A synopsis of the Afro-Malagasy species previously included in Schefflera
(Araliaceae): resurrection of the genera Astropanax and Neocussonia
Author(s): Porter P. Lowry II, Gregory M. Plunkett, Morgan R. Gostel & David G. Frodin
Source: Candollea, 72(2):265-282.
Published By: The Conservatory and Botanical Garden of the City of Geneva (CJBG)
https://doi.org/10.15553/c2017v722a4
URL: http://www.bioone.org/doi/full/10.15553/c2017v722a4
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A synopsis of the Afro-Malagasy species
previously included in Schefflera
(Araliaceae): resurrection of the genera
Astropanax and Neocussonia
Porter P. Lowry II, Gregory M. Plunkett, Morgan R. Gostel & David G. Frodin
Abstract
LOWRY, P.P., II, G.M. PLUNKETT, M.R. GOSTEL & D.G. FRODIN (2017). A synopsis of the Afro-Malagasy species previously included
in Schelera (Araliaceae) : resurrection of the genera Astropanax and Neocussonia. Candollea 72 : 265-282. In English, English abstract. DOI:
http://dx.doi.org/10.15553/c2017v722a4
Phylogenetic studies have shown that Schelera J.R. Forst. & G. Forst., the largest genus of Araliaceae, is grossly polyphyletic, representing ive distinct clades within the family, and that one of these clades, comprising species from continental
Africa, Madagascar, the Comoro Islands and the Seychelles, contains two morphologically distinct subclades. In an efort
to establish monophyletic genera among the elements that have been treated in Schelera over the last several decades,
we resurrect the genera Astropanax Seem. (15 species) and Neocussonia (Harms) Hutch. (16 species) to encompass the
members of the two “Afro-Malagasy” subclades. In this synoptical revision, we provide updated generic descriptions
along with a key to the genera, and make 26 new combinations. A lectotype is designated for each genus, a neotype is
provided for one accepted species name, and lectotypes are designated for 16 accepted species names and 12 heterotypic
synonyms. For each accepted species, full synonymy is provided along with complete citation of type specimens and an
indication of habitat and geographic range.
Keywords
ARALIACEAE – Astropanax – Neocussonia – Schelera – Taxonomy – Nomenclature
Addresses of the authors :
PPL : Missouri Botanical Garden, P.O. Box 299, St. Louis, MO, 63166-0299, U.S.A. and Institut de systématique, évolution, et biodiversité (ISYEB) (UMR
7205 – CNRS MNHN UPMC EPHE), Muséum national d’Histoire naturelle, Sorbonne Universités, C.P. 39, rue Cuvier 57, 75231 Paris CEDEX 05, France.
E-mail : pete.lowry@mobot.org
GMP : Cullman Program for Molecular Systematics, New York Botanical Garden, 2900 Southern Blvd., Bronx, NY, 10458-5126, U.S.A.
MRG : Department of Botany, National Museum of Natural History, MRC 166, Smithsonian Institution, Washington, DC 20013-7012, U.S.A.
DGF : Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom.
Submitted on September 14, 2016. Accepted on May 5, 2017.
ISSN : 0373-2967 – Online ISSN : 2235-3658 – Candollea 72(2) : 265-282 (2017)
First published online on June 24, 2017
© CONSERVATOIRE ET JARDIN BOTANIQUES DE GENÈVE 2017
266 – Synopsis of Astropanax and Neocussonia (Araliaceae)
Introduction
he species of Schelera J.R. Forst. & G. Forst. are most commonly small trees or arborescent hemi-epiphytes, typically
contributing to the understory vegetation of humid forests
throughout the tropics of both the Old and New Worlds.
Within Araliaceae, they have a coherent morphology combining the presence of palmately compound leaves with the
absence of prickles, but the simplicity of this generic deinition
belies a very complex taxonomic history (reviewed in Plunkett et al., 2005) in which no fewer than 14 other generic
names were subsumed under Schelera, largely through the
studies of Frodin (e.g., 1975, 1986, 1989, 1993, 1995), as
well as Philipson (1979, 1995) and Lowry (1989). More
recently, the advent of phylogenetic analyses using molecular
characters revealed that Schelera, as broadly circumscribed,
was polyphyletic, representing ive major clades in Araliaceae
(Plunkett et al., 2005), most only distantly related to one
another. hese ive clades did not correspond to groups recognized in any of the various taxonomic systems published
over the decades for the family, but they did align well with
the system of subgeneric groupings proposed by Frodin (see
Plunkett et al., 2005 ; later updated by Frodin et al., 2010),
which were largely based on a combination of geography
and a suite of morphological characters. he importance of
geography was also relected in the informal names for four of
the ive major clades : Neotropical Schelera, Asian Schelera,
Melanesian Schelera, and Afro-Malagasy Schelera, and even
the ifth, Schelera sensu stricto (which includes the generitype), is geographically coherent, although it overlaps in part
with Melanesian Schelera in the Paciic islands. Plunkett et
al. (2005) recommended that the three clades with the smallest
number of species, the Melanesian and Afro-Malagasy groups
along with Schelera sensu stricto, be treated as immediate
priorities for taxonomic recircumscription. Towards this end,
the phylogeny of Melanesian Schelera was evaluated using
a nearly comprehensive sample of taxa (47/49 spp. ; Plunkett & Lowry, 2012), and a subsequent synoptic revision
treated these species in six subgenera of a newly reinstated
and expanded genus Plerandra A. Gray (Lowry et al., 2013).
In parallel, Gostel et al. (2017) completed a phylogenetic
reconstruction of the Afro-Malagasy clade, paving the way
for the present study, in which we present a synopsis of the
taxonomic implications resulting from our recent indings.
Afro-Malagasy Schelera is one of several clades forming
a large polytomy at the base of the phylogenetic tree of
Araliaceae (Plunkett et al., 2005). It currently comprises
31 accepted species (Gostel et al., 2017) and, as the name
implies, these taxa are all distributed in (sub-Saharan) continental Africa and Madagascar, as well as the nearby Comoro
and Seychelles archipelagos. Seemann (1865) was the irst to
describe African species from this clade, placing them in two
genera, Astropanax Seem. and Sciodaphyllum Seem. Harms
Candollea 72, 2017
(1894-97), however, subsequently assigned all palmatelyleaved Araliaceae from Africa to Schelera. From this time
until the third quarter of the 20th Century, the diversity and
taxonomic status of species in Schelera recognized from continental Africa and Madagascar remained poorly understood,
until the eforts of Bernardi (1969, 1973) and Bamps (1974),
who provided major contributions to our understanding of
the genus from these two regions, respectively. Despite the
non-monophyly of the infrageneric groups recognized by
Bernardi (1969 ; see Gostel et al., 2017), his work remains
an important source for species delimitations, particularly
for Malagasy species. In total, Bernardi (1966, 1969, 1973)
described six new species and three new varieties of Schelera
from Madagascar and the Comoro islands, and transferred
six additional species and one variety from Cussonia hunb.
to Schelera. Bamps (1974) described two additional species
from continental Africa, bringing the total number currently
recognized from the Afro-Malagasy region to the present 31
species. Subsequent ield and herbarium studies have revealed
19 additional species that are currently being described
(Lowry et al., unpubl. data).
he recent molecular phylogenetic studies of Gostel et
al. (2017) have conirmed the monophyly of Afro-Malagasy
Schelera and provide evidence for two well supported clades,
one of which corresponds closely to Frodin’s infrageneric
group “Meiopanax” and the other to part of his broad ‘Sciodaphyllum’ group, as circumscribed in Plunkett et al. (2005)
and Frodin et al., (2010). Because the type of Schefflera
(S. digitata J.R. Forst. & G. Forst.) belongs to another clade,
all species of Afro-Malagasy Schelera must be transferred
to one or more other genus. Our phylogenetic indings lend
themselves to one of two obvious alternatives, treating the
entire clade as a single genus or recognizing each of its two
major subclades as distinct genera. In considering the overall
task of recircumscribing the > 600 currently accepted species
of Schelera that fall outside the small type clade (Schelera
s.s.), Lowry et al. (2013) suggested that decisions should be
based on three criteria : monophyly, morphological diagnosability, and geographic coherence. his approach was applied
to the synoptical revision of Melanesian Schelera, now treated
as Plerandra. hat clade comprises six subclades, each morphologically distinct (satisfying two of the three criteria for
recognition as genera), but they overlapped considerably in
their geographies, which led to the decision to treat them as
subgenera. A similar approach was used for the second largest
genus of Araliaceae, Polyscias J.R. Forst. & G. Forst. (Lowry
& Plunkett, 2010). Applying these same criteria to AfroMalagasy Schelera, the “Sciodaphyllum” and “Meiopanax”
subclades seem better treated as distinct genera. Not only
are these groups reciprocally monophyletic, but they are also
easily distinguished based on lower and leaf characters (see
below). Moreover, with only a few exceptions, these clades
Candollea 72, 2017
are geographically coherent. Only three of the 13 species
of the continental-African “Sciodaphyllum” subclade
[S. humblotiana Drake, S. monophylla (Baker) Bernardi, and
S. myriantha (Baker) Drake] occur in Madagascar, and one
[S. procumbens (Hemsl.) F. Friedmann] is restricted to the
Seychelles, while only two of the 17 species from the Malagasy “Meiopanax” subclade [S. lukwangulensis (Tennant)
Bernardi and S. umbellifera (Sond.) Baill.] are present
in continental Africa. Unfortunately, neither of the two
informal names applied by Frodin to these groups is allowable under the Code of Nomenclature (McNeill et al.,
2012). In the case of ‘Meiopanax’, an older available name,
Neocussonia (Harms) Hutch., has priority at the generic
level. For ‘Sciodaphyllum’, the problem is more complex.
Frodin envisioned “Sciodaphyllum” as an unspecialized
group (perhaps approaching the ancestral condition of
Schefflera s.l.) that is broadly distributed across Asia, the
Neotropics, and Africa. This group, however, proved to be
polyphyletic in light of phylogenetic studies (Plunkett
et al., 2005). Moreover, the type of the genus Sciodaphyl
lum (S. brownei Spreng., from Jamaica) belongs to the
Neotropical Schefflera clade (Fiaschi & Plunkett, 2011 ;
Plunkett et al., unpubl. data). Therefore, among the
Afro-Malagasy elements of “Sciodaphyllum”, the oldest
available generic name is Astropanax Seem.
he synopsis presented here thus reinstates the generic
names Astropanax and Neocussonia. Of the 31 currently
accepted species of Afro-Malagasy Schelera, 15 are placed in
Astropanax and 16 in Neocussonia, necessitating 12 and 14 new
combinations, respectively. Information on the species from
Madagascar and continental African is available on-line in the
“Catalogue of the Vascular Plants of Madagascar” (Madagascar Catalogue, 2017) and “African Plant Database” (APD,
2017), respectively. An identiication key is provided below to
distinguish Astropanax and Neocussonia.
Systematics
Key to the genera Astropanax and Neocussonia
1. Ovary 2-5-carpellate, styles 2-5 ; leaves palmately compound or unifoliolate, apex of lealets generally truncate
to emarginate, sometimes acute (N. bojeri and N. staufe
riana) or rounded (N. halleana), generally with an evident
retuse notch and almost always with a small mucro or boss
formed by an extension of the midvein ; terrestrial trees ;
Madagascar, except N. lukwangulensis and N. umbellifera in
continental Africa ..........................................Neocussonia
1a. Ovary 5-9-carpellate, styles 5-9 ; leaves palmately compound (generally unifoliolate in A. monophyllus), apex
of leaflets acute to acuminate or caudate (rounded in
A. volkensii and sometimes in A. monophyllus), rarely if ever
Synopsis of Astropanax and Neocussonia (Araliaceae) - 267
with a retuse notch, lacking a small mucro or boss ; terrestrial or hemi-epiphytic trees, or lianas ; continental Africa,
except A. humblotianus, A. monophyllus and A. myrianthus in
Madagascar, and A. procumbens in the Seychelles ..............
........................................................................ Astropanax
A. Astropanax Seem. in J. Bot. 3 : 174, 176. 1865 (Fig. 1).
Lectotypus (designated here) : Astropanax barteri Seem.
= Geopanax Hemsl. in Hooker’s Icon. Pl. 29 : tab. 2821.
1909. Typus : Geopanax procumbens Hemsl. ([ Astro
panax procumbens (Hemsl.) Lowry, G.M. Plunkett,
Gostel & Frodin).
Hermaphroditic, unarmed, glabrous (occasionally with
indument on twigs and young inflorescence branches),
terrestrial, hemi-epiphytic, evergreen trees or rarely lianas.
Stems sparsely to well branched or plants monocaulous.
Leaves alternate, palmately compound or rarely unifoliolate
(A. monophyllus) ; petioles clasping at the base with short to
long ligulate stipules ; lealets lanceolate, ovate, elliptical,
oblanceolate, or obovate, coriaceous or subcoriaceous to
membranaceous, margins entire, crenate, toothed or incised,
apex acute to acuminate or caudate (rounded in A. volkensii
and sometimes in A. monophyllus). Inflorescences terminal,
erect, paniculate or compound-umbellate, the ultimate
units racemose, spicate, umbellate or capitate ; inlorescence
axes and peduncles subtended by thin bracts (bracts rarely
lacking) ; lowers subtended by bracteoles (or lacking) ; pedicels slender to stout, unarticulated (lowers sessile in spicate
and capitate ultimate units). Ovary inferior, 5-9-carpellate,
carpels unilocular, placentation apical ; styles 5-9, free or
partially or mostly united basally to form a stylopodium or
a distinct column. Fruits drupaceous, the disc depressedconcave to lat or short-conical ; mesocarp leshy, endocarp
scleriied around each locule, forming a pyrene.
Diversity and distribution. – Astropanax comprises 15
currently recognized species, 11 of which occur in tropical
and subtropical Africa, three in Madagascar and one in the
Seychelles, along with three as yet undescribed species in
Madagascar and possibly one more from Africa. In Africa, the
genus extends from Guinea Conakry, Liberia and Sierra Leone
in the west, through the Gulf of Guinea Islands and across
Central Africa to East Africa, northward to Ethiopia, Somalia
and Sudan, and southward to Southern Africa (Mozambique,
South Africa and Swaziland and Zimbabwe). he three Malagasy species of Astropanax occur in humid forests, extending
from the far north to the extreme southeastern part of the
island, and the single member of the group in the Seychelles
is restricted to highly degraded humid forest on the islands of
Silhouette and Mahé. Some members of the genus are fairly
268 – Synopsis of Astropanax and Neocussonia (Araliaceae)
widespread, such as A. abyssinicus and A. barteri in Africa
(Bamps 1974b ; APD, 2017) and A. myrianthus in Madagascar
(Madagascar Catalogue, 2017), whereas several others have
much more restricted geographic ranges, including A. evrardii,
A. kivuensis, A stolzii and A. tessmannii in Africa, A. humblo
tianus in Madagascar, and A. procumbens in the Seychelles.
Notes. – Seemann (1865) included five species in his
new genus (A. abyssinicus, A. baikiei, A. barteri, A. elatus, and
A. mannii), the second and fourth of which have been placed
in synonymy by most recent authors (cf. Bamps 1974a, 1974b ;
Frodin & Govaerts, 2004). Our recent molecular phylogenetic studies (Gostel et al., 2017) include samples of S. barteri
and S. mannii and place both within a clade that corresponds
to Astropanax.
We have selected as the lectotype of the genus Astropanax
barteri [Barter s.n., K!] because it is the best known and most
widely distributed of the three accepted species originally
assigned to the genus by Seemann (1865).
A1. Astropanax abyssinicus (Hochst. ex A. Rich.) Seem. in J.
Bot. 3 : 177. 1865.
[
Aralia abyssinica Hochst. ex A. Rich., Tent. Fl. Abyss. 1 :
336. 1848. [ Schelera abyssinica (Hochst. ex A. Rich.)
Harms in Engl. & Prantl, Nat. Planzenfam. III(8) :
38. 1894. [ Sciodaphyllum abyssinicum (Hochst. ex
A. Rich.) Miq. in Ann. Mus. Bot. Lugduno-Batavi
1 : 26. 1863. [ Heptapleurum abyssinicum (Hochst. ex
A. Rich.) Benth. & Hook. f. ex Vatke in Linnaea 40 :
191. 1876.
Lectotypus (designated here) : ETHIOPIA. Prov. Tigray : Mt.
Scholoda, 11.VI.1837, fr., Schimper 283 (P [P00466317]! ;
isolecto- : BM [BM000902775, BM000902776]!, BR
[BR08860958]!, FI!, HAL [HAL0117394] image
seen, HOH [HOH-009899, HOH-009890] images
seen, JE [ JE00000511] image seen, K [K000350399]!,
L [L0008492, L0008493, L0008494, L0008495]!, M
[M00095801, M00104947, M00104948] images seen,
MPU [MPU023965, MPU023966] images seen, NY
[NY00274565]!, OXF!, P [P00466316, P00466318]!, TUB
[TUB003050, TUB003051, TUB003052, TUB003053]
images seen, WAG [WAG00000241] image seen).
= Paratropia elata Hook. f. in J. Linn. Soc., Bot. 7 : 196.
1864. [ Astropanax elatus (Hook. f.) Seem. in J. Bot. 3 :
177. 1865. [ Sciodaphyllum elatum (Hook. f.) Seem. in
J. Bot. 3 : 267. 1865. [ Heptapleurum elatum (Hook.f.)
Hiern in Fl. Trop. Afr. 3 : 30. 1877. [ Schelera hookeri
ana Harms in Engl. & Prantl, Nat. Planzenfam. III(8) :
38. 1894 [non S. elata (Buch.-Ham.) Harms]. Lectotypus (designated here) : CAMEROON. Southwest
Candollea 72, 2017
Region : Mt. Cameroon, II.1862, bud, Mann 1181
(K [K00350377]! ; isolecto- : K [K00350376,
K00350378]!).
= Schefflera acutifoliolata De Wild. in Rev. Zool. Bot.
Africaines 8 : 14. 1920. Lectotypus (designated here) :
D EMOCRATIC R EPUBLIC OF THE C ONGO . Prov.
Nord-Kivu : Ruwenzori, Lamia, ± 2000 m, s.d., fl.,
Bequaert 4310 (BR [BR00886094]! ; isolecto- : BR
[BR00886078]!).
Diversity and distribution. – Astropanax abyssinicus occurs in
mid- to high-altitude humid forest, gallery forest, and forestgrassland transition areas, from 1,450 to 2,800 m elevation,
and ranges from Ethiopia south to Zambia and Malawi in
eastern Africa, as well as in disjunct populations well to the
west in Cameroon (Bamps, 1974b ; APD, 2017).
Notes. – Aralia abyssinica was based on three specimens at
P, one of which has an original label in Richard’s handwriting
and has therefore been chosen as the lectotype.
Among the three specimens at Kew on which Hooker
based Paratropia elata, two bear original labels, one of which
has a leaf and the other a full inlorescence in bud. We have
selected the second of these as the lectotype.
Schelera acutifoliolata was based on two specimens at BR,
one of which is more complete, comprising an inlorescence
and leaf, and has therefore been selected as the lectotype.
A2. Astropanax barteri Seem. in J. Bot. 3 : 177. 1865.
[
Sciodaphyllum barteri (Seem.) Seem. in J. Bot. 3 : 267.
1865. [ Heptapleurum barteri (Seem.) Hiern in Fl.
Trop. Afr. 3 : 20. 1877. [ Schefflera barteri (Seem.)
Harms in Engl. & Prantl, Nat. Planzenfam. III(8) :
38. 1894.
Typus : S IERRA L EONE . Western Prov. : Sugar Loaf
Mountains, fr., Barter s.n. (holo- : K [K00350373]!).
= Astropanax baikiei Seem. in J. Bot. 3 : 177. 1865. [ Scio
daphyllum baikiei (Seem.) Seem. in J. Bot. 3 : 267. 1865.
[ Heptapleurum baikiei (Seem.) Hiern in Fl. Trop.
Afr. 3 : 30. 1877. [ Schelera baikiei (Seem.) Harms
in Engl. & Prantl, Nat. Planzenfam. III(8) : 38. 1894.
Typus : NIGERIA : sine loc., fr., Barter 1851 (holo- : K
[K00350375]!).
= Schefflera hierniana Harms in Engl. & Prantl, Nat.
Pflanzenfam. III(8) : 38. 1894. [ Heptapleurum
scandens Hiern in Fl. Trop. Afr. 3 : 30.1877 [nom.
illeg.] [non H. scandens (Blume) Seem.]. Lectotypus
(designated here) : CAMEROON. Southwest Region :
Mt. Cameroon, bud & fr., Mann 1180 (K [K00350379]! ;
isolecto- : K [00350380]!).
Candollea 72, 2017
Synopsis of Astropanax and Neocussonia (Araliaceae) - 269
A
B
C
D
E
F
Fig. 1. – Photographs of Astropanax Seem. A. Astropanax monophyllus (Baker) Lowry, G.M. Plunkett, Gostel & Frodin ; B. Astropanax myrianthus (Baker)
Lowry, G.M. Plunkett, Gostel & Frodin ; C. Astropanax barteri Seem. ; D. Astropanax mannii (Hook.f.) Seem. ; E. Astropanax stolzii (Harms) Lowry, G.M.
Plunkett, Gostel & Frodin ; F. Astropanax volkensii (Harms) Lowry, G.M. Plunkett, Gostel & Frodin.
[A : Ravelonarivo 4270 ; B : Antilahimena 4680 ; C : Texier 608 ; D : Droissart 1207 ; E : Unvouchered photo, N end of Lake Nyasa, Tanzania ; F : Lowry 4987]
[Photos : A : D. Ravelonarivo ; B : P. Antilahimena ; C : N. Texier ; D : V. Droissart ; E : Paul Latham ; F : P. Lowry]
270 – Synopsis of Astropanax and Neocussonia (Araliaceae)
= Heptapleurum dananense A. Chev. in Bull. Soc. Bot.
France 58 (Mém. 8d) : 178. 1912. [ Schelera danan
ensis (A. Chev.) Harms ex Engl. in Engl. & Drude,
Veg. Erde 9 : 778. 1921. Typus : IVORY COAST. Prov.
Danane : Pays des Dyolas, au bord de la Rivière
Boan, près de Danané (Fort Hittos), 16.IV.1895, fr.,
Chevalier 21275 (holo- : P [P00697823]! ; iso- : BR
[BR08860934]!, K [K000350374]!, MO!, NY!, P
[P00697824, P00697825]!).
= Schelera ledermannii Harms in Bot. Jahrb. Syst. 53 :
359. 1915. Lectotypus (designated here) : EQUATORIAL G UINEA . Bioko [Fernando Po] : Moka,
1200-1800 m, VIII.1911, l., Mildbraed 7107 (HBG
[HBG-502745]!).
= Schelera henriquesiana Harms ex Henriq. in Bol. Soc.
Brot. 27 : 192. 1917. Lectotypus (designated here) :
SÃO TOMÉ & PRINCIPE. São Tomé : Baté Pa, Vermelho, 670 m, VII.1885, bud & l., Moller 490 (COI
[COI00075041] image seen). Syntypi : SÃO TOMÉ &
PRINCIPE. São Tomé : sine loc., VIII. 1903, l., Henr
iques 18 (COI [COI00075042] image seen) ; Baté Pa,
600 m, VII. 1885, y. fr., Moller 569 (COI [COI00075040]
image seen) ; Baté Pa, 500 m, VIII.1885, l. & fr., Moller
701 (COI [COI00075039] image seen).
Habitat and distribution. – Astropanax barteri ranges from
Guinea Conakry, Liberia and Sierra Leone in the west of
Africa to the Gulf of Guinea Islands and Central Africa as
far east as central Democratic Republic of the Congo and
south to NE Angola (Bamps, 1974b ; APD, 2017). It occurs
in lowland to montane humid, gallery and swampy forest, from
elevations as low as 100 m and as high as 2,200 m.
Notes. – In the protologue for Astropanax barteri, Seemann
(1865) provided a description based on fruiting material along
with the following citation : “Sugar Loaf Mountains, Niger
(Barter! n. 2027)”. However, this collection, deposited at K,
comprises a sterile specimen from “Prince’s Island” (Príncipe).
In all likelihood, A. barteri was also based on Barter s.n. (also
at K), a fruiting collection from the Sugar Loaf Mountains
(“Niger” was at that time more broadly circumscribed to
include present day Sierra Leone). Given Seemann’s clear
indication that the type of his new species came from this
locality and the fact that he obviously saw fruiting material,
we have chosen to interpret the single sheet of Barter s.n. as
the holotype, following Bamps (1974).
Schelera hierniana was a replacement name for the illegitimate Heptapleurum scandens, a later homonym of H. scandens
(Blume) Seem. (Seemann, 1865). It was based on two sheets
of Mann 1180 from Cameroon, both at K, from which we have
selected the specimen in young fruit bearing the original label
Candollea 72, 2017
as the lectotype. While Frodin & Govaerts (2004) regarded
Schelera hierniana as distinct from S. barteri ([ Astropanax
barteri), we have chosen to follow Bamps (1974a, 1974b) in
treating it as a synonym, at least until more detailed morphometric and/or phylogenetic analyses can be undertaken.
Schefflera ledermannii was based on two collections at
B (Milbraed 7107 from Bioko and Ledermann 1554 from
Cameroon), both of which appear to have been destroyed.
A duplicate of Milbraed 7107 is deposited in HBG and has
been selected as the lectotype.
Schelera henriquesiana was based on four collections at
COI, one of which (Moller 490) has been selected as the
lectotype because it is more complete than the other syntypes
(Henriques 18, Moller 569 and 701).
A3. Astropanax evrardii (Bamps) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Schelera evrardii Bamps in Bull. Jard. Bot. Belg. 43 : 425.
1973.
Typus : DEMOCRATIC REPUBLIC OF THE CONGO. Prov.
Equateur : Djoa (Terrotoire Bolomba), 17.V.1958, y. fr., Evrard
4071 (holo- : BR [BR00886167]! ; iso- : K [K00350381]!).
Habitat and distribution. – Astropanax evrardii is known
from only a few localities in central Democratic Republic of
the Congo, where it grows in low-altitude humid and swampy
forest between 300 m and 450 m elevation (Bamps, 1974b ;
APD, 2017)
Notes. – This species closely resembles A. goetzenii but
difers by several morphological features (see Bamps, 1971,
1974a, 1974b), which appear to justify its recognition as a
distinct species.
A4. Astropanax goetzenii (Harms) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Schelera goetzenii Harms in Goetzen, Durch Afr., Repr.
7 : 376. 1895.
Neotypus (designated here) : DEMOCRATIC REPUBLIC OF
CONGO. Prov. Kivu-Nord : E slope of Niragongo
Volcano [= Mt. Nyiragongo], 2440 m, 6.I.1931, Burtt 3224
(K [K001208218]!).
= Schefflera stuhlmannii Harms in Bot. Jahrb. Syst. 26 :
243. 1899. Neotypus (designated here) : TANZANIA.
Morogoro Region : Morningside, Uluguru Mountains, XI.1934, bud, Bruce 240 (K! ; isoneo- : BM
[BM00129656]!, BR [BR14695070]!).
THE
= Schelera adolifridericii Harms in Wiss. Ergebn. Deut.
Zentr.-Afr. Exped., Bot. 2 : 590. 1913. Typus : RWANDA :
Bugoie, Milbraed 1478 (B†).
Candollea 72, 2017
= Schefflera mildbraedii Harms in Wiss. Ergebn. Deut.
Zentr.-Afr. Exped., Bot. 2 : 591. 1913. Typus : RWANDA.
Prov. Ouest : Rukarara [= Lukarara], Milbraed 1010
(B†).
= Schelera sycidiifolia Lebrun in Bull. Jard. Bot. Etat Bruxelles 13 : 19. 1934. Lectotypus (designated here) : DEMOCRATIC REPUBLIC OF THE CONGO. Prov. Nord-Kivu :
entre Kasindi et Lubango, chaîne W du lac Edouard,
2320 m, I.1932, bud, Lebrun 4755 (BR [BR00886166]! ;
isolecto- : BR [BR00886165]!, K [K00350386]!).
Habitat and distribution. – Astropanax goetzenii occurs
in eastern Africa, from eastern Democratic Republic of the
Congo, Burundi and Rwanda through southern Tanzania to
Mozambique and Zimbabwe. It grows in humid to seasonally
dry forest, and also in secondary forest, from 800 to 2,500 m
elevation (Bamps, 1974b ; APD, 2017).
Notes. – he holotype of Schelera goetzenii Harms (Götzen
s.n.), from what is now known as Mt. Nyiragongo in the
Democratic Republic of the Congo, was deposited at B and
is presumably destroyed. We have not been able to locate any
isotypes, so we have designated as the neotype the only other
know collection from the same mountain, Burtt 8155, which is
deposited at K.
Schelera stuhlmannii Harms was based on two collections
made by Franz Ludwig Stuhlmann, (1863-1928) in Tanzania
(Stuhlmann 8849 and 9277), both of which were deposited at B
and are therefore presumably destroyed, and neither of which
appears to have duplicates in other herbaria. While Frodin &
Govaerts (2004) regarded S. stuhlmannii as distinct from S.
goetzenii ([ Astropanax goetzenii), we have chosen to follow
Bamps (1974a, 1974b) in treating it as a synonym, at least until
more detailed morphometric and/or phylogenetic analyses can
be undertaken. We have therefore designated as a neotype the
specimen of Bruce 240 deposited at K, which corresponds well to
the protologue and is represented by two duplicates at BM and
BR. By contrast, we have refraied from designating a neotype
for S. adolifridericii Harms and S. mildbraedii Harms, whose
types was also deposited at B and for which no isotypes are
known, because we have not been able to ind any material that
appears to correspond well to the protologues.
he type collection of Schelera sycidiifolia Lebrun at BR
is represented by two sheets. We have selected the specimen
bearing the original label as the lectotype.
A5. Astropanax humblotianus (Drake) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Schelera humblotiana Drake in Grandidier, Hist. Phys.
Madagascar 35 : tab. 404. 1896.
Synopsis of Astropanax and Neocussonia (Araliaceae) - 271
Lectotypus (designated here) : MADAGASCAR : sine loc., y. fr.,
Humblot 640 (P [P00466310]! ; isolecto- : K!, LD [LD1215974,
LD1216274]!, P [P00466308, P00466309]!, W!).
Habitat and distribution. – Astropanax humblotianus has
been collected at a limited number of sites in eastern Madagascar, from Ankirindro near Maroantsetra in the northeast
to Ambalabe in the center-east of the island. It occurs in
mid-altitude humid forest, from c. 400 to 1,150 m elevation
(Madagascar Catalogue, 2017).
Notes. – From among the three specimens of Drake 640 at
P, we have selected the most complete one, with an entire adult
leaf and a portion of an inlorescence with a nearly complete
ultimate unit, as the lectotype.
A6. Astropanax kivuensis (Bamps) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Schelera kivuensis Bamps in Bull. Jard. Bot. Belg. 41 :
251. 1971.
Typus : DEMOCRATIC REPUBLIC OF THE CONGO. Prov.
Nord-Kivu : Kikoma, terr. Masisi, 1077 m, 17.II.1959, bud
& l., A. Léonard 3082 (holo- : BR [BR00886171]! ; iso- :
BR [BR00886169, BR00886170]!).
Habitat and distribution. – Astropanax kivuensis is only
known from eastern Democratic Republic of the Congo
(Bamps, 1974b ; APD, 2017), where it has been collected in
humid forest at 1,000-1,700 m elevation.
Notes. – Among the members of the genus with ultimate
inlorescence units arranged racemosely, Astropanax kivuensis is
distinctive in having 3 orders of inlorescence branching (i.e., panicles of umbellules, with a distinct primary axis), vs. just 2 orders
of branching, thus forming racemes of umbellules arising from
nothing more than a short primary axis (Bamps, 1974a, 1974b).
A7. Astropanax mannii (Hook.f.) Seem. in J. Bot. 3 : 177.
1865.
[
Paratropia mannii Hook. f. in J. Linn. Soc., Bot. 6 : 10.
1862. [ Schelera mannii (Hook. f.) Harms in Engl. &
Prantl, Nat. Planzenfam. III(8) : 38. 1894. [ Hepta
pleurum mannii (Hook. f.) Benth. in Bentham & Hook.
f., Gen. Pl. 1 : 942. 1876. [ Brassaia mannii (Hook. f.)
Hutch., Gen. Fl. Pl. 2 : 623. 1967.
Lectotypus (irst step designated by Bamps, 1974b : 133 ;
second step designated here) : EQUATORIAL GUINEA.
Bioko [Fernando Po] : sine loc., 1860, y. fr., Mann 289 (K
[K00350392]! ; isolecto- : K [K00350393, K00350394,
K00350395]! ; P [P00466313, P00466314, P00466315]!,
S [S11-26125] image seen).
272 – Synopsis of Astropanax and Neocussonia (Araliaceae)
= Schelera mannii var. lancifolia Harms in Bot. Jahrb.
Syst. 53 : 358. 1915. Lectotypus (designated here) :
EQUATORIAL GUINEA. Bioko : sine loc., bud & fl.,
Mildbraed 6410 (HBG!).
Habitat and distribution. – Astropanax mannii is restricted
to the mountains of western Cameroon and adjacent Nigeria,
as well as the Gulf of Guinea Islands of Annobón, Bioko and
São Tomé (Bamps, 1974b ; APD, 2017), where it occurs in
humid forest between 300 and 2,300 m elevation.
Notes. – Bamps (1974b) indicated Mann 289 at K as the
lectotype, citing two additional collections from Cameroon
(Mann 1182 and 2168) as “syntypes”. However, the protologue
only mentions “In Ins. Fernando Po, alt. 5000 ped.” and as a
consequence the specimens from Cameroon do not represent
original material. A total of four sheets of Mann 289 are
deposited at K ; we have further lectotypiied by selecting the
specimen bearing Mann’s original label.
he holotype of Schelera mannii var. lancifolia at B was
presumably destroyed, so we have designated the only known
duplicate, deposited at HBG, as the lectotype.
A8. Astropanax monophyllus (Baker) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Cussonia monophylla Baker in J. Linn. Soc., Bot. 20 :
155. 1883. [ Neocussonia monophylla (Baker) Hutch.,
Gen. Fl. Pl. 2 : 79. 1967. [ Schelera monophylla (Baker)
Bernardi in Candollea 24 : 97. 1969.
Typus : MADAGASCAR : sine loc., y. fr., Baron 1279 (holo- :
K [K00350555]! ; iso- : BM!, P [P00466311, P00466312]!).
Habitat and distribution. – Astropanax monophyllus is
endemic to Madagascar, where it is widely distributed in
humid forest from c. 800 to 2,500 m elevation, extending
from Montagne d’Ambre National Park in the far north to
Andohahela National Park and Tsitongambarika Reserve in
the southeastern part of the island (Madagascar Catalogue,
2017).
Notes. – Notwithstanding the species epithet, the leaves of
Astropanax monophyllus often have 2 or 3 lealets, one of which
is almost always considerably larger.
A9. Astropanax myrianthus (Baker) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Cussonia myriantha Baker in J. Linn. Soc., Bot. 20 :
157. 1883. [ Schefflera myriantha (Baker) Drake in
Grandidier, Hist. Phys. Madagascar 35 : t. 403. 1896.
[ Neocussonia myriantha (Baker) Hutch., Gen. Fl. Pl.
2 : 79. 1967.
Candollea 72, 2017
Typus : MADAGASCAR : sine loc., bud, Baron 2017 (holo- :
K [K00350556]! ; iso- : G [G00015604]!, P [P00466304]!).
= Schefflera myriantha var. attenuata Bernardi in Candollea 24 : 101. 1969. Lectotypus (designated here) :
MADAGASCAR. Prov. Toliara : massif de l’Andohahela,
haute vallée de la Sakamalio, [24°32’S 46°41’E], 16001900 m, I.1934, l., Humbert 13575 (P [P00466305]! ;
isolecto- : G [G00015603] image seen ; P [P00466306,
P00466307]!).
Habitat and distribution. – Astropanax myrianthus, as circumscribed here, is restricted to Madagascar, where it occurs in
humid forest at scattered sites from the far north (Montagne
d’Ambre National Park) to the extreme south (Andohahela
National Park), at 950 to 2,350 m elevation (Madagascar
Catalogue, 2017).
Notes. – Based on the results of our recent molecular phylogenetic work (Gostel et al., 2017), Astropanax myrianthus
is circumscribed here to comprise only the populations from
Madagascar previously included in Schelera myriantha by various
authors (e.g., Bamps, 1974a, 1974b ; Bernardi, 1969 ; Frodin &
Govaerts, 2004). While material from continental Africa long
assigned to S. myriantha is essentially indistinguishable morphologically, molecular analyses show that if a broad circumscription
of this taxon is applied, it is paraphyletic with respect to two
clearly distinct and well delimited species in Madagascar (Astro
panax humblotianus and two as yet undescribed taxa). Moreover,
the African and Malagasy material assigned to S. myriantha, as
historically circumscribed, belongs to two distinct, well-supported
clades in all molecular analyses, prompting Gostel et al. (2017)
to conclude that two separate species must be recognized, which,
despite their morphological similarity, can be distinguished
unambiguous on the basis of geography. Specimens from Africa
are therefore referred to Astropanax polysciadus (Harms) Lowry,
G.M. Plunkett, Gostel & Frodin (see below). Material from the
Comoro Islands was not available for the molecular study, so the
placement of these populations must await additional analyses
that include samples from this archipelago.
he type collection of Schelera myriantha var. attenuata
comprises three sheets at P, the most complete of which has
been selected as the lectotype.
A10. Astropanax polysciadus (Harms) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Schelera polysciada Harms in Engl., Planzenw. OstAfrikas C : 297. 1895.
Lectotypus (designated here) : T ANZANIA . Tanga
Region : über Kilema, 2000 m, II.1894, l., Volkens 1877
(K [K00350400]! ; isolecto- : BM [BM001209174] image
seen, G [G00341679] image seen).
Candollea 72, 2017
= Schelera bequaertii De Wild. in Rev. Zool. Bot. Africaines 8 : 11. 1920. Typus : DEMOCRATIC REPUBLIC
OF THE C ONGO . Prov. Nord-Kivu : Ruwenzori,
Lanuri, 25.V.1914, bud & fl., Bequaert 4405 (BR
[BR00886258]!).
= Schelera congesta De Wild. in Rev. Zool. Bot. Africaines 8 : 12. 1920. Lectotypus (designated here) :
D EMOCRATIC R EPUBLIC OF THE C ONGO . Prov.
Nord-Kivu : Ruwenzori, Butagu, 2200-2600 m,
15.IV.1914, y. fr., Bequaert 3721 (BR [BR00886263]! ;
isolecto- : BR [BR00886259]!).
– Schelera angiensis De Wild. in Rev. Zool. Bot. Africaines 9 : 35. 1921 [nom. illeg.].
= Schefflera nyasensis De Wild. in Pl. Bequaert. 4 :
348. 1928. Typus : TANZANIA. Morogoro Region :
Rungwe Mt., Mbaka River, y. fr., Stolz 1602 (holo- :
BR [BR00886260]! ; iso- : BM [BM00902771,
BM00902772] images seen ; CAS!, K [K00350401,
K00350402, K00350403, K00350404, K00350405]!,
P [P00834314]!, UPS!, Z!).
Habitat and distribution. – Astropanax polysciadus is circumscribed here to include the continental African populations previously placed in Schefflera myriantha by recent
authors (Bamps, 1974a, 1974b ; Bernardi, 1969 ; Frodin
& Govaerts, 2004), but which must be excluded from that
species (Gostel et al., 2017). Its range extends from Ethiopia
south to Malawi (Bamps, 1974b ; APD, 2017), where it occurs
in a range of habitats including humid forest, bamboo thickets,
woody ericoid formations and riverine or swampy forest at
elevations between 1,600 and 3,400 m.
Notes. – he recognition of Astropanax polysciadus, like that
of A. myrianthus, is based on a careful interpretation of total
evidence from geographic and molecular data, which support
their being treated as distinct taxa despite a lack of morphological diagnostic characteristics (Gostel et al., 2017).
The original material on which Harms based his
description of Schelera polysciada was presumably deposited at
Berlin and thus must have been destroyed, so we have selected
the duplicate at K as the lectotype because it represents the
most complete specimens among the known isotypes.
he type material of Schelera congesta at BR comprises two
specimens, one of which is fertile and bears the original label,
and has therefore been selected as the lectotype.
Synopsis of Astropanax and Neocussonia (Araliaceae) - 273
A11. Astropanax procumbens (Hemsl.) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Geopanax procumbens Hemsl. in Hooker’s Icon. Pl. : tab.
2821. 1909. [ Schelera procumbens (Hemsl.) F. Friedmann in Bull. Mus. Natl. Hist. Nat., sect. B, Adansonia
8 : 254. 1986.
Lectotypus (designated here) : S EYCHELLES . Mahé :
Cascade Estate, 450 m, II.1905, bud & l., homasset 192
(K [K000350560]! ; isolecto- : K [K000350561]!).
Habitat and distribution. – Astropanax procumbens is known
only from two islands in the Seychelles, Mahé and Silhouette.
It has become extremely rare and on Mahé it is known only
from highly secondarized mid-altitude forest dominated by
introduced cinnamon (Cinnamomum verum J. Presl).
Notes. – Among the two specimens of Thomasset 192
deposited at K, we have selected as the lectotype the one that
is fertile and bears the collector’s original, hand-written label.
A12. Astropanax stolzii (Harms) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Schefflera stolzii Harms in Bot. Jahrb. Syst. 53 : 358.
1915.
Lectotypus (first step designated by Bamps, 1974b :
135 ; second step designated here) : TANZANIA. Mbeya
Region : Rungwe Mts., [Nyassa Hochland], Kyimbila, 1600-1800 m, VII.1913, fl. & y.fr., Stolz 2044
([K00350552]! ; isolecto- : BM [BM00645576] image
seen, BR [BR00886173]!, C [C10000295] image seen,
G [G00341680] image seen, JE [ JE00000305] image
seen, K [K00350553]!, L [L0008579] image seen,
M [M0104942] image seen, MO [MO-1608570]!,
P [P00697822]!, S [S-G-5496, S08-3328] images
seen, STU [STU000492] image seen, W!, WAG
[ WAG0000242] image seen, Z [Z-000001556,
Z-000001557] images seen).
Habitat and distribution. – Astropanax stolzii is restricted to
southeastern Tanzania and extreme northern Malawi (Bamps,
1974b ; APD, 2017 ; Tropicos, 2017), where it grows in humid
forest at 1,600 to 2,360 m elevation.
Notes. – The holotype of Schefflera stolzii Harms was
presumably at B and would thus have been destroyed.
Bamps (1974b) designated the material at K as the lectotype, an unfortunate choice because the two specimens
deposited there are rather fragmentary and are not accompanied by original labels, unlike several isotypes deposited
in other herbaria in Europe that have original labels with
the name of the new species written in Harms’ hand.
274 – Synopsis of Astropanax and Neocussonia (Araliaceae)
We have further lectotypified by selecting the specimen
from Kew with young, detached fruits in a fragment
packet as the lectotype (the other specimen is sterile,
bearing only two leaves).
A13. Astropanax tessmannii (Harms) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Schelera tessmannii Harms in Bot. Jahrb. Syst. 53 : 360.
1915.
Lectotypus (designated by Bamps, 1974a : 28) : EQUATORIAL GUINEA. Río Muni : Wele Nzas Prov., Nkolentangan,
bud & l., Tessmann 344 (K [K000350396]!).
Habitat and distribution. – Astropanax tessmannii is known
from a few localities in Equatorial Guinea (Río Muni), northern Gabon, southern Cameroon, northern Republic of Congo
and north-central Democratic Republic of the Congo (Bamps,
1974b ; APD, 2017). It occurs in humid and swampy forest
from sea level to c. 750 m elevation.
Notes. – he holotype of Schelera tessmannii was presumably at B and would thus have been destroyed, prompting
Bamps (1974a) to designate the only known duplicate at K
as the lectotype.
A14. Astropanax urostachyus (Harms) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Schefflera urostachya Harms in Wiss. Ergebn. Deut.
Zentr.-Afr. Exped., Bot. 2 : 591, tab. 78. 1913. [ Schef
lera barteri var. urostachya (Harms) Tennant in Kew
Bull. 15 : 333. 1961.
Lectotypus (designated here) : Tab. 78 from the
protologue, based on the type collection, Democratic
Republic of the Congo : Muera, I.1908, Mildbraed 2185
(holotype : B†).
= Schefflera tridentata De Wild. in Rev. Zool. Bot.
Africaines 8 : 13. 1920. Lectotypus (designated here) :
D EMOCRATIC R EPUBLIC OF THE C ONGO . Prov.
Kivu-Nord : Angi, Territoire Rutshuru, 20.IX.1914,
bud, Bequaert 5790 (BR [BR00582516]! ; isolecto- :
BR [BR00582381, BR00582414]!).
Habitat and distribution. – Astropanax urostachyus is
restricted to eastern Democratic Republic of the Congo,
western Uganda and northwestern Tanzania (Bamps, 1974b ;
APD, 2017 ; Tropicos, 2017), where it occurs in humid and
gallery forests from 700 to 1,700 m elevation.
Notes. – he holotype of Schelera urostachya Harms was
presumably at B and thus would have been destroyed. No
isotypes or any other original material is known, so we have
Candollea 72, 2017
chosen to designate as the lectotype the excellent plate that
accompanies the description, which surely must have been
based on the holotype.
he type material of Schelera tridentata De Wild. comprises three specimens at BR, one of which bears Bequaert’s
original label and has thus been selected as the lectotype.
A15. Astropanax volkensii (Harms) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Heptapleurum volkensii Harms in Bot. Jahrb. Syst.
19(47) : 41. 1894. [ Schelera volkensii (Harms) Harms
in Engl. & Prantl, Nat. Planzenfam. III(8) : 36. 1894.
[ Brassaia volkensii (Harms) Hutch. in Gen. Fl. Pl. 2 :
623. 1967.
Lectotypus (designated here) : TANZANIA. Kilimanjaro Region : Kilimanjaro, Marangu, 2200 m, X.1893,
bud, Volkens 1297 (K [K00350551]! ; isolecto- : BM
[BM00902770]!). Syntypus : TANZANIA. Kilimanjaro
Region : Marangu, 2700 m, IX. 1893, l., Volkens 986 (BR
[BR08862693]!).
Habitat and distribution. – Astropanax volkensii occurs in
eastern Africa, from Ethiopia through Uganda and Kenya to
northeastern Tanzania (Bamps, 1974b ; APD, 2017 ; Tropicos,
2017), where it grows in humid forest from 1,600 to 3,250 m
elevation.
Notes. – he material at B of the two collections cited in the
protologue of Heptapleurum volkensii (Volkens 986 and 1297)
was presumably destroyed. A single duplicate of Volkens 986 is
deposited at BR, and while it is the most complete specimen
available, we have selected the sheet of Volkens 1297 at K as the
lectotype because a duplicate (albeit somewhat less complete)
is available at BM.
Unplaced name
Schelera humblotii Harms in Engl. & Prantl, Nat. Planzenfam. III(8) : 38 (1894).
Lectotypus (designated here) : U NION OF THE
COMOROS. Grand Comore : sine loc., 20.XI.1855, l.,
Humblot 1469 (P [P00209106]! ; isolecto- : BM!, P
[P00209107, P00209108]!, W!).
Notes. – his name, based on material from the Comoro
Islands, has been treated by most recent authors (e.g., Bernardi, 1969 ; Bamps, 1974b ; Frodin & Govaerts, 2004)
as a synonym of Schelera myriantha s.l., i.e., as broadly circumscribed by them to include populations from Madagascar, continental Africa and the Comoros. However, with the
separation of populations from Madagascar and continental
Candollea 72, 2017
Africa into two morphologically similar but genetically distinct species (Astropanax myrianthus and A. polysciadus, respectively) based on evidence from recent molecular phylogenetic
work (Gostel et al., 2017), the assignment of populations
from the Comoros remains uncertain. Until material from this
archipelago can be added to the molecular phylogeny of Astro
panax in order to assess whether it belongs to the Malagasy or
African clade, we are not able to determine if the name Schef
lera humblotii should be placed in synonymy under Astropanax
myrianthus or A. polysciadus.
It is not clear whether Schelera humblotii Harms was based
on material available to the author at Berlin or whether he may
have examined specimens in Paris or possibly even at BM,
although none of this material was annotated by Harms. In
any case, if material had been available at B it can be presumed
to have been destroyed, so we have selected as the lectotype
the specimen at P bearing two inlorescence branches mounted
together (the other sheets have a leaf and a single inlorescence
branch, respectively)
B. Neocussonia (Harms) Hutch., Gen. Fl. Pl., Dicot. 2 : 79.
1967.
[
Cussonia sect. Neocussonia Harms in Engl. & Prantl,
Nat. Planzenfam. III(8) : 54. 1894.
Lectotypus (designated by Hutchinson, 1967 : 79) :
N. umbellifera (Sond.) Hutch.
= Schefflera sect. Meiopanax Baill. in Hist. Pl. 7 : 162.
1879. Typus : S. umbellifera (Sond.) Baill. [[ Neocus
sonia umbellifera (Sond.) Hutch.].
Hermaphroditic, unarmed, glabrous, terrestrial, evergreen trees. Stems sparsely to moderately branched or plants
monocaulous. Leaves alternate, palmately compound or unifoliolate ; petioles clasping at the base with ligulate stipules ;
leaflets lanceolate, ovate, elliptical, oblanceolate, obovate,
obcordate, obrhombic, obdeltoid, or obhastate, coriaceous or
subcoriaceous to membranaceous, margins entire, crenate, or
serrate-crenate, apex very broadly acute to truncate, sometimes
rounded (N. halleana) or acute (N. bojeri and N. stauferiana) to
acuminate (N. lukwangulensis), generally retuse with an evident
notch, and almost always with a small mucro or minute boss
in the sinus formed by an extension of the midvein. Inflo
rescences terminal, erect, paniculate or paniculate-umbellate
(compound-umbellate in N. bojeri), the ultimate units racemose, umbellate or racemose-umbellate, with most lowers
borne in a terminal umbel and 1-several inserted along the
peduncle ; inflorescence axes usually with persistent, stiff,
triangular bracts ; lowers each subtended by a bracteole (or
bracteole lacking) ; pedicels slender to stout, unarticulated, or
lowers sessile. Ovary inferior, 2–5-carpellate, carpels unilocular, placentation apical ; styles 2–5, free, borne on a short, lat
Synopsis of Astropanax and Neocussonia (Araliaceae) - 275
to conical stylopodium. Fruits drupaceous, the disc concave
to lat, stylopodium short-conical to conical ; mesocarp leshy,
endocarp scleriied around each locule, forming a pyrene.
Diversity and distribution. – Neocussonia includes 16 currently recognized species, two in eastern and southern Africa
and 14 in Madagascar, one of which is elevated here from the
rank of variety. All of the c. 15 additional species that remain
to be described in the genus likewise occur in Madagascar. In
Africa, one species (N. lukwangulensis) is restricted to Tanzania
while the other (N. umbellifera) ranges from southern Tanzania
to South Africa (APD, 2017). he Malagasy species occur
primarily in various types of humid forest, with the notable
exception of N. bojeri, which grows in much drier Tapia forest
and wooded grassland-bushland mosaic vegetation. Several
species in Madagascar have extensive distributions (e.g.,
N. bojeri, N. longipedicellata and N. vantsilana) while many
others have very restricted ranges (in particular N. capuroniana,
N. fosbergiana, N. frodiniana, N. halleana, N. rainaliana and
N. weibeliana) (Madagascar Catalogue, 2017).
Notes. – This group was originally described by Harms
(1894-97) as a section of Cussonia, which he circumscribed
to include ive species currently recognized as Schelera bojeri,
S. monophylla, S. myriantha, S. umbellifera and Seemannaralia
gerrardii (Seem.) R. Vig., based primarily on the presence of
lowers borne in umbellules. Hutchinson (1967), in keeping
with his mechanical delimitation of genera within Araliaceae
based largely on inlorescence features, raised Harms’s section to
the rank of genus, designating Neocussonia umbellifera as the lectotype, while also including Neocussonia buchananii (a synonym
of Schelera umbellifera) but excluded Seemannaralia gerrardii.
B1. Neocussonia bojeri (Seem.) Hutch., Gen. Fl. Pl. 2 : 79. 1967.
[
Cussonia bojeri Seem. in J. Bot. 4 : 298. 1866. [ Schelera
bojeri (Seem.) R. Vig. in Ann. Sci. Nat., Bot., sér 9, 4 :
116. 1906.
Lectotypus (designated by Bernardi, 1969) : MADAGASsine loc., Bojer s.n. (G [G00015611]! ; isolecto- : K!, P
[P02503035]!).
CAR :
Habitat and distribution. – Neocussonia bojeri is widely
distributed in drier areas of central and southern Madagascar,
where it occurs in Tapia forest and wooded grassland-bushland
mosaic vegetation at elevations between 500 m and 2,300 m
(Madagascar Catalogue, 2017).
Notes. – Neocussonia bojeri is the only member of the genus
that is adapted to areas with prolonged dry periods, and its
ire-resistant habit enables it to survive even in areas that are
subjected to frequent burning.
276 – Synopsis of Astropanax and Neocussonia (Araliaceae)
B2. Neocussonia bracteolifera (Frodin) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
B4. Neocussonia favargeri (Bernardi) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
Cussonia capuroniana var. bracteolata Bernardi in Ber.
Schweiz. Bot. Ges. 76 : 362. 1966. [ Schelera capuro
niana var. bracteolata (Bernardi) Bernardi in Candollea
24 : 108. 1969. [ Schelera bracteolifera Frodin in Frodin
& Govaerts, World Checklist Bibliog. Araliaceae : 328.
2004.
Typus : MADAGASCAR. Prov. Antsiranana : Tsaratanana,
2500-2700 m, X.1912, bud, Perrier de la Bâthie 6881 (P
[P00442775]!).
Habitat and distribution. – Neocussonia bracteolifera is
known from humid forest at only two sites, the Tsaratanana
massif in northern Madagascar, where it occurs between
c. 2,300 and 2,600 m elevation, and Anjialavahely, some 50 km
to the ESE, where it was collected at about 1,550 m (Madagascar Catalogue, 2017).
Notes. – Frodin & Govaerts (2004) opted to publish
Schelera bracteolifera as a new name at the rank of species
for the taxon originally described as Cussonia capuroniana var.
bracteolata rather than making a new combination by elevating
the varietal name to the rank of species.
B3. Neocussonia capuroniana (Bernardi) Lowry, G.M. Plunkett, Gostel & Frodin, comb. nov.
[
Candollea 72, 2017
Cussonia capuroniana Bernardi in Ber. Schweiz. Bot.
Ges. 76 : 360. 1966. [ Schelera capuroniana (Bernardi)
Bernardi in Candollea 24 : 108. 1969.
Typus : M ADAGASCAR . Prov. Antananarivo : Forêt
d’Ambohitantely, sur le Tampoketsa d’Ankazobe,
25.III.1963, bud & l., Service Forestier 22650 (holo- : P
[P00442774]! ; iso- : P [P00298719]!).
Habitat and distribution. – Neocussonia capuroniana is
known only from humid forest at the Ambohitantely Reserve
and the nearby Manankazo-Ankazobe forestry station, to the
NW of the town of Ankazobe (Madagascar Catalogue,
2017).
Notes. – he holotype of Cussonia capuroniana at P bears a
type label clearly aixed by Bernardi in Geneva. he annotation made by Frodin in 2001 designating this specimen as a
lectotype was thus superluous and in any case this choice was
never published.
[
Schelera favargeri Bernardi in Ber. Schweiz. Bot. Ges.
76 : 369. 1966.
Typus : MADAGASCAR. Prov. Antsiranana : Tsaratanana,
XII.1912, bud & y. fr., Perrier de la Bâthie 3576 (holo- : P
[P00442784]! ; iso- : G [G00015609]!).
Habitat and distribution. – Neocussonia favargeri occurs
in several mountain massifs in northern Madagascar, from
Manongarivo in the west through Tsaratanana and several
nearby ranges (including Ambohimirahavavy and Mt.
Ampomotra) to Anjanaharibe-Sud Reserve and Marojejy
National Park in the east (Madagascar Catalogue, 2017).
It is found in humid forest between c. 1,200 and 2,400 m
elevation.
Notes. – he holotype of Schelera favargeri bears a type
label affixed by Bernardi in Geneva. The annotation made
by Frodin in 2001 designating this specimen as a lectotype
was thus superluous and in any case this choice was never
published.
B5. Neocussonia fosbergiana (Bernardi) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Cussonia fosbergiana Bernardi in Ber. Schweiz. Bot. Ges.
76 : 363. 1966. [ Schelera fosbergiana (Bernardi) Bernardi
in Candollea 24 : 117. 1969.
Typus : M ADAGASCAR . Prov. Antsiranana : vallée de
la Lokoho, Mt. Beondroka au N de Maroambihy, 1001450 m, 17-22.III.1949, bud, l. & y. fr, Humbert 23439
(holo- : P [P00442788]! ; iso- : G [G00015607]!, P
[P00442787, P00498704]!).
Habitat and distribution. – Neocussonia fosbergiana is known
only from Marojejy National Park and the Betsomanga massif
c. 25 km to the NNE (Madagascar Catalogue, 2017),
where it grows in humid forest and ericoid thickets between
1,200 and 1,700 m elevation.
Notes. – he holotype of Cussonia fosbergiana at P bears a
type label clearly aixed by Bernardi in Geneva, rendering the
annotation made by Frodin in 2001 designating this specimen
as a lectotype superfluous, and in any case this choice was
never published.
Candollea 72, 2017
A
Synopsis of Astropanax and Neocussonia (Araliaceae) - 277
B
C
D
E
F
G
Fig. 2. – Photographs of Neocussonia (Harms) Hutch. (including both published and new, as yet unpublished species). A. Neocussonia sp. ;
B. Neocussonia vantsilana (Baker) Lowry, G.M. Plunkett, Gostel & Frodin ; C. Neocussonia favargeri (Bernardi) Lowry, G.M. Plunkett, Gostel & Frodin ;
D. Neocussonia halleana (Bernardi) Lowry, G.M. Plunkett, Gostel & Frodin ; E. Neocussonia sp. nov. ; F. Neocussonia capuroniana (Bernardi) Lowry, G.M.
Plunkett, Gostel & Frodin ; G. Neocussonia frodiniana (Bernardi) Lowry, G.M. Plunkett, Gostel & Frodin.
[A : Lowry 7157 ; B : Lowry 7100 ; C : Callmander 384 ; D : Ravelonarivo 3340 ; E : Lowry 6705 ; F : Plunkett 2328 ; G. Lowry 4504]
[Photos : A-B, E-G : P. Lowry ; C : M. Callmander ; D : D. Ravelonarivo]
278 – Synopsis of Astropanax and Neocussonia (Araliaceae)
B6. Neocussonia frodiniana (Bernardi) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Schelera frodiniana Bernardi in Candollea 24 : 103.
1969.
Typus : M ADAGASCAR . Prov. Toliara : massif de
l’Andohahela, 1700-1950 m, I.1934, bud, Humbert
13591 (holo- : P [P00442783]! ; iso- : G [G00015608]!, P
[P00442781, P00442782]!).
Candollea 72, 2017
here are two specimens of Humbert 23495 at P, one of
which bears an annotation by Frodin made in 2001 indicating
it as the lectotype, but this designation was never published.
We have selected this specimen as the lectotype as it is the
more complete of the two sheets.
B8. Neocussonia litoralis (Bernardi) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. et stat. nov.
[
Habitat and distribution. – Neocussonia frodiniana is known
only from the upper slopes of Mt. Trafanaomby, the highest
peak in Andohahela National Park (Madagascar Catalogue, 2017), where it occurs in humid forest at an elevation
of c. 1,700-1,950 m.
Notes. – his species was originally known only from the
type material, but additional specimens were recently collected
from the same locality (Madagascar Catalogue, 2017).
B7. Neocussonia halleana (Bernardi) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Schelera halleana Bernardi in Ber. Schweiz. Bot. Ges.
76 : 371. 1966.
Lectotypus (designated here) : M ADAGASCAR . Prov.
Antsiranana : Vallée de la Lokoho, Mont Beondroka au
N de Maroambihy, 1000-1450 m, 17-22.II.1949, bud
& fl., Humbert 23495 (P [P00442780]! ; isolecto- : G
[G00015605, G00015606]!, P [P00442779]!).
Habitat and distribution. – Neocussonia halleana is known
only from Marojejy National Park in northeastern Madagascar,
where it is restricted to ericoid thicket and high altitude humid
forest on and just below the summits of Marojejy and Beondroka (Madagascar Catalogue, 2017) between c. 1,700 and
2,100 m elevation.
Notes. – he label data accompanying Cours 4388 (G,
P) indicate that it was collected on the summit of Marojejy on 28 March 1949, but give the altitude as 1000 m,
which is far below the top of the mountain. Cours and
Humbert both participated in the March 1949 expedition
to Marojejy during which this material was gathered, and
collections of many species clearly taken from the same
individuals were distributed under both of their numbers. It
may be that Cours 4388 is identical to Humbert 23495, and
indeed this interpretation was used at G, where their sheet
[G00015606] is recorded as an isotype. However, given the
diferent collection date, we have chosen not to regard this
collection as comprising type material.
Schelera vantsilana var. litoralis Bernardi in Candollea
24 : 119. 1969.
Typus : M ADAGASCAR . Prov. Toamasina : Tampolo,
Ampasina, Fénérive, 2.VII.1957, l., Service Forestier 17701
(holo- : P [P00442793]! ; iso- : P [P00442792]!, TEF
[TEF000006, TEF000007]!).
Habitat and distribution. – Neocussonia litoralis is restricted
to fewer than ten isolated areas of low elevation littoral forest
along the east coast of Madagascar, a fragmented and highly
threatened ecosystem (Consiglio et al., 2006). It ranges from
Ambatobiriry near Sambava in the north (where all remaining
forest was removed during the last few years) to Ste. Luce in
the south (Madagascar Catalogue, 2017).
Notes. – Bernardi (1969) described this taxon as a variety
of Schelera vantsilana ([ Neocussonia vantsilana), which occurs
in humid forest at mid- to high elevations, but it is more than
suiciently distinct morphologically to warrant recognition at
the species level.
From among the two sheets of Service Forestier 17701
deposited in the Paris herbarium, we have selected the one
comprising the most complete material and bearing an original
label as the lectotype. his specimen also has an annotation
by Frodin indicating that he intended to designate it as the
lectotype, although his choice was never published.
B9. Neocussonia longipedicellata (R. Vig. & Danguy ex
Lecomte) Lowry, G.M. Plunkett, Gostel & Frodin, comb. nov.
[
Cussonia longipedicellata R. Vig. & Danguy ex Lecomte,
Madag., Bois Analamaz. 117. 1922. [ Schelera longi
pedicellata (R. Vig. & Danguy ex Lecomte) Bernardi in
Candollea 24 : 114. 1969.
Lectotypus (designated here) : MADAGASCAR. Prov. Antananarivo : Forêt d’Analamazaotra, l., Service Colonial 56
(P [P00442778]! ; isolecto- : [P00442776, P00442777]!).
= Cussonia racemosa Baker in J. Linn. Soc., Bot. 20 : 156.
1883. [ Schelera macerosa Bernardi in Candollea 24 :
113. 1969 [non S. racemosa (Wight) Harms]. Typus :
Madagascar : sine loc., bud, Baron 2015 (holo :- K
[K000350557]! ; iso- : P [P00442785, P00442786]!).
Candollea 72, 2017
Habitat and distribution. – Neocussonia longipedicellata is
the most widespread and morphologically variable member
of the genus in Madagascar. It occurs in mid-elevation humid
forest (c. 800-1,700 m) from Montagne d’Ambre National
Park in the far north to Kalambatritra Reserve in the south,
with outlying populations at several sites in the west (e.g.,
Bongolava, Ambohijanahary Reserve, and in low elevation
gallery forest along the middle Mangoky River) (Madagascar Catalogue, 2017).
Note. – From among the three syntypes of Cussonia longi
pedicellata deposited at P, we have selected the most complete
as the lectotype.
B10. Neocussonia lukwangulensis (Tennant) Lowry, G.M.
Plunkett, Gostel & Frodin, comb. nov.
[
Cussonia lukwangulensis Tennant in Kew Bull. 14 : 221.
1960. [ Schelera lukwangulensis (Tennant) Bernardi in
Candollea 24 : 94. 1969.
Typus : T ANZANIA . Morogoro Region : Uluguru
[Mountains], Tanana, 4.II.1935, bud, Bruce 769 (holo- :
K [K00350588]! ; iso- : BM [BM000902773]!, BR
[BR08861689]!, P [P01816987]!).
Habitat and distribution. – Neocussonia lukwangulensis
is endemic to Tanzania, where it occurs in many of the
Eastern Arc mountains, from the Chome Forest Reserve in
the South Pare Mountains south to the Kipengere Game
Reserve and the Rungwe Forest Reserve near Mbeya in
the southwestern part of the country (Bamps, 1974b, APD,
2017 ; Tropicos, 2017). It grows in humid forest from
c. 1,350 to 2,136 m elevation.
Notes. – Harms (1899) listed Stuhlmann 9112 among the
syntypes of his Cussonia buchananii (a synonym of Neocussonia
umbellifera ; see below), but this specimen is clearly referable
to N. lukwangulensis.
B11. Neocussonia moratii (Bernardi) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
Synopsis of Astropanax and Neocussonia (Araliaceae) - 279
Notes. – his species was described based on a single lowering specimen, but has since been collected at various stages
of development on at least 10 more occasions (Madagascar
Catalogue, 2017).
B12. Neocussonia rainaliana (Bernardi) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Schelera rainaliana Bernardi in Candollea 35 : 112.
1980.
Typus : MADAGASCAR. Prov. Toliara : Fort-Dauphin,
Station Forestière de Mandena, 17.II.1972, bud, l., Debray
1730 (P [P00442772]!).
Habitat and distribution. – Neocussonia rainaliana is
restricted to low elevation littoral forest on sand at three sites
to the N of Tolagnaro in southeastern Madagascar (Antsotso,
Mandena and Ste Luce) (Madagascar Catalogue, 2017).
Commercial exploitation of ilmenite (titanium oxide) by Rio
Tinto QMM has impacted the local populations at the latter
two sites, although substantial portions of the remaining forest
(c. 660 ha at Mandena and c. 780 ha at Ste Luce) have been
formally designated as protected areas that are being managed
with support from the mining company.
Notes. – his species was described on the basis of a single,
rather fragmentary unicate specimen, but intensive botanical
inventory work in the Tolagnaro region (see Lowry et al.,
2008 ; Madagascar Catalogue, 2017) has generated nearly
30 additional collection, conirming the distinctiveness of this
taxon.
B13. Neocussonia stauferiana (Bernardi) Lowry, G.M. Plunkett, Gostel & Frodin, comb. nov.
[
Schelera stauferiana Bernardi in Candollea 24 : 110.
1969.
Typus : MADAGASCAR. Prov. Fianarantsoa : Bassin de
l’Itomampy, Mt. Papanga, près de Befotaka, [23°51’S
46°57’E], 2.XII.1928, fl., Humbert 6922 (holo- : P
[P00442796]! ; iso- : P [P00442794, P00442795]!).
Typus : MADAGASCAR. Prov. Toamasina : Zahamena,
Canton Manaka-Est, Dist. Ambatondrazaka, 17.I.1959,
l., Réserves Naturelles 10980 (P [P00442773]!).
Habitat and distribution. – Neocussonia stauferiana is fairly
widespread in Madagascar, occurring in mid-elevation humid
forest from Zahamena reserve south to Mt. Papanga and the
Ivakoany massif, with an outlying population near Andapa in
the NE (Madagascar Catalogue, 2017).
Habitat and distribution. – Neocussonia moratii occurs in
mid-altitude humid forest at several sites along the central part
of Madagascar’s eastern escarpment, from Zahamena reserve
in the north to Lakato in the south, at elevations between
c. 1,050 and 1,400 m (Madagascar Catalogue, 2017).
Notes. – The holotype of Schefflera staufferiana bears a
type label clearly aixed by Bernardi in Geneva, making the
annotation made by Frodin in 2001 designating this specimen as a lectotype superluous, and in any case this choice
was never published.
[
Schelera moratii Bernardi in Candollea 28 : 7. 1973.
280 – Synopsis of Astropanax and Neocussonia (Araliaceae)
B14. Neocussonia umbellifera (Sond.) Hutch., Gen. Fl. Pl. 2 :
79. 1967.
[
Cussonia umbellifera Sond. in Linnaea 23 : 49. 1850.
Schelera umbellifera (Sond.) Baill., Hist. Pl. 7 : 162.
1879.
[
Lectotypus (designated here) : SOUTH AFRICA. Prov.
KwaZulu-Natal : Port Natal [= Durban], bud, Guein
zius s.n. (S [S14-41368] image seen ; isolecto- : CGE!,
G [G00341683]!, LE!, P [P04352720, P04352724,
P04352725]!, PRE!, S [S14-41367] image seen, SAM!,
W!). Syntypi : SOUTH AFRICA. Prov. KwaZulu-Natal :
between Omsamculo (Umzimkulu) and Omcomas
(Umkomaas), Drège s.n. (S [S14-41370] image seen).
= Cussonia chartacea Schinz in Bull. Herb. Boissier 2 : 211.
1894. Typus : SOUTH AFRICA. Prov. KwaZulu-Natal :
Umbilo, Rehmann 8096 (not found).
= Cussonia buchananii Harms in Bot. Jahrb. Syst. 26 :
251. 1899. [ Schefflera umbellifera var. buchananii
(Harms) Tennant in Kew Bull. 14 : 220. 1960. [ Neo
cussonia buchananii (Harms) Hutch., Gen. Fl. Pl. 2 :
79 (1967). Lectotypus (designated here) : MALAWI :
Nyasaland, sine loc., 1895, bud & fl., Buchanan 295
(BM [BM00645597]! ; isolecto- : BR [BR00886172]!,
E [E00217670] image seen, K [K000350554]!, SRGH
[SRGH0106562-0] image seen).
Habitat and distribution. – Neocussonia umbellifera occurs
in East and Southern Africa, where it is widespread in humid
forest at altitudes of 200 to 2,200 m, from Zimbabwe and
Malawi in the north to Swaziland and South Africa in the
south (Bamps, 1974b ; APD, 2017).
Notes. – Cussonia umbellifera Sond. was based on two
collections from South Africa, Gueinzius s.n. from Port
Natal (Durban), and Drège s.n. from between Omsamculo (Umzimkulu) and Omcomas (Umkomaas). A large
number of specimens of this taxon made by Drège are
widely deposited in many herbaria, but they appear to
represent several separate gatherings and it is not clear
which of these gatherings might have been used by Sonder
nor which specimens actually represent each individual
gathering. We have chosen to lectotypify this name using
the fertile specimen of Gueinzius s.n. at S, which was
unquestionably seen by Sonder and thus used for his
description.
Cussonia chartacea was based on Rehmann 8096. We have
not been able to locate any material of this collection.
Cussonia buchananii was based on two collections,
Buchanan 295, which is referable to Neocussonia umbel
lifera, and Stuhlmann 9112, which represents material
of N. lukwangulensis (see above). Frodin & Govaerts
Candollea 72, 2017
(2004) placed C. buchananii in synonymy under Schefflera
umbellifera ( [ Neocussonia umbellifera), suggesting that a
lectotypification might have been made, but we cannot
find any evidence that it actually was. In order to maintain
that placement in synonymy, we have lectotypified Cussonia
buchananii using Buchanan 295. The original material on
which Harms based his description was presumably deposited at Berlin and thus must have been destroyed, so we
have selected the duplicate at BM as the lectotype because
it represents the best specimen, which reflects the fact that
material of this collection was originally distributed from
there.
B15. Neocussonia vantsilana (Baker) Lowry, G.M. Plunkett,
Gostel & Frodin, comb. nov.
[
Cussonia vantsilana Baker in J. Linn. Soc., Bot. 20 :
156. 1883. [ Schelera vantsilana (Baker) Bernardi in
Candollea 24 : 117. 1969.
Lectotypus (first step designated by Bernardi, 1969 :
133 ; second step designated here) : MADAGASCAR : sine loc.,
Baron 1016 (K [K00350559]! ; isolecto- : P [P00442789,
P00442790]!). Syntypi : MADAGASCAR : sine loc., Bojer s.n.
(K!) ; sine loc., Parker s.n. (K!).
Habitat and distribution. – Neocussonia vantsilana is fairly
widespread in Madagascar, where it occurs in mid-elevation
humid forests (c. 1,000-1,700 m) from Anjialavabe near
Andapa in the north to Sahalava forest near Kalambatritra
reserve in the south (Madagascar Catalogue, 2017).
Notes. – In the protologue, Baker (1883) cited three collections (Baron 1016, Bojer s.n., and Parker s.n.), the irst of which
was designated as the lectotype by Bernardi (1969), without,
however, indicating an herbarium. We have selected the specimen at K, which was available to Baker, as the lectotype.
B16. Neocussonia weibeliana (Bernardi) Lowry, G.M. Plunkett, Gostel & Frodin, comb. nov.
[
Schelera weibeliana Bernardi in Candollea 24 : 111.
1969.
Lectotypus (designated here) : Madagascar. Prov. Antsiranana : vallée de la Lokoho, Mt. Beondroka, au N de
Marombihy, 1000-1450 m, 17-22.III.1949, y. fr., Humbert
23433 (P [P00498716]! ; isolecto- : G [G00015601]!).
Habitat and distribution. – Neocussonia weibeliana is known
only from the type collection made in humid forest at 1,0001,450 m elevation on Mt. Beondroka in Marojejy National
Park (Madagascar Catalogue, 2017).
Candollea 72, 2017
Notes. – Bernardi based his description of Schelera weibe
liana on two duplicates of the same collection, both of which
were deposited at P at the time, one of which was subsequently
sent to G. We have chosen the specimen at P as the lectotype
as it is the more complete of the two original syntypes.
Acknowledgments
We thank Xavier Aubriot, Sven Buerki, Martin Callmander,
Alan Paton, Tariq Stévart, and Jacek Wajer for assistance in
locating and obtaining scans of types and of published papers.
We are grateful to the curators of the following herbaria for
providing access to their collections : BM, BR, CAS, FI, G,
HBG, K, L, LD, MO, NY, OXF, P, TAN, TEF, UPS, W and
Z. he authors also thank Roy Gereau and Martin Callmander
for valuable suggestions that helped improve the inal version
of the manuscript. Field and herbarium work was supported by
grants from the U.S. National Science Foundation (9981641,
0613728/0943958, and 1556139/1556327, GMP and PPL
co-PIs ; 0743355, PPL as Co-PI). Fieldwork in Madagascar
was carried out under collaborative agreements between
the Missouri Botanical Garden and the Parc Botanique et
Zoologique de Tsimbazaza and the Direction de la Recherche
Forestière et Piscicole, FOFIFA. We are grateful for courtesies extended by the Government of Madagascar (Direction
Générale des Forêts) and by Madagascar National Parks. We
are especially grateful to the many dedicated ield botanists
working in Madagascar who have diligently collected material
of Araliaceae : they are too numerous to mention, but a permanent record of their names and their valuable contributions is
documented in the thousands of collections they have made
over the years.
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