In young shoots of the tea bush, catechins amount
Schima, Gordonia and Stewartia have winged to more than 30% of dry matter weight but later
seeds, although the development of the wing dif- decrease in quantity. Caffeine and its precursors
fers in at least Schima and Polyspora (Gordonia theobromine and theophylline are restricted to
s.l.) (Tsou 1997, 1998). The shiny seeds sometimes sect. Thea; they have not been found in wild
(e.g. Pyrena ria) have a large hilum and so look species of Camellia or other Theaceae. Caffeine
remarkably like those of Sapotaceae. The seed occurs in the tea bush in quantities of 2.5-4%, and
coat is very thick for an Asterid, often being more the high content in catechins and caffeine in the
than 30 cells across. The testa is vascularised. The tea bush results from selection by man for these
exotesta is lignified or not, and the cells are polyg- substances. Triterpenes and their glycosides
(saponins) are widely distributed in the family and Prince and Parks's (2001) classification of the
found in seeds, bark, wood, and leaves. The plants family is a suggestion of how the tribal classifica-
often accumulate aluminium and fluoride. tion may look in the future:
SUBDIVISION AND RELATIONSHIPS WITHIN THE - Theaeae: Camellia, Pyrenaria, Polyspora,

FAMILY. Relationships within Theaceae are still Laplacea, Apterosperma.
rather unclear. In earlier classifications (e.g. - Gordonieae: Gordonia, Franklinia, Schima.
Melchior 1925, 1964; Airy Shaw 1936; Sealy 1958; - Stewartieae: Stewartia.
Keng 1962), various combinations of simple and/or
continuously varying characters like the winging There are perhaps even more problems with
of the seed and the relative size, distinctness and species delimitation than with generic boundaries,
connation of bracts, sepals and petals were em- as comments after the descriptions below make
phasized. More recently, Ye (1990a, 1990b) and clear.
Tsou (1998), although adding new information
(especially the latter) and thinking about the char- AFFINITIES. In the past, Theaceae s.l. (including
acters in rather different ways from these earlier Ternstroemiaceae and Sladenia) seemed to be
authors, still ended up with groupings relying a pivotal family showing relationships with a
heavily on a few characters. Thus, a variety of number of other taxa. The separate stylodia or
arrangements in the family have been proposed, style branches of some Theaceae, as well as the
Tsou (1998) suggesting there were two tribes, intergradation between prophylls, calyx, and
Camellieae and Gordonieae, the latter including corolla, suggested a rather basal position in the
Stewartiinae and Gordoniinae. Prince and Parks system for some. Although Theaceae are largely
(2001) propose a three-tribe classification, the polypetalous, the realization that their stamen
Stewartieae being sister to Gordonieae and development was centrifugal removed them
Theaeae. Their work is based on an analysis of vari- from direct links with basal polystaminate
ation in two chloroplast genes and flanking spacer angiosperms. Cronquist (1981) included 18 fami-
regions and, although apparently not that dissimi- lies in his Theales, and links between Theaceae
lar from Tsou's classification, the tree suggests a and families like Clusiaceae, also multistaminate,
major change of taxon limits in the family. In par- seemed to be established by the "intermediate"
ticular, major realignments within Gordonia s.l. Bonnetioideae, including genera like Bonnetia,
and both Gordonieae and Theaeae are implied. Kielmeyera, both with spiral leaves and the former
Gordonia lasianthus and G. brandegeei remain in with minutely serrate leaves. Wood anatomy also
Gordonieae, but Polyspora (Old World) and seemed to support such a link (Baretta-Kuipers
Laplacea (New World), both commonly included in 1976), but DNA sequence analyses have demon-
Gordonia, are separate lineages within the strated that Cronquist's Theales form three major
Theaeae. We have not adopted any tribal classifica- groups that are now included in Malpighiales,
tion below, pending more detailed molecular Malvales, and Ericales, with three other families
work and a re-evaluation of the morphological placed in the Caryophyllales and the Asterid I and
data. Nevertheless, it is almost certain that II groups respectively. Nevertheless, even after
Gordonia will not retain the circumscription it is the exclusion of Bonnetia and its allies (now in
given below. Malpighiales), of Pelliciera, Tetramerista (both in
Generic limits elsewhere in the family are a Ericales, see this volume) and Asteropeia (see vol.
matter of some dispute. The limited sampling of V of this series), all at one time or another asso-
Prince and Parks {2001) supports the broad cir- ciated with Theaceae, there is no evidence that the
cumscription of Camellia adopted below, while rest of Theaceae is monophyletic, although it is
that of Pyrenaria is supported by the largely unambiguously ericalean (Morton et al. 1996; APG
unpublished work of Yang (Yang 1988; Yang and 2003). The molecular evidence (Prince and Parks
Ming 1995a, 1995b). Hartia, with characters such 2001; and particularly the five-gene analysis of
as evergreen leaves, narrowly winged seeds, and Anderberg et al. 2002) is consistent with its sepa-
buds without scales, has often been separated from ration into three morphologically quite distinct,
Stewartia, with deciduous or semi-evergreen monophyletic groups, Ternstroemiaceae, Sladeni-
leaves, winged seeds, and buds with scales. aceae, and Theaceae s.str.
Recognition of Hartia would make Stewartia
paraphyletic (Prince and Parks 2001; Li et al. DISTRIBUTION AND HABITATS. Theaceae are
2002). scattered in the north temperate region and in the
Theaceae 467
tropics, especially in South East Asia, where there about half comes from India and China. Green teas
are numerous species in evergreen rainforest. are not fermented during the drying process, other
There are clear links between Theaceae of SE kinds of teas are; alkaloids in tea include caffeine
U.S.A. and those of E Asia. Thus, the two species of and theobromine. Teas have been used as a bever-
Stewartia from the SE U.S.A. are sister to much age in China since at least 350 B.C.E., and tea and
of the rest of the genus and had diverged from tea drinking were introduced to Japan in about
them by the late Miocene (Li et al. 2002). However, 700 c.E., the tea ceremony taking its modern form
understanding details of the biogeographical rela- in the fifteenth century. Tea drinking became
tionships within the family awaits further findings common in England as from the middle of the
on its phylogeny (see above). seventeenth century, and the import of tea was a
Franklinia alatamaha is one of the better- monopoly of the British East India Company until
known North American plants, largely because it the nineteenth century.
has become extinct since it was first discovered. It Camellia japonica, together with thousands
was first found by the Bartrams in 1765 along the of cultivars derived from it and species like C.
Alatamaha River at Fort Barrington, Georgia, saluensis, C. reticulata and C. sasanqua and their
but it had become extinct by 1790, quite possibly hybrids, is a much-grown horticultural shrub with
because of over-collection (Wood 1959). single or double white to red but often odourless
flowers; less frequently-cultivated species have
PALAEOBOTANY. Theaceae are known from the yellow flowers. The seeds of Camellia oleifera
late Cretaceous and are common in the Tertiary. and some other species yield tea oil used mainly
Leaves of the form genus Ternstroemites, common as hair oil, in cooking, and for lubrication. Other
in North America, as well as the form genera Tern- species used as ornamentals include Franklinia
stroemiacinium and Ternstroemioxylon, resemble altamaha, which grows quite readily in cultivation,
genera in both Theaceae and Ternstroemiaceae Stewartia spp., some with very beautiful bark,
(Grote and Dilcher 1989), and so are of little value and Gordonia lasianthus. The timber of Schima is
in understanding the history of the group. Seeds locally of some importance, while other genera
of Theaceae are known from the Cretaceous such as Gordonia may yield woods of minor
(Knobloch and Mai 1986). Grote and Dilcher value.
{1989, 1992, for a literature summary) show that in
the Tertiary a variety of extant and extinct genera KEY TO THE GENERA
(the latter including two genera of Theaceae from
1. Fruits drupes or leathery indehiscent capsules
the Middle Eocene Claiborne flora) were found 2. Pyrenaria
throughout the northern Hemisphere. A rather - Fruits capsular 2
generalised theaceous flower has been found in 2. Anthers often basifixed; carpels mostly 3; seeds large
Baltic amber, and a rather Schima-like wood is (usually >1 em), wingless 1. Camellia
reported from Borneo (Kramer 1974). Gordonia - Anthers often versatile; carpels mostly 5; if seeds large and
wingless, then capsule valves deciduous and seeds angular
was common in the early Miocene Brandon lignite (Pyrenaria) 3
from Vermont, U.S.A. (Tiffney 1994). Current evi- 3. Capsule valves deciduous 4
dence suggests that Camellia once grew in North - Capsules valves persistent, and/or capsules lacking clearly-
America, two genera not now known from Japan defined valves (Franklinia) 5
4. Capsule <1 em across; seeds <1 em long, reniform; calyx
once occurred there, while perhaps four genera are eventually caducous 3. Apterosperma
known from Europe, remains of both vegetative - Capsule >1 em across; seeds >1 em long, ovoid to angular;
(e.g. Kvacek and Walther 1984b) and reproduc- calyx persistent or caducous 2. Pyrenaria.
tive parts being quite common; no member of 5. Capsules (conical-)ovoid to cylindrical 6
the family is currently found there. However, as - Capsules spherical to more or less depressed-globose 7
6. Calyx at the base of the fruit; capsule with columella; seeds
generic limits of extant taxa are re-evaluated (see with wing at one end 4. Gordonia
above), the fossil record will need similar treat- - Calyx more or less investing the fruit; capsule lacking
ment; detailed studies of leaf surface may be columella, seeds wingless or with a wing all round
valuable. 7. Stewartia
7. Plant deciduous; pedicels short; capsule loculicidal, also
septicidal from the base; seeds wingless, not reniform
EcoNOMIC IMPORTANCE. The dried young shoots 6. Franklinia
of Camellia sinensis constitute perhaps the most - Plant evergreen; pedicels medium to long; capsule loculi-
important caffeine drink (Ukers 1935 for general, cidal; seeds surrounded by wing, reniform 5. Schima
if somewhat dated information). Some 2,600,000
tons are produced annually ( 1996 values), of which
Bartholomew, pers. comm.)-300 spp., E Asia to

Indomalesia, mainly S China. Species limits are
very uncertain. The genus has been divided into
four subgenera, Protocamellia Hung T. Chang {3
sections), Camellia (7 sections), Thea (L.) Hung T.
Chang {8 sections), Metacamellia Hung T. Chang
(2 sections: H.T. Chang 1981, see also H.T. Chang
and Bartholomew 1984, Ye 1988).
2. Pyrenaria Blume
Pyrenaria Blume, Bijdr.: 1119 (1827); Keng, Gard. Bull.
Singapore 33: 264-289 (1980); Hung T. Chang & Kuan,
Tutcheria Dunn (1908); Hung T. Chang & Kuan, Higher plants
of China 4: 603-607 (2000).
Sinopyrenaria Hu (1956).
Parapyrenaria H.T. Chang (1963); Ying et al., Endemic genera
seed plants China: 696-698 (1993); Hung T. Chang & Kuan,
Glyptocarpa Hu (1965).
Tree; leaves spiral, evergreen, coriaceous, serrate.

Pedicels short, bracteoles 2, rarely many, persist-
ent, rarely deciduous. Sepals 5-6, unequal; petals
5-6{ -12), slightly connate basally; stamens basally
connate, basally adnate to petals, anthers versatile;
ovary (3-)5-6(-10)-locular, with 2-3(-5) ovules
per carpel; style simple, branched, or free stylodia.
Fruit a drupe, rarely a capsule, columella +; calyx
persistent, at base of fruit, or deciduous. Seeds
Fig. 134. Theaceae. Camellia vietnamensis. A Habit. B ellipsoid, faces flattened; embryo straight, cotyle-
Stamens. C Pistil. D Fruit. E Dehiscing fruit. F Seeds. (Hu 1965) dons thin, tightly folded and twisted, endosperm
0. 42 spp., in evergreen forests, SE Asia, China
(many species) southwards toW Malesia.
1. Camellia L. Fig. 134
Camellia L., Sp. Pl.: 698 ( 1753 ); Sealy, A revision of the genus 3. Apterosperma Hung T. Chang
Camellia (1958); H.T. Chang, Acta Sci. Nat. Univ. Sunyatseni,
Monogr. Ser. 1: 1- 180 (1981); H.T. Chang & Bartholomew, Apterosperma Hung T. Chang, Acta Sci. Nat. Univ. Sunyatseni
Camellias (1984). 1976: 90 (1976); Ying et al., Endemic genera seed plants
Thea L. (1753). China: 691-692 (1993); Hung T. Chang & Kuan, Higher
Piquetia (Pierre) H. Hallier (1921). plants of China 4: 613 (2000).
Yunnanea Hu (1956).
Small tree; leaves ?spiral, evergreen, coriaceous,
Shrubs or trees; leaves spiral or distichous, ever- serrate. Pedicels medium, bracts 2, deciduous.
green, coriaceous, serrate. Pedicel none-medium; Sepals 5, equal; petals 5, slightly connate basally;
bracteoles 2 or more, not sharply differentiated stamens 22-24, anthers basifixed, ovary 5-locular,
from calyx, persistent or deciduous. Sepals 5- 21; with 3-4 ovules per carpel; style short. Capsule
petals 5-12, shortly connate basally; stamens depressed-globose, columella +; calyx and valves
with outer whorl adnate to petals in basal half, caducous. Seeds reniform, wingless. One sp., A.
connate into a tube or not, anthers more or less oblatum Hung T. Chang, S China, in open mixed
basifixed; ovary 3( -5)-locular, with 1-2{-10) forests, 600 m.
ovules per carpel; style simple, branched, or free
stylodia. Fruit a capsule, columella +; calyx per- 4. Gordonia Ellis
sistent, at base of fruit, rarely deciduous. Seeds
more or less globose, wingless; embryo straight, Gordonia Ellis, Phil. Trans. Lond. 60: 518, t. 11 (1771),
cotyledons hemispherical, endosperm +. 100 (B. nom. cons.; Keng, Gard. Bull. Singapore 37:1- 47 (1984);
Theaceae 469
Hung T. Chang & Kuan, Higher plants of China 4: 607-609

(2000).
Laplacea Kunth (1822); Kobuski, J. Arnold Arbor. 30: 166-186
(1949), 31 : 405-429 (1950).
Polyspora Sweet ex G. Don (1831).
Trees; leaves distichous or spiral, evergreen, coria-

ceous, entire or serrate. Bracteoles 2-7, at the apex
of the pedicel, deciduous. Sepals 5, unequal; petals
5(-10), slightly connate basally; stamens adnate
to the base of the corolla, base forming fleshy
pads, anthers versatile; ovary (3-)5( -10)-locular,
with (2-)3-5(-10) ovules per carpel; style simple,
or style branches more or less developed, short to
long; stigmas capitate. Capsule cylindrical, often
longitudinally angled, columella +; calyx usually
persistent, at the base. Seeds flat, with an apical
wing; embryo slightly curved, cotyledons thin, flat,
endosperm thin. 20-65 spp., perhaps 6 spp. in
China, 7-35 spp. elsewhere in Southeast Asia, 3-25
spp. New World tropics, 1 sp. U.S.A. In Polyspora
there is a gradual transition between the calyx and
corolla.
5. Schima Blume Fig. 135

Schima Reinw. ex Blume, Cat. Buitenz. 80 (1823); Bloembergen,
Reinwardtia 2: 133-183 (1952); Keng, Gard. Bull. Singapore
46: 77-87 (1994); Hung T. Chang & Kuan, Higher plants of Fig. 135. Theaceae. Schima wallichii. A Flowering branch. B
China 4: 609- 613 (2000). Stamen. C Calyx with pistil. D Calyx and central columella of
fruit. E Open fruit from above. FSeed. GFruit valve, showing
septum loosening from lower half of fruit-wall. Drawn by
Trees; leaves spiral, evergreen, coriaceous, entire S. Kirno. (Bloembergen 1952)
or crenate. Flowers often crowded near branch
apices; pedicels long, prophylls 2, deciduous, in
upper 1/3 of pedicel. Sepals 5, nearly equal; petals simple. Capsule globose, also dehiscing septici-
5, slightly connate basally, the outermost one dally from the base to half way up, columella +;
enveloping the others in bud; stamens adnate to calyx deciduous. Seeds angular; embryo straight,
the base of the corolla, anthers versatile; ovary cotyledons thin, folded, endosperm thin. One
5( -7)-locular, with 2-6 ovules per carpel; style species, F. alatamaha Marshall, SE U.S.A.
simple. Capsule subglobose, columella at most (Georgia), now known only in cultivation.
short; calyx persistent, at the base. Seeds flat,
reniform, surrounded by a narrow wing; embryo
curved, cotyledons thin, unequal, flat or somewhat 7. Stewartia L. Fig. 136
longitudinally folded, endosperm thin. 1-30 spp., Stewartia L., Sp. Pl.: 698 ( 1753 ); Spongberg, J. Arnold Arbor. 55:
India, Southeast Asia to New Guinea, esp. variable 182-214 (1974); Ye, Acta Sci. Nat. Univ. Sunyatseni 1982(4):
in China. 108- 116 (1982); Li, Acta Phytotax. Sinica 34: 48- 67 (1996);
Prince, Castanea 67: 290- 301 (2002), molec. syst.
Hartia Dunn (1902); Hung T. Chang & Kuan, Higher plants of
6. Franklinia Marshall China 4: 613-616 (2000).
Franklinia Marshall, Arbust. Am.: 48 ( 1785).

Shrubs or trees; leaves distichous, evergreen or
deciduous, often membranous, serrate. Pedicels
Suckering shrub; leaves spiral, deciduous, mem- short, prophylls 2, persistent; sepals 5, equal; petals
branous, serrate. Pedicel short, prophylls 2, decid- 5, connate basally; stamens many, filaments
uous; sepals 5; petals 5, free, stamens many, adnate connate in lower half, anthers versatile; ovary 5-
to the base of the corolla, anthers versatile; ovary locular, with 1-7 basal ovules per carpel, style
5-locular, with 6-12 ovules per carpel; style simple or branched. Capsule ovoid, usually
Chang, H.T., Bartholomew, B. 1984. Camellias. Portland:

Timber Press, London: Batsford.
Cuenoud, P., Savolainen, V., Chatrou, L.W., Powell, M.,
Grayer, R.J., Chase, M.W. 2002. Molecular phylogenetics
of Caryophyllales based on nuclear 18s rDNA and plastid
rbcL, atpB and matK DNA sequences. Am. J. Bot. 89:
132-144.
Eichler, A.W. 1875-1878. See general references.
Erbar, C. 1986. Untersuchungen zur Entwicklung der spirali-
gen Bliiten von Stewartia pseudocamellia (Theaceae). Bot.
Jahrb. Syst. 106: 391-407.
Field, B.S. 1993. Theaceae. In: Heywood, V. (ed.) Flowering
plants of the world. New York: Oxford University Press,
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Grote, P.J., Dilcher, D.L. 1989. Investigations of angiosperms
from the Eocene ofNorthAmerica: a new genus ofTheaceae
based on fruit and seed remains. Bot. Gaz. 150: 190-206.
Grote, P.J., Dilcher, D.L. 1992. Fruits and seeds of tribe Gor-
donieae (Theaceae) from the Eocene of North America.
Am. J. Bot. 79: 744- 753.
Hu, H.-H. 1965. New species and varietie[s] of Camellia and
Theopsis of China (1). Acta Phytotax. Sinica 10: 131-142,
pl. 23-28.
Keng, H. 1962. Comparative morphological studies in the
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Knobloch, E., Mai, D.H. 1986. Monographie der Friichte und
Samen in der Kreide von Mitteleuropa. Rozpr. Ustr. Ust.
Geol. 47: 1-219.
Kobuski, C.E. 1949. Studies in the Theaceae. XVIII. The West
Fig.l36. Theaceae. A-C Stewartia ovata. A Flowering branch- Indian species of Laplacea. J. Arnold Arbor. 30: 166-186.
let. B Apically dehiscent capsule with persistent style branches Kobuski, C.E. 1950. Studies in the Theaceae. XX. Notes on the
and persistent calyx. C Seed. D, EStewartia sinensis. D Pistil. South and Central American species of Laplacea. J. Arnold
E Dormant winter buds. (Spongberg 1974) Arbor. 31: 405-429.
Kondo, K. 1977. Chromosome numbers in the genus Camellia.
Biotropica 9: 86-94.
Kramer, K. 1974. Die tertiaren Holzer Siidost-Asiens (unter
enveloped by persistent calyx, columella 0. Seeds Ausschluss der Dipterocarpaceae), 2. Teil. Palaeonto-
subovoid or flattened, winged or not; cotyledons graphica B, 145: 1-150.
Kvacek, z., Walther, H. 1984a. Nachweis tertiarer Theaceen
thin and flat, endosperm at most slight. 30 spp.,
most in China, 3 spp. in Japan and Korea, 2 spp. in I. Teil - Epidermale Merkmalskomplexe rezenter Theaceae.
SE U.S.A. Feddes Repert. 95: 209- 227.
Kvacek, Z., Walther, H. 1984b. Nachweis tertiarer Theaceen
II. Teil - Bestimmung fossiler Theaceen-Sippen. Feddes
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Catling, D.M. (eds.) Wood structure in biological and tech- lichen Pfianzenfamilien, ed. 2, 21. Leipzig: W. Engelmann,
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Theaceae 471
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472 B. Stahl
Theophrastaceae
B. STAHL
Theophrastaceae Link, Handbuch 1: 440 (1829), nom. cons. filaments fused throughout their length. ]acquinia
nervosa is deciduous and leafless in the rainy
Shrubs or small trees, usually evergreen. Leaves season.
alternate, mostly arranged in one or several
pseudowhorls at branch tips, petiolate, exstipulate; VEGETATIVE MORPHOLOGY. All Theophrastaceae
blades simple, glandular-punctate, often faintly are woody, varying in habit from dwarf-shrubs
striate. Inflorescences terminal or lateral, race- (Neomezia and a few species of Clavija) to shrubs
mose, each flower subtended by a single bract. or small trees to about 15m high. Plants of
Flowers hypogynous, actinomorphic, 5( 4)- ]acquinia, Deherainia, and Votschia are more or
merous, hermaphrodite or unisexual, the aestiva- less richly branched, whereas those of Clavija,
tion imbricate; calyx persistent, lobes free to base, Neomezia, and Theophrasta are sparsely branched
the margins usually erose-ciliate; corolla sym- or unbranched. Among the latter genera,
petalous, usually firm and waxy; tube well devel- Theophrasta is always unbranched (Stahl 1987),
oped, shorter to somewhat longer than the lobes; whereas plants of Clavija and Neomezia usually
lobes oblong to suborbicular, usually somewhat produce one or a few branches (Stahl1991).
unequal in size; stamens homomerous, ante- The leaves are alternate but are usually con-
petalous; staminodes alternipetalous, fused with densed at stem apices into one or several pseud-
the corolla tube; filaments flattened, fused to the owhorls (Fig. 137A, B). A strictly alternate condi-
lower part of the corolla, basally connate or (often tion is met with in several species of ]acquinia. The
in Clavija) united for their entire length; anthers rhythmic production of leaves is clearly expressed
dithecal, basifixed, extrorsely dehiscent through also in species growing under more or less asea-
longitudinal slits, the upper and lower parts of sonal conditions and is also retained in plants cul-
thecae filled up with a white meal of calcium tivated in the greenhouse.
oxalate crystals; ovary superior to semi-inferior, During shoot dormancy the shoot apex is cov-
ovoid to subglobose, unilocular; placenta a basal ered by minute scale leaves, which are left on the
column; ovules few to numerous, anatropous, stem as the shoot prolongates. These scale leaves
bitegmic, tenuinucellate, spirally inserted on but are soft and ephemeral except in Theophrasta
not immersed into a basal column; style shorter to where they become lignified and persist on the
somewhat longer than the ovary; stigma capitate trunk as spines.
or truncate, entire or vaguely lobed. Fruit a berry The leaves lack stipules and in most taxa have a
with a dry and sometimes woody pericarp, short, but well-demarcated petiole, the lower part
indehiscent, subglobose, oblong or ovoid, yellow, of which is more or less swollen. Long petioles, to
orange or red; pulp orange or yellow, juicy and about half the length of the lamina, occur in a few
sweet. Seeds 1 to many, brown or pale brown; species of Clavija. The family exhibits a very large
endosperm abundant, hard; embryo straight, the variation in leaf size, from small and sometimes
cotyledons foliaceous or poorly differentiated. needle-shaped leaves in ]acquinia (Stahl 1995),
A neotropical family of six genera and about via medium-sized leaves in Deherainia, Neomezia,
90 species. Votschia and some species of Clavija to large or
very large leaves in Theophrasta and many species
CHARACTERS OccuRRING IN RELATIVELY FEw of Clavija (to 130 em long; Stahl1991). The lamina
GENERA AND SPECIES. Stem spiny in is simple, usually oblanceolate or elliptic, and
Theophrasta; corolla green in Deherainia; anthers mostly coriaceous or chartaceous. The leaf
distinctly produced at apex in Theophrasta and margins are entire in Deherainia, ]acquinia, and
Neomezia; fruits puberulous in Neomezia and Votschia, whereas they are spinose-serrate in
Clavija glandulifera; flowers in Clavija mostly uni- Neomezia and Theophrasta; in Clavija, species
sexual or functionally unisexual and then with the with entire, serrulate, serrate, and spinose-serrate
Theophrastaceae 473
leaf margins occur. All species of ]acquinia have in ]acquinia and absent in Clavija, Deherainia,
leaves with an apical spine (Fig. 137M), although and Theophrasta, and the strands are usually com-
in some species this is only visible in young, devel- posed of 2-4 or 4 cells. Wood rays are typically
oping leaves (Stahl1992). multiseriate, being 3-6 cells wide in Deherainia
and more than 10 cells wide in Clavija, ]acquinia,
VEGETATIVE ANATOMY. All Theophrastaceae and Theophrasta. Silica grains in the wood have
have immersed glandular trichomes consisting of been reported in some species of Clavija (ter Welle
a short stalk-cell and a multicellular, circular and 1976).
somewhat flattened head. They occur on leaves
and floral parts but are lacking on the corolla INFLORESCENCE STRUCTURE. In Deherainia,
in Clavija. Long, unbranched, glandular-tipped ]acquinia, and Votschia, the inflorescences are ter-
trichomes occur on the leaves and floral parts minal, whereas in Clavija and Neomezia they are
of Deherainia smaragdina and the lower leaf sur- produced laterally, mainly among and just beneath
face of many species of Clavija, and branched, the foliage. In Theophrasta the inflorescences
eglandular hairs are present on the leaves of some appear above the uppermost leaf whorl, but orig-
species of ]acquinia. The young shoots of inate just beneath the shoot tip.
Theophrasta, Neomezia, and many species of The inflorescence is basically racemose with few
Clavija have a very dense and often light brown to many flowers that mature successively during
pubescence of minute, eglandular, and largely a period of one to several weeks. Some ]acquinia
unbranched trichomes. Similar trichomes occur have corymbose or umbellate inflorescences. In
also on branchlets of several species of ]acquinia. Deherainia the inflorescence is usually reduced,
A curious type of branched trichomes with thick often consisting of a single flower (Lepper 1989).
cell walls provides a dense, white or rust -coloured The bracts that subtend the flowers are often
pubescence on branchlets of some Antillean shifted upward on the pedicel, particularly in
species of ]acquinia. species with long pedicels.
Transsections of the vascular strand in petioles
of Deherainia and ]acquinia are open-crescentic FLOWER STRUCTURE AND ANATOMY. The flowers
and composed of 3-5 bundles. In the other genera of Theophrastaceae are hypogynous, actinomor-
the petiolar vascular strand is circular showing phic, and 5- or (sometimes in Clavija) 4-merous.
various degrees of complexity. In size they vary from 3-4 mm diam., as in some
With the exceptions of a few species of Clavija, species of Clavija and ]acquinia, to 35-45 mm
extraxylary sclerenchyma is present in the leaves diam. in Deherainia.
of all Theophrastaceae (Votsch 1904; Stahl 1989, The calyx is persistent and usually more or less
1991). It consists of short, unbranched fibre cells greenish; the lobes are well developed and mostly
(Stahl1987), which are arranged in subepidermal erose-margined. The corolla is thick and waxy and
bundles or layers. In most species this scleren- exhibits a large variation with regard to shape and
chyma occurs near the epidermis on both sides colour. It is subrotate in Clavija, broadly campan-
of the leaf, but in some species of ]acquinia and a ulate in Deheriania, Votschia, and Neomezia,
few species of Clavija the foliar sclerenchyma is campanulate in Theophrasta and most species of
more or less sunken into the subjacent mesophyll. ]acquinia, and urceolate in some ]acquinia species.
In addition, the adaxial foliar sclerenchyma is The lobes, which are imbricate in bud, are broadly
sometimes separated from the epidermis by one oblong or subrotate and usually somewhat
or sometimes two layers of hypodermal cells. In unequal in size. Orange is the most widespread
dried leaves the extraxylary foliar sclerenchyma colour of the corolla and is found in all species of
appears as a fine striation running more or less Clavija and any species of ]acquinia. Other corolla
parallel to the lateral veins. colours are green (Deherainia), white (Jacquinia
Sclereids in the form of irregularly branched spp.), and pale yellow (Jacquinia spp., Votschia). In
cells are present in the leaves of a few species of Neomezia, the corolla is basically orange but has a
]acquinia. Stone cells occur throughout the family distinct brownish tinge, which is also observed in
in the pith of young stems and in tissues of repro- one species of Clavija. The corolla of some species
ductive organs. of ]acquinia are pale orange in bud and at the
According to Metcalfe and Chalk (1950), the beginning of anthesis become orange-red. In
wood of Theophrastaceae has small vessels Theophrasta the corolla is dull yellow or buff with
(usually less than 50 !lm diam.) with simple per- a distinct orange-brown tinge in bud. The orange
foration plates. Parenchyma is extremely sparse corolla colour is retained in properly dried her-
474 B. stahl
barium specimens. In Theophrasta and ]acquinia formation. The embryo sac is probably of the
berterii the corolla becomes dark brown and Polygonum type with ephemeral antipodes;
finally black when aged or upon drying. endosperm formation is nuclear. Some, but not
All genera have a series of staminodial all Theophrastaceae (and Myrsinaceae) have an
appendages alternating with the corolla lobes. endosperm with thickened, abundantly pitted
These appendages are completely fused with the cells walls (see Anderberg and Stahl 1995, fig. 7,
corolla, being inserted at the mouth of the tube, and Takhtajan 1992, fig. p. 56). In Deherainia,
except in Theophrasta, where they are situated Mauritzon (1936) classified this tissue as
inside the tube. In ]acquinia and Votschia the perisperm developed from the inner integument,
appendages are large and flattened and resemble but this interpretation still needs to be verified.
an extra whorl of corolla lobes; in Clavija they
are gibbous-ellipsoid, in Theophrasta gibbous and POLLEN MoRPHOLOGY. Pollen grains in the
transversely oblong, in Deherainia lanceolate to family are spheroidal and vary in size from 17 to
narrowly obovate, and in Neomezia small and 42Jlm in diam. (Erdtman 1952; Carrasquel 1970 ).
triangular. They are 3-colporate, except in several species of
The stamens are of the same number as the Clavija, which have 4-colporate grains. The exine
corolla lobes and stand in front of them. The fila- is foveolate in ]acquinia, fossulate in Deherainia,
ments are somewhat flattened and fused at the rugulate in Neomezia, Theophrasta, and Votschia,
base, where they are adnate to the corolla tube. and reticulate in Clavija. Extratectal structures
The flowers are protandrous and dehisce in the form of spinules are restricted to Clavija,
extrorsely with longitudinal slits. In all genera although they are lacking there in at least one
except Clavija, the stamens are loosely coherent species.
during the male phase and spread during the
female phase. In Clavija, the stamens of male, most KARYOLOGY. Various studies indicate a basic
hermaphrodite and functionally male flowers chromosome number of x = 18 for Theophras-
have the filaments fused into a permanent tube, taceae (Faure 1968), although 2n = 40 and 2n = 28
whereas in female flowers the non-functional have been reported in one species of Clavija and
stamens are distinct throughout anthesis. In ]acquinia respectively (Gadella et al. 1969; Hanson
Neomezia and Theophrasta, the anthers are pro- et al. 200 1).
duced at the apex. The upper and lower parts of
the thecae are filled up with a white meal of POLLINATION AND REPRODUCTIVE SYSTEMS. All
calcium oxalate druses and crystals, which at genera of Theophrastaceae seem to be insect-
anthesis is exposed at the tip and base of the pollinated. ]acquinia and Clavija, the flowers of
anther and forms a distinct visual contrast to the which have a perfumed or fruity scent respectively,
yellow pollen. First described for Deherainia by appear to be bee-pollinated, whereas Neomezia,
Chandler (1911) and Pohl (1931), these oxalate Theophrasta, and Deherainia, which have flowers
accumulations occur throughout the family. with a foetid scent, most likely are pollinated by
The gynoecium is unilocular and has a globular Diptera. However, a species of gall midges was
central placenta. Although certainly syncarpous, observed visiting flowers of Clavija in Ecuador
the number of carpels forming the gynoecium is (Gagne et al. 1997). The glandular trichomes
unknown. In all genera the ovules are densely set on floral parts may produce small quantities of
on the surface of the placenta and their number nectar, as has been observed in ]. macrocarpa
varies from 40 to 250 (Fig. 137L); only in Clavija (Vogel 1986), but nectar as free liquid inside the
are there usually less than 30 ovules, and these are corolla is lacking in all genera. Even though pollen
often rather sparsely distributed on the placenta is produced in small quantities, pollen-gathering
(Fig. 137F). bees may well be important as pollinators in
]acquinia, Clavija, and Votschia. In Deherainia,
EMBRYOLOGY. The anthers are reported to have Neomezia, and Theophrasta pollination is clearly
fibrous endothecium and a secretory tapetum based on deceit. In Theophrasta the blossoms
with uninucleate cells (Johri et al. 1992). Like in mimic fruit-bodies of mushrooms through colour,
most other members of the Primulales, the ovules scent, and in being covered by plant litter, and
in Theophrastaceae are anatropous, tenuinucellate are probably pollinated by Diptera that visit the
and bitegmic (Mauritzon 1936). The micropyle is flowers for oviposition (Stahl 1987). The role of
formed by both integuments, which are clearly dis- the calcium oxalate crystals in pollination, if any,
tinguishable only during the early stages of ovule is unknown.
Theophrastaceae 475
All genera except Clavija have hermaphrodite, (Knudsen and Stahl 1994). In Clavija the scent
protandrous flowers. In Clavija, most species have is characterised mainly by sesquiterpene hydro-
flowers that are morphologically intermediate carbons, in ]acquinia by benzenoids, phenyl
between the hermaphrodite and dioecious con- propanoids, and (in orange-flowered species)
dition and usually appear as gynodioecious, al- trimethylcyclohexanes derivatives of carotenoids,
though the female function is probably strongly and in Deherainia and Theophrasta by fatty-acid
suppressed (Stahl 1991). Studies of herbarium derivatives.
material suggest that a few species are functionally
androdioecious. Deherainia grown under green- SUBDIVISION OF AND RELATIONSHIPS WITHIN
house condition has been reported to be self- THE FAMILY. Theophrastaceae are divided into
incompatible (Chandler 1911). A poor fruit set two groups, one including the pachycaul genera
has also been observed in cultivated plants of Clavija, Neomezia, and Theophrasta, and one
Theophrasta. compnsmg the richly branched genera
Deherainia, ]acquinia, and Votschia. This subdivi-
FRUIT AND SEED. Theophrastaceae form indehis- sion is in agreement with the classifications
cent berries with dry, coriaceous or woody peri- proposed by de Candolle (1844) and Votsch
carps. The fruits are yellow or orange, globose (1904), who referred the two groups to the tribes
or ovoid, glabrous or rarely brownish-puberulous, Clavijeae and Jacquinieae respectively. It is also
and vary in size from 0.5 to 5 em diam. Each fruit consistent with a preliminary cladistic analysis
has usually less than 5 and rarely more than 10 (Stahl1991).
seeds, which are entirely or partly imbedded in an
orange and sweet-tasting placental pulp. The seeds AFFINITIES. Theophrastaceae are firmly placed in
are 0.3-10mm wide, globose-compressed, irregu- Primulales because of their 5( 4-)-merous flowers,
larly obtuse-angled, or flattened and oblong. They sympetalous corolla, single whorl of antepetalous
are light to dark brown and have a thin, smooth or stamens, and unilocular ovary with a free, central
faintly reticulate testa. The endosperm is abundant placenta. Being woody and tropical, the family has
and hard and has smooth or irregularly thickened usually been related to the Myrsinaceae. Phyloge-
cell walls. The embryo is straight and has a well- netic studies based on morphological and rbcL
developed hypocotyl and foliaceous or small and sequence data (Anderberg and Stahl 1995;
poorly differentiated cotyledons. Anderberg et al. 1998) placed Theophrastaceae as
the sister group to the rest of primuloid families.
DISPERSAL. No reports on seed dispersal in the However, in more recent work (Kallersjo et al.
family have been published; however, the juicy and 2000), Maesa (Maesaceae) replaces Theophras-
coloured fruits strongly suggest zoochory, proba- taceae as the basal family in the Primulales, and
bly involving birds as the most important vectors. Theophrastaceae with Samolus (Samolaceae) as
Collections of Clavija and Theophrasta with fruits the sister group forms the second branch of the
that evidently were pecked open by birds suggest primuloids, which is basal to the rest of the primu-
that the seeds often are deposited near the mother- loids (Myrsinaceae and Primulaceae s.str.). The
plant as the soft pulp is consumed. Other dispersal monophyly of the Theophrastaceae is well sup-
vectors that could be involved are forest rodents ported by features such as pseudowhorled leaves,
and monkeys. The brownish colour of the fruits of extraxylary foliar sclerenchyma, and anthers with
Neomezia and their position near to the ground calcium oxalate crystals (Anderberg and Stahl
suggest rodents or reptiles as dispersal agents of 1995).
this genus.
DISTRIBUTION AND HABITATS. Theophrastaceae
PHYTOCHEMISTRY. Green parts are rich in are strictly neotropical, being distributed from
saponins and the seeds have an oily endosperm northern Mexico and southern Florida to south-
lacking starch (Hegnauer 1973). The orange ern Brazil and northern Paraguay. At the generic
pigment in flowers of ]acquinia nervosa has been level the family is most diverse in and around
identified as trans-crocetindialdehyd and trans- the Caribbean. Neomezia and Theophrasta are
crocetinhalbaldehyd (Eugster et al. 1969), and it endemic to the islands of Cuba and Hispaniola
seems likely that the same carotenoids are pre- respectively; Votschia is restricted to a narrow
sent in the other orange-flowered species of the strip of land along the Caribbean side of eastern
family. The chemical composition of the floral Panama; Deherainia is distributed from eastern
scent differs to a large extent between the genera Mexico to Honduras; and ]acquinia has 12 species
476 B. Stahl
in the Mesoamerican region, about 18 in the 4. Leaf margins entire to serrate, rarely spinose-serrate;
Antilles, and a few in northern South America. flowers uni- or bisexual, sweet-scented; corolla crateriform
6. Clavija
Clavija is the only genus in the family centred on - Leaf margins usually spinose-serrate; flowers bisexual,
the South American mainland, where it is partic- with foetid odour; corolla ± campanulate 5
ularly diverse in north-western South America 5. Erect, unbranched treelets or shrubs; inflorescences many-
and the western Amazon. flowered, appearing above the foliage; corolla appendages
The family as a whole has mainly a lowland transversely oblong 4. Theophrasta
- More or less decumbent and sparsely branched dwarf-
distribution but species of ]acquinia and Clavija shrubs; flowers solitary or inflorescences few-flowered,
may reach an elevation of 2000-2500m. Species appearing among and beneath the leaves; corolla
occurring above 1500m altitude usually have a appendages triangular 5. Neomezia
restricted distribution.
Most Theophrastaceae grow in seasonally dry
1. ]acquinia L. Fig. 137M-O
areas from semideciduous and deciduous forests
to thorn scrubs. The last-mentioned vegetation ]acquinia L., Fl. Jamaica: 27 ( 1759); Mez in Pflanzenreich IV.
type is a common habitat for ]acquinia, several 236a: 28-44 (1903); Stahl, Nord. J. Bot. 9: 15-30 (1989), 15:
species of which inhabit dry coastal vegetation 493-511 (1995).
and some may even be found in the upper edge
of mangrove swamps. Although several species Richly branched shrubs and small trees. Leaves
of Clavija grow in deciduous or semideciduous small, sometimes needle-like, mostly spine-tipped,
forests, many inhabit wet, evergreen forests. Some margins entire. Racemes terminal, few- to many-
Amazonian species of Clavija are restricted to flowered. Flowers 5-merous, hermaphrodite;
habitats with temporarily inundated soils. corolla campanulate or urceolate, orange, white,
or yellowish; corolla appendages flattened and
ECONOMIC IMPORTANCE. Theophrastaceae are petaloid; stamens filaments united at base; anthers
of little economic importance. Crushed roots and obtuse-triangular. Fruit subglobose or oblong,
immature fruits of some species of ]acquinia have pericarp thick and woody or thin and brittle.
been used locally as a fish poison ("barbasco") About 32 spp., West Indies, Mexico to northern
and, in coastal Ecuador, crushed, immature fruits South America.
of ]. sprucei are used in shrimp farms to clear
baskets from unwanted fish. In the northern 2. Deherainia Decaisne
Caribbean the soft cortex of some ]acquinia
species has been used as a substitute for soap. Deherainia Decaisne,Ann. Sci. Nat. Bot. VI, 3: 138 (1876); Stahl,
Nord. J. Bot. 9: 20-21 (1989).
Flowers of ]acquinia were once used to decorate
the Maya temples, and flowers of Clavija and
Shrubs or small trees. Leaves medium-sized,
]acquinia are strung into necklaces and garlands
margins entire. Racemes terminal, few-flowered,
by indigenous people in the western Amazon and
or flowers solitary. Flowers 5-merous, hermaphro-
Mesoamerica respectively (Lundell 1960; Standley
dite; corolla broadly campanulate, green; corolla
and Williams 1966). Fruits of Clavija are eaten
appendages lanceolate or narrowly obovate;
as a "forest -snack" throughout Central and South
stamens filaments united at base; anthers obtuse-
America.
triangular. Fruit ovoid, tapering towards apex,
peri carp thin and brittle. Two spp., eastern Mexico
KEY TO THE GENERA
to Honduras.
1. Usually richly branched shrubs and trees; leaves small
(<15cm long) or very small, margins entire 2
- Sparsely branched or unbranched treelets, shrubs or 3. Votschia Stahl
dwarf-shrubs; leaves of medium size (rarely <15cm long)
to very large (to 130 em long), margins serrate, serrulate, Votschia Stahl, Brittonia 45: 204-207 (1993).
spinulose, or entire 4
2. Corolla campanulate or urceolate, orange, white or Branched shrubs. Leaves medium-sized, margins
yellowish, 0.5-1.5 em diam. at anthesis; leaves generally entire. Racemes terminal, few-flowered. Flowers
<Scm long, mostly spine-tipped l.]acquinia 5-merous, hermaphrodite; corolla broadly cam-
- Corolla broadly campanulate, green or yellow, 2.5-
4 em diam. at anthesis; leaves usually >5 em long, not panulate, pale yellow; corolla appendages flattened,
spine-tipped 3 petaloid; stamens filaments united at base; anthers
3. Corolla green; fruit ovoid, tapering towards apex obtuse-oblong. Fruits subglobose, pericarp some-
2. Deherainia what thick and woody. One sp., V. nemophila
- Corolla yellow; fruit subspherical 3. Votschia
(Pittier) Stahl, north-eastern Panama.
Theophrastaceae 477
4. Theophrasta L. Fig. 137G-L Fig. 137. Theophrastaceae. A Clavija ornata, habit. B-F C.
pungens. B Flowering shoot. C Flower, seen from above.
Theophrasta L., Sp. Pl. 1: 149 (1753); Stahl, Nord. J. Bot. 7: D Stamina! tube. E Calyx and gynoecium of bisexual or
529-538 (1987). morphologically male flower. F Placenta and ovules. G-L
Theophrasta jussieui. G Leaf. H Inflorescence. I Flower, vertical
Unbranched shrubs or treelets, the stem spiny. section. J Part of floral tube with one stamen and staminodes.
K Stamen, seen from outside. L Placenta with ovules. M-0
Leaves large, margins spinose-serrate or serrulate. ]acquinia aculeata. M Flowering shoot. N Flower. 0 Fruit.
Racemes subterminal, many-flowered. Flowers (Takhtajan 1981)
5-merous, hermaphrodite; corolla campanulate,
orange-brown to pale buff, turning brownish black
with age and upon drying; corolla appendages Unbranched or sparsely branched shrubs or small
gibbous, transversely oblong, inserted within the trees. Leaves large to very large, margins entire,
tube; stamens filaments united at base; anthers serrulate, serrate, or rarely spinose-serrate.
narrowly oblong, produced at apex. Fruit subglo- Racemes lateral, many- or few-flowered. Flowers
bose, pericarp rather thin and brittle. Two spp., 5- or 4-merous, often unisexual; corolla crateri-
Hispaniola. form, orange; corolla appendages gibbous-oblong;
stamens often fused into a tube; anthers obtuse-
triangular, rarely with short apical production.
5. Neomezia Votsch Fruit subglobose, usually glabrous, pericarp
Neomezia Votsch, Bot. Jahrb. Syst. 33: 541 (1904). mostly thin and brittle. About 55 spp., Central and
South America, Haiti.
Unbranched or sparsely branched dwarf-shrubs.
Leaves medium-sized to large, margins spinose-
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in the order Primulales, with special emphasis on the family
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478 B. Stahl
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midge (Diptera: Cecidomyiidae) from Ecuador associated 60.
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Soc. Wash. 99: 110-114. the Antilles and South America. Nord. J. Bot. 15:493-511.
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Index to Scientific Names
References to main entries in bold-faced print, to illustrations in italics.
Abdulmajidia 231 Alphitonia 335

Acaena 365 Alvimiantha 330
A. poeppigiana 365 A. tricamerata 331
Acanthogilia 306 Amblyanthopsis 278
Acanthogilioideae 306 Amblyanthus 277
Acanthothamnus 41 Amelanchier 380
Aceratium 141 Ampelozizypheae 335
Achras 402 Ampelozizyphus 335
Ackama 106 Amygdalaceae 343
Acomastylis 367, 368 Anagallis 273
Acrolasia 251 A. arvensis 268
Acrophyllum 104 Ancistrum 365
A. australe 105 Andersonia 175
Acrostemon 173 Andresia 168
Acrotriche 177 Andromeda 181
Acsmithia 101 A. polifolia 150
Actinidia 17 Andromedeae 181
A. polygama 17 Androsace 316
A. strigosa 17 Androstoma 177
Actinidiaceae 4, 7, 14 Anneslea 456
Adenilema 108 Anodopetalum 103
Adenostoma 382 Anomalanthus 173
Adinandra 457 Anthodon 55
Adolphia 328 Antholoma 139
Aegiceras 277 Anthopteropsis 186
A. corniculatum 268 Anthopterus 189
Aegicerataceae 266 Antistrophe 277
Afrardisia 275 Aosa 248
Afrocrania 88 Apatophyllum 41
Afrosersalisia 416 Aphanes 371
Agapetes 183 Aphanopetalum 109
A. scortechinii 185 Apodostigma 55
Agarista 181 Apopetalum 384
Agauria 181 Apterosperma 468
Agelaea 80 Arachnocalyx 173
Agrimonia 361 Arbuteae 169
Aistopetalum 102 Arbutoideae 169
A. viticoides 102 Arbutus 169
Alangiaceae 82 A. andrachne 149
Alangium 89 Archboldiodendron 457
A. chinense 89 A. calosericeum 457
Alchemilla 371 Archeria 174
A. rothii 371 Archerieae 174
A. subg. Alchemilla 372 Arcterica 180
A. subg. Aphanes 372 Arctostaphylos 169
A. subg. Lachemilla 372 Arctous 169
Aliciella 310 Ardisia 275
Allantoma 228 A. crenata 268
A. lineata 223 A. sect. Hymenandra 275
Allocassine 41 A. subg. Synardisia 275
Allophyllum 308 A. subg. Walleniopsis 276
Allotropa 167 Ardisiandra 272
Alniphyllum 440 A. wettsteinii 268
A. fortunei 436 Aremonia 362
480 Index to Scientific Names
Aretia 316 Brexia 42

Argania 410 Brexiaceae 29
Aria 375 Brexiella 42
Aristotelia 139 Broussaisia 212
Arnicratea 56 Bruckenthalia 173
Arnoldia 108 Bruinsmia 440
Aronia 376 Brunellia 28
A. sp. 377 B. integrifolia 27
Aruncus 356 B. latifolia 27
Asteranthos 428 B. standleyana 27
A. brasiliensis 428 Brunelliaceae 2, 26
Asterolinon 273 Bryanthus 170
Astroloma 177 B. gmelinii 149
Aubregrinia 413 Bryocarpum 318
Auerodendron 333 Budawangia 175
A. glaucescens 322 B. gnidioides 176
Aulandra 407 Bumelia 409
Auriculardisia 275 Burckella 409
Autranella 400 Burttia 78
Averrhoa 288 B. prunoides 78
Azalea 171 Butyrospermum 402
Badula 279 Caiophora 249

Baillonella 402 C. madrequisa 243
Balsaminaceae 4, 7, 20 Caldcluvia 106
Balthasaria 458 Caldcluvieae 106
Barbeyaceae 2 Callicoma 107
Barringtonia 230 Calluna 173
B. calyptrata 230 C. vulgaris 149, 155
Barringtoniaceae 221 Calluneae 173
Bartonia 251 Calopteryx 189
Bathiorhamneae 335 Calycomis 104
Bathiorhamnus 335 Camellia 468
Bauera 103 C. vietnamensis 468
Baueraceae 91 Camptotheca 87
Bejaria 170 Campylostemon 56
B. racemosa 149, 155 Canotia 42
Bejarieae 170 Canotiaceae 29
Bencomia 363 Cantua 307
Benthamidia 88 C. buxifolia 302
Bequaertia 56 C. quercifolia 307
Bequaertiodendron 417 Capurodendron 418
Berchemia 333 Caracasia 263
Berchemiella 332 Carambola 288
Bernardinia 81 Cardiandra 211
Berneuxia 120 Careya 231
Bertholletia 228 Cariniana 228
B. excelsa 229 C. pauciramosa 223
Bhesa 41 Carpenteria 211
B. archboldiana 41 Cassine 42
Bicuspidaria 251 Cassiope 170
Biophytum 288 C. lycopodioides 149
Blackallia 332 C. mertensiana 155
Bladhia 275 Cassiopeae 170
Blaeria 173 Cassiopoideae 170
Blumenbachia 248 Catha 43
Boerlagella 419 C. edulis 43
Boerlagia 419 Cavendishia 185
Bonplandia 307 Ceanothus 335
B. geminiflora 307 Celastraceae 1, 29
Bothrocaryum 88 Celastroideae 41
Brachycaulos 383 Celastrus 43
Brachyloma 178 C. sect. Gymnosporia 46
Brassiantha 42 C. scandens 44
Brayera 361 Celtidaceae 2
Brazzeia 429 Centunculus 273
Breviea 413 Cephalotaceae 2, 65
Index to Scientific Names 481
Cephalotus 67 Conandrium 281

C. follicularis 67 Condalia 333
Ceratiola 173 Condaliopsis 327
Ceratopetalum 103 Connaraceae 2, 74
Ceratostema 187 Connareae 78
Cercocarpus 360 Connaropsis 287
C. traskiae 360 Connarus 78
Cevallia 252 C. longistipitatus 78
Chaenomeles 374 Conomorpha 278
C. sect. Pseudocydonia 375 Conomyrsine 278
Chamaebatia 360 Conostephium 178
Chamaebatiaria 358 Corelliana 278
Chamaedaphne 181 Corema 173
C. calyculata 150 C. album 9
Chamaemeles 381 C. conradi 172
Chamaemespilus 375 Coridaceae 266
Chamaepericlymenum 88 Goris 272
Chamaerhodos 366 Cormus 375
Changiostyrax 440 Cornaceae 3, 82
Chaydaia 333 Cornus 88
Cheiloclinium 59 C. japonica 88
Cheilotheca 168 C. officina/is 84
Chichicaste 247 Cortusa 318
Chimaphila 167 C. matthioli 318
Chiogenes 182 Corythophora 228
Chromolucuma 413 C. alta 223
Chrysophylleae 411 Cosmelia 175
Chrysophyllum 414 Cosmelieae 175
Chydenanthus 230 Costera 189
Cladothamneae 171 Cotoneaster 380
Cladothamnus 171 C. sect. Malacomeles 380
Clavija 477 Couratari 229
C. ornata 477 C. stellata 223
C. pungens 477 Couroupita 228
Clematoclethra 17 C. guianensis 223
C. lasioclada 17 Cowania 360
Clethra 72 Craibiodendron 180
Clethraceae 4, 7, 69 Crataegeae 380
Cleyera 457 Crataegus 381
Cliffortia 363 Crateranthus 284
Cliftonia 115 C. talbotii 284
Cnestideae 80 Cremnobates 103
Cnestidium 81 Crinodendron 140
Cnestis 80 Crocoxylon 44
Cobaea 307 Croninia 178
C. penduliflora 307 Crossopetalum 43
C. scandens 307 Crumenaria 330
Cobaeoideae 306 Cryptandra 332
Coccosperma 173 Ctenardisia 277
Codia 107 Cuervea 56
Codieae 106 Cunonia 108
Coilostigma 173 Cunoniaceae 2, 91
Coleanthera 178 Cunonieae 107
Coleogyne 382 Curtisia 113
Colletia 329 C. dentata 113
C. hystrix 329 Curtisiaceae 3, 112
Colletieae 327 Cyathodes 178
Collomia 308 Cyathopsis 178
C. bijlora 302 Cybianthus 278
Colubrina 335 Cyclamen 273
C. asiatica 336 C. hederifolium 268
Coluria 368 Cydonia 375
Comarella 366 Cyrilla 115
Comarostaphylos 169 Cyrillaceae 4, 7, 114
Comarum 366
Combretodendron 230 Daboecia 173
Comomyrsine 278 D. azorica 155
D. cantabrica 149 Ebenaceae 5,7,125

Daboecieae 173 Eberhardtia 407
Dallachya 333 Ecclinusa 414
Dapania 288 Echinocarpus 139
Darlingtonia 424 Elachyptera 56
Davidia 87 Elaeagnaceae 2, 131
D. involucrata 84, 87 Elaeagnus 133
Davidiaceae 82 E. angustifolia 133
Davidsonia 103 Elaeocarpaceae 2, 135
Davidsoniaceae 91 Elaeocarpus 141
Decatoca 178 E. sericoloides 142
Decumaria 213 Elaeodendron 44
D. barbara 213 E. buchananii 45
Deherainia 476 Elaeoluma 415
Deinanthe 211 Eleutherostemon 188
Delpydora 414 Elingamita 278
Demosthenesia 189 Elliottia 171
Dendriopoterium 363 E. bracteata 155
Dendrobenthamia 88 E. pyroliflora 149
Denhamia 44 Ellipanthus 78
Deutzia 209 Embelia 273
D. paniculata 210 E. philippinensis 268
Diapensia 120 Emblemantha 276
D. lapponica 119 Emmenosperma 336
Diapensiaceae 4, 7, 117 Empedoclesia 185
Dicarpellum 44 Empetraceae 145
Dichotomanthes 378 Empetreae 172
Dichroa 212 Empetrum 172
Didonica 184 E. nigrum 172
Dimorphanthera 184 Empleuridium 45
D. amblyornidis 185 Englerophytum 417
Diogenesia 188 Enkiantheae 165
Dionysia 317 Enkianthoideae 165
Diospyros 128 Enkianthus 166
D. curranii 129 E. campanulatus 166
D. koeboeensis 129 E. paucijlorus 155
Dipholis 409 E. perulatus 166
Diplarche 170 Epacreae 175
D. multiflora 155 Epacridaceae 145
Diplarcheae 170 Epacrideae 175
Diploknema 409 Epacridoideae 174
Diploon 411 Epacris 175
Diplopanax 87 Epigaea 171
Diplycosia 182 E. asiatica 149, 155
D. malayana 150 E. repens 155
D. pinifolia 182 Epigaeae 171
Dirachma 123 Eremia 173
D. socotrana 123 Eremiella 173
D. somalensis 123 Eremotropa 168
Dirachmaceae 2, 122 Eriastrum 309
Discaria 329 Erica 173
Discocalyx 280 E. australis 149
Disterigma 187 Ericaceae 4, 7, 145
Docynia 375 Ericeae 170, 173
Docyniopsis 378 Ericinella 173
Dodecatheon 318 Ericoideae 170
D. meadia 318 Eriobotrya 376
Doerpfeldia 335 E. bengalensis 377
Doerpfeldieae 335 Eriogynia 356
Douglasia 316 E. sect. Kelseya 356
D. subg. Gregoria 316 E. sect. Petrophytum 356
Dracophyllum 177 Eriolobus 379
Dryadeae 359 E. sect. Docynia 375
Dryas 359 Erythrocoma 368
Drymocallis 366 Eschweilera 229
Dubouzetia 140 E. longipes 223
Duchesnea 366 Eubotrys 182
Euclea 129 Gordonia 468

Eucnide 251 Gouania 330
Eucryphia 104 Gouanieae 330
E. cordifolia 104 Grammadenia 279
Eucryphiaceae 91 Granitites 336
Euonymopsis 45 Grenacheria 274
Euonymus 45 Grias 227
E. americana 44 G. cauliflora 223
Eurya 457 Grisebachia 173
E. emarginata 9 Gronovia 252
Euryodendron 458 Gronovioideae 252
Exochorda 355 Grubbia 201
Exochordeae 354 G. rosmarinifolia 200
Grubbiaceae 3, 199
Fallugia 369 Guamatela 384
Faucherea 404 Gummellia 109
Fendlera 209 Gustavia 227
Fendlerella 209 G. hexapetala 223
Ficalhoa 432 Gyminda 46
Filipendula 361 Gymnosporia 46
Fittingia 280 Gymnosteris 311
Foetidia 231 G. parvula 307
Foetidiaceae 221
Foetidioideae 226 Hagenia 361
Fouquieria 197 H. abessynica 362
F. splendens 196 Halesia 441
Fouquieriaceae 4, 7, 195 Harrimanella 174
Fragaria 366 H. stelleriana 149
Frangula 332 Harrimanelloideae 174
Franklinia 469 Hartia 469
Fraunhofera 46 Hartogiella 42
Freziera 458 Hartogiopsis 47
F. carinata 458 Heberdenia 274
Freziereae 456 H. bahamensis 268
Friesia 139 Hedraianthera 47
Fuertesia 252 Heliamphora 424
F. domingensis 253 H. tatei var. neblinae 424
Helictonema 56
Galax 121 H. velutinum 57
G. urceolata 119, 120 Helinus 330
Ganua 408 Hemandradenia 78
Gaultheria 182 Hemitomes 168
G.procumbens 150 Hesperomeles 381
G. suborbicularis 150 Heteromeles 378
Gaultherieae 181 Hexaspora 47
Gaylussacia 189 Hippocratea 57
G. braziliensis 150 Hippocrateaceae 29, 55
Geissanthus 276 Hippocrateoideae 55
Geissoieae 105 Hippophae 134
Geissois 105 H. rhamnoides 133
Gentlea 276 Holodiscus 356
Geum 367 Hooglandia 102
G. subg. Oncostylus 368 Horkelia 366
G. subg. Orthostylus 367 Horkeliella 366
Gilia 310 Hornemannia 183
Giliastrum 310 Hottonia 317
Gillbeea 104 Hovenia 327
Gillenia 358 Huidobria 248
Gillenieae 358 Hulthemia 370
Glaux 273 Hulthemosa 370
Gloveria 46 Huodendron 440
Gluema 406 H. biaristatum 436
Glueminae 405 Huthia 307
Glyptocarpa 468 Hybosperma 335
Glyptopetalum 46 Hydrangea 211
Goniodiscus 46 H. serrata 211
Gonocalyx 184 Hydrangeaceae 3, 202
Hydrangeae 211 Lamellisepalum 332

Hydrangeoideae 209 Langloisia 309
Hydrocera 24 Laplacea 469
H. triflora 23 Lasiodiscus 336
Hydrostachyaceae 3, 216 Lateropora 184
Hydrostachys 219 Lauridia 47
H. imbricata 219 Lebetanthus 174
H. multijida 217 L. americanus 174
Hylenaea 57 Lecardia 52
Hymenandra 275 Lecomtedoxa 405
Hypopitys 168 Lecythidaceae 5,~221
Hypsophila 47 Lecythidoideae 226
Lecythis 229
Ibarraea 275 L. corrugata 223
Icacorea 275 L. pisonis 223, 229
Impatiens 24 Ledothamnus 170
I. cinnabarina 24 L. atroadenus 151
I. jurpia 21 L. sessiliflorus 149, 155
I. niamniamensis 22 Ledum 171
I. stuhlmanii 22 Leiophyllum 172
Impatientella 24 L. buxifolium 155
Inhambanella 405 Leiospermum 108
I. henriquesii 406 Lepidobotryaceae 1, 233
Ipomopsis 309 Lepidobotrys 235
Isonandra 408 L. staudtii 234
Isonandreae 407 Leptecophylla 179
Ituridendron 419 Leptodactylon 310
Ivesia 366 Leptostylis 416
Lepuropetalaceae 291
jacquinia 476 Lepuropetalon 294
f. aculeata 477 L. spathulatum 294
]amesia 208 Letestua 404
]. americana 208 Leucopogon 179
Jamesioideae 208 Leucosidea 361
jollydora 79 Leucothoe 182
Jollydoreae 79 L. keiskii 150
Linanthastrum 310
Kaernbachia 109 Linanthus 310
Kageneckia 354 Lindleya 354
Kalmia 172 Lissanthe 179
K. buxifolia 155 Lissocarpa 237
Kalmiopsis 172 L. benthamii 237
Kantou 405 Lissocarpaceae 4, 7, 236
Karwinskia 333 Loasa 248
K. rocana 322 Loasaceae 3,239
Kaufmannia 318 Loaseae 247
Kelseya 356 Loasoideae 247
Kentrothamnus 328 Loeselia 309
Kerria 359 Loeseliastrum 309
Kerrieae 359 Loeseneriella 57
Killipiella 187 Loheria 278
Killipiodendron 458 Loiseleuria 172
Kirengeshoma 210 Lophopetalum 47
Kissenia 247 Luetkea 356
Klaprothia 249 Lydenburgia 48
K. fasciculata 240 Lyonia 181
Klaprothieae 249 L. foliosa 150
Kokoona 47 L. ovalifolia 181
Krugiodendron 334 Lyonieae 180
Kurrimia 41 Lyonothamnus 383
Lysiclesia 185
Labisia 280 Lysimachia 272
Labourdonnaisia 404 L. nemorum 268
Labramia 404 Lysinema 176
Lachemilla 371
Lamanonia lOS Maba 128
L. speciosa 105 Macgregoria 54
Macleania 186 M. sect. Vitellariopsis 400

Macromeles 378 Moneses 167
M. corymbifera 377 Monimopetalum 49
Maddenia 374 Monoporus 280
Madhuca 408 Monotheca 409
M. aristulata 396 Monotoca 179
M. pallida 396 Monotropa 168
Maesa 257 Monotropaceae 145
M. japonica 256 Monotropastrum 168
M. lanceolata 256 Monotropeae 167
Maesaceae 4, 7, 255 Monotropoideae 166
Maesopsideae 334 Monotropsis 168
Maesopsis 334 Moraceae 3
M. eminii 334 Mortonia 49
Magodendron 418 Maya 49
Malaceae 343 Mycerinus 186
Malacomeles 380 Myginda 43
Maleae 375 Myrsinaceae 4, 7, 266
Malus 378 Myrsine 274
M. subg. Chloromeles 379 M. africana 268
M. sect. Docyniopsis 378 Mystroxylon 49
M. subg.Malus 379
M. subg. Sorbomalus 379 Niigelia 380
Manilkara 402 Nagelocarpus 173
M. discolor 403 Napoleonaea 284
Manilkarinae 402 Napoleonaeaceae 5, 7, 282
Manoteae 79 Nasa 249
Manotes 79 N. aequatoriana 250
M. macrantha 80 N. argemonoides 250
Marcetella 363 N. humboldtiana 250
Marcgravia 264 N. rudis subsp. australis 250
M. serrae 260 N. triphylla 250
M. sintenisi 260 Naumburgia 272
M. umbellata 260 Navarretia 308
Marcgraviaceae 4, 7, 258 Needhamia 176
Marcgraviastrum 264 Needhamiella 176
Margyricarpus 364 Neillia 357
Mastichodendron 409 N. thyrsijlora 357
Mastixia 87 Neillieae 357
M. pentandra 88 Neohemsleya 410
Mastixiaceae 82 Neolemonniera 405
Maurocenia 48 Neomezia 477
Maytenus 48 Nesiota 330
M. magellanica 48 Nesoluma 410
Melchiora 458 Neviusia 359
Melichrus 179 Nicobariodendron 61
Melliodendron 441 Niemeyera 415
M. xylocarpum 436 No/tea 330
Menepetalum 48 Norantea 263
Mentzelia 251 N. subg. Byrsophyllum 264
M. scabra 251 N. subg. Cochliophyllum 263
Mentzelioideae 250 N. subsect. Marcgraviastrum 264
Menziesia 171 N. subg. Pseudostachyum 264
M. lasiophylla 149 N. subg. Sacciophyllum 263
Mespilus 382 Northia 404
Micromeles 375 Notopora 184
Micropholis 413 N. schomburgkii 150
Microrhamnus 333 Novosieversia 368
Microsteris 311 Nuttalia 372
Microtropis 49 Nyssa 86
Mimusopeae 399 N. sylvatica 84
Mimusopinae 400 Nyssaceae 82
Mimusops 400
M. sect. Baillonella 402 Oemleria 372
M. sect. Inhambanella 405 Oligarrhena 176
M. sect. Lecomtedoxa 405 Oligarrheneae 176
M. schimperi 396 Omphalocarpeae 418
Omphalocarpum 419 Pentaphylacaceae 450

Omphalogramma 317 Pentaphylaceae 455
Oncostemum 279,268 Pentaphylax 456
Oncostylus 368 P. euryoides 456
Opocunonia 106 Peraphyllum 380
Oreanthes 187 Periclesia 187
Oreoherzogia 332 Peripentadenia 141
Oreorhamnus 332 Peripterygia 50
Ornithostaphylos 169 Peritassa 59
Ornithrophus 108 Pernettya 182
Orthaea 185 Pernettyopsis 182
0. apophysata 150 Perrottetia 50
Orthilia 167 Petalonema 24
Orthosphenia 49 Petalonychoideae 252
Orthurus 367 Petalonyx 253
Osmaronia 372 P. linearis 253
Osteomeles 381 P. parryi 253
Otherodendron 49 Petersianthus 230
Oubanguia 428 Petrophytum 356
0. africana 428 Philadelpheae 209
Oxalidaceae 2,285 Philadelphus 210
Oxalis 288 P. coronarius 9
0. contracta 289 P. hirsutus 210
0. lasiopetala 287 P. inodorus 210
0. leptocaulos 289 Philippia 173
Oxycoccus 183 Phlox 311
Oxydendreae 180 P. bryoides 307
Oxydendron P. longiflorum 307
0. arboreum 150 Photinia 376
Oxydendrum 180 P. davidiana 377
Phylica 329
Pachistima 50 Phyliceae 329
Pachystela 416 Phyllodoce 172
Pachystima 50 P. aleutica 155
Palaquium 407 Phyllodoceae 171
P. quercifolium 396 Phyllogeiton 333
Paliureae 327 Physocarpus 357
Paliurus 327 Pichonia 414
Pamphilia 440 Pieris 180
Pancheria 107 P. nana 150
P. elegans 107 Pierrina 429
Paphia 185 Piquetia 468
Paracryphia 109 Pileostegia 212
Parageum 367 Pityopus 168
Paranneslea 456 Plagiospermum 374
Parapyrenaria 468 Plagiopteraceae 29
Parardisia 275 Plagiopteron 57
Parastyrax 441 P. suaveolens 58
Para thesis 277 Plakothira 250
P. cubana 268 Planchonia 231
Parnassia 294 Planchonioideae 226
P. alpicola 294 Planocarpa 180
P. palustris 294 Platycalyx 173
Parnassiaceae 1, 291 Platycrater 213
Patascoya 458 Platylophus 103
Paxistima 50 Platypterocarpus 50
Payena 408 Platytheca 143
P. endertii 396 Pleiomeris 274
Pellegrinia 187 Pleiosepalum 356
Pelletiera 273 Plenckia 50
Pelliciera 298 Pleuranthodes 330
P. rhizophorae 298,299 Pleuricospora 169
Pellicieraceae 4, 7,297 Pleurostylia 50
Pentachondra 179 Plutarchia 188
Pentactina 355 Polemoniaceae 4, 7,300
Pentamerista 461 Polemonioideae 308
P. neotropica 462 Polemoniella 308
Polemonium 308 Pyrus 379

Polycardia 51 P. sect. Eriolobus 379
Polyclita 186 Pyxidanthera 121
Polylepis 364
P. quadrijuga 364 Quadripterygium 45
Polyspora 469 Quetzalia 52
Pomaderreae 331 Quillaja 384
Pomaderris 331
Pomatosace 316 Rapanea 274
Porteranthus 358 Rehderodendron 441
Potaninia 383 Reissantia 58
Potentilla 366 Reissekia 330
Potentilleae 366 Retanilla 328
Poteridium 363 Reynosia 333
Poterium 363 Rhacoma 43
Pottingeria 61 Rhamnaceae 2, 320
Pottingeriaceae 29 Rhamneae 332
Pourthiaea 377 Rhamnella 333
Pouteria 412 Rhamnidium 333
P. baillonii 396 Rhamnus 332
P. caimito 396 Rhaphidanthe 128
P. sect. Chromolucuma 413 Rhaphiolepis 377
P. laurifolia 396 R. philippinensis 377
P. multiflora 396 Rhaptopetaloideae 429
P. ucuqui 396 Rhaptopetalum 429, 428
Pradosia 415 Rhododendroideae 170
Presliophytum 248 Rhododendron 171
Primula 317 R. fictolacteum 149
P. denticulata 318 R. lapponicum 155
P. subg. Omphalogramma 317 R. scabribracteatum 171
Primulaceae 4, 7, 313 R. tanakae 149
Prinsepia 374 R. tomentosum 155
Prionostemma 58 Rhodoreae 170
Prionoteae 174 Rhodothamnus 172
Prionotes 174 R. chamaecistus 149
Pristimera 58 Rhodotypos 359
Pruneae 372 Richea 177
Prunus 373 Richeeae 177
P. arborea 373 Robsonodendron 52
Psammisia 186 Roridula 341
Psammomoya 51 R. gorgonias 9, 340
Pseudoconnarus 80 Roridulaceae 4, 7, 339
Pseudocydonia 375 Rosa 370
Pseudosalacia 51 Rosaceae 2, 343
Pseudoweinmannia 105 Roseae 370
Ptelidium 51 Roucheria 287
Pterocelastrus 50, 51 Rourea 81
P. echinatus 51 Roureopsis 81
Pterophylla 108 Royena 128
Pterospora 167 Rubeae 369
Pterosporeae 167 Rubus 369
Pterostyrax 441 Rupicola 176
Pullea 106 Ruptiliocarpon 235
Purdiaea 72 Rusbya 188
P. nutans 70 Ruyschia 263
Purpusia 366 Rzedowskia 52
Purshia 360
Putterlickia 52 Sadiria 276
Pycnandra 416 Sageretia 332
P. aff. vieillardii 396 Salacia 60
Pygeum 373 S. subg. Dicarpellum 44
Pyracantha 380 Salacicratea 60
Pyrenaria 468 Salacighia 60
Pyrola 167 Salacioideae 59
Pyrolaceae 145 Salaciopsis 52
Pyroleae 166 Salaxidae 173
Pyroloideae 166 Salaxis 173
Salvadoropsis 52 S. celastrifolia 432

Samolaceae 4, 7, 387 Sladeniaceae 5, 7, 431
Samolus 388 Sloanea 139
S. ebracteatus 388 S. rhodantha 140
S. valerandi 388 Smythea 334
Sanguisorba 363 Soldanella 318
Sanguisorbeae 361 S. montana 318
Sapota sect. Micropholis 413 Solonia 276
Sapotaceae 5,7,390 Sorbaria 358
Sarawakodendron 52 Sorbus 375
Sarcaulus 415 S. subg. Aria 376
Sarcodes 167 S. subg. Chamaemespilus 376
Sarcopoterium 363 S. subg. Cormus 376
Sarcosperma 411 S. subg. Micromeles 376
Sarcospermataceae 390 S.subg.Sorbus 376
Sarcotheca 287 S. subg. Torminaria 376
S. glauca 288 Souroubea 263
Sarracenia 424 S. sympetala 9
Sarraceniaceae 4, 7, 422 S. vallicola 263
Satyria 186 Spenceria 362
Saurauia 18 Sphenotoma 177
Scandivepres 41 Sphyrospermum 188
Schaefferia 53 Spiniluma 409
Schellenbergia 79 Spiraea 355
Schima 469 S. sect. Chamaebatiaria 358
S. wallichii 469 S. 'Gruppe' Holodiscus 356
Schismocarpus 252 S. sect. Petrophytum 356
Schistocarpaea 336 S. sect. Physocarpus 357
Schizocodon 120 S. sect. Sorbaria 358
Schizomeria 103 S. subg. Spiraeanthus 358
Schizomerieae 103 Spiraeaceae 343
Schizophragma 212 Spiraeanthemeae 101
Schwartzia 263 Spiraeanthemum 101
Scutia 332 S. katakata 102
Scyphanthus 249 Spiraeanthus 358
Scyphogyne 173 Spiraeeae 355
Scytopetalaceae 5, 7, 426 Spiraeopsis I 06
Scytopetaloideae 428 Sprengelia 175
Scytopetalum 429 Spyridium 331
S. tieghemii 428 Sredinskya 317
Semialarium 58 Stackhousia 54
Semiramisia 187 Stackhousiaceae 29, 54
Sericolea 141 Stackhousioideae 54
Shepherdia 133 Stellariopsis 366
S. canadensis 132 Stenanthemum 332
Shortia 120 Stephanandra 358
S. galacifolia 120 Stewartia 469
Sibbaldia 366 S. ovata 470
Sibbaldiopsis 366 S. pseudocamellia 464
Sibiraea 355 S. sinensis 470
Sideroxyleae 409 Stimpsonia 272
Sideroxylon 409 Stokeanthus 173
S. celastrinum 396 Stutzeria 106
S. gladuliferum 396 Stylogyne 276
S. sect. Synsepalum 416 Styphelia 180
Siegfriedia 331 Styphelieae 177
Sieversia 367, 368, 368 Styphelioideae 174
Simicratea 59 Styracaceae 5,7,434
Simirestis 59 Styrax 440
Simocheilus 173 S. leprosus 436
Sinojackia 440 S. pallidus 436
Sinopyrenaria 468 S. ramirezii 436
Sinosideroxylon 409 S. redivivus 436
Siphonandra 187 Suttonia 2 74
Siphonodon 53 Swida 88
Siphonodontaceae 29 Symphysia 183
Sladenia 432 S. racemosa 150
Sympieza 173 Tristemonanthus 59

Symplocaceae 5,~443 Trochocarpa 180
Symplococarpon 458 Trymalium 331
Symplocos 448 Tsebona 418
S. subg. Hopea 448 Tsusiophyllum 171
S. ophirensis var. cumingiana 448 Tutcheria 468
S. rigidissima 9
S. subg. Symplocos 448 Ulmaceae 2
Synardisia 275 Ulmarieae 361
Syndesmanthus 173 Urticaceae 3
Synsepalum 416 Utleya 184
S. cerasiferum 417
Systellantha 280 Vacciniaceae 145
Vaccinieae 183
Taihangia 367 Vaccinioideae 174, 180
Tapeinosperma 279 Vaccinium 183
Telemachia 44 V. oxycoccus 183
Tepuia 183 V. poasanum 150
Ternstroemia 456 Vallea 139
Ternstroemiaceae 5, 7, 450 Vauquelinia 354
Ternstroemieae 456 V. californica 355
Ternstroemiopsis 457 Vegaea 279
Tetraclis 128 Ventilagineae 334
Tetraglochin 364 Ventilago 334
Tetramerista 462 V. viminalis 322
Tetrameristaceae 4, 7, 461 Vesselowskya 107
Tetrardisia 275 Vincentella 416
Tetrasiphon 53 Vismianthus 79
Tetratheca 142 Visnea 458
Thamnus 173 Vitaliana 316
Thea 468 Vitellaria 402
Theaceae 5, 7, 463 V. paradoxa 396
Themistoclesia 188 Vitellariopsis 400
T. compacta 150 V. kirkii 401
Theophrasta 477 Votschia 476
T. jussieui 477
Theophrastaceae 4, 7, 472 Waldsteinia 369
Therorhodion 170 Wallenia 278
Thibaudia 189 W. laurifolia 268
Thibaudieae 183 Weigeltia 278
Thoracosperma 173 Weinmannia 108
Thyrosalacia 60 W. heterophylla 108
T. pararacemosa 60 Whipplea 209
Tieghemella 401 Wimmeria 54
T. heckelii 396 Windmannia 108
Tontelea 60 Woollsia 176
Torminalis 375 Woronowia 367
Torralbasia 53
Trachyphytum 251 Xantolis 417
Tremandra 142 Xerospiraea 356
T. stelligera 143 Xylococcus 169
Tremandraceae 135 Xylonymus 54
Trevoa 328 Xylopodia 250
T. quinquenervia 328 X. klaprothioides 251
Tricerma 53
Trichocephalus 329 Yinquania 88
Tricuspidaria 140 Yunckeria 277
Tridesmostemon 419 Yunnanea 468
Trientalis 272
Trimorphopetalum 24 Zenobia 181
Tripetaleia 171 Z. pulverulenta 150
T. bracteata 155 Zinowiewia 54
Tripterococcus 54 Ziziphus 327
T. brunonis 55 Zygalchemilla 371
Tripterygium 53

[the Families and Genera of Vascular Plants 6] K. Kubitzki (Auth.), Professor Dr. Klaus Kubitzki (Eds.) - Flowering Plants · Dicotyledons_ Celastrales, Oxalidales, Rosales, Cornales, Ericales (2004, Springer-Verlag

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[the Families and Genera of Vascular Plants 6] K. Kubitzki (Auth.), Professor Dr. Klaus Kubitzki (Eds.) - Flowering Plants · Dicotyledons_ Celastrales, Oxalidales, Rosales, Cornales, Ericales (2004, Springer-Verlag

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THE FAMILIES

Springer-Verlag Berlin Heidelberg GmbH

Volume I Pteridophytes and Gymnosperms

Volume II Flowering Plants. Dicotyledons. Magnoliid, Hamamelid and Caryophyllid Families

Volume III Flowering Plants. Monocotyledons: Lilianae (except Orchidaceae)

Volume IV Flowering Plants. Monocotyledons: Alismatanae and Commelinanae (except

Volume V Flowering Plants. Dicotyledons: Malvales, Capparales and Non-betalain

Volume VI Flowering Plants. Dicotyledons: Celastrales, Oxalidales, Rosales, Cornales, Ericales

VI Flowering Plants · Dicotyledons

With 137 Figures

Library of Congress Cataloging-in-Publication Data

© Springer-Verlag Berlin Heidelberg 2004

Production: PRO EDIT GmbH, Heidelberg

Hamburg, October 2003 K. KUBITZKI

Introduction to Families Treated in This Volume

Actinidiaceae S. DRESSLER and C. BAYER 14

Celastraceae M.P. SIMMONS . . • . • . • . 29

Diapensiaceae P.J. SCOTT 117

Ebenaceae B. W ALLNOFER . . . . . . . . . 125

Fouquieriaceae K. KUBITZKI 195

Grubbiaceae K. KUBITZKI 199

Hydrangeaceae L. HUFFORD ••.•• 202

Lecythidaceae G.T. PRANCE and S.A. MoRI 221

Maesaceae B. STAHL and A.A. ANDERBERG 255

Napoleonaeaceae G.T. PRANCE . 282

Oxalidaceae A.A. Cocucci 285

Parnassiaceae M.P. SIMMONS 291

Rhamnaceae D. MEDAN and C. SCHIRAREND 320

Samolaceae B. STAHL .. 387

Ternstroemiaceae A.L. WEITZMAN, S. DRESSLER and P.F. STEVENS 450

Index of Scientific Names 479

ANDERBERG, A.A. Department of Phanerogamic Botany, Swedish Museum of

HUFFORD, L. School of Biological Sciences, Washington State University,

STACE,H.M. Department of Botany, University of Western Australia,

Introduction to Families Treated in This Volume

2. Oxalidales details of how the three tremandraceous genera

sometimes nectariferous (Brown 1935 for S. Actinidiaceae, Roridulaceae and Sarraceniaceae is

mentals in Europe and North America. In the

KEY TO THE GENERA

1. Trees or shrubs; stamens basally connate with petals; Asia

1. Actinidia Lindl. Fig. 3

Woody climbers, dioecious or polygamous, ± Fig. 3. Actinidiaceae. A, F, G Actinidia strigosa. B-E A.

2. Clematoclethra (Franch.) Maxim. Fig. 4

Woody climbers, deciduous, ± pubescent; flowers

Whereas most Impatiens have mechanisms to

EMBRYOLOGY AND SEEDS. Mature pollen is 2-

PHYTOCHEMISTRY. Impatiens spp. regularly con-

DISTRIBUTION AND HABITATS. Hydrocera triflora

KEY TO THE GENERA

1. Hydrocera Blume Fig. 7

Semi-aquatic perennial herbs, rather succulent;

Selected Bibliography Larsen, K. 1981. Chromosome numbers in Impatiens from

Fig. 9. Brunelliaceae. A Brunellia standleyana, part of flores-

REPRODUCTIVE STRUCTURES. Brunellia has

sutural stigmas, the dissociation of exocarp and Selected Bibliography

Brunellia Ruiz & Pavon Fig. 10

Description as for family. A single genus of 61

or angular, rarely lacerate or irregularly lobed;

weakly-supported clade, as part of an early- none) and d-norpseudoephedrine (cathine), are

Shrubs, glabrous; stems with dark glands, termi-

Scandent shrub, glabrous. Leaves opposite on

Fig. I I. Celastraceae-Celastroideae. Bhesa archboldiana. A

5-lobed; anthers longitudinally dehiscent, introrse 7. Brexiella H. Perrier

Scandent shrubs, glabrous, rarely pubescent.