Blackwell Science, LtdOxford, UKBOJBotanical Journal of the Linnean Society0024-4074The Linnean Society of London, 2005? 2005
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168
Original Article
GERANIUM SECTIONS ANDINA AND CHILENSIA
C. AEDO
et al.
Botanical Journal of the Linnean Society, 2005, 149, 1–68. With 88 figures
Taxonomic revision of Geranium sections Andina and
Chilensia (Geraniaceae)
CARLOS AEDO*, CARMEN NAVARRO and MARÍA LUISA ALARCÓN
Real Jardín Botánico, CSIC, Plaza de Murillo 2, E-28014 Madrid, Spain
Received April 2004; accepted for publication February 2005
The sections Andina and Chilensia of Geranium from South America, Australia and New Zealand are taxonomically
revised. Fruits with ‘seed ejection-type’ dispersal have been found in all species, which confirms their classification
within subgen. Geranium. Section Andina consist of four species, while sect. Chilensia comprises 11 taxa. In sect.
Andina, G. brevicaule and G. parodii, previously considered as synonyms or recognized as infraspecific taxa, are
accepted at species level. In contrast, G. tucumanum, G. sessiliflorum ssp. novaezelandiae and 17 other taxa have
been reduced to synonyms. In sect. Chilensia, G. patagonicum, G. fiebrigianum and G. drumondii have been considered as synonyms of G. berteroanum, G. fallax and G. solanderi, respectively. Additionally, another 24 taxa have
been reduced to synonyms. Diagnostic morphological features are analysed and compared within and between the
sections. A multivariate morphometric study from 258 specimens was carried out to test the most significant characters. The chromosome number of six species (five never previously counted) was established. A key, species descriptions, a complete list of synonymy, a list of specimens examined and distribution maps are provided. Most species are
illustrated for the first time. Thirty-seven lectotypes and one neotype are designated. © 2005 The Linnean Society
of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68.
ADDITIONAL KEYWORDS: Australia – chromosome number – New Zealand – South America.
INTRODUCTION
The genus Geranium L. comprises c. 400 species in
temperate areas and tropical mountains throughout
most of the world (Aedo, Muñoz Garmendia & Pando,
1998a). A brief history of the generic delimitation and
infrageneric classification, as well as a description of
the genus, can be found in Aedo (1996). In addition, a
key for subgenera and sections can be found in Aedo,
Aldasoro & Navarro (1998b) and Aedo et al. (1998a),
respectively.
According to the currently accepted classification
(Yeo, 1984), Geranium is divided into three subgenera:
subgen. Erodioidea (Picard) Yeo, subgen. Robertium
(Picard) Rouy, and subgen. Geranium. The subgen.
Erodioidea which includes 22 species in four sections
has recently been monographed (Aedo, 1996, 2001a):
two sections are in the Mediterranean and western
Asia, one is centred in the mountains of tropical East
Africa, and one is in southern Brazil and northern
Argentina. According to Yeo’s (1984) sectional classifi*Corresponding author. E-mail: aedo@ma-rjb.csic.es
cation, the subgen. Robertium comprises 30 species in
eight sections, two of them not yet revised (Yeo, 1973,
1992, 2004; Aedo et al., 1998b).
Geranium subgen. Geranium, the largest of the
genus, comprises over 370 species; grouped in at least
ten sections. Section Tuberosa (Boiss.) Reiche (Davis,
1970), sect. Azorelloida Aedo, Muñoz Garm. & Pando,
Neoandina Aedo and Paramensia R. Knuth (Aedo,
Aldasoro & Navarro, 2002), sect. Trygonium Dumort.
(Aedo, 2003), and sect. Gracilia R. Knuth (Aedo et al.,
2003) have already been revised. Most species are in
sect. Geranium, which probably will be subdivided
when a satisfactory knowledge of subgen. Geranium is
obtained. Knuth’s (1912, 1931) subdivision (32 sections) of the genus has been questioned by numerous
authors, though without advancing an alternative
until Yeo’s (1984) review. However, Knuth’s sections
should be reconsidered when a new classification of
the whole subgen. Geranium is undertaken.
Knuth (1912) described his sect. Andina for stemless species with 1-flowered cymules. In Aedo et al.
(2002) we proposed a new section Neoandina (to
include stemless species from the Andes without
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
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C. AEDO ET AL.
turnip-shaped rootstock and 1-flowered cymules)
including all species of sect. Andina but G. sessiliflorum (which is the type of sect. Andina). According to
this preliminary view both sect. Andina and sect.
Chilensia should be united under the second name
(Aedo et al., 2002). However, in this study we preferred to maintain sect. Andina for stemless species
with turnip-shaped rootstock and 1-flowered cymules,
and include in sect. Chilensia species with turnipshaped rootstock, 2-flowered cymules, and developed
aerial stem. Geranium brevicaule which was included
by Knuth (1912) as a synonym of G. sessiliflorum, is
accepted here, while G. bangii is synonymized with
G. sessiliflorum. We also included in section Andina
two more species described after Knuth’s (1912) monograph: G. antrorsum and G. parodii.
Knuth (1912) precisely described his sect. Chilensia
as ‘Radix incrassata, napiformis. Pedunculi bi-, rarius
uniflori’. He recognized 23 species in this section, most
of them based on the limited material available at that
time, or sometimes supported only by literature
records. From those taxa we accept here G. solanderi
and G. retrorsum (included by Knuth under
G. pilosum as different varieties), G. berteroanum,
G. limae, G. magellanicum, G. albicans, G. core-core,
G. skottsbergii (named by Knuth G. ciliatum). Additionally, we include G. fiebrigianum as a synonym of
G. fallax, which was considered by Knuth (1912) as a
doubtful synonym of G. tucumanum (in sect. Rupicola
R. Knuth). The remaining species accepted by Knuth
(1912) have been here considered synonyms, mainly
of G. berteroanum and G. core-core. Later, Knuth
described two species treated here: G. tablasense
(Knuth, 1915) without any sectional assignation, and
G. venturianum (Knuth, 1936) as belonging to sect.
Palustria R. Knuth.
In addition to Knuth’s (1912) monograph two contributions concerning sections Andina and Chilensia
are relevant. Carolin (1965) studied the genus in
the South-west Pacific, and included three species
in his group IV: G. retrorsum, G. solanderi and
G. drummondii and two in his group V:
G. sessiliflorum (ssp. novaezelandiae and ssp. brevicaule) and G. antrorsum. We mainly agree with this
taxonomy, although we consider G. drummondii as a
synonym of G. solanderi, and accept G. brevicaule at
specific level (including G. sessiliflorum ssp. novaezelandiae). Barboza (1996) studied the genus in
Argentina. She accepted G. albicans, G. core-core,
G. venturianum, G. berteroanum, G. magellanicum
and G. sessiliflorum, with which we are in agreement. She also recognized G. patagonicum and
G. tucumanum which are here considered as
synonyms for G. berteroanum and G. sessiliflorum,
respectively. She did not recognize G. fallax in the
Argentinean flora but partially identified these speci-
mens as G. patagonicum, and included G. parodii as a
variety of G. magellanicum.
The large size of the genus in South America makes
it unfeasible to study Geranium as a single unit. The
two sections comprise a reasonable number of species
(15) for investigation. The species of this group have
the ‘seed ejection-type’ of fruit discharge, and a turnipshaped rootstock. They are mainly distributed in
southern South America, Australia and New Zealand.
Thus, following recent revisions of selected Geranium
groups (Aedo, 1996, 2001a, 2003; Aedo et al., 1998b,
2002, 2003), and in pursuit of a comprehensive monograph of the genus, we present here a revision of section Andina and Chilensia.
MATERIAL AND METHODS
This revision is based on 2008 herbarium specimens
from the following herbaria: AAU, AK, B, BAF, BH,
BM, BOLV, BR, C, CANB, CAS, CHR, CONC, CORD,
CTES, E, F, FI, G, GH, GOET, H, HAL, HBG, HO, JE,
JEPS, K, L, LD, LE, LIL, LP, LPB, M, MA, MERL,
MICH, MO, MPU, NSW, NY, P, PERTH, PH, RSA, S,
SGO, SI, SYD, U, UB, UC, UPS, US, W, WELT, WRSL,
WU and Z (see the Appendix).
Curators from BA, BO, GB, KIEL, LY, PAD and TO
kindly responded to our request, but they either did
not find any of the requested specimens in their herbaria or sent some digital images.
For scanning electron microscopy (SEM), samples
were glued to aluminium stubs, coated with 40–50 nm
gold, and examined with an ITACHI S3000N scanning
electron microscope at 18 kV.
Two hundred and fifty-eight specimens were used as
operational taxonomic units (OTUs). They were distributed as follows: G. antrorsum (12), G. brevicaule
(22), G. sessiliflorum (18), G. magellanicum (17),
G. parodii (12), G. limae (22), G. berteroanum (19),
G. solanderi (24), G. skottsbergii (15), G. core-core (21),
G. retrorsum (19), G. albicans (16), G. venturianum
(19), G. tablasense (4) and G. fallax (18). Unfortunately, G. tablasense is a restricted endemic species
poorly represented in the herbaria. However, we considered the sample sufficient for the purpose of this
study because it showed low variability.
In these specimens 28 quantitative characters were
measured using a Mitutoyo CD-15DC digital calliper,
and six ratios derived (Table 1). Each character was
analysed for its mean value, range, standard deviation
and significance using the STATISTICA package. To
represent the variability of each descriptor within species box-plots were prepared. These plots contained
medians and percentiles and were calculated using
the STATISTICA package. Schemes of characters used
to describe the leaf lamina and fruit are shown in
Figures 1 and 2, respectively.
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
DA1
Root 1
Plant height
Rootstock diameter
Stem diameter
Stem indumentum length
Segment width at the base
Segment width at the base/
segment length ratio
Main sinus length/main
segment length ratio
Second sinus length
Second sinus length/middle
segment length ratio
Maximum width/segment
length ratio
Segment lobes number
Stipule length
Petiole indumentum length
Bracteole length
Peduncle length
Pedicel length
Pedicel indumentum length
Sepals length
Sepal width/sepal length ratio
Mucro length
Mucro length/sepal length ratio
Sepals indumentum length
Petal length
Filament length
Anthers length
Gynoecium length
Fruit length
Mericarp length
Mericarp indumentum length
Rostrum length
Rostrum narrowed apex length
Rostrum indumentum length
Stigmatic remains length
Seed length
Eigen values
Total cumulative proportion
DA2
Root 2
Root 3
Root 1
-0.4123
-0.5638
-0.15698
-1.04432
-0.36929
-1.05164
-0.66594
0.54462
0.7883
-0.62298
-0.5263
-0.05411
1.33508
0.22161
DA3
Root 2
DA4
Root 1
Root 2
0.11652
0.2062
0.26027
0.30061
-0.0999
0.17003
0.1023
-0.2916
-0.1072
0.0391
0.7396
0.7092
0.694806
0.000749
0.232317
0.324868
0.26204
0.36399
0.1175
0.116438
-0.11424
-0.2164
-0.422419
-0.41378
-0.225
0.205198
0.3338
0.70739
0.23071
-0.8666
-0.01545
0.6403
-0.72181
-0.36895
0.2636
-0.08555
Root 3
Root 1
Root 2
-0.62897
1.63695
0.18968
1.66388
2.63491
-0.91708
0.68860
0.40489
0.85805
1.52890
2.20175
0.22849
0.08615
0.21760
-0.6579
0.07113
0.01589
0.57828
-0.42914
0.20358
-0.2452
0.083103
-1.66292
1.14659
-0.6689
0.5408
0.1643
0.3651
0.5160
0.9550
-0.9922
0.71058
0.24684
-0.45606
-0.43729
-1.18370
1.98610
-0.92374
-0.03404
-1.20622
1.37008
-0.95225
0.73328
-0.09041
-1.11325
0.46066
0.13966
0.19307
1.00342
0.13714
-0.29173
-0.08006
0.09162
-0.13715
0.29935
-0.32514
-1.00694
0.9616
0.09021
0.58256
-0.56797
0.04302
0.19848
-0.07897
0.21161
0.32183
0.34455
-0.40032
0.4506
-0.1276
-0.5332
-0.2428
0.1732
-0.3025
0.5153
-0.073476
0.01727
-0.425895
0.14898
-0.339816
-0.067035
0.037232
-0.12866
-0.49863
-1.17257
2.14319
0.20923
1.17396
0.35918
0.39129
0.43777
-0.69965
-1.13316
0.10104
0.92213
0.26579
0.35621
0.47439
-0.2099
-0.62543
0.06108
0.67617
0.4567
0.355175
-0.76174
-0.24096
-0.95354
1.13534
-0.4625
-0.5315
-0.1138
-0.0314
0.352
-0.4891
-0.7519
0.2466
0.3351
0.5562
0.331241
0.091514
0.330736
-0.180518
-0.24815
0.138551
-0.824957
-0.340464
0.202277
0.899789
0.1173
0.3478
-0.2007
6.6613
0.8525
0.178555
0.475094
0.008265
2.822998
1
0.11448
0.30293
-1.17235
-0.68674
-0.42767
-0.48717
-1.03937
1.95348
-0.16345
-0.44097
-2.54130
-2.31785
0.10335
36.46717
0.65243
-1.28503
0.35765
-1.17304
0.70108
-0.93488
1.45223
-0.46177
1.32124
-1.25432
0.42061
0.14402
-0.01565
-0.07671
19.42735
1
0.1481
0.3382
-0.5392
0.7005
0.8457
0.0388
3.5766
-0.2189
0.4385
-3.9618
1.7574
0.1135
0.1505
0.0479
123.9878
0.7055
0.37398
0.16065
-1.00800
-0.67213
0.75651
1.35594
4.52796
-0.75266
0.19180
-5.36268
-0.49508
0.29290
0.19515
1.54036
28.93481
0.87015
0.52588
-1.24594
-1.66003
-0.64995
0.47374
1.91191
-1.48949
0.19561
-0.55117
2.34770
0.15644
0.50800
0.49950
0.54997
18.13762
0.97335
-0.40459
-0.02262
0.03521
-0.06682
0.04942
-1.80264
0.51719
-0.00678
0.93351
-0.24807
-0.69179
0.95557
-0.55921
-0.35191
1.67353
-0.14956
-0.05502
-0.87705
-0.14765
-0.20452
0.27895
0.15956
-0.14657
-1.00646
-0.54845
0.06351
0.44305
0.31924
-0.02051
-0.30493
0.41139
6.07624
0.63707
0.11876
-0.35096
-0.70379
3.46148
1
-0.67952
0.49714
-0.01419
9.65938
0.50457
GERANIUM SECTIONS ANDINA AND CHILENSIA
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
Table 1. Standardized coefficients obtained in discriminant analyses (DAs) for canonical variables
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C. AEDO ET AL.
Figure 1. Scheme of characters used to describe the leaf
lamina. A, lamina length; B, lamina width; C, segment
length; D, main sinus length; E, second sinus length; F,
maximum width; G, segment width at the base.
To verify the significance of quantitative key
characters, Kruskal–Wallis ANOVA and Median tests
were carried out using the STATISTICA package. In
each step of the key two groups were built assembling
all measured specimens belonging to the species
under that entry. Thus, the Kruskal–Wallis test was
achieved on these two groups.
All taxa were differentiated using a combination of
qualitative and quantitative characters. This separation was tested by means of discriminant analysis
(DA). This method requires the a priori assignment of
OTUs to groups, and allows the determination of
whether the recognized groups are statistically definable entities or if there is too much variation within
groups to allow classification (Sneath & Sokal, 1973).
For DA the raw matrix was obtained, the results
sorted in discrete groups and calculations carried out
using the STATISTICA package. As DA is indicated
where there are a few close groups (Sneath & Sokal,
1973), we performed four different DAs to test the
most similar species: (1) sect. Andina (four species)
plus G. magellanicum; (2) G. limae, G. berteroanum
and G. solanderi; (3) G. skottsbergii, G. core-core,
G. retrorsum and G. albicans; (4) G. venturianum,
G. tablasense and G. fallax.
In order to determine chromosome number, seeds of
available species were germinated and transferred
into pots for cultivation. Approximately two months
later, young root tips were cut and incubated for 24 h
in distilled water at 4 ∞C, then fixed with a mixture
of 25% acetic acid and 75% ethanol, stained with
aceto-orcein, squashed and counted.
TAXONOMIC CHARACTERS
DURATION
Figure 2. Scheme of characters used to describe the fruit.
A, fruit length; B, rostrum length; C, rostrum narrowed
apex length; D, stigmatic remains length; E, mericarp
length; F, mericarp width.
Quantitative and qualitative characters are used in
the key, being the discriminant quantitative characters inferred from box-plots. Several interesting characters have broad ranges of variability which cause
some difficulties for their use. In order to avoid these
problems the ranges are included in parentheses in
the key and descriptions (even considering that in
some rare cases they differed considerably from the
mean values). The most frequent and useful values
are given as percentiles and are shown outside the
parentheses. Some overlapping was impossible to
avoid at least in some cases.
AND HABIT
All species of Geranium sect. Andina and Chilensia
are perennial herbaceous plants. They share a ± vertical, turnip-shaped rootstock, from which thin roots
arise. Only three species have ± thick and fleshy,
fusiform
roots:
G. fallax,
G. tablasense
and
G. venturianum. All species of sect. Chilensia have a
decumbent to erect, leaved stem, while plants of sect.
Andina are stemless.
INDUMENTUM
In the species studied here three trichome types have
been found, all of them simple and uniseriate
(Theobald, Krahulik & Rollins, 1979): (a) eglandular,
unicellular hairs of variable length (0.1–2.2 mm),
sometimes with an ornamented surface (Figs 3, 4).
According to Payne (1978) they could be included in
the ‘subulate’ type. They have been found in all species, widespread for all organs of the plant; (b) glan-
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
GERANIUM SECTIONS ANDINA AND CHILENSIA
3
5
cies: G. berteroanum, G. skottsbergii, G. venturianum,
G. fallax and G. tablasense (Fig. 3). In the three first
species the presence of glandular hairs varies. Some
specimens have only eglandular hairs while others
have also glandular hairs widespread on the stem,
inflorescence, sepals, rostrum and/or mericarps. In
contrast, in all studied specimens of G. fallax and
G. tablasense there are glandular hairs; (c) short
glandular hairs (<40 mm long), smooth, usually constituting two cells, although they sometimes have a
bicellular foot (Figs 4, 5). They are present in all species studied here but they are not included in the
descriptions because they are only evident at high
magnification.
4
LEAVES
5
The leaves in all species of sect. Andina and sect.
Chilensia are polygonal in outline, cordate, palmatifid,
with 5–7 segments. Two exceptions are found in sect.
Andina: some specimens of G. sessiliflorum showed a
leaf rounded in outline, and in G. antrorsum the leaf
base is cuneate to subtruncate. Basal leaves are in a
rosette, usually deciduous in sect. Chilensia and persistent in sect. Andina. Cauline leaves are opposite in
all species of sect. Chilensia. The middle segment is
rhombic in G. albicans, G. venturianum, G. fallax and
G. tablasense, while it is obtriangular in the remaining species of sect. Chilensia. In sect. Andina it is
usually obtriangular with the exception of some
specimens of G. sessiliflorum and G. parodii. Leaves
are particularly divided in G. skottsbergii because it
has narrow lobes and a deep secondary sinus.
INFLORESCENCE
Figures 3–5. Scanning electron micrographs showing
indumentum features of Geranium sect. Andina and sect.
Chilensia. Fig. 3. Eglandular (smooth) and glandular hairs
on sepals of G. fallax (Beck 21256, MA). Fig. 4. Eglandular
hairs (with an ornamented surface) and short glandular
hairs on pedicels of G. core-core (Aedo 7073, MA). Fig. 5.
Short glandular hairs on sepals of G. fallax (Beck 21256,
MA).
All species of sect. Chilensia have a monochasial cyme
with 2-flowered cymules arising along leafy stems.
However, in G. magellanicum cymules of the basal
part of the stem are usually 1-flowered (parts of them
arise directly from the rootstock) and cymules of the
upper part 2-flowered. Peduncles of G. venturianum
are noticeably longer, reaching 25 cm long in some
cases. In sect. Andina, 1-flowered cymules arise
directly from the rootstock. In some cases they are
bibracteate, with a short peduncle, while in others the
pedicel is borne directly from the rootstock. Some
exceptions are discussed under G. parodii and
G. sessiliflorum.
CALYX
dular hairs of variable length (0.3–1.4 mm), usually
with 2–4 cells, smooth, the foot consisting of cylindric
cells or of decussated cells; they are absent from sect.
Andina, and in sect. Chilensia restricted to five spe-
Sepals are smooth, not accrescent and 3-nerved in all
species of the section. The most taxonomically significant sepal features are the outline and the mucro
length. In G. core-core sepals are broadly ovate (with a
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
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C. AEDO ET AL.
high width/length ratio) whereas the remaining species of the sections Andina and Chilensia have lanceolate sepals (with a low width/length ratio). The
mucro is particularly long in G. antrorsum, and to a
lesser extent, in G. brevicaule. Among species of sect.
Chilensia, G. venturianum and G. tablasense show the
longer mucro and G. albicans and G. limae the shorter
ones.
versal veins at the apex) and brownish in all species
(except in G. fallax and G. solanderi that are usually
blackish). The rostrum abruptly ends in the stigmatic
remains except in three species of sect. Chilensia
(G. fallax, G. tablasense and G. venturianum), and in
one of sect. Andina (G. parodii), which have a narrowed apex. The stigmatic remains are usually glabrous, and they are noticeably long in G. tablasense
and G. venturianum.
COROLLA
The corolla of all species of sections Andina and
Chilensia are actinomorphic. The petals usually have
an entire or slightly notched apex, but G. tablasense
shows a short notch. Geranium tablasense is also
unique in having a short claw. In most species of both
sections petals are glabrous on both sides and ciliate
on the basal margin. However, G. parodii and
G. venturianum are hairy on the base of the adaxial
side. Geranium sessiliflorum, G. brevicaule and
G. tablasense usually have glabrous petals, although
in some specimens scattered cilia can be found on the
basal margin. Finally, G. antrorsum has glabrous
petals.
STAMENS,
POLLEN AND NECTARIES
In Geranium the ten stamens are arranged in two
whorls. The filaments of sect. Andina and Chilensia
are not exerted, and lanceolate. They are ciliate at
least at the base, and in some species hairy on the
abaxial side.
Pollen of all species studied here, as in almost the
entire genus (Weber, 1996; Aedo et al., 2002, 2003), is
characterized by reticulate exine ornamentation with
distinctly baculate, clavate or gemmate supratectal
elements (Figs 6–11).
The five hemispherical nectaries are arranged alternately to the external whorl of staminal filaments.
They are glabrous except in G. limae and G. parodii,
which usually exhibit a tuft of hairs at the top of each
nectary.
FRUIT
Geranium sect. Andina and sect. Chilensia are
assigned to subgenus Geranium, which exhibits the
‘seed ejection-type’ of fruit discharge (Yeo, 1984). In
this type a single seed is actively discharged by the
explosive recurvature of the awn of the fruit, which
remains together with the mericarp, attached to the
columella. Mericarps have a basal callus, on which are
arranged some hairs that prevent the seed from dropping during the pre-explosive interval (Figs 12, 13).
Mericarps have a similar size and are smooth (except
in some specimens of G. parodii which have 1–2 trans-
SEEDS
The seeds are more or less ellipsoid in the three sections. The seed-coat appears finely reticulate at a magnification of 30¥, but SEM shows a reticulate surface
due to the prominence of the outer and middle layers
of the outer integument (Figs 14–19). The outer layer
has cells with thickened walls and collapsed lumina,
forming a polygonal structure. The seed-coat is usually brownish and bears scattered stomata.
Carolin (1965) characterized G. retrorsum by its
seeds with ‘isolateral alveolae’ and G. solanderi by its
seeds with ‘large alveolae’. These species, and the
remaining of sections Chilensia and Andina, show
considerable variation in the form of cells of the seedcoat (Figs 18, 19). A broad range of variation from
square to rectangular cells can be found in the same
species, suggesting that this feature has little taxonomic value.
According to Corner (1976), seeds of Geraniaceae
have cells which contain solitary crystals and tannin
in the inner layer of the outer integument. These crystal cells are also present in the endotesta of Hypseocharis Remy and Oxalidaceae (Boesewinkel, 1988).
According to Boesewinkel & Been (1979) crystals
extend as small protrusions beyond the so-called
light-line, and reach the coat surface of the seed
(Figs 20, 21). They form bipyramidal figures produced
by calcium oxalate trihydric crystals (Netolitzky,
1926). In Geranium sect. Chilensia crystals are
present in all species. In contrast, among species
of sect. Andina, crystals are absent on the seed
coat (G. antrorsum, G. brevicaule, G. sessiliflorum)
(Fig. 22) or they are very rare (G. parodii). A preliminary study of other groups of Geranium indicates that
they are present in subgen. Erodioidea (G. reflexum
L.), as well as in other sections of subgen. Geranium
(G. rotundifolium L.). However, we found no crystal
on the seed coat of species of subgen. Robertium
(G. robertianum L.; G. macrorrhizum L.; G. reuteri
Aedo & Muñoz Garm.).
The cotyledons are always conduplicate, one half of
each cotyledon lying in the primary fold of the opposite
cotyledon. They have entire margins, truncate bases
and short petioles, as found in sect. Neoandina (Aedo
et al., 2002).
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
GERANIUM SECTIONS ANDINA AND CHILENSIA
6
7
8
9
10
11
7
Figures 6–11. Scanning electron micrographs showing pollen features of Geranium sect. Andina and sect. Chilensia. Fig.
6. Pollen grain of G. antrorsum (Woodward s.n., SYD). Fig. 7. Pollen grain of G. berteroanum (Aedo 7391, MA). Fig. 8. Pollen
grain of G. brevicaule (Morris 86146, HO). Fig. 9. Pollen ornamentation of G. core-core (Aedo 7006, MA). Fig. 10. Pollen
ornamentation of G. core-core (Aedo 7197, MA). Fig. 11. Pollen ornamentation of G. tablasense (Cárdenas 758, US).
CHROMOSOME
NUMBER
The only species of sect. Andina counted is
G. sessiliflorum (Table 2). Three counts showed
2n = 56 (one in this study and two from the literature),
and one n = 26. The last number was attributed to
G. tucumanum by Barboza (1983). This species has
been considered in this study as a synonym of
G. sessiliflorum, and the voucher of this count was
reidentified as G. sessiliflorum. Unfortunately, neither
Gauger (1937) nor Warburg (1938) indicated the origin
of their specimens, and no vouchers of these counts
were kept. Thus, some doubts about the identity of
these plants may exist, especially if we consider that
G. sessiliflorum ssp. novaezelandiae (here considered
as G. brevicaule) is commonly cultivated in European
gardens.
Six species of Geranium sect. Chilensia have
been counted, five in this study and one by Barboza
(1983) (Table 2). Barboza (1983) reported n = 14 for
G. albicans, three numbers have been found for
G. berteroanum (2n = 52, 2n = 56 and 2n = 84), one
for G. core-core (2n = 56), G. magellanicum (2n = 112),
and
G. solanderi
(2n = c. 24)
and
two
for
G. skottsbergii (2n = c. 39, 2n = 52).
Among species of Geranium the most common chromosome number is 2n = 28, particularly in subgen.
Geranium and subgen. Erodioidea. Yeo (1984) considered x = 14 as a secondary basic number, while Van
Loon (1984b) accepted x = 14 as the main basic chro-
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
8
C. AEDO ET AL.
12
13
Figures 12–13. Scanning electron micrographs showing
mericarp features of Geranium sect. Andina and sect. Chilensia. Fig. 12. Lateral view of a mericarp of G. skottsbergii
(Aedo 6838, MA). Fig. 13. Basal callus of a mericarp of
G. limae, showing hairs which prevent the seed from dropping during the pre-explosive interval (Goodspeed et al.
11526, UC).
mosome number in the genus. According to the last
view, the number 2n = 56 found in some species of sect.
Andina and Chilensia would correspond to tetraploids, 2n = 84 found in G. berteroanum to hexaploids,
and 2n = 112 found in G. magellanicum to octoploids.
The number 2n = 84 was reported previously only in
G. sanguineum L. (Van Loon, 1984a). It is a European
species also belonging to subgen. Geranium, which
seems to be no relation to G. berteroanum. The number of G. magellanicum is unusually high in Geranium. Only in two species of the subgen. Robertium
have higher chromosome numbers been recorded:
2n = 128 (Yeo, 1973).
The number 2n = 52, which has been found in
G. berteroanum, G. skottsbergii and G. sessiliflorum
seems to be restricted to American species of Geranium subgen. Geranium, and would be a tetraploid
level for a basic number x = 13. It has been reported
for six species from North America (Shaw, 1952;
Faasen & Nadeau, 1976; Taylor & Taylor, 1977; Löve &
Löve, 1982; Ward, 1984), three species from Mexico
(Beaman, De Jong & Stoutamire, 1962; Seavey, 1975),
one species from Peru (Huynh, 1965). All of those
species belong to subgen. Geranium, although their
relationships with species of section Chilensia and
Andina remain unknown. Chatterjee & Sharma
(1970) reported n = 26 in a specimen of G. robertianum
from India. However, Van Loon (1984b) exhaustively
studied this species (128 specimens counted), and
found 2n = 64 in all the cases. This suggests that the
report of Chatterjee & Sharma (1970) should be confirmed. The number 2n = c. 24 has been found in
G. solanderi. In this species a more precise count
should be made in order to determine the exact chromosome number. However, a basic number x = 12 or
Table 2. Chromosome counts in Geranium sect. Andina and sect. Chilensia. Digital images of the vouchers from CORD
were studied
Section
Species
Andina
G. sessiliflorum
G. sessiliflorum
G. sessiliflorum
G. sessiliflorum
G. albicans
G. berteroanum
G. berteroanum
G. berteroanum
G. berteroanum
G. core-core
G. core-core
G. magellanicum
G. solanderi
G. skottsbergii
G. skottsbergii
Chilensia
n
2n
56
56
26
56
14
52
56
84
84
56
56
112
c. 24
c. 39
52
Origin
Voucher
Source
cultivated
unknown
Argentina, Salta
Chile, Biobío
Argentina, Córdoba
Chile, Maule
Chile, Región Metropolitana
Chile, Biobío
Chile, Magallanes
Chile, Antofagasta
Chile, O’Higgins
Chile, Magallanes
New Zealand, Chatham Is
Chile, Valparaíso
Chile, Coquimbo
unknown
unknown
Barboza 13 (CORD)
Aedo 7500 (MA)
Barboza 9 (CORD)
Aedo 7154 (MA)
Aedo 6737 (MA)
Aedo 6913 (MA)
Aedo 7459 (MA)
Aedo 7006 (MA)
Aedo 7073 (MA)
Aedo 7445 (MA)
Lange 225 (AK)
Aedo 6821 (MA)
Aedo 6832 (MA)
Gauger (1937)
Warburg (1938)
Barboza (1983)
this study
Barboza (1983)
this study
this study
this study
this study
this study
this study
this study
this study
this study
this study
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
GERANIUM SECTIONS ANDINA AND CHILENSIA
14
15
16
17
18
19
20
21
22
9
Figures 14–22. Scanning electron micrographs showing seed features of Geranium sect. Andina and sect. Chilensia.
Fig. 14. Seed of G. core-core (Aedo 7073, MA). Fig. 15. Seed of G. brevicaule (Hynes s.n., AK). Fig. 16. Seed of G. solanderi
(Davis 8059, PERTH). Fig. 17. Seed coat of G. sessiliflorum (Aedo 7486, MA). Fig. 18. Seed coat of G. solanderi (Lange s.n.,
AK). Fig. 19. Seed coat of G. solanderi (Mueller s.n., U). Fig. 20. Seed coat of G. solanderi, showing a group of crystals
(Davis 8059, PERTH). Fig. 21. Seed coat of G. albicans, showing two crystals (Sparre 209, S). Fig. 22. Seed coat of
G. antrorsum (Woodward s.n., SYD).
x = 13, would be preliminarily attributed to this
species. In Geranium asphodeloides Burm. f. and
G. sylvaticum L. the number 2n = 24 was also counted
(Van Loon & Oudemans, 1982; Van Loon, 1984a; Baltisberger, 1991; Dmitrieva, 1986, respectively). Both
Old World species belong to different groups of subgen.
Geranium, apparently not related to species of section
Chilensia and Andina. The number 2n = c. 39 would
be a triploid for a basic number x = 13. Triploid are
rare in Geranium, and are reported in some Old World
species of subgen. Geranium (G. deprehesum (E.G.
Almq.) Lindm.; Warburg, 1938; G. platypetalum Fisch.
& C.A. Mey.; Gauger, 1937, Warburg, 1938) all of them
based in x = 14. Additionally, Van Loon (1984a) found
2n = 28 and 42 in G. macrostylum Boiss. The plants
with the chromosome number 2n = 42 had a high percentage of small, abnormal pollen grains. The case of
G. skottsbergii would be somewhat similar, but plants
with 2n = c. 39 show normal pollen grains.
HABITAT
AND DISTRIBUTION
Geranium sect. Andina comprises four species, two
from South America and two from Australia. Geranium sessiliflorum occurs along the Andes from central Peru to Patagonia over more than 5000 km.
G. parodii shows an allopatric distribution occurring
only in Sierra de Achala and neighbouring areas (central Argentina). On the other side of the Pacific Ocean,
G. brevicaule is present in New Zealand, Tasmania
and south-east Australia, whereas G. antrorsum is
restricted to the Australian Alps.
Geranium sect. Chilensia comprises 11 species, nine
from South America and two from Australia. Geranium core-core grows along the Andes from central
Ecuador to Patagonia over more than 5400 km; it is
also introduced in some temperate areas of Europe
and North America. In addition, four more or less allopatric groups of species could be recognized: (1) species
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
10
C. AEDO ET AL.
Figure 23. Box-plot representing altitude variation in species of Geranium sect. Andina and sect. Chilensia.
of coastal ranges of Peru (G. limae); (2) species of
south-east Peru, Bolivia and northern Argentina
(G. fallax, G. venturianum and G. tablasense); (3) species of north-east Argentina, Uruguay and southern
Brazil (G. albicans); and (4) species of central and
south Chile and neighbouring areas of Argentina
(G. berteroanum, G. magellanicum and G. skottsbergii). In Oceania, G. retrorsum and G. solanderi
grow sympatrically. Both species are found in southwest Australia, south and south-east Australia, Tasmania and New Zealand.
Species from sect. Andina grow in open habitats
(dunes, river shores, turfy bogs, grasslands, puna or
open forests) from sea level to c. 5000 m (Fig. 23).
Species of sect. Chilensia are common in open forest or shrublands, although they can also be found
in more exposed habitats such as roadsides, cultivated grounds, dunes, meadows, river shores or
rocky and wet slopes, from sea level to c. 4000 m
(Fig. 23).
MORPHOMETRIC ANALYSIS
Box-plots showing the variability of the 34 most discriminant characters are shown in Figures 24–57.
These characters were used to perform the discriminant analyses (see below), and the most operative ones
to build the key of the species (in conjunction with
qualitative characters).
In Table 3 are shown the results of the Kruskal–
Wallis ANOVA and Median tests for quantitative
characters used in the key of species. The groups
tested were significantly different at 99% in all cases
but three. In step 5, the character petal length was
significant at 95% in the Median test, although it
was significant at 99% in the ANOVA test. This step
was also supported by a qualitative character of petal
indumentum, and another quantitative character,
rostrum narrowed apex length, which was highly significant in both tests. In step 14, sepal mucro length
was significant at 95% in the Median test, although
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
Plant height (cm)
Stem diameter (mm)
albicans
albicans
limae
limae
limae
berteroanum
berteroanum
berteroanum
skottsbergii
solanderi
solanderi
solanderi
venturianum
venturianum
tablasense
venturianum
tablasense
tablasense
fallax
fallax
fallax
magellanicum
magellanicum
antrorsum
magellanicum
antrorsum
brevicaule
brevicaule
brevicaule
sessiliflorum
sessiliflorum
sessiliflorum
parodii
parodii
parodii
antrorsum
sessiliflorum
parodii
sessiliflorum
parodii
sessiliflorum
antrorsum
parodii
26
brevicaule
24
brevicaule
22
brevicaule
fallax
20
magellanicum
18
fallax
magellanicum
antrorsum
16
magellanicum
antrorsum
tablasense
fallax
14
venturianum
tablasense
12
solanderi
solanderi
venturianum
tablasense
8
limae
berteroanum
10
albicans
6
core-core
retrorsum
4
skottsbergii
2
0
2.4
2.2
2.0
1.8
1.6
1.4
1.2
berteroanum
1.0
limae
0.8
limae
berteroanum
0.6
albicans
0.4
retrorsum
albicans
0.2
retrorsum
0.0
–0.2
0.45
0.40
0.35
0.30
0.25
0.20
0.15
0.10
0.05
0.00
core-core
skottsbergii
core-core
solanderi
venturianum
Rootstock diameter (mm)
Stem indumentum length (mm)
11
Median
25%-75%
Non-Outlier Range
Outliers
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Median
25%-75%
Non-Outlier Range
Outliers
GERANIUM SECTIONS ANDINA AND CHILENSIA
Median
25%-75%
Non-Outlier Range
Outliers
Median
25%-75%
Non-Outlier Range
Outliers
Ratio segment width at the base/middle segment length
skottsbergii
120
albicans
100
retrorsum
80
retrorsum
60
core-core
retrorsum
40
core-core
20
skottsbergii
core-core
0
–20
5.0
4.5
4.0
3.5
3.0
2.5
2.0
1.5
1.0
0.5
0.0
–0.5
14
12
10
8
6
4
2
0
skottsbergii
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Figures 24–57. Box-plots representing the variability of quantitative characters in Geranium sect. Andina and sect.
Chilensia.
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
Segment width at the base (mm)
C. AEDO ET AL.
berteroanum
solanderi
venturianum
tablasense
venturianum
tablasense
fallax
magellanicum
antrorsum
fallax
magellanicum
antrorsum
brevicaule
sessiliflorum
parodii
Median
25%-75%
Non-Outlier Range
Outliers
Median
25%-75%
Non-Outlier Range
Outliers
Second sinus length (mm)
Ratio maximum width/segment length
Stipules length (mm)
berteroanum
solanderi
berteroanum
solanderi
venturianum
tablasense
venturianum
tablasense
venturianum
tablasense
fallax
magellanicum
antrorsum
fallax
magellanicum
antrorsum
fallax
magellanicum
antrorsum
brevicaule
brevicaule
brevicaule
sessiliflorum
parodii
sessiliflorum
parodii
sessiliflorum
parodii
22
berteroanum
solanderi
20
retrorsum
albicans
limae
18
retrorsum
albicans
limae
16
14
core-core
retrorsum
albicans
limae
12
core-core
10
skottsbergii
core-core
8
6
skottsbergii
skottsbergii
4
2
0
1.0
0.9
0.8
0.7
0.6
0.5
0.4
18
16
14
12
8
10
6
28
26
berteroanum
solanderi
brevicaule
4
1.0
retrorsum
albicans
limae
sessiliflorum
parodii
2
24
core-core
retrorsum
albicans
limae
brevicaule
0
0.9
0.8
core-core
sessiliflorum
parodii
Median
25%-75%
Non-Outlier Range
Outliers
Median
25%-75%
Non-Outlier Range
Outliers
Median
25%-75%
Non-Outlier Range
Outliers
Median
25%-75%
Non-Outlier Range
Outliers
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
fallax
magellanicum
antrorsum
0.7
venturianum
tablasense
0.6
limae
berteroanum
solanderi
0.5
retrorsum
albicans
0.4
skottsbergii
skottsbergii
core-core
Ratio main sinus length/main segment length
0.3
0.6
0.5
0.4
0.3
0.2
0.1
0.0
18
16
skottsbergii
12
Ratio second sinus length/middle segment length
14
12
10
8
6
4
2
Figures 24–57. Continued
Segment lobes number
Peduncles length (mm)
Petioles indumentum length (mm)
2.4
2.2
2.0
1.8
1.6
1.4
1.2
1.0
0.8
0.6
skottsbergii
core-core
core-core
core-core
retrorsum
retrorsum
retrorsum
albicans
limae
albicans
limae
albicans
limae
berteroanum
solanderi
berteroanum
solanderi
venturianum
tablasense
berteroanum
solanderi
venturianum
tablasense
fallax
magellanicum
fallax
magellanicum
antrorsum
antrorsum
antrorsum
brevicaule
sessiliflorum
brevicaule
sessiliflorum
brevicaule
sessiliflorum
parodii
parodii
parodii
parodii
antrorsum
brevicaule
sessiliflorum
parodii
10
sessiliflorum
parodii
venturianum
tablasense
fallax
magellanicum
8
brevicaule
sessiliflorum
berteroanum
solanderi
6
antrorsum
brevicaule
albicans
limae
4
fallax
magellanicum
core-core
retrorsum
2
venturianum
tablasense
skottsbergii
0
venturianum
tablasense
fallax
magellanicum
–2
berteroanum
solanderi
70
berteroanum
solanderi
60
albicans
limae
50
albicans
limae
40
retrorsum
retrorsum
30
core-core
20
skottsbergii
core-core
10
0
10
9
8
7
6
5
4
3
2
skottsbergii
Bracteoles length (mm)
13
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Median
25%-75%
Non-Outlier Range
Outliers
GERANIUM SECTIONS ANDINA AND CHILENSIA
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Pedicels length (mm)
Sepals length (mm)
antrorsum
0.4
skottsbergii
skottsbergii
venturianum
tablasense
fallax
magellanicum
0.2
0.0
280
260
240
220
200
180
160
140
120
80
100
60
40
0
20
–20
2.2
2.0
1.8
1.6
1.4
1.2
1.0
0.8
0.6
0.4
0.2
0.0
Figures 24–57. Continued
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
Pedicels indumentum length (mm)
1.0
0.9
0.8
0.7
0.6
C. AEDO ET AL.
tablasense
fallax
magellanicum
antrorsum
brevicaule
sessiliflorum
parodii
Median
25%-75%
Non-Outlier Range
Outliers
3.5
3.0
2.5
2.0
1.5
1.0
skottsbergii
core-core
retrorsum
albicans
limae
berteroanum
solanderi
venturianum
tablasense
fallax
magellanicum
antrorsum
brevicaule
sessiliflorum
parodii
0.5
skottsbergii
core-core
retrorsum
albicans
limae
berteroanum
solanderi
venturianum
tablasense
fallax
magellanicum
antrorsum
brevicaule
sessiliflorum
parodii
Mucro length (mm)
0.0
parodii
0.5
solanderi
venturianum
3.0
2.8
2.6
2.4
2.2
2.0
1.8
1.6
1.4
1.2
1.0
0.8
0.6
0.4
0.2
0.0
sessiliflorum
0.4
berteroanum
Median
25%-75%
Non-Outlier Range
Outliers
antrorsum
brevicaule
9
fallax
magellanicum
0.3
core-core
retrorsum
albicans
limae
Sepals indumentum length (mm)
8
tablasense
7
solanderi
venturianum
6
limae
berteroanum
5
albicans
4
retrorsum
3
core-core
2
1
skottsbergii
0.2
0.50
0.45
0.40
0.35
0.30
0.25
0.20
0.15
0.10
0.05
skottsbergii
Filaments length (mm)
14
Ratio sepal width/sepal length
skottsbergii
core-core
retrorsum
albicans
limae
berteroanum
solanderi
venturianum
tablasense
fallax
magellanicum
antrorsum
brevicaule
sessiliflorum
parodii
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Median
25%-75%
Non-Outlier Range
Outliers
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
skottsbergii
core-core
retrorsum
albicans
limae
berteroanum
solanderi
venturianum
tablasense
fallax
magellanicum
antrorsum
brevicaule
sessiliflorum
parodii
0.00
18
16
14
12
10
8
6
4
2
Figures 24–57. Continued
Ratio mucro length/sepal length
Petals length (mm)
2.0
1.8
1.6
1.4
1.2
1.0
0.8
0.6
0.4
0.2
30
parodii
9
8
7
6
5
4
3
skottsbergii
core-core
retrorsum
albicans
limae
berteroanum
solanderi
venturianum
tablasense
fallax
magellanicum
antrorsum
brevicaule
sessiliflorum
parodii
2
1
5.0
4.5
4.0
3.5
3.0
2.5
2.0
1.5
22
20
skottsbergii
core-core
retrorsum
albicans
limae
berteroanum
solanderi
venturianum
tablasense
fallax
magellanicum
antrorsum
brevicaule
sessiliflorum
parodii
Gyneceo length (mm)
15
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Median
25%-75%
Non-Outlier Range
Outliers
Median
25%-75%
Non-Outlier Range
GERANIUM SECTIONS ANDINA AND CHILENSIA
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Median
25%-75%
Non-Outlier Range
Outliers
Mericarps length (mm)
18
parodii
28
sessiliflorum
16
sessiliflorum
26
antrorsum
brevicaule
14
antrorsum
brevicaule
24
fallax
magellanicum
12
fallax
magellanicum
22
tablasense
10
tablasense
20
solanderi
venturianum
8
solanderi
venturianum
18
limae
berteroanum
6
limae
berteroanum
16
albicans
4
2
albicans
14
retrorsum
Median
25%-75%
Non-Outlier Range
Outliers
retrorsum
12
8
skottsbergii
core-core
retrorsum
albicans
limae
berteroanum
solanderi
venturianum
tablasense
fallax
magellanicum
antrorsum
brevicaule
sessiliflorum
parodii
core-core
Rostrum length (mm)
core-core
10
6
4
1.6
skottsbergii
skottsbergii
core-core
retrorsum
albicans
limae
berteroanum
solanderi
venturianum
tablasense
fallax
magellanicum
antrorsum
brevicaule
sessiliflorum
parodii
skottsbergii
Anthers length (mm)
Fruit length (mm)
1.4
1.2
1.0
0.8
0.6
0.4
0.2
0.0
Figures 24–57. Continued
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
Mericarps indumentum length (mm)
C. AEDO ET AL.
6
1.6
5
1.4
Rostrum indumentum length (mm)
4
3
2
1
0.8
0.6
0.4
5.5
3.4
5.0
3.2
antrorsum
brevicaule
sessiliflorum
parodii
venturianum
tablasense
fallax
magellanicum
berteroanum
solanderi
albicans
limae
core-core
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
3.0
4.5
2.8
4.0
3.5
3.0
2.5
2.0
2.6
2.4
2.2
2.0
1.8
1.5
1.6
1.0
1.4
parodii
antrorsum
brevicaule
sessiliflorum
venturianum
tablasense
fallax
magellanicum
berteroanum
solanderi
albicans
limae
1.0
retrorsum
parodii
sessiliflorum
brevicaule
fallax
magellanicum
antrorsum
venturianum
tablasense
berteroanum
solanderi
albicans
limae
retrorsum
core-core
skottsbergii
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
core-core
1.2
0.5
0.0
retrorsum
0.0
skottsbergii
sessiliflorum
parodii
antrorsum
brevicaule
magellanicum
solanderi
venturianum
tablasense
fallax
limae
berteroanum
albicans
retrorsum
skottsbergii
core-core
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Seeds length (mm)
Stigmatic remains length (mm)
1.0
0.2
0
–1
1.2
skottsbergii
Rostrum narrowed apex (mm)
16
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Figures 24–57. Continued
it was significant at 99% in the ANOVA test. This
step was also supported by a qualitative character,
main leaf segment shape, and another quantitative
character, petal length, which was highly significant
in both tests. Finally, in step 16 stigmatic remains
length was not significant in the Median test,
although it was significant at 99% in the ANOVA
test. This step is also supported by a quantitative
character, sepal length, which is highly significant in
both tests.
In Discriminant Analysis 1 (DA1) the four species of
sect. Andina (G. antrorsum, G. brevicaule, G. parodii
and G. sessiliflorum) plus G. magellanicum were analysed. The plot of root 1 against root 2 shows a great
separation for G. magellanicum and G. parodii OTUs
(Fig. 58). The characters contributing most to this separation were: rostrum length, fruit length, rostrum
narrowed apex, pedicel length, pedicel indumentum
length, seed length, gynoecium length, peduncle
length, rootstock diameter length and petal length.
Geranium antrorsum, G. brevicaule and G. sessiliflorum were more clearly discriminated in the plot of root
2 against root 3 (Fig. 59). In this case rostrum length,
gynoecium length and petal length were the most relevant characters (Table 1).
In DA2 G. limae, G. berteroanum and G. solanderi
were analysed (Fig. 60). The most discriminant characters were: fruit length, bracteoles length, rostrum
length, ratio main sinus length/main segment length,
peduncle length and filament length (Table 1).
In DA3 G. albicans, G. skottsbergii, G. core-core and
G. retrorsum were analysed. The plot of root 1 against
root 2 shows a great separation for G. skottsbergii and
G. albicans OTUs (Fig. 61), while OTUs of G. core-core
and G. retrorsum were more clearly discriminated in
the plot of root 2 against root 3 (Fig. 62). The most discriminant characters were gynoecium length, ratio
sepal width/sepal length, fruit length, stem diameter,
stem indumentum length and rostrum length
(Table 1).
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
GERANIUM SECTIONS ANDINA AND CHILENSIA
17
Table 3. Kruskal–Wallis ANOVA and median test results for each step of Geranium sect. Andina and Chilensia species
key. Values of Chi-square and H higher than 3.84 are significant at 95%, and higher than 6.64 at 99%. Steps without
quantitative characters are not shown; *indicates step with additional qualitative character
Valid N
Kruskal–Wallis
ANOVA
Median test
Key step
Character
group 1
group 2
Chi-square
P
H
P
3*
4
4
5*
5
6*
8*
9
9
11
11
13
14*
14
15*
15
16
16
petal length
petal length
mucro length/sepal length
petal length
rostrum apex
peduncle length
rostrum narrowed apex
second sinus/middle segment length
petal length
second sinus/middle segment length
petal length
sepal width/sepal length
petal length
mucro length
segment width at base/segment length
mucro length
sepal length
stigmatic remains length
32
15
22
173
153
170
29
19
19
119
115
21
62
64
42
42
23
19
12
17
18
4
4
19
32
15
12
15
12
19
16
16
22
22
19
18
16.50
15.18431
25.85859
4.138991
18.11675
21.34825
21.94903
9.663158
7.038755
9.083473
17.91498
22.55639
12.13781
5.568182
13.57576
21.48962
18.62434
3.634013
0.0000
0.0001
0.0000
0.0419
0.0000
0.0000
0.0000
0.0019
0.0080
0.0026
0.0000
0.0000
0.0005
0.0183
0.0002
0.0000
0.0000
0.0566
23.00764
16.48857
24.75166
11.074459
24.96222
49.86967
21.06862
15.77173
9.926733
25.92280
25.28147
24.57248
19.06564
6.231933
22.28891
24.86179
13.91956
8.802685
0.0000
0.0000
0.0000
0.0009
0.0000
0.0000
0.0000
0.0001
0.0016
0.0000
0.0000
0.0000
0.0000
0.0125
0.0000
0.0000
0.0002
0.0030
Table 4. Correct classifications and values of P obtained in discriminant analyses (DAs) of Geranium sect. Andina and
Chilensia
G. sessiliflorum
G. brevicaule
G. antrorsum
G. parodii
G. magellanicum
G. berteroanum
G. solanderi
G. limae
G. albicans
G. core-core
G. retrorsum
G. skottsbergii
G. tablasense
G. venturianum
G. fallax
DA
No. of OTUs
Correct
classifications (%)
1
1
1
1
1
2
2
2
3
3
3
3
4
4
4
14
13
4
4
8
18
14
7
15
15
14
6
4
19
18
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
In DA4 G. venturianum, G. tablasense and G. fallax
were analysed (Fig. 63). The most discriminant characters were: segment width at the base, rostrum indumentum length, stigmatic remains length, ratio main
sinus length/main segment length, bracteole length,
fruit length and sepal indumentum length (Table 1).
P
0.32558
0.30233
0.09302
0.09302
0.18605
0.46154
0.35879
0.17949
0.30000
0.30000
0.28000
0.12000
0.09756
0.46341
0.43902
The OTUs correctly classified were in all the cases of
the 100%, which strongly support the correct delimitation of the species (Table 4). It should be noted that
some qualitative characters, which are discussed
under each species, also contributed to delimitation of
the species.
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
18
C. AEDO ET AL.
14
12
DA1
10
DA1
12
8
10
6
8
4
6
Root 3
Root 2
2
0
–2
4
2
–4
0
–6
–2
–8
–4
–10
–12
–14
–20
–10
0
10
Root 1
20
40
30
10
magellanicum
antrorsum
brevicaule
sessiliflorum
parodii
10
skottsbergili
core-core
retrorsum
albicans
–6
magellanicum
antrorsum
brevicaule
sessiliflorum
parodii
–8
–15
–10
–5
0
5
Root 2
10
12
DA2
DA3
10
8
8
6
6
4
Root 2
Root 2
4
2
2
0
0
–2
–4
–2
–6
–4
–8
–6
–8
–6
–4
–2
0
Root 1
2
4
6
limae
berteroanum
solanderi
–10
–15
–5
0
5
Root 1
10
6
DA4
DA3
5
8
4
6
3
4
Root 2
2
Root 3
–10
1
2
0
0
–2
–1
–4
–2
–6
–3
–8
–4
–5
–10
–8
–6
–4
–2
0
2
Root 2
4
6
8
10
skottsbergili
core-core
retrorsum
albicans
–10
–20
–15
–10
–5
Root 1
0
5
10
venturianum
tablasense
fallax
Figures 58–63. Plots of the Discriminant Analyses in Geranium sect. Andina and sect. Chilensia.
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
GERANIUM SECTIONS ANDINA AND CHILENSIA
KEY
19
TO SPECIES
1. Plant acaulescent; hairs always eglandular
2. Pedicels with antrorse hairs; leaf cuneate to subtruncate (with basal leaf segments patent or upward) ........3. G.
antrorsum
2. Pedicels with patent to retrorse hairs; leaf cordate (with basal leaf segments downward)
3. Petals hairy on margin and base of abaxial side (9.5–)11.5–15.7 mm long; nectaries hairy; rostrum with a
narrowed apex 0.7–1.5 mm long ........................................................................................................... 4. G. parodii
3. Petals glabrous (rarely with few cilia on the basal margin), 2.8–8(-12.8) mm long; nectaries glabrous; rostrum
without narrowed apex.
4. Petals (4.5-)6–12.8 mm long; sepal mucro short (mucro length/sepal length ratio = (0.04–)0.07–
0.15(-0.19)) ................................................................................................................................ 1. G. sessiliflorum
4. Petals 2.8–5.5(-6.5) mm long; sepal mucro long (mucro length/sepal length ratio = (0.14–)0.20–
0.26(-0.37)) .................................................................................................................................... 2. G. brevicaule
1. Plant caulescent; hairs glandular or eglandular
5. Petals (11.4–)13.4–16.4(-16.9) mm long, glabrous, rarely with few cilia on the basal margin; rostrum with a narrow
apex (2–)2.3–3.8(-5) mm long ............................................................................................................. 13. G. tablasense
5. Petals (2.9–)5–8(-14.3) mm long, ciliate on the basal margin (in G. venturianum also on the base of the abaxial
side); rostrum usually without a narrow apex (in G. venturianum and G. fallax (0–)0.9–2(-2.4) mm long)
6. Peduncles (5.9–)8.2–15.1(-25.1) cm long; petals hairy on the margin and on the base of the abaxial
side ............................................................................................................................................... 14. G. venturianum
6. Peduncles (0.36–)1.68–3.8(-10) cm long; petals ciliate on the basal margin
7. Peduncles with glandular hairs
8. Middle segment rhombic; rostrum with a narrow apex (0–)0.9–1.1(-1.9) mm long ................ 15. G. fallax
8. Middle segment obtriangular; rostrum without a narrow apex
9. Basal leaves deeply divided with narrow lobes (second sinus/middle-segment length ratio =
(0.31–)0.40–0.53(-0.57)); petals (7–)7.9–10.2(-14) mm long ........................................10. G. skottsbergii
9. Basal leaves not deeply divided (second sinus/middle-segment length ratio =(0.13–)0.24–0.38(-0.42));
petals (5.1–)6.5–8(-9) mm long ..................................................................................... 5. G. berteroanum
7. Peduncles without glandular hairs
10. Plant decumbent, with cymules of the basal part of the stem usually 1-flowered
(part of them arises directly from the rootstock), and cymules of the upper part
2-flowered ............................................................................................................. 8. G. magellanicum
10. Plants usually erect to ascending with all cymules (always 2-flowered) arising along leafy stems
11. Basal leaves deeply divided with narrow lobes (second sinus/middle segment length
ratio = (0.31–)0.40–0.53(-0.57)); petals (7–)7.9–10.2(-14) mm long...............10. G. skottsbergii
11. Basal leaves not deeply divided (second sinus/middle segment length ratio = (0.13–)0.21–
0.41(-0.47)); petals (2.9–)3.5–7.5(-9.2) mm long
12. Pedicels with hairs retrorse-appressed
13. Sepals broadly ovate (width/length ratio = (0.60–)0.62–0.76(-0.84)) ... 11. G. core-core
13. Sepals lanceolate (width/length ratio = (0.26–)0.44–0.59(-0.65)) ....... 12. G. retrorsum
12. Pedicels with hairs patent
14. Petals (4–)4.5–5.1(-6.1); sepal mucro (0.2–)0.25–0.4(-0.6) mm long; main leaf
segment rhombic ..................................................................................9. G. albicans
14. Petals (3–)4.7–8(-9); sepal mucro (0.2–)0.3–0.8(-0.9) mm long; main leaf segment
obtriangular
15. Segment width at the base/middle-segment length ratio = (0.20–)0.24–
0.32(-0.42); nectaries usually hairy; sepal mucro (0.2–)0.3–0.4(-0.5) mm
long .................................................................................................... 7. G. limae
15. Segment width at the base/middle-segment length ratio = (0.11–)0.15–0.24(-0.31);
nectaries glabrous; sepal mucro (0.2–)0.4–0.8(-0.9) mm long
16. Sepals (3.2–)4.2–5.2(-6.3) mm long; stigmatic remains (0.6–)1–1.4(-1.6) mm
long......................................................................................... 6. G. solanderi
16. Sepals (4.4–)5.4–6.1(-7) mm long; stigmatic remains (1.1–)1.3–1.7(-2.1) mm
long................................................................................... 5. G. berteroanum
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
20
C. AEDO ET AL.
TAXONOMIC TREATMENT
Notes: Species of Geranium sect. Andina are recognized by their stemless habit, turnip-shaped rootstock, and 1-flowered cymules. This section includes
two species from South America and two species from
Australia and New Zealand. Knuth (1912) proposed
his sect. Andina to include all stemless species with 1flowered cymules, but some of them have been segregated in the new section Neoandina (Aedo et al.,
2002). According to this taxonomic interpretation,
stemless species with ramified rootstock should be
included in sect. Neoandina, and those with turnipshaped rootstock in sect. Andina.
Geranium caespitosum Walp. in Nov. Actorum
Acad. Caes. Leop.-Carol. Nat. Cur. 19 (suppl. 1):
315 (1843), nom. illeg., nom. E. James 1823
Type: Peru. Meyen s.n. (lectotype, here selected, K!)
Geranium bangii Hieron. in Bot. Jahrb. Syst. 21:
314 (1895)
Type: Bolivia. Oruro, Capi, Mar. 1890, Bang 788
(lectotype designated by Knuth, 1912: 85, B†;
isolectotypes, BM!, E!, F!, GH!, K!, MO!, NY!, W!)
[18∞47¢S, 68∞12¢W]
Geranium sessiliflorum var. microphyllum Kuntze,
Revis. Gen. Pl. 3(2): 33 (1898)
Type: Bolivia. Cochabamba, Tapacari, 17.iii.1892,
Kuntze s.n. (lectotype, here designated, NY!)
[17∞33¢S, 66∞28¢W]
Geranium sessiliflorum var. lanatum R. Knuth in
Bot. Jahrb. Syst. 37: 565 (1906)
Type: Bolivia. Fiebrig 3291 (lectotype, here designated, LD!) [probably from Tarija]
Geranium pflanzii R. Knuth in Engl., Pflanzenr.
IV.129 (Heft 53): 576 (1912)
Type: Bolivia. Palca-La Paz 1908, Pflanz 206
(holotype, B†; no authentic material found; photo,
G!, GH!, MO!, NY!)
Geranium sessiliflorum var. compactum R. Knuth
in Engl., Pflanzenr. IV.129 (Heft 53): 85 (1912)
Type: Bolivia. La Paz-Palca-Illimani, Hauthal 323
(lectotype, here designated, GOET!)
Geranium tucumanum R. Knuth in Engl., Pflanzenr. IV.129 (Heft 53): 147 (1912)
Type: Argentina. Sierra de Tucumán, La Ciénaga
1874, Hieronymus 584 (holotype, B†; isotype, K!)
Geranium razuhillcaënse R. Knuth in Repert. Spec.
Nov. Regni Veg. 28: 2 (1930)
Type: Peru. Ayacucho, Huanta, Mte. Razuhuillca,
Weberbauer 7495 (lectotype, here designated, F!;
isolectotypes, G!, K!)
Geranium malpasense R. Knuth in Repert. Spec.
Nov. Regni Veg. 34: 146 (1933)
Type: Bolivia. Chuquisacá, Cordillera de los
Sombreros, Cuesta Malpaso, 13.xii.1927, Troll 705
(holotype, B!; isotypes, JE! M!)
Geranium staffordianum R. Knuth in Repert. Spec.
Nov. Regni Veg. 40: 216 (1936)
Type: Peru. Cuzco, Stafford 208 (holotype, K!)
Geranium sessiliflorum var. albatum J.F. Macbr. in
Publ. Field Mus. Nat. Hist., Bot. Ser. 13(3): 533
(1949)
Type: R. Knuth in Engl., Pflanzenr. IV.129 (Heft
53): 84 fig. 15 (1912) (lectotype, here designated)
1. Geranium sessiliflorum Cav., Diss. 4: 198, tab. 77
fig. 2 (1787)
Type: Chile. Estrecho de Magallanes, Commerson
s.n. (lectotype designated by Garilleti, 1993: 91,
MA-475750!, isolectotype, P!)
Herbs 2.2–8.9 cm tall. Rootstock 2.8–18.9 mm diam.,
without fusiform roots. Basal leaves in a persistent
rosette; lamina 7.3–27.2 ¥ 8.8–31 cm, orbicular to
polygonal in outline, cordate (with basal leaf segments
downward), palmatifid (divided for 0.52–0.76 of its
I. Geranium sect. Andina R. Knuth in Bot. Jahrb.
Syst. 32: 222 (1903)
Type: G. sessiliflorum (lectotype, designated by R.
Knuth, 1912: 45)
Perennial herbs with rootstock usually vertical,
not tuberculate, turnip-shaped, without fusiformswollen roots; aerial stem absent; without vegetative
stems or stolon. Leaf lamina orbicular to polygonal in
outline, cordate (with basal leaf segments downward),
sometimes cuneate to subtruncate cordate (with basal
leaf segments patent or upward), without abscission
zone, palmatifid, hairy, not coriaceous and with nerves
not projected; segments 5–7, obtriangular or rhombic,
3–12-lobed in the distal half; cauline leaves absent;
stipules lanceolate, free, papery, reddish, hairy. Inflorescence with 1-flowered cymules arising directly
from the rootstock, usually solitary; peduncles (when
present) and pedicels hairy; bracteoles linearlanceolate, hairy; pedicel and peduncle together
usually not overtopping the subtending leaf. Sepals
ovate, smooth, not accrescent, 3-nerved, mucronate,
with scarious margins, hairy. Petals erect-patent, ±
obovate, entire, rarely emarginate, without claw, glabrous to hairy, purplish to white. Stamens ten, both
whorls bearing anthers; filaments not exserted, lanceolate yellowish, hairy. Nectaries hemispherical,
usually glabrous. Fruit of the seed-ejection type;
mericarps usually smooth, without longitudinal rib,
without basal beak, with a basal callus, without a
basal prong, usually with eglandular hairs, brownish;
rostrum with or without a narrowed apex; stigmatic
remains with five hairy or glabrous lobes. Seeds
ellipsoid, finely reticulate, brownish; hilum 1/4–1/6 as
long as the perimeter. Cotyledons entire.
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
GERANIUM SECTIONS ANDINA AND CHILENSIA
length), glabrescent to pilose (sometimes sericeous)
on both sides, with appressed, eglandular hairs; segments 5–7, rhombic to obtriangular, 0.9–4.7 mm at the
base (segment width at the base/segment length
ratio = (0.13–)0.16–0.23(-0.64)), 3–12-lobed in distal
half (second sinus length/middle-segment length
ratio = (0.16–)0.23–0.35(-0.50)); petioles to 8 cm long,
with patent to retrorse, appressed, eglandular hairs
0.2–1.7 mm long; stipules 5.5–15.8 ¥ 0.8–3.1 mm, with
eglandular hairs on abaxial surface and on the margin, glabrous adaxially. Inflorescence with cymules
1-flowered, solitary; peduncles 0–0.5(-1.4) cm long,
with retrorse, appressed, eglandular hairs 0.3–1 mm
long; bracteoles 5–6.1 ¥ 1–1.65 mm, linear-lanceolate,
with eglandular hairs on both sides and on the
margin; pedicels 0.4–4.2 cm long, with retrorse to
patent, appressed, eglandular hairs 0.3–1.2 mm long;
pedicel not overtopping the subtending leaf. Sepals
(3.8–)4.7–6.2(-7.1) ¥ 1.44–3.37 mm, lanceolate (width/
length ratio = (0.28–)0.35–0.43(-0.53)), with mucro
(0.3–)0.45–0.7(-1.1) mm long (mucro length/sepal
length ratio = (0.04–)0.07–0.15(-0.19)), with scarious
margins 0.1–0.2 mm wide, with erect-patent, eglandular hairs 0.3–0.6 mm long on the abaxial side, patent
hairs 0.9–2.8 mm long on the margin, glabrescent
adaxially. Petals (4.5–)6–12.8 ¥ 1.6–5 mm, entire,
rarely emarginate, notch 1 mm deep, without claw,
glabrous, rarely with few cilia on the basal margin,
purplish, sometimes white. Filaments 2.1–4.6 mm
long, yellowish, with eglandular hairs 0.2–0.3 mm
long on the abaxial side and margin; anthers
0.5–1.1 ¥ 0.3–0.64 mm, yellowish; pollen unknown
colour. Nectaries glabrous. Gynoecium 1.9–5 mm
long, yellowish. Fruit 6.8–19.5 mm long; mericarps
1.8–4.5 ¥ 0.9–2.25 mm, smooth, with erect-patent,
eglandular hairs 0.1–0.6 mm long, brownish;
rostrum 4.1–13 mm long, without a narrowed apex,
with erect-patent, eglandular hairs 0.2–1.2 mm long;
stigmatic remains (0.8–)1–1.5(-2) mm long, with 5
glabrous lobes. Seeds 1.1–3.3 ¥ 0.9–2.1 mm, finely
reticulate; hilum 1/4 as long as the perimeter.
Area: Central and South Peru, Bolivia, Chile and
Argentina (Fig. 64).
Habitat: Beaches, river shores, turfy bogs, natural
grasslands, open, moist puna, or clearing of Nothofagus or Polylepis forest; 0–5000 m.
Phenology: Flowering January – December.
Illustration: Figure 65.
Notes: Geranium sessiliflorum is a species distributed
over a large area of more than 5000 km between the
northernmost localities in Central Peru and the south-
21
ernmost ones in Tierra de Fuego. It displays an
uncommon variability, mainly in leaf shape and indumentum. Leaf outline may be orbicular or polygonal,
and more or less divided. Middle leaf segment varies
from obtriangular with three lobes to rhombic with 12
lobes. In extreme cases, leaves are white sericeous
(only in southern Peru, Bolivia, and northern Argentina), although they usually are pilose, with scattered
appressed hairs. A continuum of intermediates suggests that these forms do not deserve taxonomic recognition. Fruit size also shows a remarkable range of
variation, between 6.8 and 19.5 mm long, but no correlation with leaf shape or indumentum has been
found.
Geranium sessiliflorum is usually stemless with all
cymules borne on the rootstock apex. However, some
specimens have (together with cymules borne on the
rootstock) elongate internodes bearing a tuft of
stipules, from which an aggregate of cymules is born.
Sometimes a pair of leaves is also present in these
nodes. These forms have been named G. tucumanum
in North Argentina, but they are also present over all
the G. sessiliflorum area. Intermediate forms can be
found even on the same herbarium sheet. All these
data suggest that there is no basis for recognizing
G. tucumanum, which we have subsumed under
G. sessiliflorum. The type of G. malpasense has long
(12 mm) and emarginate petals. Otherwise, it is similar to other specimens of G. sessiliflorum.
Although G. magellanicum is quite similar to
G. sessiliflorum, it is easily distinguishable by its
2-flowered cymules spread on decumbent and foliate
stems. The differences between G. sessiliflorum and
the remaining species of sect. Andina are addressed in
the discussion under G. brevicaule, G. parodii and
G. antrorsum.
Representative specimens examined: ARGENTINA.
CATAMARCA: Aconquija, 27∞0¢S, 65∞55¢W, 16.ii.1951,
Brücher s.n. (LD); Andalgalá, 27∞36¢S, 66∞19¢W, .1917,
Jörgensen 1387 (MO, GH). JUJUY: 20 km W of
Humahuaca, 23∞12¢S, 65∞23¢W, 13.iii.1936, West 6335
(CAS, MO); camino Abra de Lizeite, primer arroyo
despues de Cajas, 22∞15¢S, 65∞18¢W, .xii.1979,
González 105 (LIL). LA RIOJA: Chilecito, Sierra Famatina, camino a La Mejicana, 29∞3¢S, 67∞51¢W, 6.ii.1927,
Parodi 7972 (GH); Sierra Famatina, Corral Colorado,
29∞20¢S, 67∞41¢W, 12.ii.1879, Hieronymus & Niederlein 7776 (G); MENDOZA: Atuel valley, 2–3 km N of
Campamento Atuel, 34∞45¢S, 69∞59¢W, 29.ii.1955,
Böcher et al. 1275 (MO, C); Malargüe, Paso Pehuenche, 35∞59¢S, 70∞24¢W, 26.ii.1989, Leuenberger &
Arroyo 3932 (B). NEUQUÉN: Aluminé, 10 km W de
Primeros Pinos, 38∞52¢S, 70∞35¢W, 11.xii.1985, Correa
et al. 9181 (CTES); Cerro Chapelco, 40∞14¢S, 71∞16¢W,
24.i.1966, Schojovskoy 100 (M); RÍO NEGRO: 3.3 km S
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
22
C. AEDO ET AL.
Figure 64. Area of distribution of Geranium sessiliflorum (dots) and G. parodii (triangles).
of El Foyel on route 258, 41∞35¢S, 71∞43¢W, 3.xii.1984,
Stuessy et al. 6794 (LP, CAL); Cerro Catedral, San
Carlos de Bariloche, 41∞13¢S, 71∞30¢W, 22.i.1964, Ward
s.n. (MO). SALTA: Cerro del Cajón, La Laguna, 25∞46¢S,
65∞14¢W, 24.i.1914, Rodríguez 1305 (SI, BAF, NY);
Chicoana, camino a Cachi, Piedra de Molino, 25∞10¢S,
65∞46¢W, 26.iii.1979, Cabrera et al. 30721 (SI). SANTA
CRUZ: 5 km al NE del Lago Belgrano, 47∞50¢S, 72∞9¢W,
8.xii.1980, Cei s.n. (MERL); Comandante Piedrabuena, 49∞59¢S, 68∞54¢W, 1.xii.1945, O’Donell 3930
(HBG, CAS, NY, TEX). TIERRA DE FUEGO: Cabo Domingo, 53∞41¢S, 67∞51¢W, 11.xii.1965, Goodall 193 (LP);
Cordón de los Cristales, Estancia Lago Roca, 50∞33¢S,
72∞45¢W, 26.xii.1958, James 109 (BM). TUCUMÁN:
Chicligasta, Estancia Santa Rosa, La Cueva, 27∞12¢S,
65∞30¢W, 17.ii.1926, Venturi 6902 (US); Lara, 26∞21¢S,
65∞56¢W, 21.i.1912, Rodríguez 263 (SI). BOLIVIA.
CHUQUISACA: Punilla-Chanauca, cerro Chataquila,
18∞48¢S, 64∞46¢W, 27.ii.1994, Wood 8042 (LPB). COCHABAMBA: Arque, 6 km E de Challa, 17∞37¢S, 66∞40¢W,
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GERANIUM SECTIONS ANDINA AND CHILENSIA
J
23
I
E
G
F
H
D
B
A
C
Figure 65. Geranium sessiliflorum Cav. A, habit; B–D, leaves; E, flower; F, petal; G, staminal filament; H, fruit; I,
mericarp; J, seed [A: Aedo 7500 (MA); B: Sanches 24b (LPB); C: Rossel 101 (LPB); D: Macbride 3006 (MA); E–G: Mez y
Barrera 1568 (SGO); H–J: Hohenacker 1160 (S).].
31.iii.1979, Beck 970 (LPB); bajada a Morochata km
60, Tunari, 17∞14¢S, 66∞32¢W, 19.xi.1990, Saravia 30
(LPB). LA PAZ: Bautista Saavedra, Khata, 15∞9¢S,
69∞0¢W, 6.i.1983, Menhofer 1825 (MA); Camacho,
Ambana 2 km hacia el S, 15∞28¢S, 69∞0¢W, 19.xii.1980,
Beck 4179 (LPB); Incachaca, 16∞25¢S, 68∞4¢W,
21.xii.1920, Asplund 6111 (US, NY); Ingavi, al SE de
Viacha, 16∞39¢S, 68∞18¢W, .xii.1990, Beck 20935 (MA).
ORURO: Avaroa, Challapata, 18∞54¢S, 66∞46¢W,
31.iii.1921, Asplund 3272 (S); Cordillera de Azanaque,
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
24
C. AEDO ET AL.
19∞3¢S, 66∞37¢W, 25.xii.1926, Troll 2983 (B, M).
POTOSÍ: above Animas mine, W of Chocaya, 21∞18¢S,
67∞44¢W, 21.ii.1936, West 6072 (MO, CAS, GH);
Linares, inter Kachitambo et Potosí, 19∞47¢S, 65∞49¢W,
4.iv.1934, Hammarlund 452 (S). TARIJA: Avilés, Pasto,
camino de Tajzara, 21∞55¢S, 65∞2¢W, 6.ii.1986, Bastión
749 (MA); Calderillo, 21∞48¢S, 63∞45¢W, 9.i.1904, Fiebrig 2627 (W, E, G, GH, HBG, M, P, S, U). CHILE.
AISÉN: Balmaceda, 45∞55¢S, 71∞41¢W, 11.xii.1954, Pfister s.n. (CONC); Coyhaique Alto, 45∞29¢S, 71∞36¢W,
25.i.1990, Martínez et al. 23 (SGO). BIOBÍO: termas de
Chillán, 36∞54¢S, 71∞24¢W, 27.i.2002, Aedo 7500 (MA).
COQUIMBO: Cordillera del río Mauro, 31∞58¢S, 71∞2¢W,
.ii., Reiche s.n. (SGO). La Araucania: Curacautin,
Malalcahuello, 38∞27¢S, 71∞35¢W, 4.i.1948, Pfister s.n.
(CONC); Malleco, Pino Hachado a 100 m del límite,
38∞39¢S, 70∞55¢W, 11.ii.1960, Ricardi & Marticorena
5112 (CONC). LOS LAGOS: Osorno, Parque Nacional
Puyehue, límite con Argentina, 40∞35¢S, 72∞8¢W,
14.ii.1972, Muñoz 537 (SGO); Valdivia, Quetrupillan,
39∞48¢S, 73∞14¢W, 20.iii.1982, Schlegel 7532 (SGO).
MAGALLANES: Cerro Donoso, río de las Chinas,
50∞44¢S, 72∞31¢W, 9.ii.1987, Arroyo et al. 870253
(CONC); Punta Arenas, pr. Lago Parrillar, 53∞24¢S,
71∞3¢W, 21.i.2002, Aedo 7486 (MA). MAULE: Curicó,
orillas de la laguna de Teno, 34∞59¢S, 71∞14¢W,
10.iii.1967, Marticorena & Matthei 883 (CONC);
Linares, valle Aguirre, 36∞10¢S, 70∞32¢W, 20.i.1961,
Schlegel 3554 (CONC); Talca, Paso Pehuenche,
35∞59¢S, 70∞24¢W, 30.i.1994, Villagrán et al. 8227
(CONC). SANTIAGO: Cajón del Yeso, Termas El Plomo,
33∞14¢S, 70∞13¢W, 17.i.1995, Muñoz et al. 3488 (SGO);
estación de eski Valle Nevado, 33∞20¢S, 70∞14¢W,
12.xii.2001, Aedo 7125 (MA). TARAPACÁ: Aguas Calientes, Tacora, 17∞44¢S, 69∞46¢W, 17.ix.1955, Ricardi
et al. 3374 (CONC); Parinacota, cerca de la laguna de
Cotacotani, camino a Guaneguane, 18∞10¢S, 69∞14¢W,
9.iii.1984, Arroyo 84–172 (CONC). TIERRA DE FUEGO:
Altos de Boquerón, 53∞17¢S, 69∞17¢W, 15.xii.1971,
Moore & R.N. Goodall 19015 (H); Isla Grande, 55∞9¢S,
68∞46¢W, .i., Maza s.n. (SGO). PERU. ANCASH: Carhuaz, Huascaran National Park, Quebrada Ishinca,
9∞22¢S, 77∞25¢W, 11.ii.1985, Smith et al. 9441 (MO, F,
ISC); Huaylas, Huascarán National Park, Auquispuquio area of ruins, 8∞50¢S, 77∞58¢W, 7.iv.1986, Smith
et al. 11990 (MO, F, ISC). APURIMAC: Andahuaylas,
13∞40¢S, 73∞25¢W, .ii., Pearce s.n. (BM). AREQUIPA:
Arequipa, 17∞19¢S, 70∞19¢W, 26.iii.1923, Guenther &
Buchtien 1356 (HBG); Cajatambo, 15∞35¢S, 71∞1¢W,
s.d., Weberbauer 2775 (WRSL). AYACUCHO: laguna
Yaurihuiri, about 205 km from Nazca on the road to
Abancay, 14∞38¢S, 73∞57¢W, 13.iii.1987, Brandbyge 319
(AAU); Señal Cerro Palmaderas, 80 km above Nazca
on road to Poquio, 14∞40¢S, 74∞29¢W, 11.iii.1987,
Brandbyge 275 (AAU). CUZCO: Calca, Amparaes, centre of the town, around the bridge, SW from Cuzco and
S of town, 13∞10¢S, 71∞54¢W, 13.xii.1986, Núñez 6749
(MO); Canas, Langui, 14∞25¢S, 71∞16¢W, 13.i.1975,
Chávez 2317 (MO). HUANCAVELICA : cerros de Laria, a
8 km de Conaica, 12∞33¢S, 75∞3¢W, 30.iii.1952, Tovar
912 (US); Tayacaja, Montepungo, 5 km E of Surcubamba, 12∞8¢S, 74∞41¢W, 13.i.1939, Stork & Horton
10383 (F, CAL). JUNÍN: 5 km E of Huancayo, 12∞4¢S,
75∞14¢W, 31.xii.1938, Stork & Horton 10217 (F);
between Tarma and Jauja, 11∞36¢S, 75∞37¢W,
24.iv.1929, Killip & Smith 21971 (NY, US). LIMA:
Canta, 11∞25¢S, 76∞45¢W, 2.v.1963, Acleto 735 (MA);
Rio Blanco, 11∞44¢S, 76∞15¢W, 20.iii.1923, Macbride
3006 (MA, F, G). MOQUEGUA: 5 km E of Lago Suche,
16∞56¢S, 70∞23¢W, 11.xii.1951, Pearson 20 (F, CAL);
Coalaque, 16∞49¢S, 71∞15¢W, 6.iii.1923, Guenther &
Buchtien 1318 (HBG). PASCO: Huaron, 11∞0¢S,
76∞25¢W, 12.vi.1922, Macbride & Featherstone 1148B
(F); Huayllay to Canta road, 11∞1¢S, 76∞21¢W,
6.iii.1977, Boeke 1118 (MO, NY). PUNO: 3 km SE of
Vilque, 15∞46¢S, 70∞15¢W, 11.i.1963, Iltis et al. 1373
(BH); 4 km E of Tincopalca, 15∞51¢S, 70∞45¢W,
12.i.1963, Iltis et al. 1475 (BH).
2. Geranium brevicaule Hook. f. in J. Bot. (Hooker) 1:
252 (1834)
Geranium sessiliflorum ssp. brevicaule (Hook. f.)
Carolin in Proc. Linn. Soc. New South Wales ser. 2,
89: 357 (1965)
Type: Australia. Tasmania, Gunn 256 (lectotype,
designated by Carolin, 1965: 357), K!; isolectotypes,
BM!, F!)
Geranium sessiliflorum ssp. novaezelandiae Carolin in Proc. Linn. Soc. New South Wales ser. 2, 89:
356 (1965)
Type: New Zealand. Inland Kaikoura Mts., Marlborough, 4 Dec. 1960, Hamlin 915 (holotype, WELT
11508!; isotype, UC!)
Geranium sessiliflorum var. arenarium G. Simpson
& J.S. Thomson in Trans. & Proc. Roy. Soc. New
Zealand 73: 158 (1943)
Type: New Zealand. Stewart Island, Paterson’s
Inlet, Simpson & Thomson s.n. (CHR, not located
according to Carolin, 1965: 357)
Geranium sessiliflorum var. maculatum G. Simpson & J.S. Thomson in Trans. & Proc. Roy. Soc. New
Zealand 73: 157 (1943)
Type: New Zealand. Lake Lyndon, Simpson &
Thomson s.n. (lectotype, designated by Carolin,
1965: 356, CHR 75697!)
Geranium sessiliflorum var. glabrum R. Knuth in
Bot. Jahrb. Syst. 37: 565 (1906)
Type: New Zealand. [without locality], Bastian s.n.
(lectotype, designated by Carolin, 1965: 357, K!)
Herbs 2.5–15.5 cm tall. Rootstock 3.2–10.5 mm diam.,
without fusiform roots. Basal leaves in a persistent
rosette; lamina 0.78–2.56 ¥ 0.89–3.18 cm, polygonal in
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
GERANIUM SECTIONS ANDINA AND CHILENSIA
outline, cordate (with basal leaf segments downward),
palmatifid (divided for 0.53–0.74 of its length), pilose,
with appressed, eglandular hairs; segments 5–7, obtriangular, 1.4–4.7 mm at the base (segment width at the
base/segment length ratio = (0.20–)0.24–0.35(-0.41)),
3–7-lobed in distal half (second sinus length/middlesegment
length
ratio = (0.10–)0.21–0.26(-0.35));
petioles to 13 cm long, with patent to retrorse, appressed, eglandular hairs 0.3–1.2 mm long; stipules
2.7–8.8 ¥ 0.9–2.5 mm, with scattered eglandular hairs
on both surfaces and on the margin. Inflorescence with
cymules 1-flowered, solitary; peduncles 0–1.1(-1.6) cm
long, with patent to retrorse, eglandular hairs
0.15–1.1 mm long; bracteoles 2–6.57 ¥ 0.6–1.25 mm,
linear-lanceolate, with scattered eglandular hairs on
both surfaces and on the margin; pedicels 0.33–4.3 cm
long, with patent to retrorse, eglandular hairs
0.15–1.1 mm long; pedicel not overtopping the subtending
leaf.
Sepals
(3.3–)4.4–5.7(-7.1) ¥ 1.4–
3.14 mm, lanceolate (width/length ratio = (0.31–)0.36–
0.45(-0.54)), with mucro (0.48–)0.9–1.3(-1.95) mm
long (mucro length/sepal length ratio = (0.14–)0.20–
0.26(-0.37)), with scarious margins 0.1–0.2 mm wide,
with erect-patent, eglandular hairs 0.3–0.6 mm long
on the abaxial side (patent hairs 1–1.7 mm long on the
margin), glabrescent adaxially. Petals 2.8–5.5(-6.5) ¥
1–3.1 mm, entire, without claw, glabrous, rarely with
few cilia on the basal margin, deep pink to white. Filaments 1.6–3 mm long, yellowish, glabrous on both
sides, ciliate on the basal margin, with hairs up to
0.1–0.2 mm long; anthers 0.4–0.7 ¥ 0.2–0.7 mm, yellowish; pollen unknown colour. Nectaries glabrous.
Gynoecium 2–3.4 mm long, unknown colour. Fruit
8.4–17.6 mm long; mericarps 2.2–3.58 ¥ 1–1.74 mm,
smooth, with erect-patent, eglandular
hairs
0.2–1.1 mm long, brownish; rostrum 5.1–11.7 mm
long, without a narrowed apex, with erect-patent, eglandular hairs 0.1–0.6 mm long; stigmatic remains
0.9–1.2(-1.6) mm long, with 5 hairy lobes. Seeds
1.5–2.8 ¥ 0.7–1.4 mm, finely reticulate; hilum 1/6 as
long as the perimeter.
Area: Australia (south-east Australia and Tasmania),
and New Zealand (Fig. 66).
Habitat: Sand dunes, lake shores, grasslands, rock
cranny on dry coastal hillside, or open Eucalyptus forest; 0–1300 m.
Phenology: Flowering October – June.
Illustration: Figure 67.
Notes: Carolin (1965) considered two subspecies of
G. sessiliflorum, one endemic to New Zealand and
other restricted to south-east Australia and Tasmania.
He distinguished ssp. novazelandiae by the long hairs
of the calyx tending to be inserted towards the margin
25
(also on the abaxial surface on ssp. brevicaule), the relatively shorter sepal mucro (c. 1 mm in ssp. novazelandiae, and 1 mm or longer in ssp. brevicaule), and
longer petals (c. 1.5 times as long as the sepals in ssp.
novazelandiae, and shorter than to slightly exceeding
the sepals in ssp. brevicaule). On the studied material
we are not able to find any difference in sepals indumentum or mucro length among plants from Australia
(and Tasmania) and New Zealand. Petals from New
Zealand plants are sometimes a bit longer than in
Australian ones. However, we have found a great overlap in this character, which made it impossible to distinguish between these groups. Thus, we prefer to
include all these plants in a single taxon which could
be more easily distinguished from South American
G. sessiliflorum.
Geranium brevicaule shares with G. antrorsum a
relatively longer sepal mucro. However, it can be easily distinguished by its pedicels with patent to retrorse
hairs (antrorse hairs in G. antrorsum), and by its
leaves with downward basal segments (patent or
upward in G. antrorsum). Additionally, G. antrorsum
has a longer gynoecium and rostrum than
G. brevicaule. G. sessiliflorum is a very variable species with a large geographical distribution, and
includes most of the differential characters proposed
for G. brevicaule. However, two quantitative characters permit the differentiation of the species, although
with some overlap. Petals of G. brevicaule are shorter
than in G. sessiliflorum, and the sepal mucro of
G. brevicaule is longer than in G. sessiliflorum. The
ratio between mucro length and sepal length also supports this difference.
Representative specimens examined: AUSTRALIA.
NEW SOUTH WALES: Bankstown City Council’s Native
Reserve, 7 Sylvan Grove, Picnic Point, 33∞58¢S,
150∞58¢E, 14.ii.1989, Miller 22 (NSW); Kosciusko
Hotel Dam, 36∞30¢S, 148∞12¢E, 24.i.1959, Carolin 778a
(K). TASMANIA: Ben Lomond National Park, along Rafferty Creek, 41∞34¢S, 147∞39¢E, 3.ii.1980, Noble 29008
(HO); Bluff Marshes, Drys Bluff, 41∞43¢S, 146∞48¢E,
13.i.1981, Brown 781 (HO); Fern Tree, 42∞55¢S,
147∞15¢E, 7.xii.1986, Morris 86146 (HO); Port Arthur,
43∞8¢S, 147∞50¢E, .i.1929, Ecbhin s.n. (HO); Preservation Island, 40∞29¢S, 148∞3¢E, 7.x.1979, Harris s.n.
(HO). NEW ZEALAND. NEW ZEALAND NORTH I: Blue
Creek Cave, Waikoropupu stream, 41∞20¢S, 175∞24¢E,
26.ii.1993, Lange 1998 (AK); Desert Rd., E Ruapehu,
39∞17¢S, 175∞34¢E, .i.1965, Druce s.n. (CHR); Hora
Hora, Middle Waikato, 37∞59¢S, 175∞38¢E, 16.xi.1912,
Petrie s.n. (WELT); Marton, Tyrone Farm, 39∞59¢S,
175∞22¢E, 9.iv.2001, Ogle 3823 (AK); Patateri Plateau,
38∞6¢S, 176∞4¢E, .i.1884, Cheeseman s.n. (AK). NEW
ZEALAND SOUTH I: Cass, 43∞2¢S, 171∞45¢E, 30.i.1978,
Lill s.n. (AK); Castle Hill, Canterbury, 43∞41¢S,
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
26
C. AEDO ET AL.
Figure 66. Area of distribution of Geranium brevicaule (dots) and G. antrorsum (triangles).
172∞42¢E, 31.xii.1957, Wood s.n. (AK); Christchurch,
Port hills, 43∞30¢S, 172∞42¢E, .1936, Lothian s.n. (GH);
Clutha Valley, Maori Point, near Tarras, 44∞50¢S,
169∞25¢E, 29.i.1974, Johnson & Hubbard s.n. (CHR);
Cobb reservoir, NW Nelson, 41∞6¢S, 172∞4¢E,
2.xii.1978, Gardner 2172 (L, CHR); Stewart Island,
46∞59¢S, 167∞48¢E, 16.xii.1962, White s.n. (AK).
3. Geranium antrorsum Carolin in Proc. Linn. Soc.
New South Wales ser. 2, 89: 357, pl. 6 fig. 14, pl. 7
fig. 10 (1965)
Type: Australia. New South Wales, Kosciusko
Hotel, 24.i.1959, Carolin 778b (holotype, NSW
66132; isotype, K!)
Herbs 5.5–15.3 cm tall. Rootstock 3.9–10.3 mm
diam., without fusiform roots. Basal leaves in a persistent rosette; lamina 0.9–3.3 ¥ 1.3–3.8 cm, polygonal in outline, cuneate to subtruncate (with basal
leaf segments patent or upward), palmatifid (divided
for 0.39–0.53 of its length), pilose, with appressed,
eglandular hairs; segments 5–7, obtriangular, 2.3–
6.8 mm at the base (segment width at the base/
segment length ratio = (0.16–)0.18–0.28), 3(-5)-lobed
at the apex (second sinus length/middle-segment
length ratio = (0.12–)0.15–0.16(-0.20)); petioles to
7 cm long, with patent to antrorse, appressed, eglandular hairs 0.3–1.6 mm long; stipules 5.7–14.2 ¥ 1.2–
3.5 mm, with eglandular hairs on abaxial surface
and on the margin, glabrous adaxially. Inflorescence with cymules 1-flowered, solitary; peduncles
0–1.2(-1.8) cm long, with antrorse, appressed, eglandular hairs 0.3–1 mm long; bracteoles 3.5–7.1 ¥ 0.8–
1.2 mm, linear-lanceolate, with eglandular hairs on
abaxial surface and on the margin, glabrous adaxially; pedicels 0.65–3.7 cm long, with antrorse,
appressed, eglandular hairs 0.2–0.9 mm long;
pedicel not overtopping the subtending leaf. Sepals
(4.3–)5.1–6.4(-8.2) ¥ 1.7–3.2 mm, lanceolate (width/
length ratio = (0.34–)0.38–0.43(-0.55)), with mucro
(1.2–)1.3–1.9(-2.9) mm long (mucro length/sepal
length ratio = (0.20–)0.23–0.36(-0.44)), with scarious
margins 0.1–0.2 mm wide, with erect-patent, eglandular hairs 0.5–0.8 mm long on the abaxial side
(patent hairs 1–1.4 mm long on the margin), glabrescent adaxially. Petals (3.6–)4.1–6.1(-8) ¥ 1.2–4.4 mm,
entire, without claw, glabrous, purplish. Filaments
2.4–3.6 mm long, yellowish, with eglandular hairs
0.2–0.3 mm long on the abaxial side and margin;
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GERANIUM SECTIONS ANDINA AND CHILENSIA
D
27
C
H
M
N
B
L
I
A
G
J
F
E
K
Figure 67. Geranium brevicaule Hook. f. A, habit; B–D, leaves; E, bracts; F, flower; G, flower without petals; H, sepal; I,
petal; J, stamen; K, fruit; L–M, mericarps; N, seed [A, C–N: Morris 86146 (HO); B: Hynes s.n. (AK).].
anthers 0.6–0.9 ¥ 0.4–0.7 mm, yellowish; pollen
unknown colour. Nectaries glabrous. Gynoecium
2.5–4 mm long, purplish. Fruit 13.9–16.5 mm long;
mericarps 2.6–3.9 ¥ 1.3–1.9 mm, smooth, with erectpatent, eglandular hairs 0.3–0.8 mm long, brownish;
rostrum 9–10.6 mm long, without a narrowed apex,
with erect-patent, eglandular hairs 0.2–0.9 mm long;
stigmatic remains (1–)1.4–1.7(-2.2) mm long, with 5
hairy lobes. Seeds 2.1–2.9 ¥ 1.2–1.7 mm, finely reticulate; hilum 1/6 as long as the perimeter.
Area: South-east Australia (Fig. 66).
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
28
C. AEDO ET AL.
Habitat: Dry, grassland with dominant Poa tussocks,
Eucalyptus open woodland, on granites; 1100–2000 m.
Illustration: Figure 68.
Notes: Geranium antrorsum is recognized by its
patent or upward basal leaf segments, giving a
Phenology: Flowering December – January (April).
D
C
B
E
H
A
I
J
K
M
L
F
G
Figure 68. Geranium antrorsum Carolin A, habit; B–D, leaves; E, apex of leaf in adaxial view; F, flower; G, pedicel; H,
sepal; I, petal; J, staminal filament; K, fruit; L, mericarp; M, seed [A, C, K–M: Carolin 798B (SYD); B, E: Eichler 13652
(SYD); D–F: Carolin s.n. (SYD); G–H, J: Adams 2493 (L); I: Beauglehole 33303 (SYD).].
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GERANIUM SECTIONS ANDINA AND CHILENSIA
cuneate to subtruncate aspect to the leaf. This species
has antrorse, appressed, eglandular hairs on the
pedicels. It has usually been considered as closely
related to G. brevicaule and G. sessiliflorum. However, these species have cordate leaves and pedicels
with patent to retrorse hairs. Additionally, G. antrorsum has sepals with mucro (1.2–)1.37–2.9 mm long,
which shows some overlap with G. brevicaule (with
mucro (0.48–)0.9–1.30(1.94) mm long), and none
with G. sessiliflorum (with mucro 0.3–0.7(1.1) mm
long).
Representative specimens examined: AUSTRALIA.
NEW SOUTH WALES: between Charlotte Pass and
Snowy River, 5.5 km ENE of Mt. Kosciusko, 36∞25¢S,
148∞19¢E, 5.ii.1957, Eichler 13652 (SYD); Cave Creek,
0.4 km N of the Blue Waterholes, 35∞37¢S, 148∞39¢E,
8.iv.1969, Rodd 815 (NSW); Cooleman Plain, Currango, 35∞37¢S, 148∞39¢E, .xii.1962, Walker 950 (K);
Coopers Swamp, Gourock Range, 25 mi S of Captains
Flat, Monaro Shire, 35∞56¢S, 149∞28¢E, 11.xii.1969,
Adams 2493 (L); Cotter River District, between Jack’s
Creek and the Cotter River, 35∞41¢S, 148∞50¢E,
14.xii.1960, Schodde 1220 (NSW); Nimmitable,
36∞31¢S, 149∞16¢E, 9.ii.1908, Cambage 1850 (NSW,
SYD); summit of Mt. Franklin, 35∞29¢S, 148∞46¢E,
8.xii.1984, Briggs 1760 (SYD). VICTORIA: Horsehair
Plain, 34 km from Omeo, toward Hotham Heights,
37∞2¢S, 147∞20¢E, 8.i.1969, Canning 1600 (L); Mt. Buffalo National Park, 1.6 km from The Horn turntable
toward the Chalet, 36∞39¢S, 146∞46¢E, 14.i.1969,
Canning 1764 (L); Native Dog Plain, BenambraWulgulmerang road, East Gippsland, 37∞4¢S,
148∞15¢E, 7.i.1970, Beauglehole et al. 33303 (SYD);
Nunniong Plains, East Gippsland, 37∞8¢S, 147∞56¢E,
20.i.1971, Beauglehole 36304 (SYD); Snowy Range,
Holmes Plain, Mt. Wellington, 37∞30¢S, 146∞48¢E,
27.xii.1972, Beauglehole 40759 (SYD).
4. Geranium parodii I.M. Johnst. in Contr. Gray
Herb. 81: 92 (1928)
Type: Argentina. Córdoba, Sierra de Achala,
1–4.xii.1926, Parodi 7514 (holotype, GH!)
Geranium sessiliflorum var. glabriusculum Kuntze,
Revis. Gen. Pl. 3(2): 33 (1898)
Type: Argentina. Córdoba, Sierra Achala, Pies de
los Gigantes, 3.xii.1878, Hieronymus s.n. (lectotype,
here selected, NY!) [31∞26¢S, 64∞48¢W]
Geranium magellanicum var. multifoliosum R.
Knuth in Engl., Pflanzenr. IV.129 (Heft 53): 70
(1912)
Type: Argentina. Córdoba, Sierra Achala, Cerro de
los Potrerillos, 1.ii.1877, Hieronymus 753 (lectotype, here designated, GOET!; isolectotype, F!)
Geranium magellanicum var. pumilum R. Knuth in
Engl., Pflanzenr. IV.129 (Heft 53): 70 (1912)
29
Type: Argentina. Córdoba, Sierra Achala, Esquina,
Quebrada de los Gigantes, 3.xii.1878, Hieronymus
s.n. (lectotype, here designated, G!)
Herbs 3.3–21 cm tall. Rootstock 7–14.3 mm diam.,
without fusiform roots. Basal leaves in a persistent
rosette; lamina 1.07–4.35 ¥ 1.16–4.57 cm, polygonal in
outline, cordate (with basal leaf segments downward),
palmatifid (divided for 0.54–0.79 of its length), pilose,
with appressed, eglandular hairs; segments 5–7,
rhombic to obtriangular, 1.1–2.5 mm at the base
(segment width at the base/segment length ratio =
0.08–0.17(-0.20)), 3–9-lobed in distal half (ratio second
sinus length/middle-segment length = (0.27–)0.31–
0.41(-0.44)); petioles to 13 cm long, with retrorse, ±
appressed, eglandular hairs 0.2–0.5 mm long; stipules
5.1–15.3 ¥ 1.21–2.2 mm, glabrous or with scattered
eglandular hairs on both surfaces and on the margin.
Inflorescence with cymules 1-flowered, solitary (rarely
in aggregates of 2–4 flowers); peduncles 0–9.4(-13.5)
cm long, with retrorse, ± appressed, eglandular hairs
0.1–0.4 mm long; bracteoles 2.3–5.8 ¥ 0.8–1.3 mm, linear-lanceolate, with scattered eglandular hairs on both
surfaces and on the margin; pedicels 1.45–6.20 cm
long, with retrorse, ± appressed, eglandular hairs
0.2–0.4 mm long; pedicel and peduncle together
usually not overtopping the subtending leaf. Sepals
(5.2–)6.1–7.2(-7.7) ¥ 1.8–3.4 mm, lanceolate (width/
length ratio = (0.27–)0.33–0.42(-0.50)), with mucro
(0.6–)0.62–0.98(-1.4) mm long (mucro length/sepal
length ratio = (0.08–)0.10–0.15(-0.18)), with scarious
margins 0.1–0.15 mm wide, with erect-patent, eglandular hairs 0.4–0.8 mm long. Petals (9.5–)11.5–
15.7 ¥ 3.2–9.4 mm, entire, without claw, hairy on margin and base of abaxial side, white. Filaments 4.3–
5.9 mm long, unknown colour, with eglandular hairs
0.1–0.4 mm long on the abaxial side and margin;
anthers 0.9–1.4 ¥ 0.6–0.85 mm, unknown colour; pollen unknown colour. Nectaries with a tuft of hairs at
the top, dorsally glabrous. Gynoecium 3.7–6.5 mm
long, unknown colour. Fruit 16.2–20.4 mm long; mericarps 3.2–4.1 ¥ 1.2–1.8 mm, smooth (sometimes with
1–2 transversal veins at the apex), with erect-patent,
eglandular hairs 0.35–0.6 mm long, brownish; rostrum
9.57–13.6 mm long, with a narrowed apex 0.7–1.5 mm
long, with erect-patent, eglandular hairs 0.2–0.5 mm
long; stigmatic remains (1.7–)2–2.3(-2.5) mm long,
with 5 glabrous lobes. Seeds 2.3–2.8 ¥ 1.2–1.4 mm,
finely reticulate; hilum 1/6 as long as the perimeter.
Area: Central Argentina (Córdoba and San Luis)
(Fig. 64).
Habitat: Wet grasslands; 1800–2600 m.
Phenology: Flowering November – February.
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
30
C. AEDO ET AL.
Illustration: Figure 69.
G. magellanicum (twice). However, G. parodii shows
some important characters that suggest that specific
status is more appropriate. Petals in G. parodii are
hairy on the abaxial side and on the margin, and
Notes: This endemic taxon from Sierra de Achala has
been described as a variety of G. sessiliflorum (once) or
M
K
L
F
1 cm
A
G
H
D
1 cm
1 cm
C
B
E
I
J
Figure 69. Geranium parodii I.M. Johnst. A–B, habit; C–D, leaves; E, bracts; F, flower; G, flower without petals; H, petal;
I–J, stamens; K, fruit; L, mericarp; M, seed [A, C–D, F–M: Hunziker 9622 (BH); B: Parodi 7514 (GH); E: Hunziker 1384
(CORD).].
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
GERANIUM SECTIONS ANDINA AND CHILENSIA
(9.5–)11.5–15.7 mm long. G. sessiliflorum and G. magellanicum have shorter petals (in the case of
G. sessiliflorum with some overlap), with few cilia on
the basal margin. Additionally, G. parodii has hairy
nectaries, and a fruit rostrum with a narrowed apex
0.7–1.5 mm long, whereas G. sessiliflorum and
G. magellanicum have glabrous nectaries and rostrum
without narrowed apex. Sepals of G. parodii have usually a mucro longer than in G. sessiliflorum and
G. magellanicum, and have a more homogeneous
indumentum (without longer hairs on the margin).
Geranium parodii has longer fruits, rostrum, pedicels,
and gynoecium than G. sessiliflorum.
This species shows some variation in plant size
and cymule structure. Small specimens have short
1-flowered cymules, without peduncle. In contrast,
more developed plants have cymules with longer
peduncles, which can be 1-flowered or, sometimes,
bear an aggregate of 2–4 flowers and a pair of opposite
leaves. A continuum of intermediate forms suggests
that these forms do not deserve taxonomic recognition.
Knuth (1912) described G. magellanicum var.
pumilum using heterogeneous material. For instance,
Hieronymus 449 is G. core-core and Hieronymus 585 is
G. leucanthum.
Representative specimens examined: ARGENTINA .
CÓRDOBA: Calamochita, pampa de San Miguel,
31∞43¢S, 64∞47¢W, 13.xii.1885, Kurtz 2944 (CORD);
Calamochita, Sierra Grande, al pie del Cerro Champaqui, 31∞59¢S, 64∞56¢W, 15.i.1952, Hunziker 9622
(BH); Cruz del Eye, Sierra de Achala, entre Tanti y
pampa de San Luis, 31∞19¢S, 64∞35¢W, 15.xii.1909,
Stuckert 20629 (CORD); Pampa de Achala, El Condor,
31∞35¢S, 64∞50¢W, 19.xii.1949, Meyer & Sleumer 15544
(LIL); Pies de los Gigantes, 31∞35¢S, 64∞49¢W, 9.i.1884,
Galander s.n. (NY); San Alberto, Sierra de Achala,
Cuesta del Tránsito, 31∞34¢S, 64∞50¢W, 5.i.1895, Kurtz
8339 (CORD, NY); San Javier, Cerro Champaquí,
31∞59¢S, 64∞56¢W, 14.ii.1924, Hosseus 13 (CORD). SAN
LUIS: El Monigote, 32∞48¢S, 66∞13¢W, 2.ii.1911, Pastore
36 (SI); Merlo, Cuesta M. Bayo, 32∞21¢S, 65∞1¢W,
27.xi.1910, Pastore 99 (SI).
II. Geranium sect. Chilensia R. Knuth in Engl., Pflanzenr. 54: 45, 68 (1912)
Type: G. patagonicum (lectotype, designated by R.
Knuth, 1912: 45)
Perennial herbs with rootstock usually vertical, not
tuberculate, turnip-shaped, with or without fusiformswollen roots; aerial stem leaved, herbaceous, without
vegetative stems or stolon. Leaf lamina polygonal in
outline, cordate (with basal leaf segments downward),
without abscission zone, palmatifid, hairy, not coriaceous, and with nerves not projected; segments 5–7,
obtriangular or rhombic, 3–17-lobed in the distal half;
31
cauline leaves opposite; stipules lanceolate, free,
papery, reddish, hairy. Inflorescence in monochasial
cyme with 2-flowered cymules (rarely 1-flowered),
solitary (usually on the aerial stem); peduncles and
pedicels hairy; bracteoles lanceolate to linearlanceolate, hairy; pedicel and peduncle together
usually overtopping the subtending leaf. Sepals ovate,
smooth, not accrescent, 3-nerved, mucronate, with
scarious margins, hairy. Petals erect-patent, ±
obovate, entire or emarginate, usually without claw,
hairy, purplish to white. Stamens 10, both whorls
bearing anthers; filaments not exserted, lanceolate,
yellowish or white, hairy. Nectaries hemispherical,
usually glabrous. Fruit of the seed-ejection type;
mericarps smooth, without longitudinal rib, without
basal beak, with a basal callus, without a basal prong,
usually with eglandular hairs, brownish or blackish;
rostrum with or without a narrowed apex; stigmatic
remains with 5 hairy or glabrous lobes. Seeds ellipsoid, finely reticulate, brownish; hilum 1/4–1/6 as
long as the perimeter. Cotyledons entire.
Notes: Species of Geranium sect. Chilensia are recognized by their aerial developed stem, turnip-shaped
rootstock, and 2-flowered cymules. This morphological
characterization is basically as Knuth (1912) proposed, although several taxa have been reduced to
synonyms. Consequently this section includes nine
species from South America and two species from Australia and New Zealand.
5. Geranium berteroanum Colla in Nuovo Giorn.
Lett., Sci. 24: 143 (1832). Geranium proximum
Bertero ex Steud. in Flora 39(28): 438 (1856)
Type: Chile. Valparaíso, Quillota 1829, Bertero
1020 (lectotype, here designated, BM!; isolectotype,
P!) [32∞54¢N, 71∞16¢W]
Geranium intermedium Bertero ex Colla in Nuovo
Giorn. Lett., Sci. 24: 143 (1832) [syn. subst.], nom.
illeg., non E. James (1823). Geranium collae Aedo,
Muñoz Garm. & Pando in Anales Jard. Bot. Madrid
56(2): 224 (1998)
Type: Chile. O’Higgins, Rancagua, Bertero s.n.
(lectotype, here designated, SGO-51197!)
Geranium patagonicum Hook. f., Fl. Antarct. 2: 252
(1845). Geranium dissectum var. patagonicum
(Hook. f) Speg. in Revista Fac. Agron. Univ. Nac. La
Plata 3: 500 (1897)
Type: Chile. Patagonia, Port Famine, King s.n.
(lectotype, here designated, K!) [53∞38¢S, 70∞56¢W]
Geranium apricum Phil. in Linnaea 28: 676 (1856).
Geranium berteroanum var. apricum (Phil.) Reiche
in Anales Univ. Chile 93: 578 (1895)
Type: Chile. Valdivia, San Juan, Philippi s.n. (lectotype, here designated, SGO-40601!; isolectotypes,
LE!, W!)
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
32
C. AEDO ET AL.
Geranium submolle Steud. in Flora 39(28): 438
(1856)
Type: Chile. Rancagua, montis La Leona, Bertero
294 (lectotype, here designated, P!; isolectotypes,
G! MO!)
Geranium andinum Phil. in Anales Univ. Chile 82:
727 (1893)
Type: Chile. O’Higgins, Popeta, Las Leñas, Jan.
1881, Philippi s.n. (lectotype, here designated,
SGO-51217!)
Geranium caespitosum Phil. in Anales Univ. Chile
82: 730 (1893) [syn. subst.], nom. illeg., non E.
James (1823). Geranium berteroanum var. caespitosum Reiche in Anales Univ. Chile 93: 578 (1895).
Geranium chilense Aedo & Muñoz Garm. in Kew
Bull. 52(3): 725 (1997)
Type: Chile. O’Higgins, baños de Cauquenes,
.ix.1879, Philippi s.n. (lectotype, here designated,
SGO-51204!, photo P!)
Geranium hispidum Phil. in Anales Univ. Chile 82:
732 (1893) [syn. subst.], nom. illeg., non L. f. (1782).
Geranium berteroanum var. hispidum Reiche in
Anales Univ. Chile 93: 578 (1895). Geranium neohispidum Aedo & Muñoz Garm. in Kew Bull. 52(3):
725 (1997)
Type: Chile. Santiago, Las Mercedes, October 1888,
Philippi s.n. (lectotype, here designated, SGO51214!, photo GH! photo P!)
Herbs 13.5–59 cm tall. Rootstock 5.8–20.4 mm diam.,
without fusiform roots. Stem erect to ascending, with
patent, eglandular hairs 0.4–1.9 mm long and, sometimes, patent, glandular hairs 0.3–1.1 mm long. Basal
leaves in a deciduous rosette; lamina 1.8–9 ¥ 2.6–
8.4 cm, polygonal in outline, cordate, palmatifid
(divided for 0.57–0.82 of its length), pilose, with
appressed, eglandular hairs, sometimes glabrous on
the adaxial side; segments 5–7, obtriangular, 1.8–
10.6 mm at the base (segment width at the base/
segment length ratio = (0.11–)0.17–0.22(-0.31)), 3–11lobed in distal half (second sinus length/middlesegment length ratio = (0.13–)0.24–0.38(-0.42)); petioles to 29.5 cm long, with patent, eglandular hairs
0.5–2 mm long, and sometimes, patent, eglandular
hairs 0.3–0.9 mm long; stipules 4–9 ¥ 0.6–3 mm, with
eglandular hairs on both surfaces and on the margin.
Inflorescence with cymules 2-flowered, solitary;
peduncles (0.8–)1.7–4.5(-5.8) cm long, with patent,
eglandular hairs 0.2–1.2 mm long and sometimes,
patent, glandular hairs 0.3–1.1 mm long; bracteoles
2.5–6.7 ¥ 0.4–1.1 mm, linear-lanceolate, with eglandular hairs on both surfaces and on the margin; pedicels
0.9–2.7 cm long, with patent, eglandular hairs 0.2–
1.2 mm long and sometimes, patent, glandular hairs
0.3–1.1 mm long; pedicel and peduncle together
usually overtopping the subtending leaf. Sepals
(4.4–)5.4–6.1(-7) ¥ 2–3.4 mm,
lanceolate
(width/
length ratio = (0.40)0.44–0.56(-0.68)), with mucro
(0.35–)0.6–0.8(-0.9) mm long (mucro length/sepal
length ratio = (0.07–)0.09–0.14(-0.16)), with scarious
margins 0.2–0.4 mm wide, with erect-patent, eglandular hairs 0.7–1.8 mm long and sometimes, patent,
glandular hairs 0.3–1.2 mm long. Petals (5.1–)6.5–
8(-9) ¥ 2.5–4.5 mm, entire or slightly notched, without
claw, glabrous on both sides, ciliate on the basal margin, purplish. Filaments 2.5–4.5 mm long, whitish to
yellowish, with eglandular hairs 0.1–0.4 mm long on
the abaxial side and margin; anthers 0.4–0.8 ¥ 0.3–
0.7 mm, whitish to pink; pollen yellow. Nectaries glabrous. Gynoecium 2.9–5.2 mm long, whitish to pink.
Fruit 11.4–24.6 mm long; mericarps 2.3–4.2 ¥ 1.1–
2.3 mm, smooth, with erect-patent, eglandular hairs
0.4–1.5 mm long and, sometimes, patent glandular
hairs 0.2–0.8 mm long, brownish; rostrum 8.5–
18.1 mm long, without a narrowed apex, with erectpatent, eglandular hairs 0.4–1 mm long and, sometimes, patent glandular hairs 0.3–1.1 mm long; stigmatic remains (1.1–)1.3–1.7(-2.1) mm long, with 5
hairy lobes. Seeds 1.3–3.1 ¥ 0.8–1.8 mm, finely reticulate; hilum 1/6 as long as the perimeter.
Area: Argentina and Chile (Fig. 70).
Habitat: Roadsides, dunes, meadows, shrubby areas,
and Acacia, Araucaria, Aristotelia or Nothofagus forest; 0–3800 m.
Phenology: Flowering January – December.
Illustration: Figure 71.
Notes: Geranium berteroanum is distributed over a
large area of more than 2800 km between the northernmost localities in Central Chile and the southernmost ones in Tierra de Fuego. This species varies in
the presence of glandular hairs. Some specimens have
only eglandular hairs while others also have glandular hairs widespread on the stem, inflorescence,
sepals, rostrum and/or mericarps. These glandular
forms have occasionally been misidentified as
G. fiebrigianum (included in G. fallax in this study).
Geranium berteroanum can be distinguished from
G. fallax by its leaves with an obtriangular middle
segment (not rhombic), and its rostrum fruit without a
narrow apex.
Geranium skottsbergii is also a species with eglandular and glandular forms, partially sympatric with
G. berteroanum, which can be distinguished by its
longer petals and more deeply divided leaves. Geranium solanderi is endemic to Australia and New
Zealand which strongly resembles G. berteroanum.
Both species share pedicels with patent hairs, and
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
GERANIUM SECTIONS ANDINA AND CHILENSIA
33
Figure 70. Area of distribution of Geranium berteroanum.
many quantitative characters (petal length, sepal
mucro length, etc.) have a broad overlap. However,
G. berteroanum has longer bracteoles, sepals and stigmatic remains than G. solanderi.
Colla (1832) described G. berteroanum using material collected by Bertero near Valparaíso, which was
published as ‘G. tuberosum L.’ by Bertero in a local
newspaper (Bertero, 1829). Steudel (1856) published
his G. proximum based on the same material, but
indicating two concrete specimens (Bertero 1019 and
Bertero 1020). These specimens are heterogeneous,
the first belonging to G. skottsbergii and the second to
G. berteroanum. We searched at TO where Colla
original herbarium is kept, but Dr Forneris kindly
informed us that there is no syntype there. Thus we
have selected Bertero 1020 (kept at BM) as the lectotype of G. berteroanum and G. proximum. Geranium
submolle was described by Steudel (1856) on a heterogeneous collection (Bertero 294). Most of specimens
belong to G. berteroanum, but some material of P and
G are G. skottsbergii, which must be excluded from the
type.
Gay (1845) recorded G. pyrenaicum Burm. f. from
Chile, indicating that there was only one specimen in
his herbarium. Later, this species was included by
Marticorena & Quezada (1985) in their check-list of
Chile. During the revision of sect. Batrachioidea
W.D.J. Kochy, to which G. pyrenaicum belongs, no new
data were provided (Aedo et al., 1998a) as no specimen
from Chile could be studied. Now the specimen supporting this record has been located at P (Colchagua,
.ii.1831, Gay s.n.), and identified as G. berteroanum,
which permits rejection of the presence of
G. pyrenaicum in Chile.
According to Stuessy & Marticorena (1990)
‘berteroanum’ is the correct form to latinize this
specific epithet, originally proposed as ‘Berterianum’.
Representative specimens examined: ARGENTINA.
BUENOS AIRES: Suerra de Ventana, Cerro el Abra,
38∞3¢S, 61∞59¢W, 17.xi.1969, Roivainen 376 (H); Tornquist, Sierra de la Ventana, Cerro Ventana, 38∞2¢S,
62∞0¢W, s.d., Proyecto Ventania 349b (LP). CHUBUT:
43 km S of Cholila on route 258, into Parque Nacional
Los Alerces, 42∞48¢S, 71∞43¢W, 3.xii.1984, Stuessy et al.
6806 (LP); Cushamen, Cholila, 42∞30¢S, 71∞26¢W,
5.i.1901, Illin 124 (G, BR, CORD, SP). MENDOZA: Las
Heras, entre Villavicencio y Los Hornillos, 32∞30¢S,
69∞1¢W, 26.xii.1947, Ruiz Leal 11083 (CORD, MERL);
Luján, Cerro Cachenta, 33∞2¢S, 69∞6¢W, .v.1987, Roig
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34
C. AEDO ET AL.
F
D
G
A
J
I
E
H
B
K
C
M
L
Figure 71. Geranium berteroanum Colla. A, habit; B–C, leaves; D, bracts; E, flower; F, flower without petals; G, sepal; H,
petal; I–J, stamens; K, fruit; L, mericarp; M, seed [A–E, G–M: Aedo 7391 (MA); f: Aedo 6727 (MA).].
12619 (MERL). NEUQUÉN: Aluminé, Pulmarí, 39∞5¢S,
70∞58¢W, 8.xii.1981, Cabrera et al. 32920 (SI); El Cruce,
villa La Angostura, 40∞47¢S, 71∞40¢W, 13.xii.1946,
Barba 1197 (LIL). RÍO NEGRO: 6 km de Pilcaniyen,
camino de Bariloche, 40∞45¢S, 70∞30¢W, 28.i.1944,
Nicora 3677 (SI); Arroyo Cascada, Cerro Catedral,
41∞13¢S, 71∞30¢W, 30.i.1945, Barba 146 (NY). SANTA
CRUZ: Deseado, cañadón frente a la estación de Biología
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GERANIUM SECTIONS ANDINA AND CHILENSIA
Marina, 47∞44¢S, 65∞56¢W, .x.1970, Crespo & Troncoso
1694 (SI); Estancia lago Roca, río Rico, 50∞30¢S,
72∞44¢W, 24.xii.1958, James 380 (BM). TIERRA DE
FUEGO: Fuegia, Lago Fagnano, Pumilio shore, 54∞35¢S,
67∞40¢W, 18.iii.1908, Skottsberg 247 (S, UPS); Ushuaia,
54∞50¢S, 68∞26¢W, 13.ii.1953, Ruiz Leal 14961 (CORD).
CHILE. AISÉN: Coihaique, 45∞34¢S, 72∞4¢W, 25.i.1934,
Espinosa s.n. (SGO); Los Mallines, near Balmaceda,
45∞55¢S, 71∞41¢W, 24.i.1943, Bruzzone 42 (LP). BIOBÍO:
Concepción, 36∞50¢S, 73∞3¢W, 18.xi.2001, Aedo 6913
(MA); parque Hualpén, morro Pompón, 36∞48¢S,
73∞10¢W, 21.xi.2001, Aedo 6931 (MA). COQUIMBO: cerros del Tofo, 68 km N of La Serena, 29∞26¢S, 71∞15¢W,
1.xi.1938, Worth & Morrison 16297 (K, GH, UC);
Ovalle, bosque de Talinay, S desembocadura del
Limarí, 30∞39¢S, 71∞43¢W, 18.xi.1940, Muñoz & Coronel
1267 (SGO). LA ARAUCANIA: 8 km al N de Yupehue,
playa Casas de Piedra, 38∞28¢S, 73∞30¢W, 21.xii.2001,
Aedo 7185 (MA); camino entre Cherquenco y
Melipeuco, 38∞43¢S, 71∞57¢W, 23.xii.2001, Aedo 7243
(MA). LOS LAGOS: Llancacura, 40∞18¢S, 73∞24¢W,
11.i.1961, Schlegel 3444 (CONC); Llanquihue,
Cayutué, 41∞16¢S, 72∞16¢W, .i.1973, Martínez s.n.
(CONC). MAGALLANES: Fuerte Bulnes, 53∞37¢S,
70∞56¢W, 25.iv.1951, Cekalovic s.n. (SGO); pr. Puerto
Natales, 51∞33¢S, 72∞40¢W, 19.i.2002, Aedo 7472 (MA).
MAULE: Constitución, playa el Cable, 35∞21¢S, 72∞27¢W,
14.xii.2001, Aedo 7154 (MA); Valdivia, Las Quinas,
35∞12¢S, 72∞16¢W, .i.1903, Buchtien s.n. (US). O’HIGGINS: Colchagua, Punta Lobos, 5 km al S de Pichilemu,
34∞24¢S, 72∞2¢W, 31.x.1973, Stebbins 8774 (SGO); Hacienda de Cauquenes, cajón del Ciprés, 34∞25¢S, 70∞25¢W,
.1875, Philippi s.n. (M). SANTIAGO: Colina, Baños de
Colina, 33∞11¢S, 70∞36¢W, 8.xi.2001, Aedo 6737 (MA);
puente Manzanito, 33∞20¢S, 70∞19¢W, 12.xii.2001, Aedo
7136 (MA). TIERRA DE FUEGO: Bahia Inutil, estancia
Cameron, 53∞22¢S, 69∞16¢W, 19.iii.1964, Moore 1064
(SGO); camino a Puerto Arturo, cabo Nose, 53∞44¢S,
70∞8¢W, 24.i.1994, Pisano et al. 7727 (CONC). VALPARAÍSO: Aconcagua, Petorca, carretera panamericana,
4 km antes del puente Guaquén, 32∞18¢S, 71∞24¢W,
15.x.1971, Marticorena et al. 1286 (CONC); Juncal,
Uspallata pass, 32∞49¢S, 70∞5¢W, .ii.1903, Buchtien s.n.
(US, BM, GH).
6. Geranium solanderi Carolin in Proc. Linn. Soc.
New South Wales ser. 2, 89: 350 (1965) Geranium
pilosum Sol. ex Willd., Sp. Pl. 3(1): 706 (1801) [syn.
subst.], nom. illeg., non Cav. (1788). Geranium
dissectum var. pilosum Hook. f., Fl. Tasman. 1: 56
(1855). Geranium dissectum var. australe Benth.,
Fl. Austral. 1: 296 (1863). Geranium carolinianum
var. australe (Benth.) Fosberg in Occas. Pap. Univ.
Hawaii 32: 6 (1937)
Type: New Zealand [without locality], Forster 531
(lectotype, designated by Carolin, 1965: 351, K!)
35
Geranium dissectum f. tasmanica Gand. in Bull.
Soc. Bot. France 47: 306 (1901)
Type: Australia. Tasmania, Stone Pit Chapel,
December 1875, Spicer 44 (lectotype, here designated, LY digital image!)
Geranium drummondii Carolin in Proc. Linn. Soc.
New South Wales ser. 2, 89: 353, pl. 6 fig. 4 (1965)
Type: Australia. Swan River, Drummond 4 bi (holotype, K!; isotype, E!)
Herbs 12–100 cm tall. Rootstock 2.4–19.1 mm diam.,
without fusiform roots. Stem erect to ascending, with
patent to retrorse, not appressed, eglandular hairs
0.4–1.8 mm long. Basal leaves in a ± deciduous
rosette; lamina 1.2–4.5 ¥ 1.4–5.7 cm, polygonal in outline, cordate, palmatifid (divided for 0.47–0.82 of its
length), pilose, with ± erect, eglandular hairs; segments 5–7, obtriangular, 2.2–6.8 mm at the base
(segment width at the base/segment length ratio =
(0.12–)0.14–0.24(-0.30)), 3–10-lobed in distal half
(second sinus length/middle-segment length ratio =
(0.13–)0.20–0.35(-0.47)); petioles to 18 cm long, with
patent, eglandular hairs 0.4–1.7 mm long; stipules
2.2–8 ¥ 0.5–2.1 mm, with eglandular hairs on abaxial
surface and on the margin, glabrous adaxially.
Inflorescence with cymules 2-flowered, solitary;
peduncles (0.4–)1.8–4(-6.1) cm long, with patent to
retrorse, not appressed, eglandular hairs 0.3–2 mm
long; bracteoles 1.5–7.4 ¥ 0.3–0.8 mm, lanceolate,
with eglandular hairs on abaxial surface and on the
margin, glabrous adaxially; pedicels 0.8–3.4 cm long,
with patent to retrorse, not appressed, eglandular
hairs 0.2–2 mm long; pedicel and peduncle together
usually overtopping the subtending leaf. Sepals
(3.2–)4.2–5.2(-6.3) ¥ 1.6–2.7 mm, lanceolate (width/
length ratio = (0.38–)0.43–0.51(-0.62)), with mucro
(0.2–)0.4–0.7(-0.9) mm long (mucro length/sepal
length ratio = (0.04–)0.08–0.13(-0.25)), with scarious
margins 0.1–0.2 mm wide, with eglandular hairs
0.1 mm long on the abaxial side (and eglandular hairs
0.4–1.8 mm long on the margin), glabrous adaxially.
Petals (3–)4.7–6(-8.1) ¥ 1.4–4.2 mm, entire, without
claw, glabrous on both sides, ciliate on the basal
margin, purplish. Filaments 2.4–4.5 mm long, yellowish, glabrous on both sides, ciliate on the basal
margin, with hairs up to 0.1–0.3 mm long; anthers
0.4–0.8 ¥ 0.2–0.7 mm, yellowish; pollen unknown
colour. Nectaries glabrous. Gynoecium 2.5–4.8 mm
long, yellowish. Fruit 13.1–20.6 mm long; mericarps
2.2–3.2 ¥ 1.1–1.8 mm, smooth, with erect-patent,
eglandular hairs 0.1–1.5 mm long, usually blackish;
rostrum 8.8–15.1 mm long, without a narrowed apex,
with erect-patent, eglandular hairs 0.1–1.1 mm long;
stigmatic remains (0.6–)1–1.4(-1.6) mm long, with 5
hairy lobes. Seeds 1.6–2.2 ¥ 0.9–1.6 mm, reticulate;
hilum 1/6 as long as the perimeter.
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C. AEDO ET AL.
Figure 72. Area of distribution of Geranium solanderi (Californian localities are not shown).
Area: Australia (Norfolk I, south-west Australia,
south-east Australia and Tasmania), and New
Zealand (North I, South I and Chatham Is); introduced in USA. (California) (Fig. 72).
Habitat: Dunes, lava tongues, river gorges, moist
grassy areas, shrublands, and Acacia, Agonis, Callitris, Eucalyptus or Melaleuca forest; 0–1960 m.
Phenology: Flowering January – December.
Illustration: Figure 73.
Notes: Gardner (1984) studied the variability of
G. solanderi in New Zealand, and recognized two
forms based mainly on petal length, and indumentum,
named ‘large petals’ and ‘coarse hairs’. However, this
author found that intermediate character states or a
combination of the features diagnostic of each form
were common in Australian material, which has been
confirmed in this study. All these data suggest that
such forms do not deserve taxonomic recognition.
Carolin (1965) described G. drummondii based on
three specimens from south-west Australia. These
specimens are characterized by a dense and entangled
indumentum especially on pedicels, sepals and mericarps. However intermediate forms are common in
this area and in other Australian localities. Considering these data and the great variability of G. solanderi
throughout its whole area we prefer to subsume
G. drummondii as a synonym of G. solanderi.
Geranium solanderi strongly resembles G. berteroanum, which is an endemic from the Andes. The
differences are discussed under the second species.
Additionally, it should be indicated that among the
studied material of G. solanderi no specimen with
glandular hairs has been found, whereas in
G. berteroanum specimens with glandular hairs are
common.
Representative specimens examined: AUSTRALIA .
NEW SOUTH WALES: 37 km from Corrong Victoria,
toward Omeo, 34∞11¢S, 144∞28¢E, 4.i.1969, Canning
1415 (GH); Armidale, university grounds, 33∞30¢S,
151∞39¢E, 5.xi.1958, Paterson s.n. (G); Bowral, 34∞28¢S,
150∞25¢E, 2.ii.1959, Carolin 829 (L); Bungonia state
recreation reserve, 34∞48¢S, 150∞30¢E, 13.ix.1978,
Canning 4396 (CANB); Canberra, 35∞16¢S, 149∞9¢E,
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GERANIUM SECTIONS ANDINA AND CHILENSIA
F
A
37
I
E
G
J
C
D
K
L
H
B
M
Figure 73. Geranium solanderi Carolin. A, habit; B, rootstock; C, leaf; D, peduncle; E, bracts; F, flower; G, flower without
petals; H, sepal; I, petal; J, stamen; K, fruit; L, mericarp; M, seed [A–M: Keighery & Gibson 436 (PERTH).].
12.xi.1962, McKee 9665 (L, BM). QUEENSLAND: Darling Downs District, 3 miles SE of Blaxland, 27∞12¢S,
151∞24¢E, 13.x.1940, Smith & Everist 799 (GH); Lockyer valley, 27∞25¢S, 152∞36¢E, 11.v.1943, Clemens 42/
65 (GH); Maryborough, Mary river, 25∞32¢S, 152∞42¢E,
14.x.1930, Hubbard 4367 (L); Millmerran, 27∞52¢S,
151∞16¢E, 15.iii.1931, Hubbard 5848 (K, L); Pale
Creek, 28∞17¢S, 152∞48¢E, 1.vi.1959, Carolin 1023 (G).
SOUTH AUSTRALIA: 8 km S of Lucindale, 37∞2¢S,
140∞21¢E, 19.xii.1985, Gibbons 506 (HO); Adelaide,
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C. AEDO ET AL.
34∞55¢S, 138∞36¢E, s.d., Whittaker s.n. (K); Moreton
Bay, 34∞30¢S, 136∞17¢E, s.d., Mueller s.n. (E); Mt. Lofty
range, Saunders Creek, 34∞42¢S, 139∞9¢E, 26.vii.1969,
Blaylock 1313 (AK). TASMANIA: Ball Bay, 41∞8¢S,
146∞52¢E, .xii.1955, Curtis s.n. (HO); Hobart town,
42∞52¢S, 147∞19¢E, 13.xii.1839, Le Guillou s.n. (P);
Marrawak, Green Point, 40∞54¢S, 144∞39¢E, 26.i.1969,
Canning 1850 (L); Mt. Wellington, 42∞54¢S, 147∞14¢E,
.1971, Allan s.n. (HO); Spring beach, Prosser Bay,
42∞34¢S, 147∞54¢E, 19.xii.1952, Melville 2538 (K, GH,
PERTH). VICTORIA: 21 km from Omeo, toward Mitta
Mitta, 37∞5¢S, 147∞35¢E, 6.i.1969, Canning 1525 (US);
Altona, at the mouth of Skeleton Creek, 37∞52¢S,
144∞49¢E, 21.xi.1967, Cullimore 100 (PERTH); Gramphians mts., 37∞15¢S, 142∞26¢E, .xii.1912, Tilden s.n.
(U); Hume highway, km 31 from Chiltern-Beechworth
exit to Wodonga, 36∞7¢S, 146∞53¢E, 7.x.1978, Canning
4410 (L); Port Phillip, 37∞51¢S, 144∞58¢E, .1839, Duncan s.n. (E). WESTERN AUSTRALIA: 1.7 km S intersection of Sollya road and Brockman highway, 34∞5¢S,
115∞34¢E, 20.i.1997, Godden & Day 72.4 (PERTH);
2.2 km E of Caves road along Moses rock road,
33∞46¢S, 115∞0¢E, 5.xii.1996, Casson & Annels 39.2
(PERTH); 3 km N of Yallingup, 33∞41¢S, 115∞2¢E,
1.x.1989, Keighery s.n. (PERTH); along Balladonia
road, S of Mt Rogged, E of Esperance, 33∞51¢S,
121∞53¢E, 5.xii.1971, Royce 10150 (PERTH); Ballingup, 33∞47¢S, 115∞59¢E, 28.x.1946, Royce 1330
(PERTH). NEW ZEALAND. CHATHAM IS: Lake
Huro, Mangape Stream Outlet, 43∞56¢S, 176∞31¢W,
29.iii.1996, Lange 225 (AK); NEW ZEALAND NORTH I:
Aorangi Island, Arid Point, 35∞29¢S, 174∞45¢E,
24.x.1995, Lange 3154 (AK); Auckland city, mount
Wellington, 36∞5¢S, 174∞5¢E, 3.v.1978, Gardner 1944
(AK); Bay of Islands County, Opito Bay, 35∞12¢S,
174∞3¢E, 9.x.1972, Orchard 3486 (AK); Cavalli Island
group, Haraweka Island, 34∞58¢S, 173∞58¢E, 3.i.1979,
Wright 3008 (AK); Great Barrier I., Broken Islands,
Mahuki Island, 36∞14¢S, 175∞18¢E, 2.i.1985, Wright
6887 (AK). NEW ZEALAND SOUTH I: Botanical Hill,
Nelson, 41∞16¢S, 173∞18¢E, 17.xi.1980, Gardner 2777
(AK); Christchurch city, Saint Albans, 43∞30¢S,
172∞39¢E, 1.xii.2001, Sykes s.n. (AK); Mackay’s Bluff,
41∞11¢S, 173∞22¢E, 16.xi.1980, Gardner 2775 (AK); Te
Kakaho Island, 40∞54¢S, 174∞6¢E, 26.iii.1984, Wright
6371 (AK). USA. CALIFORNIA: Humboldt Co., Trinidad, 41∞3¢N, 124∞8¢W, 11.vi.1911, Tracy 3238 (UC).
patent to retrorse, not appressed, eglandular hairs
0.5–1.2 mm long. Basal leaves in a deciduous rosette;
lamina 1.3–5.5 ¥ 1.7–6.6 cm, polygonal in outline, cordate, palmatifid (divided for 0.46–0.78 of its length),
pilose, with appressed, eglandular hairs; segments
5–7, obtriangular, 3.6–9 mm at the base (segment
width at the base/segment length ratio = (0.20–)0.24–
0.32(-0.42)), 3–11-lobed in distal half (second sinus
length/middle-segment length ratio = (0.14–)0.24–
0.29(-0.34)); petioles to 20 cm long, with patent to retrorse, appressed, eglandular hairs 0.5–1.4 mm long;
stipules 2.5–9 ¥ 0.7–3.4 mm, with eglandular hairs on
both surfaces and on the margin. Inflorescence with
cymules 2-flowered, solitary; peduncles (1.1–)2–
3.7(-6.8) cm long, with patent to retrorse, eglandular
hairs 0.4–1.6 mm long; bracteoles 2–5.9 ¥ 0.5–1.9 mm,
linear-lanceolate, with eglandular hairs on both surfaces and on the margin; pedicels 0.7–2.2 cm long,
with patent to retrorse, eglandular hairs 0.4–1.6 mm
long; pedicel and peduncle together usually not
overtopping the subtending leaf. Sepals (3.5–)4.5–
5.5(-6.3) ¥ 1.7–4.2 mm,
lanceolate
(width/length
ratio = (0.36–)0.52–0.65(-0.89), with mucro (0.2–)0.3–
0.4(-0.5) mm long (mucro length/sepal length
ratio = (0.03–)0.05–0.07(0.09)), with scarious margins
0.1–0.25 mm wide, with erect-patent, eglandular
hairs 0.3–1.7 mm long. Petals (5–)5.6–7.5(-8.2) ¥
2–4.6 mm, entire, without claw, glabrous on both
sides, ciliate on the basal margin, purplish. Filaments
2.6–4.9 mm long, yellowish, with eglandular hairs
0.1–0.4 mm long on the abaxial side and margin;
anthers 0.4–0.8 ¥ 0.3–0.6 mm, unknown colour; pollen
unknown colour. Nectaries usually with a tuft of hairs
at the top, dorsally glabrous. Gynoecium 3–5 mm long,
unknown colour. Fruit 11–21.1 mm long; mericarps
2.7–3.5 ¥ 1.3–2 mm, smooth, with erect-patent,
eglandular hairs 0.4–1.1 mm long, brownish; rostrum
7.5–15.4 mm long, without a narrowed apex, with
erect-patent, eglandular hairs 0.3–1.3 mm long; stigmatic remains 1.1–1.5(-1.8) mm long, with 5 hairy
lobes. Seeds 1.8–2.5 ¥ 1.1–1.6 mm, finely reticulate;
hilum 1/6 as long as the perimeter.
7. Geranium limae R. Knuth in Engl., Pflanzenr.
IV.129 (Heft 53): 74 (1912)
Type: Peru. Lima, San Lorenzo, July 1836, Gaudichaud 4 (lectotype, here designated, P!; isolectotypes, G!, US!)
Phenology: Flowering (April) July – November.
Herbs 8–55 cm tall. Rootstock 8.7–24.4 mm diam.,
without fusiform roots. Stem erect to ascending, with
Area: Peru (Fig. 74).
Habitat: Coastal desert, on hillsides in sandy or rocky
areas; 0–2800 m.
Illustration: Figure 75.
Notes: Geranium limae shares with G. albicans
and G. berteroanum patent hairs on peduncles and
pedicels. The differences between G. limae and
G. albicans are indicated under the first species. In
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GERANIUM SECTIONS ANDINA AND CHILENSIA
39
Figure 74. Area of distribution of Geranium limae (triangles) and G. fallax (dots).
addition, the ranges of G. limae and G. albicans are
completely nonoverlapping. Geranium limae has
sepals shorter and with a shorter mucro than
G. berteroanum and hairs of the rostrum are longer
than in G. berteroanum. The base of the middle segment is wider in G. limae than in G. berteroanum, and
the main sinus of the middle segment is not as deep as
in G. berteroanum, which gives a less deeply divided
appearance to G. limae leaves. In G. limae nectaries
usually have a tuft of hairs at the top, whereas they
are always glabrous in G. berteroanum. Glandular
hairs are always lacking in G. limae, whereas they can
be present in G. berteroanum.
Knuth (1912) indicated, probably by mistake, that
Gaudichaud collected the original material of G. limae
in 1841. Gaudichaud acted as botanist to La Bonite
during its circumnavigation of the globe in 1836–1837,
and the label of the type specimen clearly states
‘Juillet 1836’.
Representative specimens examined: PERU. ANCASH:
Santa, lomas de La Chay, 40 km N of Barranca,
10∞26¢S, 77∞57¢W, 18.ix.1938, Stork et al. 9207 (UC,
GH). AREQUIPA: Camana, ravine N of Atiquipa,
15∞47¢S, 74∞22¢W, 20.ix.1938, Worth & Morison 15665
(UC, GH); lomas de Atiquipa, cerca de Chala, entre
Nazca y Chala, 15∞48¢S, 74∞22¢W, 20.x.1946, Ferreyra 1493 (MA, GH, US). LA LIBERTAD: cerro
Campana, km 565 cerca de Trujillo, 8∞0¢S, 79∞7¢W,
18.viii.1952, Ferreyra 8620 (MA). LAMBAYEQUE:
Chiclayo, Cerro Reque, 6∞50¢S, 79∞46¢W, 29.vii.1979,
Llatos 497 (NY, GH). LIMA: 6 km E of Pachacamac,
12∞13¢S, 76∞52¢W, 6.ix.1953, Saunders 187 (BM, UC);
Amancaes, 12∞1¢S, 77∞1¢W, 23.ix.1940, Asplund
13754 (S, G, GH); Atocongo, 12∞7¢S, 76∞56¢W,
10.xi.1946, Ferreyra 1539 (MA); Chancay, 11∞33¢S,
77∞15¢W, 24.ix.1952, Ferreyra 8750 (MA); Huarochiri, Viso, 11∞48¢S, 76∞19¢W, 23.iv.1939, Goodspeed
et al. 11526 (UC, GH); isla de San Lorenzo, 12∞5¢S,
77∞13¢W, .1838, Wilkes s.n. (US); lomas de Amancaes, 12∞1¢S, 77∞1¢W, 8.x.1955, Böcher et al. 349
(C, G, GH); lomas de Lachay, 11∞17¢S, 77∞30¢W,
30.x.1976, Cerrate 6407 (MA); Lurín, 12∞16¢S,
76∞52¢W, 18.viii.1953, Ferreyra 9557 (MA); Mangomarca, 24 km NE de Lima, 11∞50¢S, 76∞56¢W,
22.vii.1956, Ferreyra 11797 (MA).
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40
C. AEDO ET AL.
C
E
D
G
J
A
H
K
I
F
B
Figure 75. Geranium limae R. Knuth. A, habit; B, leaf; C, bracts; D, flower; E, flower without petals; F, sepal; G, petal;
H, staminal filament; I, fruit; J, mericarp; K, seed [A–C, I–K: Asplund 13774 (S); D–H: Ferreyra 2445 (US).].
8. Geranium magellanicum Hook. f., Fl. Antarct. 2:
251 (1845)
Type: Chile. Strait of Magalhaens, Elisabeth
Island, Darwin s.n. (lectotype, here designated, K!)
Herbs 8–34 cm tall. Rootstock 5.9–14.1 mm diam.,
without fusiform roots. Stem decumbent, with patent
to retrorse, appressed, eglandular hairs 0.3–1.4 mm
long. Basal leaves in a ± persistent rosette; lamina
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GERANIUM SECTIONS ANDINA AND CHILENSIA
1.12–4.38 ¥ 1.3–4.4 cm, polygonal in outline, cordate,
palmatifid (divided for 0.51–0.86 of its length), pilose,
with appressed, eglandular hairs; segments 5–7, obtriangular, 1.9–5.2 mm at the base (segment width at the
base/segment length ratio = (0.10–)0.17–0.23(-0.28)),
3–9-lobed in distal half (second sinus length/middlesegment
length
ratio = (0.17–)0.25–0.34(-0.43));
petioles to 14 cm long, with patent to retrorse,
appressed, eglandular hairs 0.4–1.4 mm long; stipules
3.6–10.5 ¥ 0.9–3 mm, with eglandular hairs on both
surfaces and on the margin. Inflorescence with
cymules (1–)2-flowered, solitary (cymules of the basal
part of the stem usually 1-flowered, a part of them
arise directly from the rootstock, and cymules of the
upper part 2-flowered); peduncles (1.3–)2.8–4(-6.9) cm
long, with patent to retrorse, appressed, eglandular
hairs 0.5–1.2 mm long; bracteoles 3.1–7.2 ¥ 0.6–
1.8 mm, linear-lanceolate, with eglandular hairs on
both surfaces and on the margin; pedicels 0.54–
4.89 cm long, with patent to retrorse, appressed,
eglandular hairs 0.5–1.2 mm long; pedicel and peduncle together usually overtopping the subtending leaf
Sepals 4.7–6(-6.5) ¥ 1.9–3.4 mm, lanceolate (width/
length ratio = (0.33–)0.44–0.51(-0.62)), with mucro
(0.2–)0.4–0.5(-1) mm long (mucro length/sepal length
ratio = (0.03–)0.07–0.10(-0.16)), with scarious margins 0.2–0.3 mm wide, with erect-patent, eglandular
hairs 0.4–1.9 mm long (usually the longest on the
margin). Petals (3.9–)6.2–8(11.7) ¥ 2–5.45 mm, entire,
without claw, glabrous on both sides, ciliate on the
basal margin, purplish. Filaments 2.2–4.3 mm long,
whitish to yellowish, with eglandular hairs 0.2–
0.5 mm long on the abaxial side and margin; anthers
0.4–1 ¥ 0.4–0.8 mm, yellowish; pollen yellow. Nectaries glabrous. Gynoecium 2.9–5 mm long, pinkish.
Fruit 15.3–22.2 mm long; mericarps 3–4 ¥ 1.4–
2.1 mm, smooth, with erect-patent, eglandular hairs
0.4–1.3 mm long, brownish; rostrum 10.4–16.5 mm
long, without a narrowed apex, with erect-patent,
eglandular hairs 0.2–0.7 mm long; stigmatic remains
(1–)1.3–1.7(-2.1) mm long, with 5 hairy lobes. Seeds
2–2.6 ¥ 1.2–1.8 mm, finely reticulate; hilum 1/6 as
long as the perimeter.
Area: South Chile and South Argentina (Fig. 76).
Habitat: Bogs with Acaena or Gunnera, open grassy
slopes, and Nothofagus or Araucaria forest; 0–1200 m.
Phenology: Flowering November – April.
Illustration: Figure 77.
Notes: Geranium magellanicum is easily recognizable
by its decumbent habit and its particular inflorescence
arrangement. The cymules of the basal part of the
41
stem are usually 1-flowered, and some of them arise
directly from the rootstock. On the middle and
upper part of the decumbent stems the cymules are
2-flowered. This inflorescence structure suggests
that G. magellanicum has a close relationship to
G. sessiliflorum, and could be a link between sect.
Andina and sect. Chilensia. However, G. sessiliflorum
never has 2-flowered cymules as G. magellanicum
does. Additionally, G. magellanicum has longer fruits,
rostrum, peduncles, pedicels and pedicel indumentum
than G. sessiliflorum.
The indumentum of G. magellanicum on stems,
petioles, peduncles and pedicels varies from patent to
retrorse appressed, but no specimens with glandular
hairs have been found. Moore (1983) included
G. patagonicum as a synonym of G. magellanicum,
and consequently considered that G. magellanicum
can have glandular hairs. Barboza & Correa (1988)
and Barboza (1996) excluded glandular specimens
from the G. magellanicum concept, which is in accordance with the view of this study. Geranium berteroanum (which included G. patagonicum) can easily
be distinguished from G. magellanicum by its erect to
ascending stems with only 2-flowered cymules, and by
its longer sepal mucro. Additionally, Barboza (1996)
recognized G. magellanicum var. pumilum, which is
considered here as a different species (G. parodii). The
differences are discussed under the second species.
Representative specimens examined: ARGENTINA.
CHUBUT: río Senguerr, alrededores del lago Blanco,
45∞54¢S, 71∞15¢W, .1903, Koslowsky 12445 (CTES);
valle de la Laguna Blanca, 45∞52¢S, 71∞15¢W,
12.i.1902, Koslowsky 71 (Z, BM, CORD, K). RÍO
NEGRO: San Carlos de Bariloche, 41∞9¢S, 71∞18¢W,
20.ii.1905, Buchtien 1332 (GH). SANTA CRUZ: GüerAike, río Turbio, 51∞36¢S, 72∞17¢W, 16.ii.1993, Roig
et al. 14738 (MERL); lago Argentino, ruta nacional 40,
18 km NW del ACA Tres Lagos, orillas río Shehuen,
49∞37¢S, 71∞30¢W, 10.ii.1975, Boelcke et al. 16271 (UC);
río Gallegos, Cerro Los Conventos, 51∞54¢S, 69∞21¢W,
14.iv.1950, Sleumer 971 (LIL, G). TIERRA DE FUEGO:
Ushuaia, camino de Fique, 54∞54¢S, 68∞13¢W, 31.i.1962
(SI); Estancia Cullen, arroya Beta, 52∞44¢S, 68∞33¢W,
5.i.1972, Moore & Goodall 347 (C, SI). CHILE. AISÉN:
cerca río Coyhaique, 45∞33¢S, 72∞4¢W, 11.ii.1974,
Navas s.n. (CONC). LA ARAUCANIA: Cautín, Victoria,
along the road to the Termas, 400 m from hotel Tolhuaca, 38∞13¢S, 72∞20¢W, 14.iii.1939, Morrison & R.
Wagenknecht 17516 (BH, G, GH, MO, S, UC); volcán
Llaima, estación de esquí Las Araucarias, 38∞41¢S,
71∞50¢W, 22.xii.2001, Aedo 7224 (MA, CONC). LOS
LAGOS: Quilquico, 42∞28¢S, 73∞42¢W, 15.xii.1944,
Acevedo s.n. (SGO). MAGALLANES: Elisabeth Island,
52∞52¢S, 70∞43¢W, Thomson s.n. (E); Patagonia, isla
Contramaestre, 52∞57¢S, 70∞21¢W, 17.xii.1970, Pisano
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42
C. AEDO ET AL.
Figure 76. Area of distribution of Geranium magellanicum (dots) and Geranium albicans (triangles).
2907 (MO, CONC); peninsula de Brunswick, camino
del Club Andino, 53∞30¢S, 71∞24¢W, 8.i.1980, Dollenz
635 (GH); pr. Puerto Natales, 51∞35¢S, 72∞36¢W,
19.i.2002, Aedo 7471 (MA, CONC); Puerto Natales,
P.N. Torres del Paine, hostería Pehoe, 51∞6¢S, 73∞4¢W,
4.xi.1990, Rico 7242 (MA); Río Seco, en la playa,
53∞4¢S, 70∞52¢W, 29.i.1964, Álvarez 50 (CONC); río
Tres Paragos al S of Punta Arenas, 53∞9¢S, 70∞57¢W,
1.iii.1917, Bonarelli s.n. (SI); Sandy Point, 53∞10¢S,
70∞56¢W, 16.iii.1872, Blake s.n. (GH). TIERRA DE
FUEGO: bahía de San Gregorio, 52∞27¢S, 69∞48¢W,
17.i.2002, Aedo 7449 (MA, CONC); Barrancas del Carmen Sylva, 53∞54¢S, 69∞5¢W, 5.ii.1896, Dusén 455
(UPS, LD, S); pr. Porvenir, puente Barguetto, 53∞0¢S,
70∞7¢W, 17.i.2002, Aedo 7445 (MA); río Santa María a
60 km, camino S, 53∞24¢S, 70∞18¢W, 10.ii.1972, Pisano
3540 (GH, CONC).
9. Geranium albicans A. St.-Hil., Fl. Bras. Merid. 1:
83 (1825)
Type: Brazil. Rio Grande do Sul, Giribatubà,
Saint Hilaire 1897 bi (lectotype, here designated,
P!)
Geranium rotundifolium var. americanum A.
St.-Hil. & Naudin in Ann. Sci. Nat., Bot. (Paris) ser.
2, 18: 25 (1842)
Type: Brazil. Rio Grande do Sul, Giribatubà, Saint
Hilaire 1895 (lectotype, here designated, P!)
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GERANIUM SECTIONS ANDINA AND CHILENSIA
C
43
G
E
F
D
A
H
J
B
I
K
Figure 77. Geranium magellanicum Hook. f. A, habit; B, leaf; C, bracts; D, flower; E, flower without petals; F, sepal; G,
petal; H, stamen; I, fruit; J, mericarp; K, seed [A–K: Högberg s.n. (S).].
Geranium selloi Aedo & Muñoz Garm., Kew Bull.
52(3): 726 (1997). Geranium senecioides R. Knuth
in Engl., Pflanzenr. IV.129 (Heft 53): 73 (1912) [syn.
subst.], nom. illeg., non Dump. Course. (1802)
Type: Brazil. [without locality] Sellow 4022 (lectotype, here designated, S!; isolectotype, LE!)
Herbs 18–41 cm tall. Rootstock 5.9–17.4 mm diam.,
without fusiform roots. Stem erect to ascending, with
patent, eglandular hairs 1–2.2 mm long. Basal leaves
in a ± persistent rosette; lamina 1.6–4.2 ¥ 1.8–5 cm,
polygonal in outline, cordate, palmatifid (divided for
0.49–0.77 of its length), pilose, with ± appressed,
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44
C. AEDO ET AL.
eglandular hairs; segments 5–7, rhombic, 2.1–8 mm
at the base (segment width at the base/segment
length ratio = 0.16–0.24(-0.29)), 7–12-lobed in distal
half (second sinus length/middle-segment length
ratio = (0.25–)0.32–0.41); petioles to 12 cm long, with
patent, eglandular hairs 0.9–2.2 mm long; stipules
2–7 ¥ 0.3–1.5 mm, with eglandular hairs on both surfaces and on the margin. Inflorescence with cymules 2flowered, solitary; peduncles (0.36–)0.65–1.8(-3.1) cm
long, with patent, eglandular hairs 0.7–1.9 mm long;
bracteoles 1.6–4.8 ¥ 0.4–1 mm, linear-lanceolate, with
scattered eglandular hairs on both surfaces and on
the margin; pedicels 0.5–3.6 cm long, with patent,
eglandular hairs 0.7–1.9 mm long; pedicel and
peduncle together usually overtopping the subtending
leaf. Sepals (4.2–)5–5.6(-6.5) ¥ 2.3–3.8 mm, lanceolate
(width/length ratio = 0.50–0.61(-0.66)), with mucro
(0.2–)0.25–0.40(-0.6) mm long (mucro length/sepal
length ratio = (0.03–)0.04–0.08(-0.11)), with scarious
margins 0.1–0.2 mm wide, with erect-patent, eglandular hairs 0.5–1.5 mm long. Petals (4–)4.5–5.1(-6.1) ¥
1.7–3 mm, entire or slightly notched, without claw,
glabrous on both sides, ciliate on the basal margin,
purplish, sometimes white. Filaments 2–3.8 mm
long, yellowish, with eglandular hairs 0.2–0.3 mm
long on the abaxial side and margin; anthers 0.4–
0.7 ¥ 0.3–0.5 mm, unknown colour; pollen of unknown
colour. Nectaries glabrous. Gynoecium 2.5–4.3 mm
long, unknown colour. Fruit 14–18.9 mm long;
mericarps 2.5–3.3 ¥ 1.4–1.9 mm, smooth, with erectpatent, eglandular hairs 0.4–1.4 mm long, brownish;
rostrum 9.6–14.8 mm long, without a narrowed apex,
with patent, eglandular hairs 0.4–1.3 mm long; stigmatic remains 1–1.4(-2) mm long, with 5 hairy lobes.
Seeds 1.2–2.2 ¥ 1–1.4 mm, finely reticulate; hilum 1/6
as long as the perimeter.
Area: South Brazil, Uruguay and north-east Argentina (Fig. 76).
Habitat: Roadsides, open, sandy grounds, or grasslands; 50–940 m.
Phenology: Flowering September – December.
Illustration: Figure 78.
Notes: Geranium albicans is characterized by its
peduncles and pedicels with eglandular patent hairs,
its short petals and its short sepal mucro. Leaves are
densely hairy and the main leaf segment has a rhombic outline. Geranium albicans is sometimes confused
with eglandular forms of G. berteroanum. However,
G. berteroanum has longer petals and an obtriangular
main leaf segment, as well as a longer sepal mucro
and shorter hairs on the peduncles and pedicels.
G. core-core, a species also present in the area of
G. albicans, has shorter hairs on the stem, relatively
wider sepals and retrorse-appressed hairs on pedicels.
Representative specimens examined: ARGENTINA.
BUENOS AIRES: along road to Balcarce, 10 km N of
Mar del Plata, 38∞0¢S, 57∞32¢W, 11.xii.1938, Eyerdam
et al. 23628 (GH, UC); Junín, 34∞30¢S, 61∞12¢W,
15.x.1929, Lahitte & Clos 241 (FI); laguna de los
Padres, 37∞57¢S, 57∞44¢W, .xi.1913, Valentini 32 (SI);
Pellegrini, Salliquedó, estancia Los Gossos, 36∞19¢S,
63∞0¢W, 10.xi.1943, Cabrera 8017 (LP, SI); Pigüé,
37∞28¢S, 62∞5¢W, 10.xi.1932, Burkart 4707 (SI).
CÓRDOBA: Río Cuarto, 33∞8¢S, 64∞21¢W, 10.xii.1908,
Stuckert 19428 (CORD). ENTRE RÍOS: Concordia,
arroyo Ayui, 31∞16¢S, 57∞57¢W, 23.ix.1977, Troncoso
et al. 2090 (NY); Gualeguay, Estancia San Ambrosio,
33∞10¢S, 59∞14¢W, 21.x.1949, Burkart 18089 (CTES);
Villaguay, Jubileo, 31∞44¢S, 58∞37¢W, 24.x.1961, Pedersen 6313 (US, C, L). LA PAMPA: Chapaleufu, 35∞12¢S,
63∞32¢W, 30.xi.1983, Steibel & Troiani 7716 (CORD).
RÍO NEGRO: San Antonio, extremo N de Sierra Grande,
frente a Pueblo Viejo, 41∞35¢S, 65∞22¢W, 21.x.1979,
Correa et al. 7111 (UC). SANTA FÉ: Arroyo Frias,
32∞57¢S, 60∞40¢W, 15.i.1949, Morello 4027 (SI). BRAZIL. RIO GRANDE DO SUL: 4 km E de Piratini, 31∞27¢S,
53∞5¢W, 11.x.1972, Lindeman et al. 20673 (U, CTES);
near Caçapava do Sul, 30∞30¢S, 53∞30¢W, 9.xi.1977,
Pedersen 11965 (L, C, GH); Quinta, pr. Rio Grande,
32∞4¢S, 52∞16¢W, 8.xi.1901, Malme 269 (S). SANTA
CATARINA: 9 km N de Cruzeiro, 26 km NE de Sao
Joaquim, 28∞9¢S, 49∞45¢W, 25.xii.1982, Krapovicas &
Schinini 38296 (CTES). URUGUAY. CERRO LARGO:
Cerro Largo, Río Negro, Palleros, 32∞5¢S, 54∞52¢W,
.xii.1937, Gallinal et al. 1343 (F). FLORIDA: Timote,
Santa Clara, 33∞39¢S, 55∞47¢W, 8.x.1943, Gallinal
et al. 5290 (MA, MO, US, U). LAVALLEJA: estancia
Pororó, 34∞10¢S, 54∞54¢W, 2.xii.1955, Pedersen 3588
(C). MALDONADO: ruta 12, entre Pan de Azucar y
Minas, 34∞47¢S, 55∞14¢W, 12.x.1963, Arrillaga et al.
1616 (F). MONTEVIDEO: Montevideo, 34∞50¢S, 56∞10¢W,
Saint Hilaire s.n. (MPU). SORIANO: Juan Jackson,
Monzón-Heber, 33∞55¢S, 57∞12¢W, .x.1943, Gallinal
et al. 5354 (MA, NY).
10. Geranium skottsbergii R. Knuth in Repert. Spec.
Nov. Regni Veg. 34: 143 (1933)
Type: Chile. Coquimbo, estancia Fray Jorge, 13
Aug. 1917, Skottsberg 752 (holotype, B†; isotype,
S!) [30∞40¢S, 71∞37¢W]
Geranium ciliatum Phil. in Anales Univ. Chile 82:
727 (1893) [syn. subst.], nom. illeg., non Cav.
(1787). Geranium berteroanum var. ciliatum
Reiche in Anales Univ. Chile 93: 578 (1895).
Geranium philippii J.F. Macbr. in Candollea 6:
7 (1934). Geranium berteroanum var. philippi
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
GERANIUM SECTIONS ANDINA AND CHILENSIA
45
C
A
F
D
G
E
H
J
I
B
K
Figure 78. Geranium albicans A. St.-Hil. A, habit; B, leaf; C, bracts; D, flower; E, flower without petals; F, sepal; G, petal;
H, staminal filament; I, fruit; J, mericarp; K, seed [A–C, I–K: Wall & Sparre 209 (S); D–H: Pedersen 4756 (C).].
(J.F. Macbr.) L.E. Navas, Fl. Cuenca Santiago
Chile 2: 230 (1976)
Type: Chile. Santiago 1856, Philippi s.n. (lectotype, here designated, MA!; isolectotypes, K!, W!)
Herbs 33–70 cm tall. Rootstock 7–17 mm diam., without fusiform roots. Stem erect to ascending, with
patent to retrorse, appressed, eglandular hairs
0.3–1.7 mm long, and, sometimes, patent, glandular
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46
C. AEDO ET AL.
hairs 0.3–1.2 mm long. Basal leaves in a ± deciduous
rosette; lamina 2.5–7.2 ¥ 2.7–8.2 cm, polygonal in outline, cordate, palmatifid (divided for 0.6–0.9 of its
length), pilose, with appressed, eglandular hairs;
segments 5–7, obtriangular, 2.4–7.2 mm at the base
(segment width at the base/segment length ratio =
(0.06–)0.10–0.19(-0.23)), 3–10-lobed in distal half
(second sinus length/middle-segment length ratio =
(0.31–)0.40–0.53(-0.57)); petioles to 38 cm long, with
patent to retrorse, appressed, eglandular hairs
0.4–2 mm long, and, sometimes, patent, glandular
hairs 0.3–1.2 mm long; stipules 3.6–9 ¥ 0.6–2 mm,
with eglandular hairs on both surfaces and on the
margin. Inflorescence with cymules 2-flowered, solitary; peduncles (0.7–)1.8–4(-5.5) cm long, with patent
to retrorse, appressed, eglandular hairs 0.4–1.5 mm
long, and, sometimes, patent, glandular hairs 0.3–
1.4 mm long; bracteoles 2.3–6.1 ¥ 0.5–1.1 mm, linearlanceolate, with eglandular hairs on both sides and on
the margin; pedicels 1–3.1 cm long, with patent to retrorse, appressed, eglandular hairs 0.4–1.5 mm long,
and, sometimes, patent, glandular hairs 0.3–1.4 mm
long; pedicel and peduncle together usually overtopping the subtending leaf. Sepals 5–6(-8) ¥ 2.1–
3.8 mm, lanceolate (width/length ratio = (0.37–)0.43–
0.57(-0.63)), with mucro (0.4–)0.7–0.9(-1.2) mm long
(mucro length/sepal length ratio = (0.08–)0.11–
0.16(-0.22)), with scarious margins 0.1–0.15 mm
wide, with erect-patent, eglandular hairs 1.1–2.2 mm
long (usually the longest on the margin), and sometimes, patent, glandular hairs 0.3–1.4 mm long. Petals
(7–)7.9–10.2(-14) ¥ 4–7.2 mm, entire or slightly
notched, without claw, glabrous on both sides, ciliate
on the basal margin, purplish. Filaments 3.1–4.5 mm
long, whitish to yellowish, with eglandular hairs
0.2–0.4 mm long on the abaxial side and margin;
anthers 0.4–1.1 ¥ 0.4–0.9 mm, yellowish or purplish;
pollen yellow. Nectaries glabrous. Gynoecium 3–7 mm
long, whitish to pink. Fruit 15.3–20.1 mm long; mericarps 2.5–3.3 ¥ 1.3–2 mm, smooth, with erect-patent,
eglandular hairs 0.3–1.2 mm long, brownish; rostrum
10.4–15.3 mm long, without a narrowed apex, with
erect-patent, eglandular hairs 0.3–1.5 mm long, and,
sometimes patent glandular hairs 0.2–1.3 mm long;
stigmatic remains 1–1.6(-2.3) mm long, with 5 hairy
lobes. Seeds 1.7–2.5 ¥ 1.1–2 mm, finely reticulate;
hilum 1/6 as long as the perimeter.
Area: Central Chile (Fig. 79).
Habitat: Dunes, dry grassy slopes, shrublands, and
Peumus forests; 0–1300 m.
Phenology: Flowering January – December.
Illustration: Figure 80.
Notes: Geranium skottsbergii is easily distinguished
by its deeply divided leaves with narrow lobes and
relatively long petals. Some specimens have only
eglandular hairs on the stem, petioles and pedicels,
sometimes retrorse and appressed, sometimes more or
less patent. There are also specimens with particularly long hairs on the sepal margin. Additionally,
plants with glandular and eglandular hairs are
present over the entire area of this species. Geranium
skottsbergii is endemic to central Chile, where it may
be confused with G. berteroanum, which has shorter
petals and less divided leaves. Geranium core-core,
which is also present in this area, has leaves that are
less divided than G. skottsbergii, stems with shorter
hairs, a shorter gynoecium and relatively wider
sepals.
Representative specimens examined: CHILE. AISÉN:
Chile australis, Los Sauces, 8.viii.1896, Dusén 319 (S).
COQUIMBO: Cuesta de Cavilolén, 31∞45¢S, 71∞19¢W,
11.xi.2001, Aedo 6842 (MA); Limarí, P.N. Fray Jorge,
Quebrada El Mineral, 30∞40¢S, 71∞38¢W, 9.viii.1948,
Jiles 682 (CONC); Llanos, Quebrada el Teniente,
32∞8¢S, 71∞39¢W, 10.ix.1942, Muñoz & Pisano 3385
(SGO); Pichidangui, 32∞9¢S, 71∞30¢W, 11.xi.2001, Aedo
6832 (MA). LA ARAUCANIA: Cautín, Cunco, 38∞55¢S,
72∞2¢W, .i.1931, Barros 24699 (CONC); Cautín,
Temuco, 38∞44¢S, 72∞36¢W, .xii.1946, Gunckel 36602
(CONC); Purén, 38∞1¢S, 73∞5¢W, 1.xi.1945, Montero
4679 (CONC); Traiguén, 38∞15¢S, 72∞40¢W, 2.xi.1952,
Salazar s.n. (CONC). MAULE: Baños de Cauquenes,
35∞58¢S, 72∞21¢W, .1875, Dessauer s.n. (M); Constitución, 35∞20¢S, 72∞25¢W, .ix.1958, Barnier 286
(CONC); Zarral, Linares, Villanueva, Cerro Alto de
Caliboro, 35∞51¢S, 71∞36¢W, 12.x.1955, Aravena 16
(CONC). O’HIGGINS: montis La Leona, Rancagua,
34∞30¢S, 71∞4¢W, .v.1818, Bertero 294 (P); San
Fernando, Centinela, 34∞20¢S, 71∞28¢W, .x.1929,
Montero 1599 (CONC); San Fernando, cerro Echaurrina, 34∞32¢S, 71∞1¢W, 22.x.1929, Montero 1597
(CONC). SANTIAGO: cerro de Renca, Santiago, 33∞24¢S,
70∞45¢W, .ix.1878, Philippi s.n. (SGO); Cerro Manquehue, 33∞21¢S, 70∞36¢W, 10.ix.1949, Rodríguez s.n.
(CONC); Cerro San Cristobal, 33∞25¢S, 70∞39¢W,
16.ix.1917, Skottsberg 974 (S); El Tabo, Quebrada
Córdoba, 33∞27¢S, 71∞41¢W, .x.1964, Gunckel 42784
(CONC); entre Culiprán y Melipilla, 33∞48¢S, 71∞18¢W,
.x.1958, Bailey s.n. (SGO). VALPARAÍSO: 3 km al S de
Papudo, 32∞31¢S, 71∞28¢W, 10.xi.2001, Aedo 6824
(MA); Aconcagua, bei Zapallar, 32∞34¢S, 71∞27¢W,
29.viii.1971, Zöllner 5164 (L); Cachagua, Quebrada El
Tigre, 32∞37¢S, 71∞26¢W, 27.ix.1971, Covarrubias s.n.
(SGO); Catapilco, 32∞33¢S, 71∞16¢W, .ix.1865, Philippi
s.n. (SGO); Concón, 32∞55¢S, 71∞32¢W, .x.1884, Philippi
s.n. (SGO).
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GERANIUM SECTIONS ANDINA AND CHILENSIA
47
Figure 79. Area of distribution of Geranium skottsbergii.
11. Geranium core-core Steud. in Flora 39(28): 438
(1856), nom. cons. prop.
Type: Chile. Quillota, Bertero 1018 (lectotype, designated by Aedo, 2004; P!; isolectotype, G! MO!)
Geranium rapulum A. St.-Hil. & Naudin in Ann.
Sci. Nat., Bot. (Paris) ser. 2, 18: 25 (1842), nom.
rej. prop.
Type: Brazil. Rio Grande do Sul, Gaudichaud
1204 (lectotype, designated by Aedo, 2004; P!;
isolectotype, K!)
Geranium commutatum Steud. in Flora 39(28):
439 (1856)
Type: Chile. prope urbem Valdivia, Lechler 259
(lectotype, here designated, W!; isolectotype, FI
photo! K! LE! P!)
Geranium ochsenii Phil. in Linnaea 28: 676
(1856). Geranium commutatum var. ochsenii
(Phil.) Reiche in Anales Univ. Chile 93: 577 (1895)
Type: Chile. Los Lagos, Valdivia, San Juan,
Philippi s.n. (lectotype, here designated, W!;
isolectotypes, LE!, SGO-51240!, photo P!)
Geranium moorei Phil. in Anales Univ. Chile 82:
728 (1893)
Type: Chile. Maule, Curicó, Todos los Santos, September 1882, E. Moore s.n. (lectotype, here designated, SGO-40572!, photo P!)
Geranium squamosum Phil. in Anales Univ. Chile
82: 733 (1893)
Type: Chile. Bíobio, Talcahuano, pr. San Vicente,
Philippi s.n. (lectotype, here designated, SGO51249!)
Geranium melanopotamicum Speg. in Anales Mus.
Nac. Hist. Nat. Buenos Aires ser. 2, 4: 254 (1902)
Type: Argentina. Río Negro, Carmen de Patagones, Feb. 1898, Spegazzini s.n. (lectotype, here
designated, LP digital image!; isolectotype, LPS
photocopy!)
Geranium argentinum R. Knuth in Engl., Pflanzenr. IV.129 (Heft 53): 77 (1912)
Type: Argentina. Córdoba, San Francisco 1871,
Lorentz 271 (lectotype, here designated, CORD
digital image!; photograph of B specimen, F! G!
GH!)
Geranium subsericeum R. Knuth. in Engl., Pflanzenr. IV.129 (Heft 53): 49 (1912)
Type: Argentina. Río Negro, 21.xi.1874, Berg 39
(lectotype, here designated, LP digital image!)
Geranium herrerae R. Knuth in Repert. Spec. Nov.
Regni Veg. 28: 1 (1930)
Type: Peru. Moquegua, Torata, 17–18.iii.1925,
Weberwauer 7479 (neotype, here designated, F!;
isotypes, G! K! NY! S!)
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48
C. AEDO ET AL.
G
A
E
D
I
K
L
J
C
M
B
H
F
Figure 80. Geranium skottsbergii R. Knuth. A–B, habit; C, leaf; D, bracts; E, peduncle; F, flower; G, flower without petals;
H, sepal; I, petal; J, staminal filament; K, fruit; L, mericarp; M, seed [A, F, H–J, L–M: Aedo 6807 (MA); B: Aedo 6832 (MA);
C–E, G, K: Aedo 6839 (MA).].
Herbs 15–75 cm tall. Rootstock 4–18.1 mm diam.,
without fusiform roots. Stem erect to ascending, with
retrorse, appressed, eglandular hairs 0.2–0.5 mm
long. Basal leaves in a deciduous rosette; lamina 1.56–
4.75 ¥ 1.7–5.6 cm, polygonal in outline, cordate, palmatifid (divided for 0.57–079 of its length), pilose, with
appressed, eglandular hairs; segments 5–7, obtriangular, 2–8.5 mm at the base (segment width at the base/
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GERANIUM SECTIONS ANDINA AND CHILENSIA
segment length ratio = (0.14–)0.18–0.27(-0.31)), 3–11lobed in distal half (second sinus length/middlesegment
length
ratio = (0.18–)0.25–0.37(-0.39));
petioles to 17 cm long, with retrorse, appressed, eglandular hairs 0.2–0.5 mm long; stipules 3.5–8.2 ¥ 0.5–
2.5 mm, with eglandular hairs on abaxial surface and
on the margin, glabrous adaxially. Inflorescence with
cymules 2-flowered, solitary; peduncles (0.6–)1.3–
2.8(-5.1) cm long, with retrorse, appressed, eglandular hairs 0.2–0.6 mm long; bracteoles 2–4.5 ¥ 0.4–
0.9 mm, linear-lanceolate, with eglandular hairs on
abaxial surface and on the margin, glabrous adaxially;
pedicels 1.1–3.1 cm long, with retrorse, appressed,
eglandular hairs 0.2–0.6 mm long; pedicel and
peduncle together usually overtopping the subtending
leaf. Sepals 3–4.2(-5.5) ¥ 2–4.3 mm, ovate (width/
length ratio = (0.60–)0.62–0.76(-0.84)), with mucro
(0.4–)0.6–0.9(-1.1) mm long (mucro length/sepal
length ratio = (0.07–)0.13–0.20(-0.32)), with scarious
margins 0.1–0.2 mm wide, with antrorse, ± appressed,
eglandular hairs 0.1–1.1 mm long on the abaxial side,
glabrous adaxially. Petals (3.1–)4.1–6(-7.1) ¥ 1.7–
3.5 mm, entire or slightly notched, without claw, glabrous on both sides, ciliate on the basal margin, purplish. Filaments 1.8–3.7 mm long, whitish, glabrous
on both sides, ciliate on the basal margin, with hairs
up to 0.1–0.2 mm long; anthers 0.3–0.8 ¥ 0.3–0.6 mm,
whitish to pink; pollen yellow. Nectaries glabrous.
Gynoecium 2.3–4 mm long, whitish. Fruit 9.5–19 mm
long; mericarps 2.6–3.2 ¥ 1.2–2.3 mm, smooth, with
erect-patent, eglandular hairs 0.2–1.4 mm long,
brownish; rostrum 9.1–14.5 mm long, without a
narrowed apex, with erect-patent, eglandular hairs
0.1–0.7 mm long; stigmatic remains (0.9–)1.1–
1.2(-1.5) mm long, with 5 hairy lobes. Seeds 1.8–
2.9 ¥ 1–1.6 mm, finely reticulate; hilum 1/6 as long as
the perimeter.
Area: Ecuador, Peru, Bolivia, South Brazil, Chile and
Argentina; introduced in Great Britain and USA
(California) (Fig. 81).
Habitat: Roadsides, cultivated grounds, beaches,
grassy hillsides, along water courses, or shrublands;
0–4000 m.
Phenology: Flowering January – December.
Illustration: Figure 82.
Notes: Geranium core-core is a species native to South
America and readily distinguishable by its peduncles
and pedicels with retrorse-appressed, eglandular
hairs, by its leaves not deeply divided and short petals.
It is quite similar to G. retrorsum, a species from Australia and New Zealand. The most noticeable differ-
49
ence is the sepal shape: G. retrorsum has lanceolate
sepals (low width/length ratio), whereas G. core-core
has broadly ovate sepals (high width/length ratio).
Additionally, G. core-core has longer fruits and a
longer rostrum than G. retrorsum.
Two sheets with original material (Bertero 1018) of
G. core-core are kept at G. One of them is G. core-core,
and the other is G. skottsbergii. Following our typification the first becomes the isolectotype of G. corecore, while the second should be excluded from the
type concept of this species. The type of G. herrerae
was destroyed during World War II, and no duplicate
of this collection has been found. Thus we have
selected as the neotype a collection by Weberwauer
also from south Peru, which was handwritten by
Knuth as G. herrerae. Geranium rapulum was published 14 years before G. core-core, but was never used
in major publications. On the contrary, G. core-core
has been used in many floristic publications from
Chile and Argentina. Therefore it has been proposed
that the name G. core-core be conserved in order to
maintain nomenclatural stability (Aedo, 2004).
Representative specimens examined: ARGENTINA.
BUENOS AIRES: Villarino, 60 km W de Pedro Luso,
39∞0¢S, 62∞45¢W, .xii.1964, Fabris 5614 (LP). CHUBUT:
de Esquel a Trevelín, 43∞4¢S, 71∞28¢W, 57.ii.1944,
Nicora 3843 (SI); Los Rápidos, 42∞10¢S, 71∞39¢W,
24.i.1945, Castellanos 151 (K). Córdoba: Calamochita,
río San Miguel, camino a Yacanto, 31∞43¢S, 64∞47¢W,
22.i.1969, Krapovickas & Cristobal 14672 (SI); El
Salto, cañada del río Los Chorrillos, Sierras de Córdoba, 32∞7¢S, 64∞13¢W, 27.xi.1939, Bridasoli 264 (LP).
MENDOZA: Capital, 32∞53¢S, 68∞49¢W, 29.x.1933, Ruiz
Leal s.n. (MERL); Chacras de Coria, 33∞0¢S, 68∞52¢W,
12.iv.1954, Ruiz Leal 643 (MERL). NEUQUÉN: Aluminé, 39∞13¢S, 70∞56¢W, 4.xii.1979, Söyrinki s.n. (H);
Huiliches, Quilquihue, 40∞3¢S, 71∞7¢W, 9.ii.1966, Ruiz
Leal 24635 (CORD). RÍO NEGRO: Bariloche, lago
Nahuel Huapí, 41∞2¢S, 71∞8¢W, .1926, Edwards s.n.
(BM); Choele Choel, 39∞18¢S, 65∞39¢W, .ii.1948,
Burkart 15905 (SI). SAN JUAN: Calingasta, 31∞20¢S,
69∞24¢W, .xi.1941, Rodrigo 3038 (LP); Zonda, Estancia
Maradona, Puesto La Ciénaga, 31∞32¢S, 68∞44¢W,
22.i.1986, Guaglianone et al. 1407 (SI). SANTA CRUZ:
Puesto San Julián, 49∞17¢S, 67∞43¢W, .vii.1925, Blake
33 (BM, K). BOLIVIA. COCHABAMBA: between monte
Puncu and Sehuencas, 17∞31¢S, 65∞13¢W, 4.ii.1995,
Wood 9304 (LPB); Pocona, 16∞19¢S, 70∞43¢W,
10.xi.1978, Steinbach 8669 (K, BM, E, G, GH, NY, S).
LA PAZ: Calacoto, 16∞31¢S, 68∞5¢W, 1.i.1957,
Cañigueral 229 (LPB); Inquisivi, Quime Lower
Bridge, near the lower bridge in Quime about 0.5 km
W of Hospital San Antonio, 16∞59¢S, 67∞13¢W,
23.xi.1987, Lewis 871073 (LPB, MO). BRAZIL. without locality, .1815, Sellow (BM). CHILE. AISÉN: Chile
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50
C. AEDO ET AL.
Figure 81. Area of distribution of Geranium core-core (Californian and European localities are not shown).
Chico, 46∞33¢S, 71∞44¢W, 12.xii.1954, Pfister s.n.
(CONC). ANTOFAGASTA: Toconao, 23∞11¢S, 68∞0¢W,
28.xi.2001, Aedo 7006 (MA). ATACAMA: Huasco, río
Laguna Grande, entre la Junta de Valeriano y Las
Papas, 28∞51¢S, 70∞7¢W, 18.i.1983, Marticorena et al.
83303-A (CONC); km 8 río Chollay, 29∞5¢S, 70∞8¢W,
17.i.1994, Arancio et al. 94114 (CONC). BIOBÍO: Concepción, Mirador Alemán, 36∞50¢S, 73∞2¢W, 4.xi.2001,
Aedo 6728 (MA); Talcahuano, 36∞43¢S, 73∞6¢W,
3.xi.2001, Aedo 6722 (MA). COQUIMBO: Choapa, Illapel, 31∞38¢S, 71∞10¢W, .xi.1989, Chiuminato 23
(CONC); Illapel, Cuesta Espino, 31∞21¢S, 71∞3¢W,
19.x.1945, Biese 2011 (S, BM, NY). JUAN FERNÁNDEZ
IS.: Isla de Más a Fuera, quebrada de La Colonia,
33∞45¢S, 80∞46¢W, 27.xi.1965, Muñoz & Sierra 7028
(CONC); Isla de Más a Tierra, plazoleta del Yunque,
33∞38¢S, 78∞52¢W, 28.xii.1954, Skottsberg 33 (S). La
Araucania: Coi-Coi, La Lobería, 38∞50¢S, 73∞28¢W,
21.xii.2001, Aedo 7197 (MA); Puerto Saavedra,
38∞42¢S, 73∞15¢W, 22.xii.2001, Aedo 7203 (MA). LOS
LAGOS: Chiloe, playa Puñihuil, 41∞55¢S, 74∞1¢W,
6.i.2002, Aedo 7387 (MA); pr. Futaleufú, 43∞10¢S,
71∞45¢W, 5.i.2002, Aedo 7375 (MA). MAULE: Paso
Nevado, 35∞43¢S, 71∞10¢W, 13.xii.2001, Aedo 7153
(MA); Constitución, 35∞20¢S, 72∞25¢W, 17.x.1919,
Holway 123 (US). O’HIGGINS: Baños de Cauquenes,
34∞15¢S, 70∞34¢W, .1872, Reed s.n. (CONC); Rancagua,
34∞30¢S, 71∞4¢W, .1828, Bertero s.n. (SGO). SANTIAGO:
puente Manzanito, 33∞20¢S, 70∞19¢W, 12.xii.2001,
Aedo 7130 (MA); Aguas Negras, El Volcan, 33∞50¢S,
70∞12¢W, .ii.1967, Richter s.n. (CONC). TARAPACÁ:
Arica, quebrada de Socoroma, 18∞15¢S, 69∞38¢W,
5.v.1972, Ricardi et al. 152 (CONC). VALPARAÍSO:
Aconcagua, La Ligua, 32∞27¢S, 71∞13¢W, .i.1970,
Martínez s.n. (CONC); Aconcagua, Zapallar, 32∞33¢S,
71∞28¢W, .ii.1899, Johow s.n. (CONC). ECUADOR.
AZUAY: Cuenca, 2∞52¢S, 78∞58¢W, 17.ix.1918, Rose
et al. 22876 (US, GH, NY). LOJA: Cajanuma, 16 km al
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GERANIUM SECTIONS ANDINA AND CHILENSIA
51
I
H
G
A
F
J
L
M
E
C
K
B
D
Figure 82. Geranium core-core Steud. A, habit; B–D, leaves; E, bracts; F, flower; G, flower without petals; H, sepal; I,
petal; J, staminal filament; K, fruit; L, mericarp; M, seed [A, C, E–M: Aedo 7387 (MA); B: Aedo 7006 (MA); D: Aedo 7375
(MA).]
S de Loja, 4∞4¢S, 79∞11¢W, 16.vii.1947, 1635 (NY); Loja,
3∞59¢S, 79∞12¢W, 14.vi.1946, Espinosa 562 (NY).
TUNGURAHUA: Ambato a Huachi, 1∞17¢S, 78∞37¢W,
29.x.1944, Acosta Solís 8884 (F); entre Casigana y El
Sueño, al SW de Ambato, 1∞15¢S, 78∞37¢W, 14.xii.1944,
Acosta Solís 9303 (F). GREAT BRITAIN. Kent,
Bracken Hill, near Sevenoaks, garden of Mc Clintock,
51∞15¢N, 0∞12¢E, .ix.1979, Mc Clintock s.n. (P). PERU.
APURIMAC: Abancay, Ampuy, 13∞38¢S, 72∞52¢W,
12.ii.1939, Stork et al. 10628 (G). AREQUIPA: Arequipa,
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52
C. AEDO ET AL.
16∞23¢S, 71∞32¢W, 15.viii.1923, Guenther & O. Buchtien 691 (HBG); S slopes of Chachani mountain,
16∞11¢S, 71∞31¢W, .iii.1920, Hinkley 66 (US, GH, NY).
JUNÍN: 19 km NE of Tarma on road to San Ramon,
11∞25¢S, 75∞41¢W, 17.xii.1978, Dillon & Turner 1331
(C, F); Contumazá, Singarrán, El Molino-San Martín,
7∞22¢S, 78∞48¢W, 1.xi.1979, Sagástegui 9343 (NY).
LIMA: Huarochiri, Chicla, 11∞42¢S, 76∞16¢W, 6.vi.1940,
Asplund 11455 (S); Langa, 12∞7¢S, 76∞26¢W,
15.vii.1838, Barclay 2356 (BM). USA. CALIFORNIA:
Alameda Co., San Leandro Bay Shoreline, Oakland,
37∞48¢N, 122∞16¢W, 25.viii.1991, Ertter 10725 (UC);
Contra Costa Co., Mount Diablo Regional Park, Pine
Pond, 37∞51¢N, 121∞55¢W, 8.iv.1987, Ertter 17410 (MA).
12. Geranium retrorsum L’Hér. ex DC., Prodr. 1: 644
(1824). Geranium dissectum var. retrorsum (L’Hér.
ex DC.) Hook. f., Fl. Nov. – Zel. 1: 39 (1852). Geranium pilosum var. retrorsum (L’Hér. ex DC.) Jeps.,
Man. Fl. Pl. Calif. 589 (1925)
Type: New Zealand. [without locality], Banks s.n.
(lectotype, designated by Carolin, 1965: 349, G;
isolectotype, BM!)
Geranium australe Nees in Lehm., Pl. Preiss. 1:
162 (1845), nom. illeg., non Dum. Cours. (1805).
Geranium pilosum var. australe (Nees) Ostenf. in
Biol. Meddel. Kongel. Danske Vidensk. Selsk. 3:
721 (1921)
Type: Australia. [without locality], Preiss 1907
(lectotype designated by Carolin, 1965: 349, LE!)
Geranium dissectum var. patulum Sol. ex Hook. f.,
Handb. N. Zeal. Fl. 36 (1864). Geranium pilosum
var. grandiflorum R. Knuth in Engl., Pflanzenr.
IV.129 (Heft 53): 75 (1912), nom. illeg.
Type: New Zealand. [without locality], Banks &
Solander s.n. (lectotype, here designated, BM!)
Herbs 12–39 cm tall. Rootstock 4.9–15.7 mm diam.,
without fusiform roots. Stem erect to ascending,
with retrorse, appressed, eglandular hairs 0.2–0.9 mm
long. Basal leaves in a ± deciduous rosette; lamina
0.99–4.08 ¥ 1.6–5.3 cm, polygonal in outline, cordate,
palmatifid (divided for 0.56–0.84 of its length), pilose,
with appressed, eglandular hairs; segments 5–7, obtriangular, 1.2–5.6 mm at the base (segment width at the
base/segment length ratio = (0.07–)0.14–0.21(-0.34)),
3–9-lobed in distal half (second sinus length/middlesegment length ratio = (0.24–)0.31–0.37(-0.45)); petioles to 23 cm long, with retrorse, appressed, eglandular hairs 0.2–0.9 mm long; stipules 2–6.5 ¥ 0.5–
1.5 mm, with eglandular hairs on abaxial surface and
on the margin, glabrous adaxially. Inflorescence with
cymules 2-flowered, solitary; peduncles (0.6–)1.3–
3.1(-8.4) cm long, with retrorse, appressed, eglandular hairs 0.2–0.6 mm long; bracteoles 1.4–4.2 ¥ 0.3–
0.9 mm, linear-lanceolate, with eglandular hairs on
abaxial surface and on the margin, glabrous adaxially;
pedicels 0.6–3.7 cm long, with retrorse, appressed,
eglandular hairs 0.2–0.6 mm long; pedicel and peduncle together usually overtopping the subtending leaf.
Sepals (3.2–)4.9–5.5(-6.3) ¥ 1.5–3.5 mm, lanceolate
(width/length ratio = (0.26–)0.44–0.59(-0.65)), with
mucro (0.2–)0.4–0.6(-0.9) mm long (mucro length/
sepal length ratio = (0.05–)0.07–0.11(0.16)), with
scarious margins 0.1–0.2 mm wide, with antrorse, ±
appressed, eglandular hairs 0.1–1.1 mm long on the
abaxial side, glabrous adaxially. Petals (2.9–)3.5–
5.3(-9.2) ¥ 1.4–4.5 mm, entire or slightly notched,
without claw, glabrous on both sides, ciliate on the
basal margin, purplish. Filaments 2.2–4.7 mm long,
whitish, glabrous on both sides, ciliate on the basal
margin, with hairs up to 0.1–0.15 mm long; anthers
0.4–1 ¥ 0.2–0.6 mm, yellowish; pollen unknown
colour. Nectaries glabrous. Gynoecium 2–4.5 mm long,
yellowish. Fruit 11–18 mm long; mericarps 2.5–3.5 ¥
1.3–1.7 mm, smooth, with erect-patent, eglandular
hairs 0.3–1.2 mm long, brownish; rostrum 7.8–
13.2 mm long, without a narrowed apex, with erectpatent, eglandular hairs 0.1–0.7 mm long; stigmatic
remains (0.6–)0.9–1.1(-1.3) mm long, with 5 hairy
lobes. Seeds 1.5–2.3 ¥ 1–1.4 mm, finely reticulate;
hilum 1/6 as long as the perimeter.
Area: Australia (south-west Australia, south-east
Australia and Tasmania), and New Zealand; introduced in USA (Hawaii) (Fig. 83).
Habitat: Roadsides, dry rocky or sandy grasslands,
dunes, shrublands and open forest with Eucalyptus,
Acacia or Agonis; 0–1280 m.
Phenology: Flowering January – December.
Illustration: Figure 84.
Notes: As previously stated G. retrorsum is quite
similar to G. core-core. The differences between the
two species have been indicated under the second
species. Geranium retrorsum is sympatric to
G. solanderi, from which it can be distinguished by its
retrorse-appressed hairs on pedicels. Additionally,
G. retrorsum has shorter hairs on stem, petioles,
pedicels and sepals than G. solanderi.
Geranium retrorsum was recorded in California
(Aedo, 2001b; previous authors there indicated) by
mistake. Most of the specimens supporting this
record were re-examined and are now identified as
G. core-core, as they have broadly ovate sepals. Geranium retrorsum was also reported as introduced in
Europe (Channel Islands) (Kent, 1959; Lousley,
1962). Yeo (2002) re-identified specimens from this
area as G. herrerae. We essentially agree with this
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GERANIUM SECTIONS ANDINA AND CHILENSIA
53
Figure 83. Area of distribution of Geranium retrorsum (Hawaiian localities are not shown).
opinion, although the prior name for this taxon is
G. core-core.
Representative specimens examined: AUSTRALIA.
NEW SOUTH WALES: 15 miles E of Guyra, on Ebor
road, 30∞12¢S, 151∞40¢E, 9.i.1958, Vickery 42756 (K);
21.6 km from Grenfell on road to Forbes, 33∞44¢S,
148∞6¢E, 27.ix.1978, Moore 8013 (CANB); Armidale,
30∞30¢S, 151∞39¢E, .iii.1907, Stopford s.n. (BM).
QUEENSLAND: Gilruth Plains, C’mulla, 28∞7¢S,
145∞58¢E, 9.x.1941, Allen s.n. (CANB); Stanforth,
.xi.1943, Clemens s.n. (GH). SOUTH AUSTRALIA: 2 km
SW of Evondale Homestead on the Coorong road,
36∞29¢S, 139∞48¢E, 1.x.1982, Donner 9045 (L); 25 km E
of Kingston, Reedy Creek Range, 36∞57¢S, 140∞3¢E,
7.x.1982, Donner 9203 (GOET). TASMANIA: Hedgero,
Launceston, 43∞30¢S, 146∞2¢E, 27.x.1943, Curtis s.n.
(K); Prime Seal island, Furneaux group, 40∞3¢S,
147∞45¢E, s.d., Whinray 393 (CANB). VICTORIA:
Buckland river, Nelson’s Creek, 36∞51¢S, 146∞51¢E,
1.xii.1972, Canning 3276 (CANB); Dookie Agricultural
College, hill 4 km S of Mt Major, 36∞24¢S, 145∞41¢E,
26.viii.1992, Crawford 1807 (CANB). WESTERN AUSTRALIA: along Esperance Ravensthorpe road, 33∞40¢S,
120∞47¢E, 25.ix.1968, Wilson 7862 (PERTH); 1 km
along Nun Road from junction with Don Road, NE
of Busselton, 33∞21¢S, 116∞24¢E, 5.x.1996, Crowley
661 (PERTH); 16 km NE of Jerramungup, 33∞56¢S,
118∞55¢E, 18.viii.1974, Newbey 4298 (PERTH) .
NEW ZEALAND. NEW ZEALAND NORTH I.: Auckland
Ecological Region, Inner Gulf Islands Ecological
district, Tiritiri Matangi Island, 36∞36¢S, 174∞53¢E,
15.x.1971, Esler & Scott s.n. (AK); Aupouri Ecological
region, Rangaunu Heads, 34∞57¢S, 173∞15¢E, .x.1897,
Matthews s.n. (AK); Coromandel Co., southernmost
of Ngamotukaraka Island, 36∞42¢S, 175∞23¢E,
28.viii.1983, Wright 5709 (AK, G). NEW ZEALAND
SOUTH I.: Aniseed valley, Nelson, 41∞17¢S, 173∞14¢E,
15.xi.1980, Gardner 2772 (AK); Braeview Crescent,
Dunedin city, 45∞53¢S, 170∞28¢E, .ii.1979, Esler s.n.
(AK); Canterbury Land District, Canterbury Plains,
Leeston, 43∞49¢S, 172∞15¢E, 10.xii.1994, Nielsen 10091
(C). USA. HAWAII: Hawaii I, Humuula, saddle
between Mauna Lea & Mauna Kea, 19∞42¢N,
155∞30¢W, 12.vi.1950, Rose 50H8 (CAS); Waimea,
20∞1¢N, 155∞39¢W, 4.iv.1948, Hosaka 3590 (P).
13. Geranium tablasense R. Knuth in Meded. RijksHerb. 27: 68 (1915)
Type: Bolivia. Cochabamba, Tablas, .v.1911, Herzog 2181 (lectotype, here selected, L!; isolectotype,
S!) [17∞05¢S, 65∞59¢W]
Herbs 37–55 cm tall. Rootstock 8–8.2 mm diam., with
fusiform roots. Stem erect, with patent to retrorse, not
appressed, eglandular hairs 0.3–1.2 mm long, and
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54
C. AEDO ET AL.
E
I
H
D
G
A
F
B
J
L
C
K
Figure 84. Geranium retrorsum L’Hér. ex DC. A, habit; B–C, leaves; D, bracts; E, flower; F, flower without petals; G, sepal;
H, petal; I, staminal filament; J, fruit; K, mericarp; L, seed [A, D–L: Mueller s.n. (NY); B: Wright 5709 (AK); C: Carter 22
(K).]
patent, glandular hairs 0.4–0.9 mm long. Basal leaves
in a ± deciduous rosette; lamina 3.6–5.4 ¥ 5.1–6.2 cm,
polygonal in outline, cordate, palmatifid (divided for
0.52–0.61 of its length), pilose, with appressed, eglan-
dular hairs (young leaves sericeous abaxially); segments 5–7, rhombic, 3.2–4.6 mm at the base (segment
width at the base/segment length ratio = 0.09–
0.12(-0.14)), 10–14-lobed in distal half (second sinus
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GERANIUM SECTIONS ANDINA AND CHILENSIA
length/middle-segment length ratio = (0.24–)0.27–
0.37(-0.42)); petioles to 8.2 cm long, with patent,
eglandular hairs 0.3–1.3 mm long; stipules 7.5–9.8 ¥
2.7–3.2 mm, glabrous or with scattered eglandular
hairs on abaxial surface and on the margin, glabrous
adaxially. Inflorescence with cymules 2-flowered, solitary; peduncles (3.7–)4–7(-8) cm long, with patent to
retrorse, not appressed, eglandular hairs 0.2–0.6 mm
long, and patent, glandular hairs 0.4–0.9 mm long;
bracteoles 2.5–6.7 ¥ 0.6–1.8 mm, lanceolate, with scattered eglandular hairs on abaxial surface and on the
margin, glabrous adaxially; pedicels 2.8–4.6 cm long,
with patent to retrorse, not appressed, eglandular
hairs 0.2–0.6 mm long, and patent, glandular hairs
0.4–1.3 mm long; pedicel and peduncle together
usually overtopping the subtending leaf. Sepals
7.4–8 ¥ 2.4–3.3 mm, lanceolate (width/length ratio =
0.32–0.38(-0.41)), with mucro (0.7–)0.75–1.3(-1.4) mm
long (mucro length/sepal length ratio = (0.09–)0.10–
0.16(-0.17)), with scarious margins 0.3–0.4 mm wide,
with erect-patent, eglandular hairs 0.5–1.4 mm
long, and patent, glandular hairs 0.4–1 mm long on
the abaxial side, glabrous adaxially. Petals
(11.4–)13.4–16.4(-16.9) ¥ 7.4–8.2 mm,
emarginate,
notch 1.5–1.9 mm deep, with a claw 1–3.5 mm long,
glabrous, rarely with few cilia on the basal margin,
purplish. Filaments 6.5–8 mm long, yellowish, glabrous on both sides, ciliate on the basal margin, with
55
hairs up to 0.1–0.2 mm long; anthers 1.1–1.8 ¥ 0.8–
1.2 mm, unknown colour; pollen unknown colour. Nectaries glabrous. Gynoecium 7–8.2 mm long, unknown
colour. Fruit 24.2–27.7 mm long; mericarps 3.2–4 ¥
1.6–1.8 mm, smooth, with erect-patent, eglandular hairs
0.3–0.9 mm long, brownish; rostrum 17–21.1 mm
long, with a narrowed apex (2–)2.3–3.8(-5) mm long,
with erect-patent, eglandular hairs 0.4–0.9 mm long;
stigmatic remains (2.6–)2.8–3.4(-3.8) mm long, with 5
glabrous lobes. Seeds 2.1–2.2 ¥ 1.1–1.5 mm, finely
reticulate; hilum 1/4 as long as the perimeter.
Area: Central Bolivia (Fig. 85).
Habitat: Rocky and wet slopes; 2800–3400 m.
Phenology: Flowering March – May.
Illustration: Figure 86.
Notes: Geranium tablasense shares with G. venturianum and G. fallax leaves with rhombic segments and
rostrum with a narrowed apex. It can be recognized by
its longer petals, which shows some overlap with
G. venturianum but none with G. fallax. Additionally,
petals of G. tablasense are glabrous or rarely with few
cilia on the basal margin, while they are hairy on the
basal margin in G. venturianum (and base of the abax-
Figure 85. Area of distribution of Geranium venturianum (dots) and G. tablasense (triangles).
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56
C. AEDO ET AL.
A
B
H
D
G
E
F
C
I
K
J
L
Figure 86. Geranium tablasense R. Knuth. A, habit; B, leaf; C, bracts; D, flower; E, flower without petals; F, sepal; G–H,
petals; I, staminal filament; J, fruit; K, mericarp; L, seed [A, C–I: Steinbach 9576 (PH); B: Steinbach 9576 (U); J–L: Herzog
2181 (L).].
ial side) and G. fallax. Geranium venturianum has
longer peduncles and shorter staminal filaments and
rostrum with a shorter narrowed apex than
G. tablasense. Geranium tablasense also has leaves
with narrow segments at the base, and longer fruits
than G. venturianum and G. fallax.
Representative specimens examined:
BOLIVIA.
COCHABAMBA: Chapare, La Aduana, 17∞32¢S, 65∞13¢W,
10.iii.1929, Steinbach 9576 (E, NY, PH, S, U);
Incachaca, 17∞14¢S, 66∞28¢W, .iii.1941, Cárdenas 2180
(GH, PH, S, U); km 80 road to Chimoré-Cochabamba,
16∞59¢S, 65∞7¢W, .iv.1939, Cárdenas 758 (US).
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GERANIUM SECTIONS ANDINA AND CHILENSIA
14. Geranium venturianum R. Knuth, Repert. Spec.
Nov. Regni Veg. 40: 218 (1936)
Type: Argentina. Tucumán, Cerro del Campo, 15
Dec. 1928, Venturi 7723 (lectotype, here designated, S!; isolectotypes, BM! K! MO! SI)
Herbs 27–88 cm tall. Rootstock 6.9–13.6 mm diam.,
with fusiform roots. Stem erect, with patent, eglandular hairs 0.1–0.9 mm long, and, sometimes, patent,
glandular hairs 0.2–0.7 mm long. Basal leaves in a ±
deciduous rosette; lamina 3.2–7.5 ¥ 4–9.98 cm, polygonal in outline, cordate, palmatifid (divided for 0.50–
0.66 of its length), pilose, with appressed, eglandular
hairs (young leaves sericeous abaxially); segments
5–7, rhombic, 2.4–12.6 mm at the base (segment
width at the base/segment length ratio = (0.08–)0.14–
0.23(-0.30)), 8–16-lobed in distal half (second sinus
length/middle-segment length ratio = (0.23–)0.25–
0.31(-0.49)); petioles to 16.5 cm long, with patent,
eglandular hairs 0.2–1 mm long, and sometimes,
patent, glandular hairs 0.2–0.7 mm long; stipules
3.2–16.7 ¥ 1–5 mm, with eglandular, and sometimes
glandular, hairs on both surfaces and on the margin.
Inflorescence with cymules 2-flowered, solitary;
peduncles (5.9–)8.2–15.1(-25.1) cm long, with patent,
eglandular hairs 0.2–0.7 mm long, and sometimes,
patent, glandular hairs 0.2–0.8 mm long; bracteoles
2.6–8.9 ¥ 0.5–1.2 mm, lanceolate, with eglandular,
and sometimes glandular, hairs on both surfaces and
on the margin; pedicels 1.25–4.66 cm long, with
patent, eglandular hairs 0.2–0.7 mm long, and sometimes, patent, glandular hairs 0.2–0.8 mm long;
pedicel and peduncle together usually overtopping
the subtending leaf. Sepals (5.5–)6.5–8.2(-8.9) ¥ 1.8–
3.7 mm, lanceolate (width/length ratio = (0.27–)0.35–
0.43(-0.49)), with mucro (0.6–)1–1.5(-2.1) mm long
(mucro length/sepal length ratio = (0.10–)0.12–
0.22(-0.26)), with scarious margins 0.2–0.3 mm wide,
with erect-patent, eglandular hairs 0.3–0.8 mm
long, and sometimes, erect-patent, glandular hairs
0.2–0.8 mm long. Petals (10–)11–11.8(-14.3) ¥ 5–
10.7 mm, entire, without claw, hairy on margin and
base of abaxial side, purplish. Filaments 4–7.1 mm
long, yellowish, with eglandular hairs 0.2–0.3 mm
long on the abaxial side and margin; anthers 0.5–
1.7 ¥ 0.4–1 mm, unknown colour; pollen unknown
colour. Nectaries usually glabrous. Gynoecium 4.5–
7 mm long, unknown colour. Fruit 19.4–26.7 mm
long; mericarps 3–3.7 ¥ 1.3–2.2 mm, smooth, with
erect-patent, eglandular hairs 0.3–1.1 mm long and,
sometimes, patent glandular hairs 0.2–0.7 mm long,
brownish; rostrum 12.8–19.3 mm long, with a narrowed apex 1–2.4 mm long, with erect-patent, eglandular hairs 0.3–0.8 mm long and, sometimes, patent
glandular hairs 0.2–0.7 mm long; stigmatic remains
(3–)3.2–3.6(-5) mm long, with 5 hairy lobes. Seeds
57
1.9–2.4 ¥ 1.2–1.5 mm, finely reticulate; hilum 1/4 as
long as the perimeter.
Area: Bolivia, and North Argentina (Fig. 85).
Habitat: Meadows, grassy slopes, and shrublands;
1000–3200 m.
Phenology: Flowering October – April.
Illustration: Figure 87.
Notes: Geranium venturianum is easily recognizable
by its longer peduncles, which decrease in length
towards the apex of the inflorescence. Thus, in the
description of G. venturianum and key we consider
peduncles of the basal part of the inflorescence.
Although a small overlap in peduncle length exists
between G. venturianum and G. tablasense, length of
petals, staminal filaments, fruit and rostrum apex permits a satisfactory discrimination between the species. Geranium venturianum varies in the presence of
glandular hairs. Some specimens have only eglandular hairs while others also have glandular hairs widespread on stem, inflorescence, sepals, rostrum and/or
mericarps. These glandular forms have occasionally
been misidentified as G. fiebrigianum (included in
G. fallax in this study) or G. patagonicum (included in
G. berteroanum in this study). Geranium venturianum
can be distinguished from G. berteroanum by its
leaves with a rhombic middle segment (not obtriangular) and its rostrum fruit with a narrow apex. The best
way to distinguish G. venturianum from G. fallax is
the length of peduncles, clearly longer in the first species. Additionally, G. venturianum has longer fruits,
narrow apex of the rostrum, stigmatic remains,
bracteoles, sepals and petals than in G. fallax.
Representative specimens examined: ARGENTINA.
CATAMARCA: Andalgalá, Esquina Grande, 27∞17¢S,
65∞59¢W, 12.ii.1910, Jörgensen 1562 (US, S); C. och
Tucuman, 27.xi.1946, Wall s.n. (S); Chabarilla,
.iv.1952, Brücher 9440 (M). JUJUY: Capital, cerro
Zapla, camino de la antena, 24∞15¢S, 65∞7¢W,
21.xi.1986, Ahumada 5283 (CTES, LIL); Capital, lagunas de Yala, 24∞7¢S, 65∞28¢W, 26.iii.1979, Cabrera et al.
30670 (SI); Capital, Sierra de Zapla, Mina 9 de Octubre, subida a la antena, 24∞15¢S, 65∞7¢W, 23.i.1975,
Zuloaga & Deginani 197 (LP); Santa Bárbara, subida
al Centinela, 24∞16¢S, 64∞22¢W, 11.xii.1983, Rotman
935 (CTES, LIL, SI). SALTA: Capital, quebrada de San
Lorenzo, 26∞6¢S, 64∞41¢W, 13.x.1926, Venturi 5070 (US,
S, SI); Chicoana, Cuesta del Obispo, 26∞6¢S, 64∞41¢W,
10.iv.1982, Novara 2660 (CORD, S); Chicoana, Quebrada de Escoipe, cerca de Agua Negra, 25∞8¢S,
65∞41¢W, 2.xii.1960, Ruiz Leal 21231 (CORD, MERL);
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58
C. AEDO ET AL.
C
F
A
D
E
H
J
G
I
L
K
B
Figure 87. Geranium venturianum R. Knuth. A–B, habit; C, bracts; D, peduncle; E, flower; F, flower without petals; G,
sepal; H, petal; I, staminal filament; J, fruit; K, mericarp; L, seed [A, C–I: Wood et al. 15287 (LPB); B: Burkart et al. 30494
(CORD); J–L: Venturi 2667 (S).].
Guachipas, Alemania, 25∞36¢S, 65∞37¢W, 6.xii.1929,
Venturi 9848 (NY, K, MO, S); Rosario de Lerma, quebrada del Toro, ruta 51, km 28, unos 3 km al W de
Campo Quijano, 24∞55¢S, 65∞39¢W, 26.iv.1987, Novara
6575 (G). Tucumán: Burruyacú, Alto de Medina,
26∞22¢S, 65∞30¢W, .i.1924, Venturi 2667 (S); Chicligasta, La Alvaresada, 26∞42¢S, 65∞26¢W, 27.xi.1946,
Sparre 938 (S); Chicligasta, Las Pavas, 27∞15¢S,
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GERANIUM SECTIONS ANDINA AND CHILENSIA
65∞52¢W, 22.xi.1926, Venturi 4650 (GH, BR, CAS, NY,
S, W); La Banderita, 27∞17¢S, 65∞54¢W, .ii.1950,
Brücher 9396 (M); Tafí, La Hoyada, 26∞40¢S, 65∞31¢W,
.i.1923, Venturi 1979 (MA, GH); Tafí, Sierra de
Tucumán, Cuesta de Siambón, 26∞42¢S, 65∞26¢W,
18.iii.1872, Lorentz s.n. (CORD). BOLIVIA. CHUQUISACA: Azurduy, on descent from Cordillera de los
Sombreros towards Tarvita, 19∞49¢S, 64∞32¢W,
4.xii.1999, Wood et al. 15287 (LPB). COCHABAMBA:
between Lopez Mendoza and Montepuncu, 17∞32¢S,
65∞22¢W, 16.xii.1994, Wood 8955 (LPB, S).
15. Geranium fallax Steud. in Flora 39(28): 439 (1856)
Type: Peru. Puno, Tabina & Lechler 1907 (lectotype, here designated, P!; isolectotypes, GOET! K!)
[14∞8¢S, 69∞32¢W]
Geranium fiebrigianum R. Knuth in Bot. Jahrb.
Syst. 37: 560 (1906)
Type: Bolivia. Tarija, Calderillo, Fiebrig 2631
(lectotype, here designated, P!; isolectotypes, BM!
E! G! GOET! K! M! US! W!) [21∞48¢S, 63∞45¢W]
Geranium titicacaense R. Knuth in Repert. Spec.
Nov. Regni Veg. 45: 60 (1938)
Type: Peru. Moquegua, Carumas, 21.ii.-6.iii.1925,
Weberbauer 7307 (holotype, B†; isotypes, BM! G!
K! S!) [16∞49¢S, 70∞43¢W]
Herbs 18–85(-100) cm tall. Rootstock 4.2–12.7 mm
diam., with fusiform roots. Stem erect to ascending,
with patent, eglandular hairs 0.4–1.4 mm long, and,
sometimes, patent, glandular hairs 0.4–0.7 mm
long. Basal leaves in a deciduous rosette; lamina
2.4–8.1 ¥ 1.9–8.9 cm, polygonal in outline, cordate,
palmatifid (divided for 0.43–0.67 of its length), pilose,
with appressed, eglandular hairs; segments 5–7,
rhombic, 3.2–10.6 mm at the base (segment width
at the base/segment length ratio = (0.10–)0.13–
0.18(-0.24)), 4–17-lobed in distal half (second sinus
length/middle-segment length ratio = (0.17–)0.21–
0.28(-0.32)); petioles to 16.5 cm long, with patent,
eglandular hairs 0.4–1.6 mm long, and sometimes,
patent, glandular hairs 0.4–0.8 mm long; stipules
3.6–13.4 ¥ 0.9–5 mm, with eglandular hairs on abaxial surface and on the margin (rarely glabrous),
glabrous adaxially. Inflorescence with cymules 2flowered, solitary; peduncles (1.5–)3.1–5.1(-10) cm
long, with patent, eglandular hairs 0.3–1.1 mm long,
and usually, patent, glandular hairs 0.4–1.2 mm
long; bracteoles 2–5.5 ¥ 0.5–1.48 mm, linear-lanceolate,
with eglandular hairs on abaxial surface and on the
margin, glabrous adaxially; pedicels 2.6–4.2 cm long,
with patent, eglandular hairs 0.3–1.1 mm long, and
patent, eglandular hairs 0.4–1.2 mm long; pedicel and
peduncle together usually overtopping the subtending leaf. Sepals (5.2–)5.7–6.5(-8.2) ¥ 2.2–3.7 mm, lanceolate (width/length ratio = (0.32–)0.42–0.49(-0.59)),
59
with mucro (0.3–)0.7–0.9(-1.3) mm long (mucro
length/sepal length ratio = (0.05–)0.09–0.15(-0.22)),
with scarious margins 0.1–0.2 mm wide, with erectpatent, eglandular hairs 0.4–0.9 mm long, and
patent, glandular hairs 0.4–1.2 mm long on the
abaxial side, glabrous adaxially. Petals (4–)4.7–
7.5(-9.9) ¥ 1.9–6.3 mm, entire or slightly notched,
without claw, glabrous on both sides, ciliate on the
basal margin, purplish. Filaments 2–5.3 mm long,
yellowish, with eglandular hairs 0.2–0.3 mm long on
the abaxial side and margin; anthers 0.4–1.2 ¥
0.4–0.9 mm, unknown colour; pollen unknown colour.
Nectaries glabrous. Gynoecium 2.2–5 mm long,
unknown colour. Fruit 18.1–23.8 mm long; mericarps 2.8–4 ¥ 1.3–2 mm, smooth, with erect-patent,
eglandular hairs 0.4–0.9 mm long and, patent glandular hairs 0.4–0.9 mm long, brownish to blackish;
rostrum 13.2–17.4 mm long, with a narrowed apex
(0–)0.9–1.1(1.9) mm long, with erect-patent, eglandular hairs 0.2–0.9 mm long and, patent glandular
hairs 0.4–1.2 mm long; stigmatic remains (1.2–)1.4–
1.7(-4) mm long, with 5 hairy lobes. Seeds 1.9–2.7
¥ 0.9–1.5 mm, finely reticulate; hilum 1/6 as long as
the perimeter.
Area: South Peru, Bolivia and north and central
Argentina (Fig. 74).
Habitat: Roadsides, cultivated grounds, rocky and
grassy slopes, river shores, shrublands and Alnus,
Eugenia, Podocarpus or Polylepis forest; 1200–
4000 m.
Phenology: Flowering October – May.
Illustration: Figure 88.
Notes: Geranium fallax, as previously mentioned, is
quite similar to G. venturianum and G. tablasense.
However, this species always has glandular hairs,
sometimes restricted to pedicels, sepals and fruits, but
usually spread over all the inflorescence and stem. In
G. fallax petals are shorter than in G. venturianum
and G. tablasense, and glabrous on both sides, with
cilia on the basal margin, while they are glabrous in
G. tablasense, and hairy on the margin and base of the
abaxial side in G. venturianum. Additionally, G. fallax
has a narrowed apex of the rostrum that is shorter
than G. venturianum and G. tablasense.
The specimens from Bolivia were usually identified as G. fiebrigianum, which is considered in this
study as a synonym of G. fallax. Other specimens
have occasionally been misidentified as G. patagonicum (included in G. berteroanum in this study),
which has leaves with obtriangular segments (not
rhombic).
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60
C. AEDO ET AL.
F
H
A
I
E
G
J
L
B
C
K
D
Figure 88. Geranium fallax Steud. A–B, habit; C, leaf; D, bracts; E, flower; F, flower without petals; G, sepal; H, petal; I,
staminal filament; J, fruit; K, mericarp; L, seed [A, C–L: Beck 13831 (MA); B: Troncoso 11307 (CORD).].
Representative specimens examined: ARGENTINA.
CATAMARCA: Ambato, Los Varela, 1 km W, camino a
Humaya, 27∞49¢S, 65∞58¢W, 28.ii.1996, Saravia et al.
13733 (CTES); Andalgalá, 27∞36¢S, 66∞19¢W, .v.1915,
Jörgensen 1194 (SI, MO); cerca Belen, 27∞38¢S,
67∞1¢W, .1879, Schickendantz 138 (CORD); El Rodeo,
Sierra de Ambato, 28∞34¢S, 66∞7¢W, 12.i.1957,
Calderón 1282 (SI); las Granadillas, 27∞28¢S, 67∞13¢W,
.ii.1872, Lorentz 572 (GOET). CÓRDOBA: Alta Gracia,
Cascada Grande, 31∞40¢S, 64∞26¢W, 31.i.1944, Pierotti
s.n. (U); Calamochita, Sierra Grande, al pie del Cerro
Champaqui, 31∞59¢S, 64∞56¢W, 14.i.1952, Hunziker
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GERANIUM SECTIONS ANDINA AND CHILENSIA
9610 (BH); Pampa de Achala, 31∞35¢S, 64∞50¢W,
2.iv.1944, Rentzell 15162 (SI, GH); Pumilla, Cruz
Grande, 7.i.1951, Santa 3657 (LIL). JUJUY: Capital,
entre León y Nevado de Chañi, Mesada, 24∞4¢S,
65∞46¢W, .iii.1963, Fabris et al. 4212 (LP);
Humahuaca, 23∞12¢S, 65∞20¢W, 7.ii.1941, Hunziker
1302 (NY); Tilcara, quebrada de Huichaira, 23∞29¢S,
65∞31¢W, 14.ii.1959, Fabris & Marchionni 1873 (LP);
Tumbaya, Volcán, ladera E al S del pueblo, 23∞43¢S,
65∞39¢W, 13.ii.1985, Kiesling et al. 5142 (SI). LA
RIOJA: Famatina, Rodeo de las Vacas, 29∞4¢S, 67∞39¢W,
.ii.1913, Flossdorf s.n. (SI); Famatina, Sierra Famatina, cueva de Noronha, 28∞58¢S, 67∞42¢W, .ii.1980, Cei
s.n. (MERL); Velazco, 29∞4¢S, 67∞4¢W, 6.iii.1944, Soriano 932 (SI). SALTA: Cerro del Cajón, La Laguna,
25∞46¢S, 65∞14¢W, 20.i.1914, Rodríguez 1272a (SI);
Chicoana, 14 km SW of San Fernando de Escoipe
along the road to Cuesta del Obispo, 3 km W of San
Martín, 25∞10¢S, 65∞50¢W, 10.ii.1993, Till 10174 (WU);
Guachipas, arroyo Querusillas, 25∞44¢S, 65∞20¢W,
7.ii.1983, Novara & Neumann 3194 (LP); Las Juntas,
23∞7¢S, 64∞33¢W, .xii.1896, Bruch s.n. (LP). SAN JUAN:
Jachal, Bella Vista, El Salto, 31∞8¢S, 69∞27¢W,
1.ii.1987, Kiesling & Megioli 6680 (SI); Sarmiento, río
Bachongo, 31∞58¢S, 68∞54¢W, 23.i.1986, Guaglianone
et al. 1479 (SI); Sarmiento, río Santa Rosa, a 5 km
desembocadura río Los Leones, 31∞59¢S, 68∞56¢W,
24.i.1986, Guaglianone et al. 1497 (SI). SAN LUIS:
Pringles, Carolina, 32∞49¢S, 68∞9¢W, 5.ii.1984, Del
Vitto & Wittenstein 644 (MERL); San Francisco,
33∞30¢S, 65∞16¢W, 4.xi.1958, Ruiz Leal s.n. (MERL);
sierra del Morro, 32∞53¢S, 64∞59¢W, .1913, Pastore 23
(SI). TUCUMÁN: camino de Acheral a Tafi del Valle,
27∞7¢S, 65∞26¢W, .i.1944, O’Donell 1390 (U); La Banderita, 27∞17¢S, 65∞54¢W, .ii.1950, Brücher 9394 (M);
La Ciénaga, 26∞46¢S, 65∞39¢W, 14.ii.1905, Lillo 4031
(UC). BOLIVIA. COCHABAMBA: Ayopaya, 10 km NW
de Independencia, 17∞7¢S, 66∞52¢W, 7.v.1988, Beck &
Seidel 14427 (MA, GH, PH, UC); Calomi, 17∞0¢S,
65∞30¢W, 8.i.1949, Brooke 5081 (BM, GH, PH, UC);
Chapare, Incachaca, 17∞14¢S, 66∞28¢W, 28.ii.1929,
Steinbach 9509 (MO, E, GH, NY, PH, UC); Quillacollo,
17∞30¢S, 66∞30¢W, 25.iii.1990, Hensen 702 (MA, GH,
PH, UC). LA PAZ: Inquisivi, cliffs 0.5 km SE of Jardin
Botanico de Quime, 16∞59¢S, 67∞13¢W, 17.ii.1990,
Lewis 37100 (LPB); Larejaca, Sorata cerro del Imimapi, 15∞46¢S, 68∞39¢W, .ii.18??, Mandon 778 (W, S);
Murillo, 1 km NW of Ovejuyo, 16∞32¢S, 68∞3¢W,
2.iv.1982, Solomon 7422 (LPB, MO, NY); Omasuyos,
de Huarina unos 10 km hacia Tiquina, 16∞12¢S,
67∞37¢W, 3.v.1999, Beck 22987 (MA); Sud Yungas,
19 km E of pass between Mururata and Illimani,
16∞34¢S, 67∞45¢W, 14.iii.1986, Solomon 15121 (LPB, G,
MO, NY, U). POTOSÍ: Quijarro, camino de Potosí a
Khucho Ingenio, 5 km antes de llegar a Khucho Ingenio, 19∞55¢S, 65∞39¢W, 16.i.1988, Schulte 92 (MA);
61
Saavedra, Cantón, Tuero Saavedra, Comunidad
Despensa, 19∞20¢S, 65∞20¢W, s.d., Romero 74 (LPB).
Tarija: Cercado, 10 km NW of Tomatas, 5 km N of
Tarija, on road through Erquis, Angosturas de Erquis,
21∞28¢S, 64∞50¢W, 9.v.1983, Solomon 10592 (MO, NY);
Cercado, Cuesta de Sama, 21∞29¢S, 65∞2¢W, 22.ii.1986,
Bastión 796 (U); Méndez, Sama, 21∞29¢S, 65∞1¢W,
22.ii.1986, Ehrich 139 (LPB). PERU. Arequipa: Arequipa, 16∞53¢S, 71∞20¢W, 4.iv.1937, Stafford 625 (K,
BN). CUZCO: inter oppidum Calca et pagum Huaman
(Choqque), 13∞18¢S, 71∞43¢W, 20.v.1934, Hammarlund
615 (NY). PUNO: Pillahuata, cerro de Cusilluyoc,
13∞7¢S, 71∞24¢W, 3.v.1925, Pennell 14133 (F).
EXCLUDED OR DUBIOUS NAMES
Geranium albicans var. glanduliferum Hieron., nom.
nud., in sched. (CORD!)
Geranium brevipes L’Hér. ex DC., Prodr. 1: 639 (1824),
nom. nud., pro syn.
Geranium core Kostel., Allg. Med.-Pharm. Fl. 1900
(1836)
Type: Feuillé, Journal des Observationes Physiques,
Mathematiques et Botaniques 3: tab. 16 (1725) (lectotype, here designated)
According to Knuth (1912) this name should be considered as a synonym of G. rotundifolium L. Dr J. Hadinec has gone carefully through Kosteletzky’s herb.
material of Geranium. He kindly informed us that
there is no specimen at PRC with any indication that
it could serve as type material of G. core Kostel. The
name is also absent from the first list of PRC material
made after the incorporation of Kosteletzky’s herbarium into PRC. Thus the drawing by Feuillé becomes
the only original element. This drawing shows a
perennial species with turnip-shaped rootstock, which
probably belongs to sect. Chilensia. However, there
is no other feature which permits a more exact
identification.
Geranium dissectum var. tuberosum F. Muell. ex R.
Knuth in Engl., Pflanzenr. IV.129 (Heft 53): 75 (1912),
nom. nud., pro syn. (and in sched., W!)
Geranium geissei R. Knuth in Engl., Pflanzenr. IV.129
(Heft 53): 70 (1912)
Type: Chile. [without more exact location specification], Geisse s.n. (holotype, B†; photo F! G! GH! NY!)
The type of this species was destroyed during World
War II. Unfortunately, no duplicate of this collection
has been found. We located some specimens collected
by Geisse at SGO, none connected to this name. Photographs of the type kept at some herbaria do not permit a clear identification, but strongly suggest that
G. geissei could be a synonym of G. berteroanum.
Geranium herrerae R. Knuth in Herrera, Chlor. Cuzco.
151 (1926), nom. nud.
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
62
C. AEDO ET AL.
Geranium magellanicum var. typicum R. Knuth in
Engl., Pflanzenr. IV.129 (Heft 53): 70 (1912), nom.
inval.
Geranium neesianum Carolin in Jessop, List Vasc. Pl.
South Australia: 35 (1983), nom. nud.
Geranium patulum Sol. in G. Forst., Fl. Ins. Austr. 91
(1786), nom. nud.
Geranium pilosum Sol. in G. Forst., Fl. Ins. Austr. 91
(1786), nom. nud.
Geranium proximum Bertero ex Steud., Nomencl. Bot.
ed. 2, 1: 679 (1840), nom. inval., sine descr.
Geranium rotundifolium var. affine Bertero ex R.
Knuth in Engl., Pflanzenr. IV.129 (Heft 53): 76 (1912),
nom. nud.
Geranium rotundifolium var. minor Bertero ex R.
Knuth in Engl., Pflanzenr. IV.129 (Heft 53): 76 (1912),
nom. nud.
Geranium santacruzense R. Knuth in Repert. Spec.
Nov. Regni Veg. 18: 289 (1922)
Type: Argentina. Santa Cruz 1902, Cáceres 12395
(holotype, B†; isotype, CORD)
Unfortunately we can not examine the type of the
species. Barboza (1996) synonymized G. santacruzense
to G. sessiliflorum, which seems suitable considering
Knuth’s description.
Geranium sessiliflorum var. typicum R. Knuth in Bot.
Jahrb. Syst. 37: 565 (1906), nom. inval.
Geranium sessiliflorum f. albiflorum Kuntze, Revis.
Gen. Pl. 3(2): 33 (1898), nom. nud., in sched. (NY!)
Geranium solanderi var. grandis Carolin in Proc.
Linn. Soc. New South Wales ser. 2, 89: 353 (1965)
Type: Australia. Ebor Gorge, New England,
2.i.1959, Carolin 766 (holotype, NSW)
Unfortunately we cannot examine the type of this
variety. Carolin (1965) stated that var. grandis has a
tap-root not napiform, which suggest that it could
belong to another section.
ACKNOWLEDGEMENTS
The authors wish to thank G. Barboza, G. Guignard,
J. Hadinec, L. Iharlegui and J. Stepanek for help in
locating some type or critical material, F. Muñoz
Garmendia for nomenclatural advice, C. Marticorena and R. Rodríguez for kind assistance during
our field trip in Chile, M. Jerez and A. Martín for
their technical support and S. Castroviejo for uncompromising support. We thank J.J. Aldasoro for his
critical review of the manuscript. We are also grateful to the curators of the cited herbaria for kind
assistance during our visits and for loan of specimens. This work was partly financed by the Spanish
Government through the research projects REN200204634-C05-05/GLO, PR2001-0342, CGL20004-00172/
BOS and by the European Union through the
COLPARSYS program.
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INDEX TO SCIENTIFIC NAMES
The accepted names are in bold type, while taxa that
are not treated here or are synonyms are not in bold type.
Geranium L., 2
Geranium subgen. Erodioidea (Picard) Yeo, 1
Geranium subgen. Robertium (Picard) Rouy, 1
Geranium sect. Andina R. Knuth, 20
Geranium sect. Azorelloida Aedo, Muñoz Garm. &
Pando, 1
Geranium sect. Batrachioidea W.D.J. Koch, 33
Geranium sect. Chilensia R. Knuth, 31
Geranium sect. Gracilia R. Knuth, 1
Geranium sect. Neoandina Aedo, 1
Geranium sect. Palustria R. Knuth, 2
Geranium sect. Paramensia R. Knuth, 1
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
64
C. AEDO ET AL.
Geranium sect. Rupicola R. Knuth, 2
Geranium sect. Trygonium Dumort., 1
Geranium sect. Tuberosa (Boiss.) Reiche, 1
Geranium albicans A. St.-Hil., 42
Geranium albicans var. glanduligerum Hieron., 61
Geranium andinum Phil., 32
Geranium antrorsum Carolin, 26
Geranium apricum Phil., 31
Geranium argentinum R. Knuth, 47
Geranium asphodeloides Burm.f., 9
Geranium australe Nees, 52
Geranium bangii Hieron., 20
Geranium berteroanum Colla, 31
Geranium berteroanum var. apricum (Phil.) Reiche, 31
Geranium berteroanum var. caespitosum Reiche, 32
Geranium berteroanum var. ciliatum Reiche, 44
Geranium berteroanum var. hispidum Reiche, 32
Geranium berteroanum var. philippi (J.F. Macbr.) L.E.
Navas, 44
Geranium brevicaule Hook. f., 24
Geranium brevipes L’Hér. ex DC., 61
Geranium caespitosum Phil., 32
Geranium caespitosum Walp., 20
Geranium carolinianum var. australe (Benth.) Fosberg, 35
Geranium chilense Aedo & Muñoz Garm., 32
Geranium ciliatum Phil., 44
Geranium collae Aedo, Muñoz Garm. & Pando, 31
Geranium commutatum Steud., 47
Geranium commutatum var. ochsenii (Phil.) Reiche, 47
Geranium core Kostel., 61
Geranium core-core Steud., 47
Geranium deprehesum (E.G. Almq.) Lindm., 9
Geranium dissectum f. tasmanica Gand., 35
Geranium dissectum var. australe Benth., 35
Geranium dissectum var. patagonicum (Hook. f.)
Speg., 31
Geranium dissectum var. patulum Sol. ex Hook. f., 52
Geranium dissectum var. pilosum Hook. f., 35
Geranium dissectum var. retrorsum (L’Hér. ex DC.)
Hook. f., 52
Geranium dissectum var. tuberosum F. Muell. ex R.
Knuth, 61
Geranium drummondii Carolin, 35
Geranium fallax Steud., 59
Geranium fiebrigianum R. Knuth, 59
Geranium geissei R. Knuth, 61
Geranium herrerae R. Knuth, 47, 61
Geranium hispidum Phil., 32
Geranium intermedium Bertero ex Colla, 31
Geranium limae R. Knuth, 38
Geranium macrorrhizum L., 6
Geranium macrostylum Boiss., 9
Geranium magellanicum Hook. f., 40
Geranium magellanicum var. multifoliosum R. Knuth,
29
Geranium magellanicum var. pumilum R. Knuth, 29
Geranium magellanicum var. typicum R. Knuth, 62
Geranium malpasense R. Knuth, 20
Geranium melanopotamicum Speg., 47
Geranium moorei Phil., 47
Geranium neesianum Carolin, 62
Geranium neohispidum Aedo & Muñoz Garm., 32
Geranium ochsenii Phil., 47
Geranium parodii I.M. Johnst., 29
Geranium patagonicum Hook. f., 31
Geranium patulum Sol., 62
Geranium pflanzii R. Knuth, 20
Geranium philippii J.F. Macbr., 44
Geranium pilosum Sol., 62
Geranium pilosum Sol. ex Willd., 35
Geranium pilosum var. australe (Nees) Ostenf., 52
Geranium pilosum var. grandiflorum R. Knuth, 52
Geranium pilosum var. retrorsum (L’Hér. ex DC.)
Jeps., 52
Geranium platypetalum Fisch. & C.A. Mey., 9
Geranium proximum Bertero ex Steud., 31, 62
Geranium pyrenaicum Burm. f., 33
Geranium rapulum A. St.-Hil. & Naudin, 47
Geranium razuhillcaënse R. Knuth, 20
Geranium reflexum L., 6
Geranium retrorsum L’Hér. ex DC., 52
Geranium reuteri Aedo & Muñoz Garm., 6
Geranium robertianum L., 6
Geranium rotundifolium L., 6
Geranium rotundifolium var. affine Bertero ex R.
Knuth, 62
Geranium rotundifolium var. americanum A. St.-Hil.
& Naudin, 42
Geranium rotundifolium var. minor Bertero ex R.
Knuth, 62
Geranium sanguineum L., 8
Geranium santacruzense R. Knuth, 62
Geranium selloi Aedo & Muñoz Garm., 43
Geranium senecioides R. Knuth, 43
Geranium sessiliflorum Cav., 20
Geranium sessiliflorum f. albiflorum Kuntze, 62
Geranium sessiliflorum ssp. brevicaule (Hook. f.)
Carolin, 24
Geranium sessiliflorum ssp. novaezelandiae Carolin,
24
Geranium sessiliflorum var. albatum J.F. Macbr., 20
Geranium sessiliflorum var. arenarium G. Simpson &
J.S. Thomson, 24
Geranium sessiliflorum var. compactum R. Knuth, 20
Geranium sessiliflorum var. glabriusculum Kuntze, 29
Geranium sessiliflorum var. glabrum R. Knuth, 24
Geranium sessiliflorum var. lanatum R. Knuth, 20
Geranium sessiliflorum var. maculatum G. Simpson &
J.S. Thomson, 24
Geranium sessiliflorum var. microphyllum Kuntze, 20
Geranium sessiliflorum var. typicum R. Knuth, 62
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
GERANIUM SECTIONS ANDINA AND CHILENSIA
Geranium skottsbergii R. Knuth, 44
Geranium solanderi Carolin, 35
Geranium solanderi var. grandis Carolin, 62
Geranium squamosum Phil., 47
Geranium staffordianum R. Knuth, 20
Geranium submolle Steud., 32
Geranium subsericeum R. Knuth, 47
Geranium sylvaticum L., 9
Geranium tablasense R. Knuth, 53
Geranium titicacaense R. Knuth, 59
Geranium tucumanum R. Knuth, 20
Geranium venturianum R. Knuth, 57
APPENDIX
INDEX
TO NUMBERED COLLECTIONS CITED
The numbers in parentheses refer to the corresponding species in the text. Three mixed collections have
been located: Bertero 294 (including G. berteroanum,
and G. skottsbergii), Lorentz 572 (including G. fallax
and G. sessiliflorum), and Sleumer 454 (including G.
core-core, and G. berteroanum).
Acleto 735 (1); Acosta Solís 8884 (11), 9303 (11);
Adams 2493 (3); Aedo 3867 (2), 4475 (2), 6722 (11),
6727 (5), 6728 (11), 6737 (5), 6772 (10), 6779 (10), 6807
(10), 6819 (10), 6821 (10), 6822 (10), 6824 (10), 6832
(10), 6838 (10), 6839 (10), 6842 (10), 6913 (5), 6931 (5),
6935 (11), 6937 (5), 6939 (5), 7006 (11), 7073 (11), 7125
(1), 7130 (11), 7136 (5), 7148 (5), 7153 (11), 7154 (5),
7165 (5), 7174 (11), 7185 (5), 7197 (11), 7203 (11), 7205
(11), 7224 (8), 7243 (5), 7296 (5), 7375 (11), 7387 (11),
7391 (5), 7414 (5), 7445 (8), 7449 (8), 7459 (5), 7460 (5),
7461 (5), 7468 (5), 7471 (8), 7472 (5), 7482 (5), 7486 (1),
7500 (1); Aguilar 93 (1); Ahumada 5283 (14); AlemánFernández 2810 (1); Allen A61 (6); Álvarez 50 (8);
Ambrosetti 1611 (5), 1659 (5); Ambrosetti & Méndez
613 (1), 4105 (1); Anderson 18 (5), 49 (9), 3344 (11);
Andreas 93 (10), 159 (11), 422 (5), 438 (5), 562 (5), 748
(11), 749 (11), 841 (10); Angulo 115 (11); Annels 1943
(12), 3481 (12); Aplin 6425 (6), 6478 (6), 6528 (6);
Aravena 16 (10); Arancio et al. 94114 (11); Arrillaga
et al. 1616 (9); Arroyo 84-172 (1), 841138 (1); Arroyo
et al. 841055 (1), 870253 (1), 92-151 (1), 92-82 (1);
Arroyo & Squeo 850810 (1), 850939 (1); Asplund 2161
(1), 2430 (1), 2801 (1), 2884 (1), 3272 (1), 3488 (15),
3677 (1), 460 (15), 6111 (1), 10958 (11), 11455 (11),
13754 (7), 13774 (7), 1886 (1); Atanasio 2096 (11);
Atkins 297 (6); Avila 61 (1).
Baeza 302 (5); Baeza et al. 11705 (11); Baeza &
Peñailillo 11433 (11); Baldeón 132 (7); Baldeón et al.
67 (7); Bang 788 (1); Barba 146 (5), 1702 (5), 193 (11),
218 (5), 387 (5), 507 (5), 74 (5), 793 (5), 1197 (5); Barclay 2356 (11); Barco 311 (5); Barnier 286 (10), 307 (5);
Barros 1932 (11), 24699 (10); Bastión 749 (1), 796 (15);
Baxter et al. 79 (5); Bayer 56 (5), 117 (11), 157 (11);
65
Beamish 1565 (3); Beauglehole 12520 (6), 36304 (3),
40759 (3); Beauglehole et al. 33303 (3); Beck 244 (1),
244b (1), 970 (1), 1315 (15), 2331 (15), 2401 (15), 2441
(1), 3828 (1), 3888 (15), 4103A (1), 4179 (1), 4363 (1),
4405 (15), 8662 (15), 8976 (11), 13817 (15), 13831 (15),
14356 (15), 14635 (15), 17242 (1), 20935 (1), 21256
(15), 21990B (1), 22401 (15), 22688 (1), 22987 (15),
22989 (15), 24386 (1); Beck et al. 11712 (11); Beck &
Seidel 14427 (15), 14576 (15); Bernardello & Moscone
577 (11); Bertero 294 (5), 294 (10), 296 (11), 1018 (11),
1018 (10), 1019 (10), 1020 (5), 1460 (11); Biese 115 (5),
2011 (11); Billiet 3554 (5); Blake 33 (11); Blaylock
1313 (6); Bliss et al. 2364 (11); Bobadilla 126 (11), 137
(11), 234 (11); Böcher et al. 349 (7), 1275 (1); Boeke
1118 (1); Boelcke 12706 (5); Boelcke et al. 13560 (11),
13695 (11), 13782 (1), 16271 (8); Boelcke & Hunziker
3513 (5); Boelcke & Correa 5216 (11), 5890 (5), 11439
(5); Boffa 42 (9); Bohlen 689 (1); Borge 165 (5); Brandbyge 275 (1), 319 (1), 329 (1), 498 (1); Bravo 490 (10);
Bridarolli 1416 (15); Bridasoli 264 (11); Bridges 218
(10), 609 (10), 777 (5); Briggs 1760 (3); Briggs & Zich
2632A (12), 2639 (12), 2640 (12); Brooke 5081 (15),
5182 (1), 5350 (1), 6250 (1); Brown 781 (2), 5224 (12);
Brücher 9394 (15), 9396 (14), 9401 (15), 9437 (1), 9440
(14), 9441 (15), 9496 (1); Brunel 202 (1); Bruzzone 42
(5); Buchtien 607 (15), 611 (15), 675 (15), 1332 (8),
1332 (5), 4128 (1), 4602 (15), 8734 (1), 8968 (1); Budún
1506 (1); Burkart 4707 (9), 9468 (11), 15570 (9), 15905
(11), 18089 (9), 27507 (5); Burkart et al. 30494 (14);
Burkart & Troncoso 11307 (15); Burlidge & Gray 4184
(12); Burmeister 38 (1); Burtt Davy 1650 (11).
Cabrera 5987 (5), 8017 (9); Cabrera & Sagastegui
19274 (9); Cabrera et al. 15275 (1), 17303 (14), 18314
(1), 18364 (1), 18398 (1), 21290 (14), 22007 (1), 22018
(1), 22921 (1), 25830 (14), 27207 (1), 30670 (14), 30721
(1), 30859 (15), 31665 (1), 32920 (5); Cabrera & Fabris
16499 (9); Cabrera & Crisci 19177 (1); Cabrera &
Frangi 20694 (1); Cabrera & Job 32 (5); Cabrera &
Kiesling 20164 (14); Calderón 1282 (15); Cambage
1850 (3); Cameron 387 (2), 388 (2), 8342 (12), 9952 (6);
Cañigueral 229 (11), 230 (15), 255 (1); Canning 1415
(6), 1458 (6), 1525 (6), 1600 (3), 1764 (3), 1850 (6), 3161
(12), 3276 (12), 4396 (6), 4410 (6), 5805 (12), 6672A
(12); Cantino 120 (5); Cárdenas 588 (1), 758 (13), 2180
(13), 6028 (1); Carolin B78 (3), B103 (3), 761 (3), 778a
(2), 784 (6), 798B (3), 818 (6), 829 (6), 861 (6), 862 (6),
865 (6), 938 (6), 1023 (6), 2069 (6), 2070 (6); Carrasco
238 (5), 276 (11); Casson & Annels 39.2 (6); Castellanos 151 (11), 728 (4), 14743 (1); Castillón 3485 (1);
Ceballos et al. BO-58 (1), 482 (1); Cerrate 6407 (7);
Chávez 2317 (1); Chiuminato 23 (11), Claren 11443
(1), Claude Joseph 147 (10), 201 (10), 692 (10), 1397
(5), 2067 (10), 2172 (10), 2734 (11), 2825 (10), 2825a
(10), 3091 (5), 3647 (10); Clemens 42/65 (6), Collier
1123 (2), Comber 833 (5), Condit 130 (11); Conrad
2669 (1), 2690 (1); Constable 19190 (6); Copley 537
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
66
C. AEDO ET AL.
(12); Cordini 14 (5), 139 (1); Correa et al. 5858 (1),
7111 (9), 9181 (1); Corvas 398 (5); Coveny 8731 (6),
10216 (6), 10289 (6), 11530 (3); Cranfield 26 (6), 7612
(6), 10388 (12); Craven 2071 (6); Crawford 1807 (12),
2137 (12), 2447 (12), 3221 (12), 6419 (12), 13517 (6);
Crespo et al. 2222 (11); Crespo & Troncoso 1694 (5);
Crowley 661 (12); Croxford 7921 (12), 7924 (12), 8100
(12); Cruz et al. 90 (1); Cullimore 100 (6); Cusato &
Rossow 4114 (5), 4131 (5); Cutler 7664 (1).
Davis 4168 (12), 8059 (6), 8170 (12), 8242 (12), 8424
(12); Davis et al. 1552 (1); Dawson & Nuñez 70 (9); Del
Vitto & Wittenstein 644 (15); Dempster 4478 (11);
Descole 3140 (5); Dessauer 1450 (10), 1451 (11);
DeVore 1301 (11); DeVore & Baeza 1631 (5); Dewing
612 (6); Diers 811 (1); Dillon et al. 3142 (1); Dillon &
Turner 1331 (11); Dollenz 84 (8), 635 (8), 1432 (8);
Donat 223 (5); Donner 9045 (12), 9203 (12); Drummond 4bis (6), 501 (6); Duarte 24 (11), 55 (11); Duek
et al. 40 (5); Dusén 77 (8), 138 (1), 233 (5), 319 (10), 455
(8), 5645 (5), 6246 (8).
Ehrich 139 (15); Eichler 13652 (3), 19916 (12);
Ellenberg 197 (1), 394 (1), 615 (1), 632 (1), 4035 (1);
Elliot 89 (11), 227 (11), 261 (5); Elmer 4046 (11); Elvebakk 289 (5); Ertter 10725 (11), 17410 (11); Espinosa
562 (11); Eyerdam 10259 (5); Eyerdam et al. 23628 (9),
24140 (8).
Fabris 1093 (5), 5614 (11); Fabris et al. 3959 (1),
4137 (1), 4212 (15); Fabris & Marchionni 1873 (15);
Fabris & Solbrig 216 (11), 890 (5); Fernández Casas &
Charpin 6616 (1); Ferraro et al. 4705 (5); Ferreyra
1493 (7), 1539 (7), 2445 (7), 4029 (7), 8620 (7), 8750 (7),
8839 (7), 9557 (7), 11797 (7), 11811 (7); Fiebrig 2627
(1), 2631 (15), 3290 (1), 3291 (1); Finckh 258 (1); Flühmann 261a (5), 2126 (5); Fortunato et al. 1872 (1); Fosberg 30464 (2); Frangi et al. 50 (1), 79 (1); Fries 1024
(1), 1034 (1); Frödin 7 (11); Furlong 89c (8), 91 (1), 92a
(5), 92b (5), 92c (5).
Gallinal et al. 782 (9), 1343 (9), 1766 (9), 5290 (9),
5354 (9); Garaventa 4929 (10); García 311 (11); 2343
(1); García et al. 911 (1), 934 (1); Gardner 1635 (12),
1944 (6), 2172 (2), 2219 (6), 2419 (6), 2635 (6), 2645 (6),
2647 (6), 2712 (6), 2772 (12), 2775 (6), 2777 (6), 2801
(2), 3376 (6), 3851 (12), 3899 (12), 3904 (6), 3913 (6),
4000 (6), 4061 (12), 4289 (12), 6066 (6), 6598 (12), 6919
(6), 7504 (12); Gardner & Knees 6188 (8); Garraty 8
(6); Gaudichaud 179 (10), 4 (7), 1204 (11); Gay 783 (1),
785 (10); Gentry 44178 (1); Germann 17 (1); Gerold
209 (1); Giardelli 199 (15); Gibbons 506 (6); Gibson &
Lyons 534 (12), 1541 (6); Giovanelli 13594 (5); Gleisner 208 (5); Godden & Day 72.4 (6); Godley 1185 (5);
Godman 22 (12); Gómez-Sosa & Múlgura 154 (1), 181
(1); González 105 (1); Goodall 193 (1), 1220 (1), 1604
(5), 2404 (1), 3594 (5); Goodspeed 23308 (15); Goodspeed et al. 11526 (7); Grant 7445 (7); Gray 5479 (6),
5824 (6), 5891 (12); Guaglianone et al. 1407 (11), 1479
(15), 1497 (15); Guenther & Buchtien 690 (11), 691
(11), 1318 (1), 1345 (1), 1348 (1), 1356 (1), 2036 (1);
Gunckel 380 (5), 508 (5), 2672 (10), 10814 (5), 13762
(11), 14604 (11), 14716 (5), 15324 (5), 16557 (11),
21592 (5), 22266 (10), 22286 (5), 26378 (5), 27185 (5),
27362 (11), 34705 (5), 35696 (10), 36602 (10), 37498
(5), 37619 (11), 38542 (10), 39187 (11), 39986 (11),
40546 (10), 41438 (5), 42211 (10), 42649 (5), 42784
(10), 42817 (11), 46921 (5); Gunn 209 (6), 256 (2), 453
(6); Gusinde 257 (5); Gutiérrez 3 (10),131 (5).
Haegi 819 (12); Hahn 12-80bis (5); Hamlin 915 (2);
Hammarlund 190 (1), 452 (1), 615 (15); Hammersley
983 (12); Harshberger 1061 (10); Hauthal 155 (1), 323
(1); Hawkes et al. 3418 (1); Healy 72 (12); Heins 522
(5); Helms 39 (12), 1426 (2); Hensen 702 (15), 1418 (1);
Herrera 2383 (1); Herzog 758 (1), 2181 (13); Hicken
25571 (9); Hieronymus 449 (11), 753 (4); Hieronymus
& Niederlein 7776 (1); Hieronymus & Lorentz 42 (1);
Hinkley 66 (11); Hollermayer 542 (5), 576 (11); Holway
19 (10), 123 (11), 261 (5); Hoogland 3107 (6); Hoogte &
Roersch 1216 (1), 2142 (1), 2356 (1), 2356b (1), 2407
(1); Hoover 6960 (11); Hosaka 3590 (12); Hosking 655
(12); Hosseus 13 (4); Hoyle 1558 (6); Hubbard 4367 (6),
5848 (6); Huidobro 1162 (9); Humbert 30780 (1); Hunziker 1302 (15), 1384 (4), 9610 (15), 9622 (4); Hunziker
& Caso 6160 (1); Hutchinson & Tovar 4220 (1).
Illin 6 (5), 36 (5), 98 (5), 124 (5); Iltis et al. 1373 (1),
1475 (1).
Jackson 100 (6); Jackson & Canning 88 (3); Jaffuel
861 (1), 869 (10), 917 (10), 1371 (11), 3872 (1), 3994 (5);
Jaffuel & Pirion 3055 (10), 3873 (5); James 109 (1), 380
(5), 1122 (5); Jiles 659 (10), 664 (10), 682 (10), 1750 (5);
Jiménez 15182 (1); Johns 82-65 (1), 82-91 (1); Jordan
151 (1); Jordan et al. 19 (1); Jörgensen 1194 (15), 1387
(1), 1387b (1), 1561 (15), 1562 (14); Junge 965 (5), 973
(5),1855 (5), 2910 (5).
Kalela 578 (1), 832 (5), 1085 (5), 1461 (5); Kalenborn 80 (1); Kausel 4351 (11); Keighery 5294 (12),
5984 (12), 6248 (6), 8485 (12), 10523 (12), 11246 (12),
11917 (12), 15099 (6); Keighery & Alford 1650 (12),
1818 (12); Keighery & Gibson 299 (6), 393 (6), 436 (6),
467 (12), 812 (12), 852 (6); Kiesling et al. 663 (15), 672
(15), 820 (14), 1602 (1), 5142 (15), 5217 (1); Kiesling &
Megioli 6680 (15); Killip & Smith 21971 (1); Killip &
Pisano 39714 (11); King 434 (11); King et al. 192 (1);
Kirk 305 (2); Kohler 90 (10); Koslowsky 71 (8), 12445
(8); Krapovicas & Schinini 38296 (9); Krapovickas
7427 (4); Krapovickas et al. 47286 (1); Krapovickas &
Cristobal 14672 (11), 16197 (9), 20526 (15); Kunkel
252 (5), 375 (11), 595 (5); Kurk 1011 (2); Kurtz 2944
(4), 6584 (15), 8339 (4), 8388 (15),11186 (11).
Lagiglia 2011 (11); Lahitte & Clos 126 (9), 194 (9),
241 (9), 256 (9); Lammers et al. 6394 (11); Landero 703
(1), 753 (1); Landrum 8181 (5); Lange 225 (6), 1903
(12), 1998 (2), 2981 (12), 2986 (6), 2987 (12), 3154 (6),
3418 (2), 3718 (6); Lanza 7559 (5); Lara 1661 (1), 1707
(1); Lechler 259 (11), 1033 (5), 1160 (1), 1907 (15), 1955
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
GERANIUM SECTIONS ANDINA AND CHILENSIA
(1), Ledezma 327 (11); Leibold 3110 (10); Leuenberger
& Arroyo 3932 (1); Levi 363 (10), 364 (10), 1330 (10);
Lewis 37063 (1), 37100 (15), 871073 (11); Liberman
412 (1), 879 (1), 918 (15); Lillo 2776 (15), 4031 (15);
Lindeman et al. 20673 (9); Llatos 497 (7); Looser 206
(10), 2231 (11); López 224 (11), 745 (5), 11404 (11);
López & Peñailillo 11548 (11), 11620 (11); Lorentz 572
(1), 572 (15), 582 (1); Lorentz & Hieronymus 42 (1),
582 (1), 584 (1); Lotti 150 (5); Lourteig & Buchinger
123 (1); Lovis 463 (12), 635 (2), 715 (2); Lupo 94 (1);
Lynd 77 (6).
Macbride 3006 (1); Macbride & Featherstone 608
(1), 808 (1), 1148 (1), 1148A (1), 1148B (1); Magens 842
(1); Mahu 250 (10), 270 (10), 497 (11), 3211 (5), 4521
(5), 7610 (11), 9064 (11), 9802 (5), 9915 (11), 9940 (5),
10025 (11), 10028 (11), 10037 (10); Makinson 1036 (3);
Malme 269 (9); Mandon 778 (15), 780 (1), 782 (1), 784
(1); Marco 586 (5); Marticorena 1617 (11); Marticorena
et al. 108 (1), 462 (11), 1009 (11), 1273 (1), 1282 (10),
1286 (5), 1396 (1), 1571 (11), 1652 (11), 1858 (11),
83303-A (11), 9150 (11); Marticorena & Ugarte 9059
(11), 9201 (11); Marticorena & Matthei 883 (1); Marticorena & Rodríguez 10018 (5); Martínez et al. 23 (1);
Maruñak et al. 227 (15); Mason 4276 (11); Matthei &
Rodríguez 546 (1), 579 (1), 614 (8); May 756 (6); McKee
9665 (6); McLean 13 (11); Meebold 4748 (2), 4748b (2),
5982 (3), 18138 (2); Melville 2538 (6); Menhofer 1767
(1), 1825 (1), 1882 (1); Merxmüller 24866 (5), 24920
(1), 25016 (1); Mexia 4078 (11), 7981 (5); Meyer 7454
(5), 8215 (5); Meyer & Sleumer 15544 (4); Mez y Barrera 1568 (1); Meza 1 (11), 338 (5); Meza & Águila
6481 (11); Meza & Villagrán 979 (10), 998 (10); Meza &
Barrera 1650 (5); Michener & Bioletti 2004 (11); Middleton 193 (12); Miers 816 (11); Miller 22 (2); Moandon
781 (1); Montero 191 (5), 1122 (5), 1586 (10), 1597 (10),
1599 (10), 1994 (5), 2199 (1), 2286 (11), 2356 (5), 4679
(10), 5607 (11), 6564 (10), 8 (11), 8423 (5), 8778 (10),
9162 (1), 9467 (5), 9701 (5),10308 (5), 10612 (5), 10629
(5), 11070 (5), 12307 (10); Moore 1064 (5), 2104 (5),
8013 (12), 8357 (2), 8740 (12); Moore et al. 8302 (1),
8315 (1), 8318 (1); Moore & Goodall 51 (8), 347 (8),
19015 (1); Morello 4027 (9); Moreno & Tonini 117 (5),
408 (5); Morris 86146 (2); Morrison & R. Wagenknecht
17516 (8); Moscal 2430 (6); Mueller 7122 (12); Muñoz
162 (11), 537 (1), 793 (11), 2142 (10); Muñoz & Coronel
1267 (5); Muñoz et al. 3488 (1); Muñoz & Pisano 3385
(10); Muñoz & Sierra 7028 (11); Muruaga et al. 13 (15).
Navarro 566 (1), 683 (1); Navas 1499 (11); Neger
13319 (5); Newbey 4298 (12), 6234 (12); Nicora 1555
(15), 1713 (15), 3677 (5), 3843 (11); Nielsen 10091 (12);
Noble 29008 (2), 29164 (2); Novara 2660 (14), 6575
(14), 9209 (15); Novara & Neumann 3194 (15); Núñez
6749 (1); Núñez et al. 9007 (1); Núñez & W. Loayza
8962 (1).
O’Donell 1390 (15), 2217 (5), 3930 (1), 5296 (1);
Ochoa 607 (11); Ochrens 865 (11); Ogle 3 (6), 3823 (2);
67
Olea 170 (15); Olrog 60 (15); Orbigny 1457 (1), 1504
(1); Orchard 3486 (6); Ostria 203 (1), 314 (1).
Pacheco & Valdebenito 6255 (11); Parks 8360 (11);
Parodi 7411 (9), 7514 (4), 7972 (1), 9666 (15), 10385
(9); Parra 210 (10); Pastore 23 (15), 36 (4), 99 (4); Pearson 20 (1); Pedersen 3588 (9), 4756 (9), 6313 (9), 11965
(9), 13203 (9), 14389 (1); Pedley 944 (6); Pennell 12384
(5), 12440 (1), 12575 (5), 12804 (5), 12824 (11), 13439
(1), 14133 (15), 14245 (11), 14247 (11); Pestalozzi 184
(1), 828 (1); Pettersson 197 (1); Peyton et al. 105 (1);
Pfister 7221 (1), 8765 (1); Philipson 10099 (2); Pierotti
1045 (1), 1327 (15); Pisano 1192 (11), 2425 (8), Pisano
2907 (8), 3540 (8), 4250 (5), 4271 (5); Pisano et al. 4058
(5), 4829 (1), 4917 (5), 6800 (5), 7361 (8), 7727 (5);
Pisano & Henríquez 6969 (1); Podestá 37 (1); Poeppig
173 (10); Preiss 1907 (12); Proyecto Ventania 247 (9),
349b (5), 794 (9), 805 (9), 913 (9), 2731 (9); Pullen 4068
(6), 10240 (6).
Racovitza 47 (5), 336 (5); Rahn 162 (7); Ralston 89
(6); Rattenbury 332 (2); Recabarren 53 (5); Rechinger
63048 (5), 64207 (5), 64298 (5); Reed 73 (11), 216 (11);
Rentzell 15162 (15); Ricardi 645 (11), 673 (11); Ricardi
et al. 152 (11), 3374 (1), 340 (1), 951 (1); Ricardi &
Marticorena 5024 (1), 5024b (1), 5112 (1), 5631/1792
(1), 5779/1939 (5); Ricardi & Matthei 116 (1), 269 (8),
488 (1); Rico 7242 (8), 7270 (5); Ríos 36 (11); Riquelme
138 (5); Roch 825 (12); Rodd 815 (3); Rodd & Briggs
2193 (6), 2222 (6); Rodrigo 584 (11), 3038 (11);
Rodríguez 263 (1), 400 (15), 1272 (15), 1272a (15),
1305 (1), 1305b (1); Rodríguez & A. Marticorena 2971
(1); Rodríguez & C. Marticorena 1460 (5); Rodríguez &
Baeza 2399 (11), 2400 (11); Roig 1047 (15), 4656 (11),
11824 (11), 12619 (5); Roig et al. 13307 (5), 13801 (5),
14033 (5), 14407 (5), 14738 (8); Roivainen 376 (5),
1017 (1), 1099 (5), 1776 (5), 2906 (11); Romero 74 (15);
Rondanelli & Humaña 11500 (11), 11512 (11); Rose
50H8 (12), 746 (12), 19150 (10); Rose et al. 22876 (11);
Rossel 101 (1); Rotman 274 (11), 278 (5), 935 (14);
Royce 1330 (6), 2323 (6), 3098 (6), 4377 (12), 10050
(12), 10150 (6); Rudolph 3705 (11); Ruiz & Lammers
8036 (11); Ruiz Leal 643 (11), 6996 (5), 8402 (11),
11083 (5), 14961 (5), 21231 (14), 23079 (5), 24635 (11);
Ruiz Leal & Roig 20562 (9).
Sagástegui 9343 (11); Saint Hilaire 1895 (9),
1897bis (9), 2296 (9); Sanches 24b (1); Santa 3657 (15);
Santesson 1214 (11), 1288 (1), 1452 (1); Saraventa 18
(10); Saravia 30 (1), 14814 (1), 15064 (1); Saravia et al.
13031 (15), 13716 (15), 13733 (15); Saunders 187 (7);
Savatier 117 (1); Schickendantz 116 (1), 138 (15);
Schilling 10062 (10); Schinini et al. 34490 (1), 34518
(1), 34541 (1), 34619 (1); Schlegel 33 (10), 137 (11), 185
(11), 788 (10), 1120 (10), 1460 (10), 1771 (11), 2741
(11), 2770 (5), 3099 (5), 3444 (5), 3554 (1), 3582 (11),
4008 (5), 5852 (1), 7532 (1); Schmit et al. 453 (1), 453b
(1); Schodde 1003 (12), 1220 (3); Schojovskoy 100 (1);
Schreiter 963 (1); Schulte 92 (15), 144 (1); Schwein-
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68
68
C. AEDO ET AL.
furth 19 (2); Scolnik 202 (11); Seabrook 358 (12); Seibert 15 (1), 76 (1), 232 (5), 270 (5); Seki 66 (11); Sellow
4022 (9); Sepúlveda 8 (5); Skottsberg 33 (11), 247 (5),
277 (5), 396 (11), 397 (11), 752 (10), 823 (1), 974 (10);
Sleumer 229 (15), 262 (1), 454 (5), 454 (11), 654 (1),
971 (8),1073 (5), 1924 (1), 2173 (15); Smith 552 (6),
3028 (1), Smith et al. 5633 (1), 9441 (1), 9790 (1),
10352 (1), 11838 (1), 11990 (1); Smith & Everist 799
(6); Smith & Sparre 150 (11), 180 (11); Sobel 2649 (1);
Solomon 4931 (1), 4962 (1), 7422 (15), 9632 (1), 9712
(15), 10592 (15), 11795 (1), 13022 (1), 13076 (15),
13284 (15), 14903 (15), 15121 (15), 15181 (15), 15813
(1), 15846 (1), 16448 (15), 17863 (15); Solomon et al.
13144 (1); Solomon & Stein 11637 (1); Solomon &
Kuijt 11509 (1); Solomon & Uehling 12116 (1);
Solomon & Chevalier 16591 (1); Solomon & King
15939 (15); Soriano 932 (15), 2860 (5), Sota 2982 (14);
Sparre 938 (14), 1839 (11), 1881 (5), 2567 (11), 3042
(5), 4177 (11), 4984 (11), 6088 (1), 9968 (5), 10018 (5),
10024 (5); Sparre & Constance 10867 (5); Sparre &
Skottsberg 112 (11); Sparre & Smith 181 (5), 186 (1),
308 (11), 427 (1); Spegazzini 42 (5); Stafford 458 (1),
484 (1), 624 (11), 625 (15), 654 (1), 747 (1), 1101 (1);
Stajsic 1165 (12); Stebbins 8774 (5); Stebbins &
Robres 8609 (5); Steibel & Troiani 7716 (9); Steinbach
5914 (1), 8669 (11), 8981 (15), 9509 (15), 9528 (1), 9576
(13); Stopford 41 (6); Stork 10939 (1); Stork et al. 9207
(7), 9255 (7), 9288 (7), 10628 (11); Stork & Horton
10217 (1), 10383 (1); Streimann 551 (6); Stuckert 1708
(11), 1835 (15), 8751 (15), 10337 (4), 10824 (15), 10889
(4), 15275 (11), 19428 (9), 20629 (4); Stuessy et al.
5383 (11), 5489 (11), 6794 (1), 6806 (5), 11571A (11),
11585 (11); Subils & Barboza 3270 (15); Sublis & Barboza 4092 (5); Swenson 371 (11); Symon 10459 (12),
11063 (12), 11923 (12).
Taylor et al. 10379 (5); Taylor & Gereau 10987 (5);
TBPA 1815 (5); Teiller et al. 2298 (11); Teillier et al.
2299 (1); Thompson 776 (3), 2288 (3), 2398 (3), 2435
(3); Tilden 687 (12); Till 10174 (15); Tovar 912 (1);
Tracy 2977 (11), 2990 (11), 3238 (6), 3239 (11), 9453
(6), 12251 (11), 15892 (11); Troll 705 (1), 2983 (1),
3101 (1); Troncoso 1254 (9); Troncoso et al. 2090 (9);
Tsujii 277 (5), 643 (1); Türpe 351 (15); Tutin 1025
(1).
Ugarte 45 (5), 89 (5), 115 (11), 144 (5), 181 (5), 190
(5), 210 (11); Ugent 4776 (15).
Valentin 141 (1); Valentini 32 (9); Valenzuela 1044
(1), 1100 (1), 36992 (5); Valla et al. 3293 (11); van Balgooy 4316 (2); van Steenis 23034 (6); Vargas 846 (1),
926 (1),4999 (15); Venturi 1913 (14), 1979 (14), 2667
(14), 2984 (14), 3089 (14), 3178 (1), 3269 (15), 4026
(15), 4036 (14), 4650 (14), 4734 (15), 5070 (14), 6902
(1), 7723 (14), 8626 (1), 9355 (1), 9848 (14); Vickery
42756 (12); Villagrán et al. 8227 (1), 8572 (5), 9366
(11); Villagrán & Águila 6037 (11); Villavicencio 353
(1); Vinet 28A (5).
Walker 167 (12), 950 (3), 5377 (12); Wall & Sparre
209 (9); Wawra 422 (7), 2730 (10); Webb 7629 (2);
Weber 2471 (12); Weberbauer 363 (1), 2775 (1), 7307
(15), 7326 (1), 7479 (11), 7495 (1); Weddell 1997 (1),
4018 (1), 4395 (1); Weisser 352 (5), 9952 (11), 9954 (5);
Weldt-Rodríguez 726/21 (11); Werdermann 363 (11),
560 (5), 1359 (1); West 4795 (1), 6072 (1), 6335 (1);
Wheder 2875 (12); Wheeler 3311 (12), 3493 (6); Whibley 4261 (12); Whinray 393 (12); Wilson 81-122 (2),
1926 (6), 4003 (12), 4243 (6), 7862 (12); Witte 20 (5);
Wood 8042 (1), 8955 (14), 9304 (11); Wood et al. 15287
(14); Worth 9117 (7); Worth & Morrison 15665 (7),
16297 (5), 16430 (5), 16461 (10), 16654 (10); Wright
3008 (6), 5221 (6), 5709 (12), 6371 (6), 6887 (6), 9531
(6), 10461 (12).
Ysern 3935 (11).
Zardini 317 (1); Zardini et al. 1909 (1); Zich & Sutton 227A (6); Zollitsch 103 (5); Zöllner 2508 (1), 5150
(10), 5164 (10), 5206 (10), 5933 (1), 6692 (5), 7263 (5),
7343 (5); Zuloaga & Deginani 197 (14).
© 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 1–68