Studies in Neotropical Araliaceae. I. Resurrection of the genus Sciodaphyllum P. Browne to accommodate most New World species previously included in Schefflera J. R. Forst. & G. Forst. PORTER P. LOWRY II1,2, GREGORY M. PLUNKETT3, M. MARCELA MORA1, ASUNCIÓN CANO , PEDRO FIASCHI5, DAVID G. FRODIN6, ROY E. GEREAU1, ÁLVARO IDÁRRAGA-PIEDRAHÍTA 7,8 , JÁIDER JIMÉNEZ-MONTOYA8, J. MOISÉS F. MENDOZA9, DAVID A. NEILL10, ORLANDO RIVERA-DIAZ11, AND CARLOS RODRIGUES-VAZ2,3 4 1 Missouri Botanical Garden, 4344 Shaw Blvd, St. Louis, Missouri 63110, USA; e-mail: Pete.Lowry@mobot.org 2 Institut de Systématique, Évolution, et Biodiversité (ISYEB), Unité Mixte de Recherche 7205, Muséum National d’Histoire Naturelle/Centre National de la Recherche Scientifique/Sorbonne Universités/École Pratique des Hautes Études, C.P. 39, 57 rue Cuvier, 75005, Paris, France 3 Cullman Program for Molecular Systematics, New York Botanical Garden, Bronx, New York 10458-5126, USA 4 Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Avda. Arenales 1256, Lima 11, Perú 5 Departamento de Botânica, Universidade Federal de Santa Catarina, Florianópolis, Santa Catarina 88040900, Brazil 6 Royal Botanic Garden, Kew, Richmond, Surrey TW9 3AE, United Kingdom 7 Fundatión Jardín Botánico de Medellín “Joachím Antonio Uribe”, Calle 73 No. 51D-14, Medellín, Colombia 8 Herbario Universidad de Antioquia (HUA), Universidad de Antioquia, Medellín, AA 1226, Colombia 9 Herbario del Oriente Boliviano, Museo de Historia Natural Noel Kempff Mercado, Av. Irala 565, Casilla 2489, Santa Cruz, Bolivia 10 Universidad Estatal Amazónica, Puyo, Pastaza, Ecuador 11 Universidad Nacional de Colombia, Facultad de Ciencias, Instituto de Ciencias Naturales, Bogotá, Colombia Abstract. Phylogenetic studies have shown that Schefflera, the largest genus of Araliaceae, is highly polyphyletic, comprising five non-sister clades within the family, and that the generic name must be restricted to a small group of Pacific Island species. In an effort to establish monophyletic genera for the other elements still remaining in Schefflera, phylogenetic analyses of the large Neotropical clade have revealed five morphologically and geographically coherent groups. Here, we reinstate the genus Sciodaphyllum, which corresponds to the largest of these five groups, presenting a synopsis of the 131 currently recognized species, including 109 new combinations, one replacement name (nomen novum), and three new heterotypic synonyms. The re-establishment of Sciodaphyllum opens the way for describing an estimated 200+ new species in the genus, nearly all of which are from the Northern and Central Andes. Keywords: Combinatio nova, nomenclature, nomen novum, taxonomy. David G. Frodin is deceased. This paper is dedicated to his memory. Porter P. Lowry II, Gregory M. Plunkett, and M. Marcela Mora, conceived and developed the work presented in this paper. The other authors provided equal input and are listed in alphabetical order. Brittonia, DOI 10.1007/s12228-019-09593-w ISSN: 0007-196X (print) ISSN: 1938-436X (electronic) © 2019, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A. BRITTONIA [VOL Resumen. Estudios filogenéticos han demostrado que Schefflera, el género más grande de Araliaceae, es altamente polifilético, con cinco clados no hermanos dentro de la familia y que el nombre genérico se debe restringir solo a un pequeño grupo de especies de las islas del Pacífico. En un esfuerzo por establecer géneros monofiléticos para los otros elementos que aún permanecen en Schefflera, los análisis filogenéticos del gran clado neotropical han revelado cinco grupos morfológicamente y geográficamente coherentes. Aquí, resucitamos el género Sciodaphyllum que corresponde al clado más grande de estos cinco grupos, presentando una sinopsis de las 131 especies actualmente reconocidas, incluyendo 109 nuevas combinaciones, un nombre nuevo (nomen novum) y tres nuevos sinónimos heterotípicos. El restablecimiento de Sciodaphyllum abre el camino para describir un estimado de más de 200 especies nuevas en el género, casi todas de las cuales son del norte y centro de los Andes. Schefflera J.R. Forst. & G. Forst. (Araliaceae) has long been circumscribed to comprise a group of mostly trees or arborescent hemi-epiphytes occurring throughout the tropics of both the Old and New Worlds. A total of 582 species were recognized by Frodin and Govaerts (2003, publ. 2004), and an additional 44 species have been published in the genus since then. Together they form a morphologically coherent group defined by a combination of palmately compound leaves with ligulate stipules, pedicels that are unarticulated below the ovary, and the absence of prickles. However, the simplicity of this generic definition conceals a complex taxonomic history (reviewed in Plunkett et al., 2005; Frodin et al., 2010) in which 19 other generic names have been regarded as synonyms of Schefflera in the circumscription adopted by Frodin (e.g., 1975, 1986, 1989, 1993, 1995), and followed by Philipson (1979, 1995) and Lowry (1989). More recently, phylogenetic analyses using DNA sequence data have revealed that this broadly delimited concept of Schefflera represents a polyphyletic group, comprising five major clades, none of which form sister pairs within Araliaceae (Plunkett et al., 2005). None of these five clades correspond to groups recognized in the various taxonomic systems of Araliaceae published over the last 150 years, but they align remarkably well with the system of subgeneric groupings proposed by Frodin (see Plunkett et al., 2005; later updated by Frodin et al., 2010) based largely on a combination of geography and a suite of morphological characters. The importance of geography is also reflected in the informal names assigned to four of the five clades: Afro-Malagasy Schefflera, Asian Schefflera, Melanesian Schefflera, and Neotropical Schefflera, and even the fifth clade, Schefflera sensu stricto (which includes the generic type), is geographically coherent, although in the Pacific islands it overlaps in part with Melanesian Schefflera. Following in-depth phylogenetic analyses of two of these groups (Plunkett & Lowry, 2012; Gostel et al., 2017), members of the Melanesian clade were placed in an expanded and recircumscribed Plerandra A. Gray (Lowry et al., 2013) and the species belonging to the Afro-Malagasy clade were transferred to Astropanax Seem. and Neocussonia (Harms) Hutch. (Lowry et al., 2017). However, species belonging to the two largest clades in the Neotropics (~400 species) and Asia (>600 species) must likewise be transferred out of Schefflera. Several years ago, we initiated a project that aims to clarify relationships among members of the Neotropical Schefflera clade and to improve our understanding of species within the group. This has involved a comprehensive review of Schefflera from the New World, including: 1) a thorough analysis of the available literature and type specimens; 2) examination of thousands of collections deposited in major herbaria with holdings from the Neotropics; 3) targeted field work focusing on the northern and central Andean countries of Bolivia, Colombia, Ecuador and Peru, where the genus is especially rich yet also poorly collected and under-studied; and 4) expanded phylogenetic analyses using sampling that is representative of the full morphological diversity and geographic range of the group. To date, this work has confirmed that, with some notable exceptions, most currently recognized species should be retained and it has also revealed that earlier projections of ca. 200 new, as-yetundescribed species are almost surely an underestimate. Moreover, two studies that focused more intensively on Neotropical Schefflera (Fiaschi & Plunkett, 2011; Plunkett et al., in press) have 2019] J. R. FORST. & G. FORST : STUDIES IN NEOTROPICAL ARALIACEAE. I. RESURRECTION... shown that the species of this group, while monophyletic, form five morphologically distinctive clades, each of which is strongly supported in the trees based on molecular evidence. Four of these clades correspond closely to morphological groupings recognized by Frodin et al. (2010), informally named Cephalopanax, Crepinella, Didymopanax, and Sciodaphyllum (the latter three names all based on previously recognized genera). The fifth clade, Gleasonia, was only recently identified on the basis of molecular evidence (Fiaschi & Plunkett, 2011). The Neotropical species of Schefflera are distributed unevenly among these five major clades. Cephalopanax and Gleasonia are the smallest, with 3 and 2 currently recognized species, respectively, whereas Crepinella has 33 currently accepted species, and 37 species were recognized in the recent revision of the Didymopanax group (Fiaschi & Plunkett, 2018). By contrast, the Sciodaphyllum clade is much larger, with 131 currently recognized species. The Cephalopanax group is largely restricted to high-elevation humid forests and subpáramo vegetation in the Andes. Each of the two species of the Gleasonia group is endemic to a single island in the greater Antilles (Hispaniola and Puerto Rico), where they grow in humid forest. Crepinella is centered in the Guiana Shield, where all but a few species are restricted to the region’s distinctive tepuis. Didymopanax occurs primarily in Brazil, Colombia, and southern Venezuela, as well as adjacent areas of Amazonian Bolivia, Ecuador and Peru, although a single widespread species, S. morototoni (Aubl.) Maguire, Steyerm. & Frodin, extends as far north as Mexico and the Caribbean basin. Didymopanax species are found at low and mid-elevation sites in both humid forests and drier environments (e.g., Brazil’s Cerrado and campos rupestres) (Fiaschi & Plunkett, 2018). The last group, Sciodaphyllum, is the most geographically widespread, with representatives occurring from low to high elevation sites, primarily in humid forests, ranging from Nicaragua in Central America and the Caribbean Islands southward as far as Bolivia, with its greatest concentration of species in the central and northern Andes. Because Schefflera must now be restricted to a small number of species in the Pacific, it can no longer accommodate the Neotropical taxa, which must be transferred elsewhere. Several options are available. The entire Neotropical clade could be treated in a single, highly diverse and rather heterogeneous genus, but doing so would obscure the significant morphological and geographic distinctions that exist between its five constituent clades. A second, more nuanced approach would similarly recognize a single genus but would formally treat each of the five clades as a subgenus, as has been done for Plerandra (Lowry et al., 2013) and also for Polyscias J.R. Forst. & G. Forst., the second largest genus of Araliaceae (Lowry & Plunkett, 2010). Based on discussions among the coauthors of this paper, primarily during a symposium organized in October, 2018, at the XII Congreso Latinoamericano de Botánica in Quito, Ecuador, we have chosen instead to adopt a third option in which each of the five clades is recognized as a separate genus. This is consistent with the strategy proposed by Lowry et al. (2013), who suggested that the delimitation of genera within Schefflera sensu lato should be based on three criteria: monophyly, morphological diagnosability, and geographic coherence, as has already been applied to the Melanesian clade (Plerandra) and the Afro-Malagasy clade (Astropanax and Neocussonia). Recognizing five genera in the Neotropics has the further advantage of maintaining a certain degree of nomenclatural stability. Generic names are already available for three of the five clades (Crepinella Marchal, Didymopanax Decne. & Planch., and Sciodaphyllum P. Browne), and combinations already exist within these three genera for 38 of their 205 currently recognized species. Moreover, once a new generic system to accommodate members of the Neotropical Schefflera clade has been adopted, the 200+ new species that have already been identified from this clade can be described and placed in well-delimited, monophyletic genera, avoiding the unnecessary publication of names in Schefflera, which would inevitably have to be transferred elsewhere at a later date. In this paper we formally resurrect the generic name Sciodaphyllum (heretofore treated in synonymy under Schefflera), which we circumscribe to include all of the Neotropical species belonging to the clade of the same name (Plunkett et al., in press), the largest of the five clades of Neotropical Schefflera. Sciodaphyllum is thus restricted to the Neotropics, and all species from elsewhere in the world that have at one time or another been included in Sciodaphyllum belong to other genera. In the present synopsis we make all of the required nomenclatural changes to insure that each accepted species has a valid name in Sciodaphyllum. In subsequent papers we will treat the members of the four other clades of BRITTONIA Neotropical Schefflera by re-establishing two genera, Crepinella (Lowry et al., 2019) and Didymopanax (Fiaschi et al., in press), and publishing two new genera to accommodate the members of the Cephalopanax and Gleasonia groups. Results In the synopsis presented below, we list all of the 131 currently recognized species of Sciodaphyllum, including 21 that already have names in the genus, 109 that require new combinations, and one that needs a replacement name. The recognition of species largely follows Frodin and Govaerts (2003, publ. 2004), with a few exceptions in cases where species they regarded as distinct are treated by us as synonyms of other taxa or, conversely, where we recognize species whose names were treated by them as synonyms. Frodin & Govaerts (2003, publ. 2004) provided a complete synonymy for each species they accepted. In the present synopsis we have therefore refrained from repeating full synonymy and only list the basionym for each new combination made, the name in Schefflera, and a limited number of other synonyms that are still being used, along with new synonyms in the few cases where we have adopted a broader delimitation than that used by Frodin & Govaerts (2003, publ. 2004). The list presented below also includes new combinations for six species described in Schefflera during the past 15 years (Ramírez-Padilla, 2004; Dorr & Stergios, 2009; Jiménez-Montoya & IdárragaPiedrahíta, 2018). In keeping with the approach adopted for the synopsis of Polyscias (Lowry et al., 2010), we have refrained from indicating types for each name treated here, primarily because many will require lectotypification, including all 31 names published by Harms, whose types were destroyed at Berlin, a situation that will necessitate extensive research in dozens of herbaria to locate original material, which is beyond the scope of this paper. Taxonomy and nomenclatural novelties Sciodaphyllum P. Browne, Civ. Nat. Hist. Jamaica, 190. 1756. Actinophyllum Ruiz & Pav., Fl. Peruv. Prodr., 51. 1794. Cotylanthes Calest., Webbia 1: 100. 1905, nom. inval. [not accepted by the author at the time of publication, cf. Art. [VOL 36.1 of the International Code of Nomenclature (Turland et al., 2018)]. Diversity and distribution.—A genus of 131 currently recognized species and more than 200 new species that remain to be described, more than 50 of which are currently being prepared for publication (see Mora et al., in press). Sciodaphyllum is restricted to the Neotropics, where it ranges from Nicaragua to Bolivia and is also present in the Lesser Antilles, with members occurring in a wide range of humid or seasonally moist habitats, from sea level to nearly 3500 m elevation. The number of species of Sciodaphyllum varies significantly from country to country. Table 1 indicates the number of currently recognized species recorded from each country (or island) in which the genus is known. Colombia has the greatest number of species of Sciodaphyllum (50), followed by Venezuela (26), Ecuador (23), Panama (21), Peru (19) and Bolivia (10). The remaining countries and islands have fewer than 10 representatives of the genus. It is not surprising that Colombia and Venezuela top this list because detailed taxonomic work has been conducted on the group relatively recently in both countries. In Colombia, José Cuatrecasas published 28 species of Schefflera that are now included in Sciodaphyllum (Cuatrecasas 1946, 1951), and several additional species have been described over the past 15 years (Ramírez-Padilla, 2004; Jiménez-Montoya & Idárraga-Piedrahíta, 2018, 2019). A detailed treatment of Araliaceae in Venezuela resulted in the description of seven new species of Schefflera, published by Maguire et al. (1984), after which Frodin (1993) added two more species. Similarly, the comparatively large number of species in Panama (21) and to a lesser extent Costa Rica (9) reflects the fact that a comprehensive revision of Araliaceae, including the description of 16 new species now placed in Sciodaphyllum, was published by Cannon and Cannon (1989) in preparation for their treatment of the family for Flora Mesoamericana (Cannon & Cannon 2009). By contrast, much less work has been published on Araliaceae in the other countries where Sciodaphyllum has been recorded. It is thus noteworthy that Ecuador ranks third in terms of species richness, with 23 currently recognized species, despite very little taxonomic work on the group in this country. Colombia and Venezuela also top the list of countries with the largest number of endemic 2019] J. R. FORST. & G. FORST : STUDIES IN NEOTROPICAL ARALIACEAE. I. RESURRECTION... TABLE 1. NUMBER OF CURRENTLY RECOGNIZED SPECIES OF SCIODAPHYLLUM PER COUNTRY OR ISLAND AND NUMBER OF ENDEMIC SPECIES PER COUNTRY/ISLAND. Country Colombia Venezuela Ecuador Panama Peru Bolivia Costa Rica Brazil Guyana Jamaica Martinique Dominica Grenada Guadeloupe Montserrat Nicaragua Trinidad Number of currently recognized species of Sciodaphyllum Number of country/island endemics 50 26 23 21 19 10 9 6 3 2 2 1 1 1 1 1 1 35 19 3 13 11 8 3 0 1 2 1 0 0 0 0 0 0 species of Sciodaphyllum (Table 1), containing 35 and 20 endemics, respectively, followed by Panama with 13, again reflecting the fact that the group has been studied recently in these countries. By contrast, only three currently recognized species are endemic to Ecuador, in part reflecting the fact that many species of Sciodaphyllum in this rather small country (by South American standards) also occur in Colombia to the north or Peru to the south. The number of endemics in Peru (11) and Bolivia (10) is also quite small given their comparatively large areas. However, the true level of species diversity in these three countries, as well as in Colombia, is clearly much higher as many additional species with restricted ranges remain to be described. A careful review of specimens at herbaria within each of these countries, as well as at institutions in North America with significant holdings from the Neotropics, has revealed at least 50 well delimited new species from Ecuador, 30 from Colombia, and 15 from Peru, the first of which are now being published (Mora et al., in press). Herbarium work has not yet been undertaken in Bolivia, but it seems certain that many novelties will be found there as well. Notes.— When Browne described Sciodaphyllum in The Civil and Natural History of Jamaica (1756), he did not include the type of any previously or simultaneously published names, nor did he mention any species. Indeed, no binomials were published in Sciodaphyllum until nearly 70 years later when Sprengle (1824, publ. 1825) published S. brownei Spreng. Browne did, however, provide an illustration of his new genus, which clearly represents original material, and an uncited Swartz collection from Jamaica deposited at the Swedish Museum of Natural History (S) may also be original material. According to Article 10.4 of the International Code of Nomenclature (Turland et al., 2018), the type of a name of a genus may be a specimen or illustration, preferably used by the author in the preparation of the protologue, other than the type of a name of an included species, by and only by conservation. Since no species were included in Sciodaphyllum by Browne, it will therefore be necessary to publish a formal proposal in Taxon to typify the generic name by conservation, and to have that proposal approved at the next International Botanical Congress, which we are now proceeding to initiate. The generic name Schefflera J.R. Forst. & G. Forst., published in 1776, was conserved against the older name Sciodaphyllum, which dates from 1756, based on a proposal by Smith (1941) that was adopted in 1950. However, with the reinstatement of Sciodaphyllum and the transfer of its members out of Schefflera, conservation of the former name no longer comes into play. Sciodaphyllum acuminatum (Pav.) Poir. in J.B.A.M. de Lamarck, Encycl. 6: 746 (1805). Actinophyllum acuminatum Pav., Mem. Real Acad. Méd. Madrid 1: 196 (1797). Schefflera BRITTONIA [VOL acuminata (Pav.) Harms in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. 3(8): 36 (1894). Peru. (Dominica, Grenada, Guadeloupe, Martinique, Montserrat). Schefflera microcephala Harms, Bot. Jahrb. Syst. 42: 148 (1908), syn. nov. Sciodaphyllum awa (Ram.-Padilla) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera awa Ram.-Padilla, Revista Acad. Colomb. Ci. Exact. 28: 483. 2004. Colombia and Ecuador. Sciodaphyllum albocapitatum (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera albocapitata M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 54 (1989). Panama. Sciodaphyllum allocotanthum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Didymopanax allocotanthus Harms, Notizbl. Bot. Gart. Berlin-Dahlem 11: 292 (1931). Schefflera allocotantha (Harms) Frodin in D. Frodin, R. Govaerts et al., World Checklist Bibliog. Araliaceae: 322 (2003 publ. 2004). Bolivia. Sciodaphyllum angulatum (Pav.) Poir. in J.B.A.M. de Lamarck, Encycl. 6: 745 (1805). Schefflera angulata (Pav.) Harms in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. 3(8): 36 (1894). Ecuador and Peru. Sciodaphyllum aquaverense (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera aquavarensis M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 54 (1989). Panama. Sciodaphyllum archeri (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera archeri Harms, Notizbl. Bot. Gart. BerlinDahlem 13: 446 (1937). Panama and Colombia. Sciodaphyllum ayangannense (Maguire, Steyerm. & Frodin) Lowry, G.M. Plunkett & M . M . M o r a , c o m b . n o v. S c h e f f l e r a ayangannensis Maguire, Steyerm. & Frodin, Mem. New York Bot. Gard. 38: 63 (1984). Guyana. Sciodaphyllum bangii (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera bangii Harms, Notizbl. Bot. Gart. BerlinDahlem 11: 285 (1931). Bolivia. Sciodaphyllum bejucosum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera bejucosa Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 545 (1946). Colombia. Sciodaphyllum bifidum (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera bifida M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 51 (1989). Panama. Sciodaphyllum bifurcatum (Idárraga & JiménezMont.) Idárraga & Jiménez-Mont., comb. nov. Schefflera bifurcata Idárraga & Jiménez-Mont., Brittonia 70: 317. 2018. Colombia. Sciodaphyllum argophyllum (Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera argophylla Frodin, Novon 3: 369 (1993). Venezuela. Sciodaphyllum blepharidophyllum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera blepharidophylla Harms, Notizbl. Bot. Gart. Berlin-Dahlem 13: 447 (1937). Panama, Colombia and Ecuador. Sciodaphyllum asymmetricum (Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera asymmetrica Frodin, Novon 3: 371 (1993). Brazil, Venezuela. Sciodaphyllum bogotense (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera bogotensis Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 544 (1946). Colombia. Sciodaphyllum attenuatum (Sw.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Panax attenuatus Sw., Prodr.: 54 (1788). Schefflera attenuata (Sw.) Frodin, Proc. Acad. Nat. Sci. Philadelphia 141: 315 (1989). Lesser Antilles Sciodaphyllum bonitum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera bonita Cuatrec., Revista Acad. Colomb. Ci. Exact. 8: 313. 1951. Colombia. 2019] J. R. FORST. & G. FORST : STUDIES IN NEOTROPICAL ARALIACEAE. I. RESURRECTION... Sciodaphyllum breviramum (Jiménez-Mont. & Idárraga) Jiménez-Mont. & Idárraga, comb. nov. Schefflera brevirama Jiménez-Mont. & Idárraga, Brittonia 70: 319. 2018. Colombia. Sciodaphyllum brownei Spreng., Syst. Veg. 1: 953 (1824). Aralia sciodaphyllum Sw., Prodr.: 55 (1788). Schefflera sciodaphyllum (Sw.) Harms in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. 3(8): 37 (1894). Jamaica. Schefflera stearnii R.A. Howard & Proctor, J. Arnold Arbor. 39: 105 (1958). Sciodaphyllum buchtienii (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera buchtienii Harms, Notizbl. Bot. Gart. BerlinDahlem 11: 286 (1931). Bolivia. Sciodaphyllum caducum (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera caduca M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 47 (1989). Panama. Sciodaphyllum cajambrense (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera cajambrensis Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 539 (1946). Colombia. Sciodaphyllum calycinum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera calycina Cuatrec., Revista Acad. Colomb. Ci. Exact. 8: 312 (1951). Colombia. Sciodaphyllum calyptricuspidatum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera calyptricuspidata Cuatrec., Revista Acad. Colomb. Ci. Exact. 8: 314 (1951). Colombia. Sciodaphyllum capitulispicatum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera capitulispicata Cuatrec., Revista Acad. Colomb. Ci. Exact. 8: 312 (1951). Colombia and Ecuador. Sciodaphyllum cartagoense (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera cartagoensis M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 54 (1989). Costa Rica. Sciodaphyllum chartaceum A.C. Sm., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1562 (1938). Schefflera brenesii A.C. Sm., Trop. Woods 66: 5 (1941), non Schefflera chartacea Merr. Costa Rica and Panama. Sciodaphyllum chococolum (Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera chococola Frodin in D. Frodin, R. Govaerts et al., World Checklist Bibliog. Araliaceae: 331 (2003 publ. 2004). Schefflera macrophylla Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 540 (1946), nom. illeg., non Schefflera macrophylla (Dunn) R.Vig. Colombia and Ecuador. Sciodaphyllum cicatricatum (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera cicatricata M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 58 (1989). Panama. Sciodaphyllum ciliatum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera ciliata Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 539 (1946). Colombia. Sciodaphyllum clausum (Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera clausa Frodin, Novon 3: 376 (1993). Venezuela. Sciodaphyllum coclense (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera coclensis M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 32 (1989). Panama. Sciodaphyllum concolor (Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera concolor Frodin, Novon 3: 379 (1993). Venezuela (Amazonas), Brazil. Sciodaphyllum connatum (Jiménez-Mont. & Idárraga) Jiménez-Mont. & Idárraga, comb. nov. Schefflera connata Jiménez-Mont. & Idárraga, Brittonia 70: 321. 2018. Colombia. Sciodaphyllum contractum (Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera contracta Frodin, Novon 3: 380 (1993). Venezuela. BRITTONIA Sciodaphyllum cracens (Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera cracens Frodin, Novon 3: 381 (1993). Venezuela. Sciodaphyllum crassilimbum (Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera crassilimba Frodin, Novon 3: 381 (1993). Venezuela. Schefflera guanayensis subsp. sipapoensis Maguire, Steyerm. & Frodin, Mem. New York Bot. Gard. 38: 67 (1984). Sciodaphyllum decagynum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera decagyna Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 545 (1946). Colombia. Sciodaphyllum dielsii (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera dielsii Harms, Notizbl. Bot. Gart. BerlinDahlem 12: 693 (1935). Ecuador and Peru. Sciodaphyllum diguanum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera diguana Cuatrec., Revista Acad. Colomb. Ci. Exact. 8: 314 (1951). Colombia. Sciodaphyllum digynum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera digyna Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 539 (1946). Colombia. Sciodaphyllum diplodactylum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera diplodactyla Harms, Notizbl. Bot. Gart. Berlin-Dahlem 14: 337 (1939). Colombia, Ecuador and Peru. Sciodaphyllum dolichostylum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera dolichostyla Harms, Bot. Jahrb. Syst. 42: 152 (1908). Peru. Sciodaphyllum duidae (Steyerm.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera duidae Steyerm., Fieldiana, Bot. 28: 444 (1952). Venezuela, Brazil. Sciodaphyllum elachistocephalum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera elachistocephala Harms, [VOL Repert. Spec. Nov. Regni Veg. 43: 110 (1938). Colombia. Sciodaphyllum epiphyticum (A.C. Sm.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera epiphytica A.C. Sm., Ann. Missouri Bot. Gard. 28: 437 (1941). Costa Rica, Panama, Colombia and Ecuador. Sciodaphyllum euryphyllum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera euryphylla Harms, Bot. Jahrb. Syst. 42: 151 (1908). Peru. Sciodaphyllum ferrugineum (Willd. ex Schult.) Decne. & Planch., Rev. Hort., IV, 3: 107 (1854). Aralia ferruginea Willd. ex Schult., Syst. Veg. 6: 71 (1820). Schefflera ferruginea (Willd. ex Schult.) Harms in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. 3(8): 36 (1894). Colombia and Ecuador. Sciodaphyllum fragrans (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera fragrans Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 537 (1946). Colombia. Sciodaphyllum glabratum (Kunth) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Panax glabratus Kunth in F.W.H. von Humboldt, A.J.A. Bonpland & C.S. Kunth, Nov. Gen. Sp. 5: 10 (1821). Schefflera glabrata (Kunth) Frodin, Proc. Acad. Nat. Sci. Philadelphia 141: 316 (1989). Trinidad, Venezuela. Sciodaphyllum guanayense (Maguire, Steyerm. & Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera guanayensis Maguire, Steyerm. & Frodin, Mem. New York Bot. Gard. 38: 66 (1984). Venezuela. Sciodaphyllum herthae (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera herthae Harms, Notizbl. Bot. Gart. BerlinDahlem 14: 338 (1939). Ecuador. Sciodaphyllum herzogii (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera herzogii Harms, Meded. Rijks-Herb. 29: 6 (1916). Bolivia. Sciodaphyllum heterotrichum Seem., J. Bot. 3: 268 (1865). Schefflera heterotricha (Seem.) R. 2019] J. R. FORST. & G. FORST : STUDIES IN NEOTROPICAL ARALIACEAE. I. RESURRECTION... Vig., Ann. Sci. Nat., Bot., IX, 9: 343 (1909). Colombia. Schefflera fontiana Cuatrec., Collect. Bot. (Barcelona) 7: 223 (1968). Sciodaphyllum huilense (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera huilensis Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 545 (1946). Colombia. Sciodaphyllum humboldtianum Decne. & Planch. ex Seem., J. Bot. 3: 268 (1865). Schefflera humboldtiana (Decne. & Planch. ex Seem.) Frodin in D. Frodin, R. Govaerts et al., World Checklist Bibliog. Araliaceae: 346 (2003 publ. 2004). Colombia. Sciodaphyllum inambaricum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera inambarica Harms, Bot. Jahrb. Syst. 42: 150 (1908). Peru. Sciodaphyllum institum (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera instita M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 58 (1989). Costa Rica. Sciodaphyllum jauaense (Maguire, Steyerm. & Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera reticulata subsp. jauaensis Maguire, Steyerm. & Frodin, Mem. New York Bot. Gard. 38: 59 (1984). Schefflera jauaensis (Maguire, Steyerm. & Frodin) Frodin, Novon 3: 389 (1993). Venezuela. Sciodaphyllum jefense (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera jefensis M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 43 (1989). Panama. Sciodaphyllum karstenianum Marchal, Bull. Acad. Roy. Sci. Belgique, II, 47: 93 (1879). Schefflera karsteniana (Marchal) Harms in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. 3(8): 36 (1894). Venezuela. Sciodaphyllum kuntzei (Harms ex Kuntze) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Didymopanax kuntzei Harms ex Kuntze, Revis. Gen. Pl. 3(2): 115 (1898). Schefflera kuntzei (Harms ex Kuntze) Frodin, Monogr. Syst. Bot. Missouri Bot. Gard. 45: 1253 (1993). Bolivia and Peru. Sciodaphyllum lancifoliolatum (Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera lancifoliolata Frodin in D. Frodin, R. Govaerts et al., World Checklist Bibliog. Araliaceae: 348 (2003 publ. 2004). Schefflera lanceolata Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 544 (1946), nom. illeg., non S. lanceolata Ridl. (1920). Colombia. Sciodaphyllum lasiogyne (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera lasiogyne Harms, Repert. Spec. Nov. Regni Veg. 15: 345 (1918). Colombia, Ecuador. Sciodaphyllum latiligulatum (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera latiligulata M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 43 (1989). Panama. Notes.—Frodin & Govaerts (2003, publ. 2004) treated Schefflera latiligulata as a synonym of S. sphaerocoma, but it can easily be distinguished by its strongly conduplicate leaflets with impressed secondary venation on the adaxial surface. Sciodaphyllum lilacinum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera lilacina Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 543 (1946). Colombia. Sciodaphyllum macphersonii (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera macphersonii M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 34 (1989). Panama. Sciodaphyllum magnifolium (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera magnifolia Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 544 (1946). Colombia and Ecuador. Sciodaphyllum maguireorum (Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. BRITTONIA [VOL Schefflera maguireorum Frodin, Novon 3: 391 (1993). Venezuela. Revista Acad. Colomb. Ci. Exact. 28: 482. 2004. Colombia. Sciodaphyllum manus-dei (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera manus-dei Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 540 (1946). Colombia. Sciodaphyllum nebularum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Didymopanax nebularum Harms, Notizbl. Bot. Gart. Berlin-Dahlem 11: 293 (1931). Schefflera nebularum (Harms) Frodin in D. Frodin, R. Govaerts et al., World Checklist Bibliog. Araliaceae: 360 (2003 publ. 2004). Bolivia. Sciodaphyllum marahuacense (Maguire, Steyerm. & Frodin) Lowry, G.M. Plunkett & M . M . M o r a , c o m b . n o v. S c h e f f l e r a tremuloidea var. marahuacensis Maguire, Steyerm. & Frodin, Mem. New York Bot. Gard. 38: 63 (1984). Schefflera marahuacensis (Maguire, Steyerm. & Frodin) Frodin, Novon 3: 392 (1993). Venezuela. Sciodaphyllum marginatum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera marginata Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 538 (1946). Colombia. Sciodaphyllum mathewsii Seem., J. Bot. 3: 268 (1865). Schefflera mathewsii (Seem.) Harms in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. 3(8): 36 (1894). Peru. Sciodaphyllum meiurophyllum (Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera meiurophylla Frodin, Novon 3: 393 (1993). Venezuela. Sciodaphyllum minutiflorum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera minutiflora Harms, Bot. Jahrb. Syst. 42: 153 (1908). Colombia, Ecuador and Peru. Sciodaphyllum monospermum (Maguire, Steyerm. & Frodin ex Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera monosperma Maguire, Steyerm. & Frodin ex Frodin, Novon 3: 393 (1993). Brazil, Guyana and Venezuela. Sciodaphyllum monzonense (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera monzonensis Harms, Bot. Jahrb. Syst. 42: 154 (1908). Peru. Sciodaphyllum munchiquense (Ram.-Padilla) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera munchiquensis Ram.-Padilla, Sciodaphyllum nephelophilum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera nephelophila Harms, Notizbl. Bot. Gart. Berlin-Dahlem 11: 286 (1931). Bolivia. Sciodaphyllum octostylum (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera octostyla M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 38 (1989). Panama. Sciodaphyllum panamense (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera panamensis M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 34 (1989). Panama. Sciodaphyllum paniculitomentosum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera paniculitomentosa Cuatrec., Revista Acad. Colomb. Ci. Exact. 8: 315 (1951). Colombia. Sciodaphyllum pardoanum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera pardoana Harms, Bot. Jahrb. Syst. 42: 149 (1908). Peru. Sciodaphyllum paruanum (Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera paruana Frodin, Novon 3: 397 (1993). Venezuela. Sciodaphyllum patulum Rusby, Mem. Torrey Bot. Club 3(3): 41 (1893). Schefflera patula (Rusby) Harms in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. 3(8): 37 (1894). Ecuador, Peru and Bolivia. Schefflera moyobambae Harms, Bot. Jahrb. Syst. 42: 154 (1908). 2019] J. R. FORST. & G. FORST : STUDIES IN NEOTROPICAL ARALIACEAE. I. RESURRECTION... Schefflera yuncacoyae Harms, Bot. Jahrb. Syst. 42: 155 (1908). Schefflera stilpnophylla Harms, Notizbl. Bot. Gart. BerlinDahlem 15: 52 (1940), syn. nov. Notes.—Frodin & Govaerts (2003, publ. 2004) recognized Schefflera stilpnophylla as a distinct species, but careful examination of the isotype at MO (the holotype at B was destroyed) as well as recent collections made in the same part of eastern Ecuador reveal that, while these plants are somewhat more robust than typical material of Sciodaphyllum patulum from southern Ecuador, Peru and northwestern Bolivia, they fall within the range of morphologies documented in these areas and nearly match several collections from central Peru. Sciodaphyllum pedicellatum (Pav.) Poir. in J.B.A.M.de Lamarck, Encycl. 6: 746 (1805). Actinophyllum pedicellatum Pav., Mem. Real Acad. Méd. Madrid 1: 195 (1797). Schefflera pedicellata (Pav.) Harms in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. 3(8): 37 (1894). Ecuador and Peru. Sciodaphyllum pedicelligerum (Maguire, Steyerm. & Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera pedicelligera Maguire, Steyerm. & Frodin, Mem. New York Bot. Gard. 38: 63 (1984). Venezuela. Sciodaphyllum pentadactylum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera pentadactyla Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 537 (1946). Colombia and Ecuador. Sciodaphyllum pentandrum (Pav.) Poir. in J.B.A.M. de Lamarck, Encycl. 6: 747 (1895). Actinophyllum pentandrum Pav., Mem. Real Acad. Méd. Madrid 1: 197 (1797). Schefflera pentandra (Pav.) Harms in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. 3(8): 36 (1894). Ecuador and Peru. Sciodaphyllum peruvianum (Aspl.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera peruviana Aspl., Svensk Bot. Tidskr. 26: 193 (1932). Peru. Sciodaphyllum pittieri (Marchal) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Didymopanax pittieri Marchal ex T. Durand & Pittier, Bull. Acad. Roy. Sci. Belgique 30: 280 (1891 publ. 1892). Schefflera rodriguesiana Frodin, Proc. Acad. Nat. Sci. Philadelphia 141: 317 (1989). Costa Rica and Panama. Sciodaphyllum planchonianum Marchal, Bull. Soc. Roy. Bot. Belgique 19: 92 (1880). Schefflera planchoniana (Marchal) Harms in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. 3(8): 36 (1894). Ecuador. Sciodaphyllum pubens (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera pubens M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 51 (1989). Costa Rica. Sciodaphyllum quindiuense (Kunth) DC., Prodr. 4: 261 (1830). Aralia quindiuensis Kunth in F.W.H. von Humboldt, A.J.A. Bonpland & C.S. Kunth, Nov. Gen. Sp. 5: 8 (1821). Schefflera quindiuensis (Kunth) Harms in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. 3(8): 37 (1894). Colombia. Sciodaphyllum ramosissimum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera ramosissima Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 545 (1946). Colombia. Sciodaphyllum reticulatum (Gleason) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Didymopanax reticulatus Gleason, Bull. Torrey Bot. Club 58: 435 (1931). Schefflera reticulata (Gleason) Maguire, Steyerm. & Frodin, Mem. New York Bot. Gard. 38: 57 (1984), nom. illeg., non S. reticulata Philipson. Schefflera sessiliflora Frodin, Novon 3: 398 (1993). Guyana, Venezuela. Sciodaphyllum robustum A.C. Sm., Brittonia 2: 254 (1936). Schefflera robusta (A.C. Sm.) A.C. Sm., Trop. Woods 66: 5 (1941). Costa Rica and Panama. Sciodaphyllum rubiginosum (Marchal) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Didymopanax rubiginosus Marchal in C.F.P. von Martius, Fl. Bras. 11(1): 234 (1878). BRITTONIA Schefflera rufa Frodin in D. Frodin, R. Govaerts et al., World Checklist Bibliog. Araliaceae: 372 (2003 publ. 2004). Venezuela. Sciodaphyllum sachamatense (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera sachamatensis Cuatrec., Revista Acad. Colomb. Ci. Exact. 8: 315 (1951). Colombia and Ecuador. Sciodaphyllum samarianum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera samariana Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 538 (1946). Colombia. Sciodaphyllum sandianum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera sandiana Harms, Bot. Jahrb. Syst. 42: 153 (1908). Peru. Sciodaphyllum sapoense (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera sapoensis M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 38 (1989). Panama. Sciodaphyllum sararense (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera sararensis Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 544 (1946). Colombia. Sciodaphyllum seibertii (A.C. Sm.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera seibertii A.C. Sm., Trop. Woods 66: 5 (1941). Costa Rica and Panama. Notes.—Frodin & Govaerts (2003, publ. 2004) treated Schefflera seibertii as a synonym of S. sphaerocoma (Benth.) Harm s [ ≡ Sciodaphyllum sphaercomum Benth.], but it clearly differs in having acuminate (vs. rounded) corollas in bud. Sciodaphyllum silvaticum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera silvatica Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 540 (1946). Colombia. Sciodaphyllum sipapoense (Maguire, Steyerm. & Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera sipapoensis [VOL Maguire, Steyerm. & Frodin, Mem. New York Bot. Gard. 38: 59 (1984). Venezuela. Sciodaphyllum sodiroi (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera sodiroi Harms, Repert. Spec. Nov. Regni Veg. 15: 246 (1918). Ecuador. Sciodaphyllum sonsonense Jiménez-Mont., Caldasia 41: 314 (2019). Colombia. Sciodaphyllum sphaerocomum Benth., Bot. Voy. Sulphur: 102 (1844). Schefflera sphaerocoma (Benth.) Harms in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. 3(8): 37 (1894). Nicaragua, Costa Rica, Panama, Colombia and Ecuador. Sciodaphyllum lehmannii Harms, Bot. Jahrb. Syst. 20 (49): 69 (1895). Schefflera lehmannii (Harms) Harms, Bot. Jahrb. Syst. 20 (49): 69 (1895). Sciodaphyllum nicaraguense Standl., J. Wash. Acad. Sci. 17: 316 (1927). Schefflera nicaraguensis (Standl.) A.C. Sm., Trop. Woods 66: 5 (1941). Schefflera minutiflora Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 543 (1946), nom. illeg., non S. minutiflora Harms (1908). Schefflera multiflora Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 542 (1946), nom. illeg., non S. minutiflora Merr. (1915). Schefflera sanquininensis Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 542 (1946), syn. nov. Sciodaphyllum sprucei Seem., J. Bot. 3: 268 (1865). Brassaia sprucei (Seem.) Hutch., Gen. Fl. Pl. 2: 623 (1967). Schefflera sprucei (Seem.) Harms in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. 3(8): 36 (1894). Brazil, Ecuador and Peru. Sciodaphyllum steyermarkii (Frodin) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera steyermarkii Frodin, Novon 3: 399 (1993). Venezuela. Sciodaphyllum systylum Donn. Sm., Bot. Gaz. 31: 113 (1901). Schefflera systyla (Donn. Sm.) R.Vig., Ann. Sci. Nat., Bot., IX, 9: 363 (1909). Costa Rica and Panama. Sciodaphyllum tamanum (Steyerm.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. 2019] J. R. FORST. & G. FORST : STUDIES IN NEOTROPICAL ARALIACEAE. I. RESURRECTION... Schefflera tamana Steyerm., Fieldiana, Bot. 28: 444 (1952). Venezuela. Sciodaphyllum ternatum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera ternata Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 546 (1946). Colombia and Ecuador. Sciodaphyllum tipuanicum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera tipuanica Harms, Notizbl. Bot. Gart. Berlin-Dahlem 11: 287 (1931). Bolivia. Sciodaphyllum tremuloideum (Maguire, Steyerm. & Frodin) Lowry, G.M. Plunkett & M . M . M o r a , c o m b . n o v. S c h e f f l e r a tremuloidea Maguire, Steyerm. & Frodin, Mem. New York Bot. Gard. 38: 60 (1984). Brazil, Venezuela. Sciodaphyllum trianae Planch. & Linden ex Marchal, Bull. Acad. Roy. Sci. Belgique, II, 47: 94 (1879). Schefflera trianae (Planch. & Linden ex Marchal) Harms in H.G.A. Engler & K.A.E. Prantl, Nat. Pflanzenfam. 3(8): 36 (1894). Colombia. Sciodaphyllum trollii (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera trollii Harms, Notizbl. Bot. Gart. BerlinDahlem 11: 287 (1931). Bolivia. Sciodaphyllum troyanum Urb., Symb. Antill. 5: 451 (1908). Schefflera troyana (Urb.) A.C. Sm., Trop. Woods 66: 5 (1941). Jamaica. Sciodaphyllum urbanianum (Marchal ex. Urb.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Didymopanax urbanianus Marchal ex Urb., Bot. Jahrb. Syst. 15: 326 (1893). Schefflera urbaniana (Marchal ex Urb.) Frodin, Proc. Acad. Nat. Sci. Philadelphia 141: 318 (1989). Martinique. Sciodaphyllum vanderwerffii (Dorr & Stergios) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera vanderwerffii Dorr & Stergios, J. Bot. Res. Inst. Texas 3: 605 (2009). Venezuela. Sciodaphyllum vasquezianum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera vasqueziana Harms, Notizbl. Bot. Gart. Berlin-Dahlem 13: 448 (1937). Colombia. Sciodaphyllum velutinum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera velutina Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 538 (1946). Colombia. Sciodaphyllum venezuelense Lowry, G.M. Plunkett & M.M. Mora, nom. nov. Schefflera pittieri Harms, Repert. Spec. Nov. Regni Veg. 23: 299 (1927), non Sciodaphyllum pittieri (Marchal) Lowry, G.M. Plunkett & M.M. Mora (see above). Venezuela. Notes.—A replacement name (nomen novum) is required for Schefflera pittieri Harms because the combination in Sciodaphyllum has been used above for another species based on Didymopanax pittieri Marchal ex T. Durand & Pittier. Sciodaphyllum viguierianum (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera viguieriana Harms, Bot. Jahrb. Syst. 42: 150 (1908). Peru. Sciodaphyllum violaceum (Cuatrec.) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera violacea Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 543 (1946). Colombia. Notes.—Frodin & Govaerts (2003, publ. 2004) indicated that this species occurs in Ecuador, but we adopt a narrower circumscription that only includes material from Colombia. Sciodaphyllum weberbaueri (Harms) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera weberbaueri Harms, Bot. Jahrb. Syst. 42: 151 (1908). Peru. Sciodaphyllum whitefoordiae (M.J. Cannon & Cannon) Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera whitefoordiae M.J. Cannon & Cannon, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 47 (1989). Panama. Sciodaphyllum yurumanguine (Cuatrec.) BRITTONIA Lowry, G.M. Plunkett & M.M. Mora, comb. nov. Schefflera yurumanguinis Cuatrec., Revista Acad. Colomb. Ci. Exact. 6: 539 (1946). Colombia. Acknowledgments As this paper was being finalized, our coauthor, friend and colleague David Frodin passed away. His vast knowledge of Schefflera and his never-ending enthusiasm and willingness to share his insights will be missed by us all. The authors thank the curators of the following herbaria for the loan of specimens or for providing access to their collections: CAUP, COL, CUVC, CUZ, F, FMB, HUA, ICESI, JAUM, K, LOJA, MEDEL, MO, MOL, NY, PSO, QCA, QCNE, VALLE, and USM. We are also grateful to Roy Gereau for advice on several matters of typification, and George Schatz for interesting discussions on delimiting genera in the Neotropical Schefflera clade. This work was supported by grants from the U.S. National Science Foundation (DEB 0943958, 1556327 and 1556139) and the National Geographic Society (CRE 5793-96, 7415-03, 8355-07). PF thanks the Brazilian National Council for Scientific and Technological Development (CNPq) for a productivity grant (PQ 306228/2016-5). Literature cited Browne, P. 1756. The civil and natural history of Jamaica. London. Cannon, M. J. & J. F. M. Cannon. 1989. Central American Araliaceae – a precursory study for the Flora Mesoamericana. Bulletin of the British Museum (Natural History). 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