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PhytoKeys 63: 63–76 (2016)
A new species in the tree genus Polyceratocarpus (Annonaceae)...
doi: 10.3897/phytokeys.63.6262
RESEARCH ARTICLE
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A new species in the tree genus Polyceratocarpus
(Annonaceae) from the Udzungwa Mountains
of Tanzania
Andrew R. Marshall1,2, Thomas L.P. Couvreur4,5,6, Abigail L. Summers1,2,3,
Nicolas J. Deere1,7, W.R. Quentin Luke8, Henry J. Ndangalasi9,
Sue Sparrow3, David M. Johnson10
1 CIRCLE, Environment Department, University of York, York, UK 2 Flamingo Land Ltd., Kirby Misperton,
North Yorkshire, UK 3 Biology Department, University of York, York, UK 4 Institut de Recherche pour le Développement, UMR-DIADE, BP 64501, F-34394 Montpellier cedex 5, France 5 Université de Yaoundé I, Ecole
Normale Supérieure, Département des Sciences Biologiques, Laboratoire de Botanique systématique et d’Ecologie, B.P. 047, Yaoundé, Cameroon 6 Netherlands Centre for Biodiversity Naturalis (section NHN), Biosystematics Group, Wageningen University, the Netherlands 7 Durrell Institute of Conservation and Ecology, School
of Anthropology and Conservation, University of Kent, Canterbury, UK 8 East African Herbarium, National
Museums of Kenya, Nairobi, Kenya 9 Department of Botany, University of Dar es Salaam, Dar es Salaam,
Tanzania 10 Department of Botany & Microbiology, Ohio Wesleyan University, Delaware, Ohio 43015 USA
Corresponding author: Andrew R. Marshall (andy.marshall@york.ac.uk)
Academic editor: M.A. Jaramillo | Received 30 October 2014 | Accepted 14 May 2015 | Published 1 June 2016
Citation: Marshall AR, Couvreur TLP, Summers AL, Deere NJ, Luke WRQ, Ndangalasi HJ, Sparrow S, Johnson
DM (2016) A new species in the tree genus Polyceratocarpus (Annonaceae) from the Udzungwa Mountains of Tanzania.
PhytoKeys 63: 63–76. doi: 10.3897/phytokeys.63.6262
Abstract
Polyceratocarpus askhambryan-iringae, an endemic tree species of Annonaceae from the Udzungwa Mountains of Tanzania, is described and illustrated. The new species is identified as a member of the genus
Polyceratocarpus by the combination of staminate and bisexual flowers, axillary inflorescences, subequal
outer and inner petals, and multi-seeded monocarps with pitted seeds. From P. scheffleri, with which it has
previously been confused, it differs in the longer pedicels, smaller and thinner petals, shorter bracts, and
by generally smaller, less curved monocarps that have a clear stipe and usually have fewer seeds. Because P.
askhambryan-iringae has a restricted extent of occurrence, area of occupancy, and ongoing degradation of
its forest habitat, we recommend classification of it as Endangered (EN) on the IUCN Red List.
Keywords
East Africa, Eastern Arc, endemism, Ndundulu, Polyceratocarpus
Copyright Andrew R. Marshall et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Introduction
The Eastern Arc Mountains of Kenya and Tanzania are well known for their high
levels of biodiversity and endemism across many plant and animal groups (Lovett and
Wasser 1993, Myers et al. 2000, Newmark 2002, Burgess et al. 2007, Dawson and
Gereau 2010). The 13 mountain blocs making up the chain are somewhat isolated
from one another, and each block exhibits its own unique suite of species. Many of the
forests occupying these mountains are under threat from expanding agriculture and
human population increase: an estimated 2.79 Mha of forest was lost between 1908
and 2000 (Willcock et al. 2016), with just 10% of the Eastern Arc Mountains still
forested (Platts et al. 2011). These development pressures have added urgency to the
inventory of the unique biota of this region.
Within the Eastern Arc chain, the 19,375 km2 Udzungwa Mountains form the
largest mountain bloc (Platts et al. 2011) and comprise the largest area of forest in
the Eastern Arc Mountains, totaling around 1,600 km2 (Marshall et al. 2010). The
Udzungwa Mountains have enormous biodiversity value, with the highest plant species richness, the highest endemic vertebrate species richness, and the second highest
endemic plant species richness of all mountain blocs in the region (Platts et al. 2010;
Rovero et al. 2014). The distinctive character of Udzungwa biodiversity has been recently highlighted by high-profile mammal species discoveries including the kipunji
monkey (Rungwecebus kipunji; Davenport et al. 2006) and Udzungwa elephant shrew
(Rhynchocyon udzungwensis; Rovero et al. 2008).
Species of the flowering plant family Annonaceae are prominent among the understory trees and woody climbers of the Eastern Arc Mountains. In these mountains,
there are 50 known Annonaceae species, 16 of which are endemic, including 12 out of
127 reported endemic tree species (R.E. Gereau, unpubl. data).
The genus Polyceratocarpus Engl. & Diels (Annonaceae subfamily Malmeoideae,
tribe Piptostigmateae, Chatrou et al. 2012) is distinguishable from other African genera of the family by the combination of a tree habit, percurrent tertiary veins of the
leaves, axillary (sometimes cauliflorous) inflorescences, occurrence of both staminate
and bisexual flowers, petals of the outer and inner whorls roughly equal in size and
shape, numerous monocarps with multiple seeds arranged in a single row, and pitted
seeds with spiniform ruminations (Couvreur et al. 2009, Couvreur et al. 2012). Engler
and Diels (1900) published the genus based on P. scheffleri Engler & Diels, collected
in the Usambara Mountains. Since that time seven additional species have been added
to the genus, all from western and central Africa.
Over the last 30 years a number of Polyceratocarpus specimens have been collected
from the Udzungwa Mountains to the south and west of the range of P. scheffleri. It
has become clear that these specimens differ consistently from P. scheffleri and other
congeners by a combination of vegetative, floral, and fruit characters, and they are
described here as a new species.
A new species in the tree genus Polyceratocarpus (Annonaceae)...
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Taxonomic treatment
Polyceratocarpus askhambryan-iringae A.R. Marshall & D.M. Johnson, sp. nov.
urn:lsid:ipni.org:names:77155232-1
Figs 1–3
Diagnosis. This species may be distinguished from other species of Polyceratocarpus by
the combination of glabrous non-glaucous leaves with finely reticulate to weakly scalariform tertiary venation, pedicels 15–22 mm long, broadly ovoid buds, chartaceous petals
10–17 mm long, 5 to 18 carpels/monocarps, and relatively large torulose monocarps.
Type. Marshall 2117 (holotype K; isotypes DSM, MO, NHT), Tanzania, Iringa Region: Ndundulu Forest, Kilombero Nature Reserve, Udzungwa Mountains,
07°48'S, 36°31'E (WGS84), 1490 m, 30 May 2011.
Description. Monopodial tree to 20 m tall, 4.0–25.4 cm diam.; bark smooth,
sparsely lenticellate, often with weak horizontal striations and pits on large trees,
grey-brown; branches spirally arranged on trunk, branching from half to two fifths of
the height of the main stem, perpendicular but sinuous and drooping slightly; twigs
longitudinally rugulose, inconsistently marked with small but prominent lenticels,
glabrous, brown. Leaves: petiole 4–9 mm long, 1.6–3.3 mm thick, roughened, black,
glabrous; lamina narrowly to broadly elliptic-oblong to oblanceolate, or rarely obovate, (5.0-)9.0–25.7 by (3.7-)4.9–8.6(-11.6) cm, chartaceous to coriaceous, greenish
gray in sicco, glabrous on both surfaces, base rounded and minutely subcordate, apex
acuminate with the acumen 12–20 mm long or occasionally obtuse, midvein plane to
slightly impressed above, raised below, secondary veins 9 to 17 per side, diverging at
45–60° from midrib, eucamptodromous to weakly brochidodromous, slightly raised
to slightly impressed above, raised below, tertiary veins finely reticulate to somewhat
scalariform, indistinct to slightly raised above, raised and conspicuous below. Inflorescences 1- or 2-flowered, axillary or occasionally ramiflorous, forming tubercles on
leafless growth; pedicels 15–29 mm long 1–3 mm diam., finely appressed-puberulent,
bearing a bract 0.8–1 mm long 1/4–2/5 of the distance above pedicel base. Flowers
bisexual or staminate, buds broadly ovoid; sepals 3, valvate, crescent-shaped, 2–3.5
mm long, partially connate at the base so that as corolla expands the calyx becomes
discoid to triangular with diameter of 7–8 mm, appressed-puberulent abaxially; petals in two whorls of 3, pale yellow in vivo; outer petals occasionally tinged pink on
abaxial surface in vivo, valvate, spreading horizontally and recurving at anthesis, narrowly elliptic to elliptic or oblong-elliptic, 10–16 mm long by (5-)8–11 mm wide,
coriaceous, apex obtuse, sparsely pubescent adaxially, ferruginous appressed-puberulent but becoming glabrate and verrucose abaxially; inner petals sometimes with
a pale brownish-yellow median stripe abaxially, valvate, erect at anthesis with the
apices recurved, narrowly elliptic-lanceolate, elliptic, or elliptic-oblanceolate, 11–17
mm long × 5–7 mm wide, coriaceous, external surface marked by a broad flattened
ridge that narrows from base to apex, glabrous adaxially, appressed-puberulent with
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Figure 1. Known distribution of Polyceratocarpus askhambryan-iringae in the Eastern Arc Mountains
(EAM). EAM boundary and forest cover (green) derived from Platts et al. (2011). Black boundaries
within EAM boundary show protected areas. Pale green areas in the lower maps show degraded forest
with canopy <10%.
A new species in the tree genus Polyceratocarpus (Annonaceae)...
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trichomes densest along ridge and at apex abaxially, verrucose on both surfaces, apex
acute; stamens ca. 200, 2.0–2.8 mm long, clavate, apex of connective obliquely truncate, pale brown with orange apex, glabrous?; carpels 5 to 18, oblong, 2.9–4.0(-6.0)
mm long by 0.9–1.1(-2.3) mm wide, densely pale brown/ferruginous-puberulous;
stigma bilobed, capitate, 1 mm in diam., glabrous, ovules ca. 10, uniseriate; torus
subglobose to broadly pyriform to oblate, 4.4–4.9 mm long by 2.6–5.4 mm diam.,
3–8 mm diam. at base. Pedicel of fruit 20–44 mm long by 3–7 mm diam., weakly
longitudinally rugulose, glabrate; torus of fruit ellipsoid to broadly pyriform, 7–15
mm diam. × 8–12 mm long, grey-brown. Monocarps up to 18 per fruit, green (rarely
with orange or vinaceous tinge) in vivo, dark brown when dried, weakly (to strongly)
recurved-falciform, (1.9) 6.0–8.6 cm by 0.7–2.2 cm, torulose, minutely verrucose,
glabrate or with a few scattered hairs, base sub-sessile or short-stipitate, stipe 1–11
mm long, 2–6 mm thick, apex rounded or sometimes short-beaked. Seeds 1–15 per
monocarp, 15 mm long by 13 mm wide by 10 mm thick, arranged in a single [or two
irregular?] rows, flattened-ellipsoid, pitted, with spiniform ruminations (fig. 3) and
raphe/antiraphe sunken in a circumferential groove.
Distribution. Polyceratocarpus askhambryan-iringae is endemic to the Udzungwa Mountains of Tanzania. It is known from Mwanihana Forest in the Udzungwa
Mountains National Park, Ndundulu Forest in the Kilombero Nature Reserve, and
the Uzungwa Scarp Forest Reserve (Fig. 1).
Habitat and ecology. Inhabits montane forest on brown sandy loam soils (pH
range 4–5 measured in Ndundulu Forest). Mean annual rainfall of collection localities
approximately 1500–2000 mm/yr (Marshall, Ndangalasi, unpubl. data). Thirty-eight
mature individuals were found mostly on slopes or ridge-tops at elevations 1090–
1540m. Mature flowers were collected in May, November, and December, fruits in
February and May-October.
Associated taxa recorded with Polyceratocarpus askhambryan-iringae include the
following: (1) Ndundulu Forest: Allanblackia ulugurensis Engl., Alsodeiopsis schumannii Engl.; Anisotes pubinervius (T.Anderson) Heine, Anthocleista grandiflora Gilg,
Beilschmiedia kweo (Mildbr.) Robyns & Wilczek, Bertiera pauloi Verdc., Cassipourea
gummiflua Tul.; Cassipourea malosana Alston; Chlorophytum brachystachyum Baker,
Cleistanthus polystachyus Hook.f. ex Planch., Clerodendrum cephalanthum Oliv., Coffea sp., Cola greenwayi Brenan, Cola stelechantha Brenan; Craterispermum longipedunculatum Verdc.; Diospyros abyssinica (Hiern) F.White, Drypetes gerrardii Hutch.;
Englerina sp. nov.; Garcinia buchananii Baker; Garcinia volkensii Engl.; Grewia mildbraedii Burret; Justicia rodgersii Vollesen; Lasiodiscus usambarensis Engl.; Maytenus
undata (Thunb.) Blakelock; Monodora globiflora Couvreur; Monanthotaxis schweinfurthii Engl. & Diels; Myrianthus holstii Engl., Ocotea usambarensis Engl.; Ochna
holstii Engl.; Parinari excelsa Sabine, Peddiea fischeri Engl., Pavetta nitidissima Bridson, Plectranthus leptophyllus (Baker) A.J.Paton, Rinorea angustifolia Baill. subsp. ardisiiflora (Oliv.) Grey-Wilson; Rinorea sp.; Rytigynia lichenoxenos (K.Schum.) Robyns
subsp. glabrituba Verdc.; Sclerochiton obtusisepalus C.B.Clarke; Solanecio epidendricus
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Figure 2. Polyceratocarpus askhambryan-iringae drawings of A tree architecture B fresh fruits C fresh
flower below D fresh ramiflorous flower buds E–F dry and fresh bisexual flower (one petal removed)
G fresh bisexual flower above H dried stamens I–J fresh and dry carpels lacking stigmas K dried carpel
with stigma, plus photographs of L fresh leaves M fruit and N flower. Drawings by Sue Sparrow, A by
Andrew Marshall, E and K by Andrew Brown, from the following specimens: Marshall 2070 (B); Marshall 2117 (C-E and G-L) and Luke 11279 (F). Scale bars: 20 mm unless stated.
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(Mattf.) C.Jeffrey; Strombosia scheffleri Engl.; Strychnos mellodora S.Moore; Strychnos
mitis S.Moore; Strychnos sp. nov; Syzygium guineense DC.; Tabernaemontana stapfiana
Britten; Tarenna pavettoides (Harv.) Sim; Thalictrum rhynchocarpum Quart.-Dill. &
A.Rich.; Uvariopsis lovettiana Couvreur & Q.Luke; Vepris stolzii I.Verd.; Vernonia
calvoana Engl. subsp. leucocalyx (O.Hoffm.) C.Jeffrey; Vernonia luhomeroensis Q.Luke
& Beentje; Vernonia sp. nr. pteropoda Oliv. & Hiern; Warneckea sp. nov.; Xymalos
monospora Baill.; Zanthoxylum paracanthum (Mildbr.) Kokwaro; Zehneria sp. nr. oligosperma C.Jeffrey. (2) Uzungwa Scarp Forest Reserve: Cassipourea gummiflua Tul;
Cleistanthus polystachyus Hook.f. ex Planch.; Craterispermum longipedunculatum Verdc.; Diospyros uzungwaensis Frim.-Møll. & H.J.Ndangalasi; Drypetes gerrardii Hutch.;
Lasiodiscus usambarensis Engl.; Psychotria megalopus Verdc.; Tabernaemontana stapfiana Britten; Tarenna uzungwaensis Bridson. (3) Mwanihana: Acalypha psilostachya
Hochst. ex A.Rich. var. psilostachya; Anisotes pubinervius (T.Anderson) Heine; Caloncoba welwitschii Gilg; Chrysophyllum gorungosanum Engl.; Coffea mufindiensis Hutch.
ex Bridson subsp. mufindiensis; Dorstenia sp. aff tenuiradiata Mildbr.; Isoglossa lactea
Lindau ex Engl. subsp. lactea; Isolona linearis Couvreur; Newtonia buchananii (Baker)
G.C.C.Gilbert & Boutique; Ochna holstii Engl.; Parinari excelsa Sabine; Phyllopentas
ulugurica (Verdc.) Kårehed & B. Bremer; Kedrostis sp.; Polystachya sp aff. canaliculata
Summerh.; Raphidiocystis chrysocoma (Schumach.) C.Jeffrey; Selaginella kraussiana
(Kunze) A.Braun; Stellaria mannii Hook.f.; Tricalysia aciculiflora Robbr.; Uvariopsis
lovettiana Couvreur & Q.Luke; Vepris nobilis (Delile) Mziray; Zanthoxylum paracanthum (Mildbr.) Kokwaro.
Additional specimens examined. TANZANIA. Iringa Region, Kilolo District:
east Udzungwa National Park, forest south of Mwanihana hill c. 2 km S of last camping site of Mwanihana trail, 1400 m, 07°48'S, 36°49'E, Couvreur 101 (DSM, OWU,
WAG); Mwanihana Forest above Sanje village, 1220 m, no grid reference, Lovett 222
(K); Udzungwa Mountains National Park, 1200 m, 07°48'S, 36°49'E, Luke 7738
(EA, K); Udzungwa Mountains National Park, 1440 m, 07°42'S, 36°52'E, Luke
11279 (EA, NHT, MO, K); Ndundulu FR, Camp 589-Camp 590, 07°47'S, 36°29'E,
1440 m, Luke et al. 10366 (MO); Kilombero Nature Reserve, Ndundulu Forest, 1540
m, 07°48'S, 36°31'E (WGS84), Marshall 2036, 2070, (NHT, MO, K); Uzungwa
Scarp Forest Reserve, Uluti, 1534 m, 08°14'S, 36°01'E, Ndangalasi HJN 392 (DSM,
OWU); Uzungwa Scarp Forest Reserve, Ilutila, 1709 m, 08°13'S, 36°01'E, Ndangalasi
393 (DSM, OWU); Udzungwa, Kilombero FR, W of Ruaha River, 1700 m, Rogers
& Hall 2300 (K); Mwanihana Forest above Sanje village, 1400 m, 07°50'S, 36°49'E,
Thomas 3656 (MO); Mwanihana Forest above Sanje village, 1400 m, 07°50'S,
36°49'E, Thomas 3698 (MO, WAG). Morogoro Region, Kilombero District: SonjoMwanihana trail, 1090 m, 07°48'S, 36°51'E, Luke 5051 (EA, K).
An additional specimen from Iringa Region (Nyambanitu Forest, Ede 65, K), may
also represent this species but bears only an old fruit pedicel lacking monocarps. Further potential Polyceratocarpus collections from Iringa Region (Lulanda Forest Reserve:
Gereau 2651, 2664, 2665, MO; Lovett 2256, MO, WAG; Luke & Luke 12779, EA
& K), were identified as neither P. askhambryan-iringae nor P. scheffleri, while another
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Figure 3. SEM photograph of Polyceratocarpus askhambryan-iringae dried seed cross-section, showing
spiniform ruminations.
from Morogoro Region was not considered to be from this genus at all (Kimboza Forest Reserve: Parry 1816, TFD; cited Verdcourt 1971).
Additional field notes. Slash dry, slightly stringy, pale yellow (to pale peach), occasionally streaked yellow-brown, dark brown at outer edge formed by the colour of
the inner bark, potpourri aroma. Leaf lamina dark green above, mid-green with greyish
tinge below, turning greenish-grey when dried, new flush pinkish; petiole initially pale
green in vivo, becoming roughened grey-brown with age; midrib yellow-green above
and below in vivo. Flower buds broadly ovoid, green or pale brown-yellow with occasional pink tinge at apex in vivo; sepals yellowish-green in vivo.
Etymology. This new species of Polyceratocarpus was named by Askham Bryan
College and Iringa International School as part of a rainforest education program.
Conservation status. Our IUCN Red List assessment for P. askhambryan-iringae
was based on “area of occupancy” (AOO), “extent of occurrence” (EOO; IUCN 2012)
and the level of threat. The 38 observed mature P. askhambryan-iringae stems were
found at four collection localities, with only 112 km between the most distant individuals. The four collection localities represented three different levels of governmental protection, including Forest Reserve (Uzungwa Scarp), Nature Reserve (Kilombero, KNR; and also Uzungwa Scarp proposed status) and National Park (Udzungwa
Mountains; UMNP), with only UMNP having the maximum level of protection
under Tanzanian law (IUCN category II; Dudley 2008). At the time of survey, the
greatest threat to P. askhambryan-iringae was widespread timber-felling observed in
Uzungwa Scarp Forest Reserve. Conversely, threats to P. askhambryan-iringae in KNR
and UMNP were much lower, both comprising only very occasional removal of polesized trees by villagers, and potential damage from an increasing elephant population
(Marshall et al. 2012). KNR was further threatened by a lack of formal ranger patrols
and rapid population expansion in the nearest villages. Given that the sources of threat
were closely associated with the three protected areas, for the purpose of Red List
assessment we considered three rather than four threat-defined “locations” (IUCN
2012). In calculating AOO, we used 10 km2 grid cells so that the projected area was
A new species in the tree genus Polyceratocarpus (Annonaceae)...
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not extrapolated far beyond the expected habitat tolerance of the species. We estimated
an AOO of 300 km2 and an EOO of 1,410 km2, with EOO, AOO, habitat quality and
the number of mature individuals, all presumed declining as a result of timber-felling
in USFR. Given this continuing decline, plus an EOO of less than 5,000 km2, an
AOO of less than 500 km2, and a population at no more than five locations, the species qualified firmly as endangered on the IUCN Red List (IUCN 2012), EN B1ab(i,
ii,iii,v)+2ab(i,ii,iii,v). Within the Udzungwa Mountains, closed-canopy forest was not
extensive at elevations suitable for P. askhambryan-iringae, and hence we expect that
future expeditions will not expand the AOO or EOO of P. askhambryan-iringae above
the IUCN endangered threshold. However, the Rubeho and Mahenge mountain blocs
adjacent to Udzungwa were more poorly known, and may contain suitable habitat in
which this species might also be found.
Discussion
A) Systematics
Polyceratocarpus askhambryan-iringae is similar to P. scheffleri and has previously been
confused with it. As far as is known, however, P. scheffleri is confined to the Usambara
Mountains and P. askhambryan-iringae to the Udzungwa Mountains; previous reports
of P. scheffleri from the Udzungwa Mountains (e. g. Lovett et al. 1988, Couvreur et
al. 2006, Eastern Arc Mountains & Coastal Forests CEPF Plant Assessment Project
2009) have been based on misidentifications of P. askhambryan-iringae. To facilitate
separation of the two East African species, their differences are contrasted in the following key:
1
–
Lamina rounded and minutely subcordate at base, with tertiary veins reticulate to weakly scalariform; pedicel 15–29 mm long, bearing a bract 0.8–1 mm
long; sepals connate only at base, distinct, crescent-shaped; petals 10–17 mm
long; monocarps up to 18 per fruit, (1.9–) 6–8.6 cm long, weakly recurvedfalciform, short-stipitate (1–11 × 2–6 mm); seeds per monocarp up to 15 ....
......................................................Polyceratocarpus askhambryan-iringae
Lamina cuneate to broadly cuneate at base, with tertiary veins distinctly scalariform; pedicel 10–16 mm long, bearing a bract 7–9 mm long; sepals connate into a disc with indistinguishable sepal lobes; petals 18–32 mm long;
monocarps 5 to 33, 6–20 cm long, strongly curved, sessile; seeds per monocarp up to ca. 25 ................................................ Polyceratocarpus scheffleri
The genus Polyceratocarpus outside East Africa has a Guineo-Congolean distribution, with the other seven species scattered from Côte d’Ivoire to northern Angola and
the Democratic Republic of the Congo. Polyceratocarpus askhambryan-iringae and P.
scheffleri both differ from most of the more western species in having larger numbers of
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carpels and large strongly torulose monocarps. The only other species with such a high
number of carpels (18 to 20) is Polyceratocarpus laurifolius Paiva from northern Angola,
but that species differs from P. askhambryan-iringae in having densely pubescent twigs,
a distinctly cuneate base to the leaf, a larger bract on the pedicel, and larger sepals.
Polyceratocarpus laurifolius also differs from P. askhambryan-iringae in having larger
petals of the staminate flowers than of the bisexual ones (Paiva 1966); in P. askhambryan-iringae no petal dimorphism between staminate and bisexual flowers was seen.
Polyceratocarpus askhambryan-iringae (as “Polyceratocarpus sp.”) was one of four
Polyceratocarpus species included in the phylogenetic analysis of Couvreur et al. (2009)
focused on the phylogeny of several closely related African genera of Annonaceae.
In this analysis P. askhambryan-iringae appeared as sister to the other three species
of Polyceratocarpus sampled (P. microtrichus (Engl. & Diels) Ghesq. & Pellegr., P.
parviflorus (Baker f.) Ghesq., and P. pellegrinii Le Thomas) forming a monophyletic
group with strong bootstrap and posterior probability support. The other five species
remain to be sampled. The genus itself, however, was nested within the western and
central African genus Piptostigma Oliv., to which it is morphologically dissimilar, so
additional sampling is needed.
B) Regional endemism and biodiversity
East Africa is an area of both high endemism and high diversity for Annonaceae, with 28
genera and 85 species known from Tanzania alone (Couvreur et al. 2006). In particular,
the Eastern Arc Mountains form an area of high species endemism for East African Annonaceae. Furthermore, many Annonaceae genera represented in these mountains have
main areas of diversity in the Guineo-Congolean region and are represented in the Eastern Arc Mountains by endemic taxa. In addition to Polyceratocarpus, the genera Annickia
Setten & Maas, Greenwayodendron Verdc., Isolona Engl., Monodora Dunal, Uvariodendron (Engl. & Diels) R.E. FR., and Uvariopsis Engl. all follow this pattern (Verdcourt
1971, 1986, Couvreur et al. 2006, Couvreur 2009, Couvreur and Luke 2010).
Polyceratocarpus askhambryan-iringae also adds to the growing list of species unique
to the Udzungwa bloc, including various recently discovered plants (e.g. Luke and
Beentje 2003; Knox et al. 2004). There are now 71 known endemic plant species from
the Udzungwa Mountains (comprising 15 trees, including P. askhambryan-iringae);
among the Eastern Arc Mountains this is second only to the Uluguru Mountains (86
endemic species [14 trees]; R.E. Gereau, unpubl. data). P. askhambryan-iringae is the
second large tree (≥20m) and the third endemic Annonaceae species to be described
from the Udzungwa Mountains over the last 20 years, following the respective discoveries of Omphalocarpum strombocarpum Y.B.Harv. & Lovett (Harvey and Lovett
1998), Toussaintia patriciae Q.Luke & Deroin (Deroin and Luke 2005) and Monodora
globiflora Couvreur (Couvreur et al. 2006). Given further new species descriptions in
preparation and the larger size of the Udzungwa Mountains, it is likely to be the most
important Eastern Arc Mountain bloc for tree species endemism. More plant species
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73
are known from the Udzungwa Mountains than other Eastern Arc Mountain blocs, and
while this is partly due to increased survey effort over other blocs, recent projections by
distribution models estimate that the number of endemic/threatened plant taxa will not
be superseded following increased exploration of other blocs (Platts et al. 2010).
The four P. askhambryan-iringae localities are coextensive with other recent discoveries of rare and endemic Eastern Arc animals, e.g. the Critically Endangered
endemic Sanje mangabey (Cercocebus sanjei; Mwanihana and Uzungwa Scarp), the
Critically Endangered kipunji monkey (Rungwecebus kipunji; Ndundulu; also found
on Rungwe Mountain), the endemic Udzungwa forest partridge (Xenoperdix udzungwensis; Ndundulu), the endemic rufous-winged sunbird (Nectarinia rufipennis;
Mwanihana, Ndundulu and Uzungwa Scarp), the endemic Udzungwa elephant shrew
(Rhynchocyon udzungwensis; Ndundulu and Mwanihana), further emphasizing the exceptional local biodiversity value. The exceptional biodiversity of these areas led to the
incorporation of Mwanihana forest into the Udzungwa Mountains National Park in
1991, and later incorporation of Ndundulu forest into the Kilombero Nature Reserve
in 2007 (Marshall et al. 2007).
The discovery of P. askhambryan-iringae further highlights the need for improved
conservation of Uzungwa Scarp Forest Reserve, one of the most important forests in
the region for primates and birds (Dinesen et al. 2001), where herpetofauna endemism
has been estimated at eight times that of the Eastern Arc as a whole (Menegon, unpublished data). Uzungwa Scarp FR is home to the Kihansi spray toad (Nectophrynoides
asperginis), which was extinct in the wild, but was successfully reintroduced in October
2012 following zoo conservation breeding (Channing et al. 2009; Gereau et al. 2014).
We hope that our discovery of yet another new endemic will add weight to ongoing
proposals for gazettement of Uzungwa Scarp Forest Reserve as a Nature Reserve, the
highest designation of protected area possible under the Tanzania Forestry Service.
Acknowledgements
Thanks to COSTECH and TAWIRI for permits. The collection expedition to Ndundulu Forest was primarily funded by Flamingo Land Ltd., through the Udzungwa Forest Project. Preliminary surveys of Ndundulu were funded by HMJ Design and Mazuri
Foods Ltd., through the British and Irish Association of Zoos and Aquaria Mammal
Working Group, and by the Wildlife Conservation Society. Collections from the Uzungwa Scarp Forest Reserve were funded by Danish International Development Agency
(DANIDA) through the ENRECA Project at the Department of Botany, University of
Dar es Salaam. Thanks to National Museums of Kenya staff especially John Kimeu, for
herbarium support. Roy Gereau (Missouri Botanical Garden) is thanked for comments
on the manuscript. Nancy Murray (Ohio Wesleyan University) for helpful comments
on the manuscript. Phil Roberts (University of York) is thanked for administrating the
production of illustrations, and Andrew Brown for producing two of the sketches. We
also thank the many field assistants who helped with collections, and with expedition
74
Andrew R. Marshall et al. / PhytoKeys 63: 63–76 (2016)
logistics, especially Richard Mgata from Udekwa village, who climbed tall trees in pursuit of vital material for this description. TLPC thanks the Netherlands Organization
for Scientific Research (NWO), Hugo de Vries Fonds and Air France-KLM for financial
help in Tanzania in 2006. Frank Mbago is thanked for assistance while in the field with
TLPC. We also thank the United Bank of Carbon for funding to develop a schools’
rainforest education program around this species, which led to the chosen species name.
And last but not least we thank all the participating schools, colleges, youth groups and
people who helped the success of this program, particularly Katie Boswell, Amber Hall,
Joshua Sammy, Sharon Sheppard and the First Sherburn Rainbows and Brownies.
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