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A synopsis of the Afro-Malagasy species previously included in Schefflera (Araliaceae): resurrection of the genera Astropanax and Neocussonia

A synopsis of the Afro-Malagasy species previously included in Schefflera (Araliaceae): resurrection of the genera Astropanax and Neocussonia

Profile image of Morgan GostelMorgan Gostel

Phylogenetic studies have shown that Sche era J.R. Forst. & G. Forst., the largest genus of Araliaceae, is grossly polyphy- letic, representing ve 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 e ort to establish monophyletic genera among the elements that have been treated in Sche era 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.

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Straddling the Mozambique Channel: molecular evidence for two major clades of Afro-Malagasy Schefflera (Araliaceae) co-occurring in Africa and Madagascar

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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 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. 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. References APD [African Plant Database] (2017). African Plant Database. Conservatoire et Jardin botaniques de la Ville de Genève & South African National Biodiversity Institute [www.ville-ge.ch/musinfo/ bd/cjb/africa/recherche.php]. Bamps, P. (1971). Deux Araliacées nouvelles du Congo-Kinshasa. Bull. Jard. Bot. Belg. 41 : 251. Synopsis of Astropanax and Neocussonia (Araliaceae) - 281 Bernardi, L. (1973). Araliacearum Madagascariae et Comores epilogus. 3. Species nova Schelerarum. Candollea 28 : 7-11. Bernardi, L. (1980). Synopsis Araliacearum Madagascariae et Comorarum Insularum (auxilio methodi “Ferulago”). Candollea 35 : 117-132. Consiglio, T., G.E. Schatz, G. McPherson, P.P. Lowry II, J. Rabenantoandro, Z.S. Rogers, R. Rabevohitra & D. Rabehevitra (2006). Deforestation and plant diversity of Madagascar’s littoral forests. Conservation Biol. 20 : 1799-1805. Fiaschi, P. & G.M. Plunkett. (2011). Monophyly and phylogenetic relationships of Neotropical Schelera (Araliaceae) based on plastid and nuclear markers. Syst. Bot. 36 : 806-817. Frodin, D.G. (1975). Studies in Schelera (Araliaceae) : the Cephaloschelera complex. J. Arnold Arbor. 56 : 427-448. Frodin, D.G. (1986). Studies in Schelera (Araliaceae), II. Northern Luzon (Philippines) species of the Heptapleurum group. Proc. Acad. Nat. Sci. Philadelphia 138 : 403-425. Frodin, D.G. (1989). Studies in Schelera (Araliaceae), IV. Synopsis of the Formenkreis comprised of Didymopanax attenuatus (Sw.) El. Marchal and allied species, with nomenclatural changes. Proc. Acad. Nat. Sci. Philadelphia 141 : 313-319. Frodin, D.G. (1993). Studies in Schelera (Araliaceae), VI. New species and subordinate taxa in the Venezuelan Guayana and immediately adjacent areas. Novon 3 : 367-403. Frodin, D.G. (1995). Neotropical montane Araliaceae : an overview. In : Churchill., S.P. et al. (ed.), Biodiversity and conservation of Neotropical montane forests : 421-431. New York Botanical Garden. Frodin, D.G. & R. Govaerts (2004). World Checklist and Bibliography of Araliaceae. Royal Botanic Gardens, Kew. Frodin, D.G., P.P. Lowry II & G.M. Plunkett (2010). Schelera (Araliaceae) : taxonomic history, overview and progress. Pl. Div. Evol. 128 : 561-595. Gostel, M.R., G.M. Plunkett & P.P. Lowry II (2017). Straddling the Mozambique Channel : molecular evidence for two major clades of Afro-Malagasy Schelera (Araliaceae) co-occurring in Africa and Madagascar. Pl. Ecol. Evol. 150 : 87-108. Bamps, P. (1974a). Araliaceae. Fl. Afrique Centrale. Meise. Harms, H. (1894-1897) Araliaceae. In : Engler, A. & K. Prantl, Nat. Planzenf. III(8) : 1-62. Leipzig, Wilhelm Engelmann. Bamps, P. (1974b). Contributions à l’étude des Araliacées africaines. Bull. Jard. Bot. Belg. 44 : 101-139. Harms, H. (1899). Araliaceae africanae. In : Engler, A. (ed.), Beiträge zur Flora von Afrika. XVII. Bot. Jahrb. Syst. 26 : 240-252. Bernardi, L. (1966). Species novae (13) et nomina mutata (2) in Araliae familia insulae Madagascariae. Ber. Schweiz. Bot. Ges. 76 : 352-395. Hutchinson, J. (1967). he genera of lowering plants vol. 2. Oxford University Press, Bernardi, L. (1969). Araliacearum Madagascariae et Comores exordium. 1. Revisio et taxa nova Schelerarum. Candollea 24 : 89-122. Lowry, P.P., II (1989). A revision of Araliaceae from Vanuatu. Bull. Mus. Natl. Hist. Nat., Sect. B, Adansonia 11 : 117-155. 282 – Synopsis of Astropanax and Neocussonia (Araliaceae) Lowry, P.P., II & G.M. Plunkett. 2010. Recircumscription of Polyscias (Araliaceae) to include six related genera, with a new infrageneric classiication and a synopsis of species. Plant Div. Evol. 128 : 55-84. Lowry, P.P., II, G.M. Plunkett & D.G. Frodin (2013). Revision of Plerandra (Araliaceae). I. A synopsis of the genus with an expanded circumscription and a new infrageneric classiication. Brittonia 65 : 42-61. Lowry, P.P., II, F. Randriatafika & J. Rabenantoandro (2008). Conservation status of vascular plant species from the QMM/Rio Tinto mining area at Mandena, Tolagnaro (Fort Dauphin) region, southeast Madagascar. Madagascar Conservation Developm. 3 : 55-63. Madagascar Catalogue (2017). Catalogue of the Plants of Madagascar. Missouri Botanical Garden, St. Louis & Antananarivo [www.tropicos.org/project/mada]. McNeill, J.M., F.R. Barrie, W.R. Buck, V. Demoulin, W. Greuter, D.L. Hawksworth, P.S. Herendeen, S. Knapp, K. Marhold, J. Prado, W. F. Prud’Homme van Reine, G.F. Smith, J. H. Wiersema & N.J. Turland (2012). International Code of Nomenclature for algae, fungi and plants (Melbourne Code) adopted by the Eighteenth International Botanical Congress Melbourne, Australia, July 2011. Regnum Veg. 154. Candollea 72, 2017 Philipson, W.R. (1979). Araliaceae. In : van Steenis C.G.G.J. (ed.), Fl. Males. I(9) : 1-105. Philipson, W.R. (1995). Araliaceae. In : Conn, B.J. (ed.), Handbook of the lora of Papua New Guinea 3 : 1-48. Melbourne University Press. Plunkett, G.M. & P.P. Lowry II (2012). Phylogeny and diversiication in the Melanesian Schelera clade (Araliaceae) based on evidence from nuclear rDNA spacers. Syst. Bot. 37 : 279-291. Plunkett, G.M., P.P. Lowry II, D.G. Frodin & J. Wen (2005). Phylogeny and geography of Schelera : pervasive polyphyly in the largest genus of Araliaceae. Ann. Missouri Bot. Gard. 92 : 202-224. Tropicos (2017). Missouri Botanical Garden, Saint Louis [http:// www.tropicos.org].

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