Evolutionary Relationships in Afro-Malagasy Schefflera (Araliaceae) Based on Nuclear and Plastid Markers

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Theses and Dissertations Graduate School

2010

Evolutionary relationships in Afro-Malagasy Schefflera (Araliaceae) based on nuclear and plastid markers

Morgan Gostel Virginia Commonwealth University

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EVOLUTIONARY RELATIONSHIPS IN AFRO-MALAGASY SCHEFFLERA

(ARALIACEAE) BASED ON NUCLEAR AND PLASTID MARKERS

A thesis submitted in partial fulfillment of the requirements for the degree of M.S. Biology at Virginia Commonwealth University.

MORGAN ROBERT GOSTEL B.S. Biology, Virginia Commonwealth University, 2008

Director: DR. GREGORY M. PLUNKETT AFFILIATE RESEARCH PROFESSOR, DEPARTMENT OF BIOLOGY, VIRGINIA COMMONWEALTH UNIVERSITY AND DIRECTOR, CULLMAN PROGRAM FOR MOLECULAR SYSTEMATICS, THE NEW YORK BOTANICAL GARDEN

Co-Director: DR. RODNEY J. DYER ASSOCIATE PROFESSOR, DEPARTMENT OF BIOLOGY

Virginia Commonwealth University Richmond, Virginia July 2010

iii

Acknowledgements

I have been tremendously fortunate in my life to be taught by truly gifted teachers – assets that are simultaneously the most important and undervalued in our world. I would like to extend my deepest gratitude to my friend and advisor, Dr. Gregory M. Plunkett, who has taught me that patience and diligence together with enthusiasm are necessary to pursue what we are most passionate about and who has provided me with the most exciting opportunities in my life.

Thanks also to Dr. Rodney J. Dyer who was kind enough to offer me a good home in his lab here at VCU which provided me with the workspace I needed to complete this project and good laughter and spirits. I also thank the rest of my committee, Dr. James M. Turbeville and Dr.

David J. Edwards for their time and assistance throughout this thesis. Special thanks go to Dr.

Porter P. Lowry II of the Missouri Botanical Garden for sharing his expertise in Araliaceae and the flora of Madagascar in general throughout this project while in the herbarium in Paris, during fieldwork in Madagascar, and remotely via telephone or email and also for providing his home for lodging during my work in Paris. Additional thanks to Dr. Antoine N. Nicolas for sharing his laboratory expertise and his assistance in the lab. I would also like to thank Bernard J. De

Villiers from the University of Johannesburg for providing me with African materials and Carlos

Rodrigues at the Campus Universitaire de Jussieu in Paris for his assistance with herbarium materials at the Museé Nationale d’Histoire Naturelle. Thanks to Missouri Botanical Garden employees in Madagacar for their assistance in field collections, including Richard

Razakamalala, Charles Rakotovao, Desiré Randrianasolo, Patrice Andrianjafy, Roger Bernard,

Ornella, Jaqueline, and Roger. Institutional thanks to the New York Botanical Garden, the

Missouri Botanical Garden, the Museé Nationale d’Histoire Naturelle in Paris, the Jardin

Botanique Nationale de Belgique, and the National Science Foundation for funding. Finally, I would like to thank my friends and family for their continuing love and support.

Table of Contents

Acknowledgements...... ii

List of Tables...... v

List of Figures...... vi

Abstract ...... 1

Introduction ...... 3

Materials and Methods...... 10

Results...... 14

Discussion ...... 18

Literature Cited...... 28

Tables...... 34

Figures...... 40

Vita...... 66

List of Tables

Table 1: Species used for DNA samples used in this study, including geographic range and

voucher information ...... 34

Table 2: Oligonucleotide primers used for PCR amplification and DNA sequencing...... 39

vii

List of Figures

Figure Legends: ...... 40

Figure 1: The strict consensus of 51,600 trees resulting from the maximum parsimony (MP) analysis of 123 sequences from the nuclear ETS rDNA spacer. Tree length = 279 steps, CI =

0.675, RI = 0.954. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. Parsimony bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as “<”...... 45

Figure 2: The strict consensus of 96,600 trees resulting from the maximum parsimony (MP) analysis of 123 sequences from the nuclear ITS rDNA spacer. Tree length = 271 steps,

CI = 0.708, RI = 0.959. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. Parsimony bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as “<”...... 46

Figure 3: The strict consensus of 100,000 trees resulting from the maximum parsimony (MP) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers. Tree length =

565 steps, CI = 0.665, and RI = 0.951. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. Parsimony bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as “<”...... 47

Figure 4: The strict consensus of 87,700 trees resulting from the maximum parsimony (MP) analysis of 123 sequences from the combined plastid markers, trnK-rps16, rpl32-trnL, and ndhF- rpl32. Tree length = 655 steps, CI = 0.612, RI = 0.918. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. Parsimony bootstrap percentages are

viii provided above the branches; bootstrap values less than 50% are recorded as “<”. Gray arrows indicate specimens whose placement has moved considerably from other analyses ...... 48

Figure 5: The strict consensus of 94,200 trees resulting from the maximum parsimony (MP) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers and combined plastid markers, trnK-rps16, rpl32-trnL, and ndhF-rpl32. Tree length = 1,238 steps, CI = 0.608,

RI = 0.929. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. Parsimony bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as “<”...... 49

Figure 6: The best tree (log likelihood = –2,279.056) based on maximum likelihood (ML) analysis of 123 sequences from the nuclear ETS rDNA spacer. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. ML bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as “<”...... 50

Figure 7: The best tree (log likelihood = –2,612.7117) based on maximum likelihood (ML) analysis of 123 sequences from the nuclear ITS rDNA spacer. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. ML bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as “<”...... 51

Figure 8: The best tree (log likelihood = –5,075.1868) based on maximum likelihood (ML) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. ML bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as

“<”...... 52

Figure 9: The best tree (log likelihood = –8,530.1115) based on maximum likelihood (ML) analysis of 123 sequences from the combined plastid markers, trnK-rps16, rpl32-trnL, and ndhF-

ix rpl32. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. ML bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as “<”. Gray arrows indicate specimens whose placement has moved considerably from other analyses...... 53

Figure 10: The best tree (log likelihood = –14,328.1040) based on maximum likelihood (ML) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers and combined plastid markers, trnK-rps16, rpl32-trnL, and ndhF-rpl32. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. ML bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as “<”...... 54

Figure 11: The majority-rule tree based on the Bayesian-inference (BI) analysis of 123 sequences from the nuclear ETS rDNA spacer. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. BI posterior probabilities are provided above the

Branches...... 55

Figure 12: The majority-rule tree based on the Bayesian-inference (BI) analysis of 123 sequences from the nuclear ITS rDNA spacer. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. BI posterior probabilities are provided above the

Branches...... 56

Figure 13: The majority-rule tree based on the Bayesian-inference (BI) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. BI posterior probabilities are provided above the branches...... 57

Figure 14: The majority-rule tree based on the Bayesian-inference (BI) analysis of 123 sequences from the combined plastid markers, trnK-rps16, rpl32-trnL, and ndhF-rpl32. Brackets to the

x right of taxon labels correspond to informal clade names discussed in the text. BI posterior probabilities are provided above the branches. Gray arrows indicate specimens whose placement has moved considerably from other analyses...... 58

Figure 15: The majority-rule tree based on the Bayesian-inference (BI) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers and combined plastid markers, trnK-rps16, rpl32-trnL, and ndhF-rpl32. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. BI posterior probabilities are provided above the branches ...... 59

Figure 16: The strict consensus tree resulting from maximum parsimony (MP) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers with geographic locality mapped onto branches using MacClade (version 4.08). DIVA (version 1.1a) output is provided above branches to indicate ancestry for divergence nodes. “A” corresponds to continental Africa and “B” corresponds to Madagascar...... 60

Figure 17: The strict consensus tree resulting from maximum parsimony (MP) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers with carpel number states mapped onto branches...... 61

Figure 18: The strict consensus tree resulting from maximum parsimony (MP) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers with leaf composition character states mapped onto branches ...... 62

Figure 19: The strict consensus tree resulting from maximum parsimony (MP) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers with inflorescence arrangement states mapped onto branches ...... 63

Figure 20: The strict consensus tree resulting from maximum parsimony (MP) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers with pedicel lengths mapped onto branches...... 64

Figure 21: The strict consensus tree resulting from maximum parsimony (MP) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers with bract persistence mapped onto branches...... 65

Abstract

EVOLUTIONARY RELATIONSHIPS IN AFRO-MALAGASY SCHEFFLERA (ARALIACEAE) BASED ON NUCLEAR AND PLASTID MARKERS

By Morgan Robert Gostel, M.S. Biology

A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science at Virginia Commonwealth University.

Virginia Commonwealth University, 2010.

Thesis Director: Dr. Gregory M. Plunkett, Affiliate Research Professor, Department of Biology and Director, Cullman Program for Molecular Systematics, The New York Botanical Garden Thesis Co-Director: Dr. Rodney J. Dyer, Associate Professor, Department of Biology

The genus Schefflera is the largest in Araliaceae, with approximately 900 species. Recent studies have shown that Schefflera is polyphyletic and represents no fewer than five distinct clades, each corresponding to a specific geographic region including Asia, continental Africa and

Madagascar, Melanesia, the Neotropics, and a small clade distributed throughout several islands in the insular Pacific Ocean. The Afro-Malagasy clade contains nearly 50 species distributed throughout tropical, sub-Saharan Africa, Madagascar, the Comoros, and the Seychelles islands.

Previous studies have suggested that this group is monophyletic, identifying two smaller subclades within Afro-Malagasy Schefflera corresponding roughly to informal groups identified as “Meiopanax” and “Sciodaphyllum” on the basis of morphology. Using sequence data from nuclear rDNA spacers and plastid markers derived from 32 of the 48 currently circumscribed species of Afro-Malagasy Schefflera, this study tested the monophyly of Afro-Malagasy

Schefflera and of each of its two proposed subclades. Trees based on this molecular data were used to examine patterns of morphological evolution and biogeography among species in the clade. Results support the monophyly of Afro-Malagasy Schefflera and both subclades, which

correspond closely to “Meiopanax” and “Sciodaphyllum” which are herein referred to as

Neocussonia and Astropanax, respectively. Additional interspecific relationships were examined, which provides evidence for hybridization among several species. Schefflera myriantha, the most widely distributed species of Afro-Malagasy Schefflera, is paraphyletic with respect to two other species, S. humblotiana and S. monophylla. Many morphological features historically used to distinguish species of Afro-Malagasy Schefflera appear to be evolutionarily labile, with a history of gains and losses (e.g., reduction in leaflet number, which occurs independently in both subclades). Biogeographic analyses suggest an African ancestry for the entire Afro-Malagasy

Schefflera clade, and for both subclades, with two independent divergence events to Madagascar.

INTRODUCTION

Taxonomic History. The genus Schefflera represents the most speciose of the 41 genera currently recognized in the angiosperm family Araliaceae (Frodin and Govaerts 2003). Current estimates of species diversity in Araliaceae indicate greater than 1,600 species, with Schefflera comprising at least half of those (Plunkett et al. 2005). The taxonomic history of Schefflera dates back to the type collection of S. digitata, an endemic of New Zealand, from Forster & Forster

(1775). Since its original description, Schefflera has been subject to a turbulent taxonomic history. The generic definition has been modified several times, primarily to place emphasis on different morphological features and often to include (or “lump”) previously distinct genera that share morphological characteristics. The early efforts to broaden the circumscription of the genus were undertaken by Baillon (1878, 1879), Harms (1894-1897) and Viguier (1906, 1909). More recently, similar approaches have been undertaken by Frodin (1975), who greatly expanded the genus to include all woody araliads having palmately compound leaves, ligulate stipules, and valvate petal aestivation, while lacking armaments and articulated pedicels (Frodin 1975). Since its implementation, this most recent definition of Schefflera has resulted in explosive growth of the genus, due in large part to inclusion of segregate genera and supplemented by focused studies in the field and herbaria.

Molecular phylogenetic studies of Schefflera have advanced the understanding of this complex genus. Recent, phylogenetic analyses of Araliaceae and, more specifically, Schefflera have uncovered extensive polyphyly in the genus (Plunkett et al. 2004, 2005). Investigations by

Plunkett et al. (2005) identified five large, distinct clades of Schefflera, distributed across the phylogenetic tree of Araliaceae. These five clades correspond strongly to particular geographic areas, and have been informally named Asian Schefflera, Neotropical Schefflera, Melanesian (or

Pacific) Schefflera, Afro-Malagasy Schefflera, and Schefflera sensu stricto (Plunkett et al. 2005).

Reconstruction of these relationships was based on a representative but somewhat limited sampling of species from throughout the genus designed to test the monophyly of Schefflera, to identify subclades, and, once identified, to initiate more extensive studies of relationships within each clade. To that end, several research projects focused on recircumscription of these clades are currently under study (e.g., Melanesian and Neotropical Schefflera). The goal of the present study is to clarify relationships within the Afro-Malagasy Schefflera clade.

Evidence from the most recent studies of Schefflera (Plunkett et al. 2005) suggest that the

Afro-Malagasy clade is part of a large basal polytomy in Araliaceae. Eight additional small lineages have also been placed in this basal polytomy, and include: Seemannaralia + Cussonia,

Cheirodendron + Raukaua, Schefflera sensu stricto, Osmoxylon, Harmsiopanax, Astrotricha,

Cephalaralia, and Motherwellia (Plunkett et al. 2005), together with three much larger clades

(Aralia-Panax, the Polyscias-Pseudopanax clade, and the Asian Palmate clade). One of the smaller clades, comprising Cussonia and the monotypic Seemannaralia, is endemic to Africa. At present, a phylogenetic study is underway for Cussonia, a genus of approximately 20 species (De

Villiers and van Wyk 2006; De Villiers unpublished), but resolution of Afro-Malagasy

Schefflera relative to other clades in the basal polytomy is beyond the scope of this study, which is focused exclusively on relationships within the clade itself.

The early taxonomic history of Afro-Malagasy Schefflera dates to the 19th Century, to

Seemann’s (1865) study, which initially described African collections in the genera Astropanax and Sciadophyllum. It was not until the end of the 19th Century that Harms (1894) began placing

African araliads in Schefflera based on an expanded definition that encompassed the morphological features of several segregate genera. Much later, in the mid-20th Century,

Bernardi and Bamps revisted the African and Malagasy species of this genus. Bernardi’s (1969) work focused on endemics from Madagascar and the nearby Comoro islands. His treatment of the genus described three new species as well as two new varieties of Schefflera and moved ten other species to Schefflera from Cussonia. Two species (S. umbellifera and S. bojeri) had been previously assigned to Schefflera by Baillon (1878) and Viguier (1906), respectively. By contrast, Bamps’ (1974) study of Schefflera focused exclusively on representatives from continental Africa. In Bamps’ work, two new species were described, adding to the two already recognized in Schefflera. Together, these studies resulted in the recognition of a total of 29 species of Afro-Malagasy Schefflera. It is from this foundation that the contemporary systematic studies of Afro-Malagasy Schefflera now emerge. The past three decades have brought the number of hypothesized species of Afro-Malagasy Schefflera to 48 according to Lowry (pers. comm.).

Presently, Frodin (in Plunkett et al. 2005) places the species of Schefflera from Africa and Madagascar within two subgenera on the basis of morphological characteristics, informally named “Meiopanax” and “Sciodaphyllum”. By contrast, Bernardi (1969) identified three series of Malagasy Schefflera, namely Racemosae, Myrianthae, and Anticipes. The nine representatives sampled in Plunkett et al. (2005) provided evidence for two subgroups, but this sampling was too limited to draw more detailed conclusions. The first clade corresponds closely with Frodin’s

“Meiopanax” group, which is here referred to as Neocussonia based on a genus recognized by

Hutchinson (1967) for Malagasy species formerly placed in Cussonia (Schefflera bojeri, S. monophylla, and S. myriantha) and one from Africa (S. umbellifera). The second clade corresponds to Frodin’s widespread “Sciodaphyllum” group, which he recognized on the basis of a “generalized” or “unspecialized” morphology. However, “Sciodaphyllum” itself is

polyphyletic and the type species belongs to another clade (Neotropical Schefflera), therefore species of Afro-Malagasy Schefflera belonging to this group are referred to as Astropanax based on a generic group established by Seemann (1865). Currently, 32 species are recognized in the

Neocussonia clade, while 16 have been placed in Astropanax (Lowry, pers. comm.).

Geography. The geographic distribution of these species raises questions regarding the origin and diversification of Afro-Malagasy Schefflera. Madagascar, the fourth largest island in the world, is the primary center of diversity of Afro-Malagasy Schefflera, with 33 of the 48 species. The island is separated from the African continent by the Mozambique Channel and lies approximately 400 km to the east of Mozambique and Tanzania. The Comoros are a small network of islands and islets that lie between Africa and the northern tip of Madagascar, having a single species of Schefflera (S. myriantha, which is also found in both Madagascar and continental Africa). The Seychelles, another small network of islands located to the north of

Madagascar in the Indian Ocean, also has a single species of Schefflera (S. procumbens).

Another 15 species are distributed across the African continent’s sub-Saharan tropics. Both subgroups, Neocussonia and Astropanax, contain species endemic to both the continent and the islands to the east. Neocussonia, however, is more diverse in Madagascar, whereas Astropanax is better represented in Africa. The geographic origin of these groups, either in Africa or

Madagascar, remains a fundamental question in understanding the diversification in this clade.

Originally part of Gondwanaland, present-day Madagascar, India and the Seychelles are estimated to have separated from Africa between 165 and 175 mya (Besse & Courtillot 1988,

Schettinot & Scotese 2005, Ali & Aitchison 2008) and tectonic movement between the two landmasses has been shown to have largely stopped approximately 125 mya (Rabinowitz et al.

1983). Later separation of India and what now comprises the Seychelles from Madagascar has been dated to roughly 88 mya (Storey et al. 1995). The ability to test hypotheses regarding biogeographic divergence among related plant groups has been facilitated by an abundance of paleomagnetic and geophysical data and has contributed to a growing interest in phytogeography

(Schatz 1996, Yuan et al. 2005). Recent molecular phylogenetic reconstruction has provided evidence for both vicariance and dispersal events in the floras endemic to landmasses once a part of Gondwanaland and this has fueled debate regarding patterns of biogeography (Crisp & Cook

2007, Donoghue & Smith 2004, McGlone 2005). Historically, phylogenetic divergence in many elements of the southern hemisphere flora and fauna has been attributed to Gondwanan vicariance, but more recent studies have challenged these conclusions (Sanmartín & Ronquist

2004). In particular, the application of molecular-dating techniques suggests long distance dispersal or a combination of vicariance and dispersal events is often responsible for much of the diversity once attributed to the appearance of geological barriers (Cook & Crisp 2005, Barker et al. 2007, Weeks et al. 2005, Zerega et al. 2005).

Broader phylogenetic connections between species of Schefflera from Africa-Madagascar to those species from Malesia and the Pacific can be largely ruled out based on previous studies, which demonstrate that these taxa belong to three unrelated clades (see Plunkett et al. 2005). A better point of comparison within Araliaceae for the biogeography of Afro-Malagasy Schefflera can be found in the study of Polyscias by Plunkett et al. (2004), which focused on phylogenetic relationships among the species from the Indian Ocean Basin (IOB). IOB Polyscias is distributed across a region that includes continental Africa, Madagascar, the Comoros, and the Mascarene islands. Of particular note is one clade identified as the “Polyscias fulva group”, representing 10 species endemic to Madagascar (3 spp.), the Comoros (1 sp.) and Africa (6 spp.) (Plunkett &

Lowry 2010, Plunkett et al. 2004). Within the Polyscias fulva clade, it was suggested that multiple dispersal events between Africa and Madagascar were likey responsible for its diversity

(Plunkett & Lowry 2010); similar hypotheses will be tested in the present study among species of Afro-Malagasy Schefflera.

Habit & Morphology. Afro-Malagasy Schefflera have been described as trees, shrubs, or lianas, often epiphytic, with some species reaching heights of 30 m (Tennant 1961, Bamps

1974). Historically, Afro-Malagasy Schefflera have been identified by a mixture of morphological features including both umbellate and racemose inflorescence arrangement, 2–3,

4–5, or 6–9 carpels, simple or palmately compound leaves, as well as other combinations of leaf and inflorescence characters (Bamps 1979, Bernardi 1980). Many combinations of states for these characters are present in species informally grouped in both Neocussonia and Astropanax.

Assignment of the morphological character states listed above has, in several cases, failed to reflect reliable species definitions for some taxa in Astropanax and Neocussonia. Bernardi

(1974) noted problems with species delimitations in his treatment of the group, which emphasized leaf shape. In particular, Bernardi noted the great variation in the leaf shape of

Schefflera longipedicellata not only among distinct specimens, but also in the same individual.

Bernardi (1969) also suggested the possibility that hybridization between two Malagasy species,

S. longipedicellata and S. monophylla, may have produced the character states exhibited by S. staufferana. A survey of morphological characters among the species of Afro-Malagasy

Schefflera will help to identify the sources of confusion leading to these ambiguous determinations and may assist in revising species circumscriptions that reflect the full range of variation in each operational taxonomic unit.

Character-state mapping may help to identify morphological features defining subclades and to test whether some character states share a single evolutionary origin (e.g., the reduction of digitately compound to simple leaves) or if these features arose multiple times during the evolutionary history of the clade. A deeper understanding of other character states, even if homoplasious, may prove useful as features for the construction of species-identification keys.

Objectives & Scope. The findings outlined in Plunkett et al. (2005) suggest that the species of Schefflera from Africa and Madagascar form a monophyletic group. The current study employs a greatly expanded sampling of the species from this region to test this finding and to explore further species-level phylogenetic relationships. Previous studies have demonstrated the utility of molecular data in phylogenetic reconstruction among the species and genera of

Araliaceae, using multiple molecular markers from both the nuclear (ITS and ETS) and plastid

(notably, trnL-trnF) genomes (e.g., Eibl et al. 2001; Plunkett et al. 2004, 2005; Tronchet et al.

2005; Nicolas & Plunkett 2009). A repeated problem in species-level molecular phylogenetics of plants has been the scarcity of informative plastid markers, resulting in poorly resolved trees at this level (Miller et al. 2009). In response to this deficit, recent studies have identified several plastid sequence regions that accumulate mutations rapidly enough to resolve relationships among taxa at the species level in many groups of angiosperms (Shaw et al. 2005, 2007; Miller et al. 2009). Several of these plastid markers were selected as candidates for this study, but three intergenic spacers were ultimately chosen for sequencing across all sampled taxa, trnK-rps16, ndhF-rpl32, and rpl32-trnL. In addition to these three markers, two nuclear spacers (ITS and

ETS) have been identified in previous studies that exhibit sufficient variation to help resolve interspecific relationships in Araliaceae (e.g., studies of Polyscias and Meryta; see Eibl et al.

2001, Plunkett & Lowry 2010, Tronchet et al. 2005) and these were also used in the present study. Sequences for all five of these markers were derived from a representative sample of 18 of the 48 currently recognized species of Afro-Malagasy Schefflera, with a primary objective to test the presumed monophyly of Afro-Malagasy Schefflera under the current system of classification, to explore patterns of interspecific relationships, to test biogeographic hypotheses regarding ancestral origins, and to identify morphological features that can be used in diagnosing subgeneric and interspecific groups of taxa.

METHODS & MATERIALS

Sampling. Comprehensive sampling of all 48 species of Afro-Malagasy Schefflera was attempted, but the lack of material from 16 species (viz., Schefflera abyssinica (Hochst ex. A.

Rich) Harms, S. “ambrensis” Lowry ined., S. “decaryi” Lowry ined., S. “gentryi” Lowry ined., S. evrardii P. Bamps, S. hierniana Harms, S. “kalambatrensis” Lowry ined., S. “lewisiae” Lowry ined., S. “masoalensis”, S. “perieri” Lowry ined., S. procumbens (Hemsl.) F. Friedmann, S.

“ranomafanensis” Lowry ined., S. “sainteucei” Lowry ined., S. stuhlmannii Harms, S. urostachya

(Engl.) Harms, and S. weibeliana Bernardi) resulted in an incomplete but still representative sampling. Of the 16 species left unsampled, ten are currently known only from the type specimen. A total of 160 accessions representing the remaining 32 species were included in this study (see Table 1). Most material was available from fresh field collections, dried on silica gel, but 25 of the 160 samples were derived from older herbarium specimens. Many taxa are represented by multiple specimens to test for monophyly and/or the potential for interspecific hybridization. Outgroup taxa were selected on the basis of relationships observed in previous

studies (i.e., Plunkett et al. 2005) and included species of Cussonia, Schefflera sensu stricto,

Osmoxylon, Astrotricha, and the monotypic Seemannaralia.

Extraction, amplification and sequencing. For each accession, total DNA was extracted using the DNeasy Plant extraction kit (QIAGEN Inc.) or a modification of the protocol described by Alexander et al. (2007). Selected DNA regions were amplified using the polymerase chain reaction (PCR) for each accession with a combination of existing and newly developed primers for each spacer (see Table 2 for list of primers). PCR reaction conditions included 0.5 µL of both forward and reverse primers (5 µM), 0.5 µL spermidine (4 mM), 2 µL total DNA, and 5 µL of either the Jumpstart REDTaq ReadyMix (SigmaAldrich) or the GoTaq Green Master Mix

(Promega Corp.) Taq polymerase mixes. With the exception of three modifications for trnK- rps16 and ndhF-rpl32, all thermocycler protocols for PCR amplification included a pre-soak step of 4 min at 94ºC, followed by 35 cycles of 30 sec at 94ºC (denaturation), 1 min at 52ºC

(annealing), and 50 sec at 72ºC (extension), and then a single post-soak of 72ºC for 4 min. Due to lower primer melting temperatures, thermocycler protocols were slightly modified for both ndhF-rpl32 and trnK-rps16 by lowering the annealing temperature to 48ºC (for trnK-rps16) or

50ºC (for ndhF-rpl32) and by extending the annealing time to 90 sec (for both markers). In addition, the extension time was modified to 135 sec for ndhF-rpl32 due to the increased length of this marker. PCR products were purified using 1.5 µL exonuclease I and 3 µL shrimp alkaline phosphatase per 5 µL of product (USB Corp.). Purified PCR products were sequenced directly using a thermocycler program of 20 sec at 94ºC, 15 sec at 55ºC, and 1 min at 60ºC for 30 cycles.

Sequencing reactions were carried out using DYEnamic ET Terminators (GE Healthcare, Inc.) or

BigDye Terminator (vers. 3.1, Applied Biosystems Corp.) and then purified using the

MultiScreen384 SEQ filtration system (Millipore Corp.) or the BigDye XTerminator purification kit (Applied Biosystems Corp.). Capillary gel electrophoresis of cleaned products was performed on a MegaBACE 1000 DNA Sequencing System (GE Healthcare, Inc.) or an ABI 3730 DNA

Analyzer (Applied Biosystems Corp.) and then assembled and edited using the Sequencher 4.7 software package (Gene Codes Corp.). Sequence alignment was adjusted manually following an initial alignment using ClustalX (Thompson et al. 1997). Informative indels resulting from sequence alignment were coded as binary (presence/absence) characters according to the method provided by Giribet and Wheeler (1999). All sequences will be deposited in the GenBank database.

Phylogenetic Analyses. Three distinct approaches to phylogenetic analysis were used in this study, including maximum parsimony (MP), maximum likelihood (ML), and Bayesian inference (BI). Five separate datasets were used, including (1) ITS, (2) ETS, (3) combined ITS +

ETS, (4) the plastid markers (treated together), and (5) a set combining sequences from the two nuclear and three plastid markers. To test for congruence among the separate datasets, the incongruence length difference (ILD) test of Farris et al. (1995) was performed using the partition homogeneity test in PAUP* 4.0b10 (Swofford 2002). Three partitions were established, representing the ITS, ETS, and combined plastid markers. An ILD test was performed comparing all three partitions simultaneously, as well as separate, pairwise ILD tests. For model based approaches (ML & BI), jModelTest (Posada 2008) was used to select the most appropriate model of sequence evolution and to maximize computational accuracy.

Maximum parsimony analyses were performed using PAUP* and a two-step protocol modified from Plunkett et al. (2005). In the first step, a heuristic search of 1,000 replicates was

generated by random, stepwise addition and TBR branch swapping, but saving no more than 100 trees per replicate. The strict consensus from this initial search was then loaded as a topological constraint for a second heuristic search that was performed following the same protocol as the first step (1,000 reps, saving 100 trees per replicate) but saving only shortest-length trees that did not agree with the topological constraint. If no additional shortest-length trees were recovered, the strict consensus from the first analysis was used as a conservative estimate of phylogenetic relationships. Bootstrap values were calculated from 1,000 replicates using PAUP*.

The model-based analyses were performed using either GARLI (version 0.96; Zwickl

2006) for ML or MrBayes (version 3.2.1; Huelsenbeck and Ronquist 2001) for BI. ML analyses were performed applying mostly the default parameters in GARLI and MrBayes, namely the

GTR + ! + I model using four variable rate categories. For ML analyses, the maximum number of generations was set to 5,000,000, saving the ML tree with the best score. ML bootstrap values were also calculated using GARLI (with the same parameters). Bayesian inference used model of sequence evolution closest to that indicated by jModelTest as selected by the Akaike information criterion (AIC), with chains run for 10 million generations for the combined nuclear + plastid dataset and 4 million generations for all other datasets. Markov Chain Monte Carlo (MCMC) was implemented using four chains, sampling every 1,000 generations.

Morphological character assessment. Morphological data were collected from specimens deposited at Musée National d’Histoire Naturelle in Paris (P) and the Jardin

Botanique National de Belgique (BR). The following five characters were coded as binary states: inflorescence arrangement (0: umbellate, 1: racemose), number of carpels (0: 2–3, 1: 4–5, and 2:

! 6), leaf morphology (0: exclusively unifoliolate, 1: 1–3 leaflets, 2: >3 leaflets), pedicel length

(0: absent, 1: 0.1–5mm, 2: ! 5mm), and retention of bracts (0: bracts persistent, 1: bracts caducous). Other characters (e.g., bract size and petiole length) were identified as potentially informative, but omitted from formal character analysis due to limited access to herbarium materials. Morphological character states were mapped onto the combined ITS + ETS tree using

MacClade version 4.08 (Maddison & Maddison 2005).

DIVA. Biogeographic relationships were explored using DIVA 1.1a (Ronquist 1996,

1997), which assigned ancestral areas to internal nodes on a fully resolved ML phylogeny produced from a reduced sample set (decreasing the number of terminals, but maintaining the basic topology), using two areas of endemism (continental Africa and Madagascar). Presence or absence of taxa in each area of endemism was coded as a binary character.

RESULTS

Sequence characteristics. A total of 160 accessions was sequenced for each of the five

DNA spacer regions employed in this study, although four specimens did not amplify for the ndhF-rpl32 marker and one specimen did not amplify for the trnK-rps16 marker. After examining the sequences for redundancy, 37 samples were removed because they shared identical sequences with others in the dataset. The remaining 123 accessions were used for phylogenetic analyses. Uncorrected pairwise distances were calculated between all sequences for each molecular marker (treating the three plastid markers separately). Pairwise distances were also calculated for plastid markers using datasets with and without coded gaps.

The length of the ETS sequences ranged from 375 to 449 nucleotides, with a total aligned length of 474 characters (103 of which were parsimony informative). Nucleotide percentages for

ETS were A = 17.7%, C = 28.6%, G = 24.5%, and T = 29.1%. Average pairwise distance among

ingroup taxa was 4.1% for ETS sequences, the greatest distance between two ingroup taxa was

9.7% (between Schefflera goetzenii 3503 and S. “vohimenensis” 2378), while 204 pairwise combinations provided distance values of zero.

The length of the ITS sequences ranged from 594 to 662 nucleotides, with a total aligned length of 675 characters (95 of which were parsimony informative). The nucleotide composition for ITS was A = 21.8%, C = 30.9%, G = 28.8%, and T = 18.5%. Average pairwise distance among ingroup taxa was 2.5% for ITS sequences, the greatest distance between two ingroup taxa was 5.5% (between Schefflera myriantha 1498 and S. vantsilana 152), while 124 pairwise combinations provided distance values of zero.

Aligned plastid sequences produced fewer parsimony-informative characters relative to the nuclear spacers. The length of the trnK-rps16 sequences ranged from 818 to 1,099 nucleotides and produced an aligned total of 1,188 nucleotide characters and 6 coded indels (for a total of 36 parsimony-informative characters). Nucleotide percentages for trnK-rps16 sequences were A = 41.9%, C = 14.9%, G = 14.6%, and T = 28.6%. Average pairwise distance among ingroup taxa was 0.5% for trnK-rps16 sequences, the greatest distance between two ingroup taxa was 3.8% (between Schefflera tessmannii 2107 and S. “floretii” 7172), while 1,262 pairwise combinations provided distance values of zero. One specimen (Schefflera “floretii”

7160) did not amplify for trnK-rps16 and this missing data was coded as ambiguous (N).

Length of the rpl32-trnL sequences ranged from 664 to 895 nucleotides and an aligned total of 944 nucleotide characters and 4 coded indels (for a total of 47 parsimony-informative characters). Nucleotide percentages for rpl32-trnL sequences were A = 35.7%, C = 14.7%, G =

12.5%, and T = 37%. Average pairwise distance among ingroup taxa was 0.6% for rpl32-trnL sequences, the greatest distance between two ingroup taxa was 2.5% (between Schefflera

monophylla 409 and S. fosbergiana 3337), while 2,016 pairwise combinations provided distance values of zero.

Sequence lengths for ndhF-rpl32 ranged from 779 to 1,250 with a total of 1,389 aligned nucleotide characters and 13 coded indels (for a total of 68 parsimony-informative characters).

Nucleotide percentages for ndhF-rpl32 sequences were A = 39.6%, C = 12.6%, G = 12.4%, and

T = 35.5%. Average pairwise distance among ingroup taxa was 0.8% for ndhF-rpl32 sequences, the greatest distance between two ingroup taxa was 2.4% (between Schefflera staufferana 7082 and S. myriantha 4988), while 639 pairwise combinations provided distance values of zero. Four specimens did not amplify for ndhF-rpl32 and this missing data was coded as ambiguous (N)

(including Schefflera humblotiana 3883, S. stolzii 3577, S. barteri 5208, and S. barteri 16998).

Significant incongruence among the three partitions established in this study could not be rejected based on results of the ILD test (p = 0.01), suggesting that the partitions may not be combinable. Nonetheless, to explore the effects of combining the datasets, all three partitions were concatenated and analyzed simultaneously. The combined ITS + ETS datasets consisted of

1,149 aligned characters, 198 ("17.2%) of which were considered parsimony informative.

MP analysis of ETS sequences produced 51,600 total trees, each of 279 steps (CI =

0.675, RI = 0.954, Fig. 1). MP analysis of the ITS dataset produced 96,600 trees, each of 271 steps (CI = 0.708, RI = 0.959, Fig. 2). The combined ITS + ETS dataset produced a total of

100,000 best trees, each of 565 steps in length (CI = 0.665, RI = 0.951, Fig. 3). The greatest resolution among all datasets in this study was provided by this combined ITS + ETS dataset and for this reason the strict consensus tree from this dataset will be used for most of the discussions and for character-state mapping.

Given the low sequence variation of the three plastid markers (all linked on the non- recombining plastid genome), the three plastid spacers were analyzed together. They yielded a total of 3,521 characters, 168 ("4.7%) of which were parsimony-informative, including 23 indels that were identified as potentially informative and coded as binary characters in the MP analyses

(these indels were omitted from model-based analyses). MP analysis of the combined plastid dataset produced 87,700 best trees, each of 633 steps (CI = 0.612 excluding uninformative characters, RI = 0.918, Fig. 4).

The dataset combining all nuclear (ITS + ETS) and plastid sequences yielded a total of

4,693 characters, 366 ("7.8%) of which were parsimony informative when coded indels were included. MP analysis of the combined nuclear + plastid dataset resulted in 94,200 best trees, each of 1,238 steps in length (CI = 0.608, RI = 0.929, Fig. 5).

Comparison of the 88 models tested in jModeltest resulted in the selection of the TIM + #

+ I model of sequence evolution based on the AIC. Both model-based approaches (ML and BI) produced the same topology with only minor variation in branch support. Likelihood for the ML trees were –2,279.056 for ETS (Fig. 6), –2,612.7117 for ITS (Fig. 7), –5,075.1868 for combined

ITS + ETS (Fig. 8), –8,530.1115 for plastid (Fig. 9), and –14,328.1040 for combined nuclear + plastid markers (Fig. 10). No variation among tree topologies was found across the 10 runs conducted using GARLI for any of the datasets. The trees resulting from the Bayesian analyses were highly similar to the topologies from the ML searches (Fig. 11–15).

Morphological Character Mapping: Results of morphological character state mapping are provided in Figures 16–21. Morphological characters were mapped on to the phylogeny

resulting from the ML analysis of the combined nuclear (ITS + ETS) dataset. No clear patterns of morphological evolution emerge as a result of this character mapping.

DIVA: Results from the DIVA analysis favored a two-dispersal scenario, with continental Africa being assigned as ancestral for both subgroups Neocussonia and Astropanax

(Fig. 16).

DISCUSSION

Afro-Malagasy Schefflera Clade. This study helps to advance the recommendations set forth by Plunkett et al. (2005), whose findings first illustrated extensive polyphyly in the genus

Schefflera as currently circumscribed. Due to the size and complexity of Schefflera, they suggested that recircumscription of the genus should proceed by testing relationships in each of the five geographically distinct clades individually, beginning with in-depth studies of the smaller clades, including Pacific Schefflera (approx. 40-50 spp.), Afro-Malagasy Schefflera

(approx. 48 spp.), and Schefflera § Schefflera (8 spp.), and broad surveys of the larger Asian and

Neotropical clades (c. 200–400 species each). This paper explores relationships among the

African and Malagasy species of Schefflera with an enhanced sampling that represents nearly

70% of the species diversity of this clade. This study confirms the monophyly of the species from Africa and Madagascar in a single clade of Afro-Malagasy Schefflera and this monophyly is strongly supported (BS = 100%, PP = 1.0). Since the publication of most recent studies of

Afro-Malagasy Schefflera (Bernardi 1979, Bamps 1974), no fewer than 17 new species have been proposed (Lowry, pers. comm.) and of these, seven new species have been included here.

Within the Afro-Malagasy clade, additional interspecific relationships are also evident.

Bernardi’s (1969) treatment of three series (Racemosae, Myrianthae, and Anticipes) is not supported, but Frodin’s (see Plunkett et al. 2005) suggestion that African and Malagasy

Schefflera represent two morphologically distinct groups is maintained by the presence of two monophyletic groups corresponding to Frodin’s “Sciodaphyllum” and “Meiopanax” (BS =

100%, PP = 1.0). These groups will be referred to as Astropanax and Neocussonia throughout the rest of this paper. In the discussions below, the infrageneric system developed by Frodin for species belonging to this clade will be used as a point for comparison for results from the current study.

Neocussonia Clade. Phylogenetic reconstruction placed 22 of the 48 species of Afro-

Malagasy Schefflera tested in this study in the Neocussonia subclade (Figs. 1–3, 5–8, and 10, BS

= 100%, Figs. 11–13 and 15, PP = 1.0). This subclade corresponds closely to the “Meiopanax” group of Baillon (1880) and Frodin (see Plunkett et al. 2005). The generic name Neocussonia was first used by Hutchinson (1967) to describe several species of Schefflera endemic to

Madagascar and this name is applied here because it has taxonomic priority over “Meiopananx” at the generic and subgeneric level. Of the species in Neocussonia, only two are African endemics (S. lukwangulensis and S. umbellifera), while the remaining species are all endemic to

Madagascar. Resolution among the species in this clade varies by both markers and, in some cases, by analysis type.

Within Neocussonia, two large clades are resolved in most topologies, which are referred to as the “Anticipes” clade and the “Palmate-Vantsilana” clade (Figs. 1–3, 5–8, 10–13, and 15).

Both of these clades as well as the remaining, less resolved species in Neocussonia are placed in a large polytomy at the base of the clade, sister to Schefflera umbellifera (Fig. 3, BS = 66% and

Fig. 15, BS = 0.94). In several cases, different samples of the same species are found in more than one subclade, and this pattern (together with other lines of evidence) suggests the possibility of interspecific hybridization.

Schefflera umbellifera is resolved as sister to the rest of Neocussonia in the parsimony trees (Fig. 1, BS = 77%, Fig. 3, BS = 66%, Fig. 6, BS = 62%, Fig. 8, BS = 60%, Fig. 11, PP =

0.69, Fig. 13, BS = 0.81, Fig. 15, BS = 0.94). Morphologically, this species is distinct from others in Neocussonia in its leaflet margins, which are serrated margins rather than entire or crennate. Similarities exist between the umbellate inflorescence in Schefflera umbellifera, which has long, narrow axes, and the inflorescences of three other species in Neocussonia (S. lukwangulensis, S. frodiniana, and S. rainaliana). Geographically, S. umbellifera is distinct from other species in Neocussonia with the exception of S. lukwangulensis, which is also endemic to continental Africa. The two model-based analyses of combined plastid datasets place S. umbellifera further within Neocussonia but branch support for this placement is low (Fig. 9, BS

= 55% & Fig. 14, PP = 0.71).

The “Palmate-Vantsilana” clade is so named because of the shared tendency of palmately compound leaves among its species together with the inclusion of S. vantsilana, whose epithet is also the vernacular Malagasy name for most Schefflera species. Within this clade, there are several subclades containing more than one species, which is evident in the clades containing specimens identified as Schefflera longipedicellata and S. vantsilana as well as those combining both S. “floretii” and S. “vohimenensis”. A third species, S. “rabenantoandroi”, is also found in the “Palmate-Vantsilana” clade with consistently high support (Fig. 3, BS = 86%, Figs. 12, 13, and 15, PP = 1.0). Schefflera “rabenantoandroi” has been elevated to the species level by Lowry

(pers. comm.) from a variety recognized by Bernardi (S. vantsilana var. litoralis) due to its

distinctive morphology (e.g., large palmately compound leaflets, long petiole and retuse leaflet apex) and distribution in low-elevation littoral forests (compared to other species of the

“Palmate-Vantsilana” clade, which are typically found at altitudes greater than 1,000 m).

Bayesian inference places the “Palmate-Vantsilana” clade within a larger monophyletic group

(Fig. 15, PP = 0.84) comprising subclades containing S. macerosa (PP = 1.0) and other specimens identified as S. longipedicellata (PP = 1.0).

In assessing the apparent polyphyly of some of the species of the “Palmate-Vantsilana” clade several factors must be considered, including sympatric distributions and overlapping morphological character states. For example, samples of S. “vohimenensis” and S. “floretii” appear together in two separate clades with low to moderate branch support (Fig. 3, BS = 62%,

Fig. 13, PP = 1.0). These two “hypothesized species”are sympatric but were considered distinct because Schefflera “floretii” has palmately compound leaves with 3–5 obdeltoid leaflets, whereas Schefflera “vohimenensis” has unifoliate or palmately compound leaves with no more than three leaflets, which are larger, more coriaceous, and have different leaf shapes (obovate) than S. “floretii”. There are three possible explanations for this polyphyly, (1) hybridization between the two species which may have blurred species distinctions, (2) faulty species circumscriptions that do not correctly capture the full range of character states for each putative species, thus separating them erroneously, or (3) incorrect species identifications for some of the samples included in this study.

Additional polyphyly in the “Palmate-Vantsilana” clade is shown for a subclade including specimens identified as S. longipedicellata and S. vantsilana (Fig. 3, BS = 68%, Fig.

13, PP = 0.99). All specimens identified as S. longipedicellata in this clade come from the same locality, while the specimens identified as S. vantsilana cover a much wider geographic range.

Sympatry between the two suggests possible hybridization, or perhaps that the definition for S. vantsilana should be expanded to include greater variation in leaflet shape. As currently circumscribed, S. vantsilana has larger leaflets than S. longipedicellata with a strongly retuse apex and an overall obdeltoid leaf shape.

The other subclade nested within Neocussonia is labeled “Anticipes”, a reference to the similarities of this clade to the composition of species comprising Bernardi’s (1974) series of the same name. This clade is not well resolved in MP analyses but morphological features and BI analyses provide moderate to strong support (Fig. 12, PP = 0.96, and Fig. 15, PP = 0.97). Within

“Anticipes”, a subclade containing three species, Schefflera halleana, S. favargeri, and S.

“humbertii”, is supported in the majority of topologies, with moderate to strong support (Fig. 3,

BS = 60%, Fig. 5, BS = 76%, Fig. 13, PP = 0.96, and Fig. 15, PP = 1.0). Elements of this clade are also supported by plastid data (Figs. 4, 9, and 14). In most trees, the “Anticipes” clade includes six additional species, Schefflera bojeri, S. staufferana, and S. antoetrensis (Figs. 3, 8,

10, 12, 13, 15), as well as S. capuroniana, S. moratii, and S. andohahelensis (Figs. 8, 10, 13, 15).

The majority of these species (all but S. bojeri) share unifoliolate leaves.

Three proposed new species also fall within the “Anticipes” clade, including Schefflera

“humbertii”, S. “andohahelensis”, and S. “antoetrensis”. There does not appear to be sufficient evidence to retain S. “humbertii” as separate from S. favargeri since these two species are both sympatric and not morphologically distinct, but there is strong support for maintaining the two remaining new species in this clade. Schefflera “andohahelensis” is the southernmost species in this clade, with a distribution limited to the Vohimena mountain range near Madagascar’s southern coast. This species, along with S. “antoetrensis”, produces copious amounts of thick, milky latex when cut. Schefflera “antoetrensis” is distinct from S. “andohahelensis” in its more

limited range to a single locality much further north near Antoetra in central Madagascar. The original collection for this species came from a single specimen collected by Capuron, which was placed in the morphologically similar S. favargeri, but Capuron’s notation on the specimen regarding the abundance of sap suggested a potential distinction from that species.

The remaining five species in Neocussonia, Schefflera rainaliana, S. frodiniana, S.

“purpuristyla”, S. bracteolifera, and S. lukwangulensis, are not well resolved in either the

“Anticipes” or “Palmate-Vantsilana” clades. Schefflera rainaliana and S. frodiniana are resolved together in most analyses (Fig. 3, BS = 70%, Fig. 8, BS = 84%, Fig. 13, PP = 1.0). Both S. rainaliana and S. frodiniana have an umbellate inflorescence with long, narrow axes, although S. rainaliana is unifoliolate and S. frodiniana has palmately compound with 3–5 leaflets. Similarly,

S. bracteolifera and S. “purpuristyla” are resolved together in many topologies (Fig. 3, BS =

78%, Fig. 8, BS = 87%, Fig 13, PP = 1.0). The formation of a clade between S. bracteolifera and

S. “purpuristyla” is surprising as the two species differ considerably in their morphologies. S. bracteolifera is unifoliolate, while S. “purpuristyla” is palmately compound with 5–7 leaflets.

Astropanax clade. Ten species were placed in the Astropanax clade (Figs. 1–3 and 5–10,

BS = 100%, Figs. 11–13 and 15, PP = 1.0) and of these, all but three are endemic to continental

Africa. This clade corresponds closely to the African elements of Frodin’s “Sciodaphyllum”, but

Plunkett et al. (2005) demonstrated that “Sciodaphyllum” was polyphyletic. Frodin (see Plunkett et al. 2005) recognized the group on the basis of a “generalized” morphology that includes terminal paniculate inflorescences, leaves crowded at the end of stems, limited branching, and non-ruminate endosperm, features shared among many geographically diverse species of

Schefflera in Africa, Asia, and the Neotropics and also across many other Araliaceae. Before

being transferred to Schefflera, several of these species were placed in the genus Astropanax by

Seemann (1865). Because the type species of “Sciodaphyllum” is placed among species in the

Neotropical Schefflera, Afro-Malagasy species of this group are referred to using Seemann’s original Astropanax. Of the Afro-Malagasy species that Frodin placed in “Sciodaphyllum”, only

S. moratii does not fall in the Astropanax clade, but is placed instead among the species of the

Neocussonia clade. This finding is consistent with the morphological characteristics of S. moratii, which has a racemose inflorescence arrangement and strictly unifoliolate leaves.

Perhaps most surprising in this clade is the paraphyly of Schefflera myriantha with respect to two Malagasy species, S. monophylla and S. humblotiana. Of these, S. monophylla represents the most morphologically diverse species in this clade, but is geographically restricted to Madagascar. Conversely, S. myriantha has the broadest geographic range (continental Africa,

Comoro islands, and Madagascar) among species of Afro-Malagasy Schefflera, but morphologically, the specimens from Africa are nearly indistinguishable from those distributed throughout the Comoros and Madagascar. Both the separate and combined analyses of the nuclear and plastid datasets indicate paraphyly in S. myriantha with strong branch support (Fig.

3, BS = 96%, Fig. 8, BS = 95%, Figs. 12–15, PP = 1.0). Lower branch support in plastid and combined nuclear + plastid topologies appears to result from inconsistent placement of a single specimen of S. monophylla (Fig. 5, BS = 58%, Fig. 10, BS = 65%). The lack of morphological differences in S. myriantha from Africa vs. Madagascar suggests the need for a more comprehensive study of specimens in light of the molecular divergence demonstrated by this study. The morphological distinction between S. monophylla and S. humblotiana is clear.

Schefflera monophylla, despite its epithet, is typically not truly unifoliolate, but instead has a large central leaflet with two much smaller lateral leaflets that are sometimes scarcely evident

but only rarely absent. Morphologically, S. humblotiana is the most distinctive of these three species, possessing extremely long, narrow leaflets. Due to the molecular divergence among species belonging to what are labeled the “myriantha-monophylla” and “African myriantha” clades, African S. myriantha may warrant recognition as a new species, but this decision must be delayed until specimens from the Comoros (currently lacking) can be included in a formal study.

There is strong support for another clade in Astropanax referred to as the “Goetzenii” clade, which includes three species, Schefflera goetzenii, S. barteri, and S. tessmannii (Fig. 5, BS

= 96%, Figs. 13 and 15, PP = 1.0). Resolution is poor within the clade, and while there are subclades, they vary in topology across datasets. In some cases, S. goetzenii is sister to S. barteri and S. tessmannii (Fig. 5, BS = 80%), while other trees place specimens of each species in a polytomy (Fig. 3, BS = 90%, Figs. 13 and 15, PP = 1.0). Each of the three species in the

“Goetzenii” clade is highly distinct morphologically. Schefflera goetzenii has caducous bracts and flowers with six-carpellate ovaries, palmately compound leaves with 6–7 narrowly obovate leaflets and a racemose inflorescence arrangement. Both S. barteri and S. tessmannii have long, persistent bracts but differ in carpel number (S. barteri is 7–9 carpellate, S. tessmannii is 5–6 carpellate) and number of leaflets (S. barteri has 5–8 leaflets, S. tessmannii has 6-8 leaflets), however, S. barteri and S. tessmannii both share inflorescence arrangement features with S. goetzenii with a paniculate-racemenose inflorescence. Geographically these species represent a large portion of tropical Africa from western sub-Saharan Africa and São Tomé to east-central sub-Saharan Africa. Poor sampling in this clade is likely attributable to the lack of resolution.

The remaining four species in the Astropanax clade form a basal polytomy with the

Schefflera myriantha-S. monophylla complex and the “Goetzenii” clade, and include S. volkensii,

S. mannii, S. kivuensis and S. stolzii. Differences in resolution among species belonging to

Astropanax may be due to missing data from ndhF-rpl32 for Schefflera humblotiana, S. stolzii, and two samples of S. barteri. On the basis of morphology (and to some degree geography) several additional species may also belong to Astropanax, but these species (Schefflera abyssinica, S. evrardii, S. hierniana, S. procumbens, S. stuhlmannii, and S. urostachya) were not available for sampling. Despite this limitation, sampling of Astropanax relative to Frodin’s Afro-

Malagasy “Sciodaphyllum” resulted in "62% species coverage and differed only in the placement of Schefflera “moratii”.

Geography. Results of the DIVA study suggests an African origin for the entire Afro-

Malagasy Schefflera clade and an African origin for both the Neocussonia and Astropanax clades

(Fig. 16). In Neocussonia, the African species, Schefflera umbellifera, is sister to the remaining members of the clade. A second Neocussonia species, S. lukwangulensis, is also endemic to continental Africa, but its placement in a polytomy renders biogeographic inference equivocal.

Two scenarios are possible – either a single divergence from continental Africa to Madagascar or a divergence to Madagascar followed by a secondary dispersal back to Africa. In Astropanax, the ancestral area is continental Africa. A single divergence event in Astropanax appears to have led to the presence of S. humblotiana, S. monophylla and Malagasy representatives of Schefflera myriantha. Conclusions regarding biogeographic hypotheses for the Comoro Islands were not possible due to the unavailability of material representing specimens of S. myriantha from the

Comoros.

Morphological patterns. All species in the Neocussonia clade share either 2–3 or 4–5- carpellate ovaries, whereas species in the Astropanax clade share 4–5 or 6 or more carpels (Fig.

17). A trend towards increases in carpel number is evident in each clade. In Astropanax there is a single increase from the ancestral state of 4–5 carpels to 6 or more carpels, but this is restricted to the “Goetzenii” clade. In Neocussonia, there are multiple independent increases in carpel number from 2–3 (the ancestral state) to 4–5 carpels in S. staufferana, in S. frodiniana, and again in S. moratii. Palmately compound leaves are ancestral in both Neocussonia and Astropanax, with reduction in leaflet number arising independently in each clade. There is a single reduction

(S. monophylla) in Astropanax (Fig. 18), but several reductions of leaflet number in the

Neocussonia clade. There are nine instances of leaflet reduction from plurifoliolate to unifoliolate leaves, and in at least one case what appears to be reduction to simple leaves due to loss of the leaflet articulation between the petiole and the petiolule (S. antoetrensis).

Inflorenscence structure is highly labile in both Neocussonia and Astropanax (Fig. 19).

Umbellate inflorence arrangement appears ancestral in Neocussonia, with racemose inflorenscences common among several species. Species in Astropanax have a primarily racemose inflorescence arrangement with the large exception found in the “African myriantha” and “myriantha-monophylla” clades. Three species in Astropanax (Schefflera stolzii, S. volkensii, and S. mannii) possess spikes, having completely lost the pedicels in their flowers, and this trait is shared with outgroup members in the genus Cussonia. Other species in Astropanax have longer pedicels, primarily among species of S. myriantha and S. monophylla (Fig. 20). Species in

Neocussonia appear to have undergone several reductions in pedicel length, although no species in this clade have completely lost this character. The persistence or loss of bracts in mature inflorescences is also highly labile in both Neocussonia and Astropanax (Fig. 21).

Conclusions. All phylogenic trees produced as a result of this study confirm previous findings of monophyly among species of Afro-Malagasy Schefflera (Plunkett et al. 2005). The study of this clade represents part of a larger effort to recircumscribe all five clades that comprise the polyphyletic genus Schefflera, and a parallel effort to recircumscribe Polyscias, the second largest genus in Araliaceae. However, unlike Schefflera, which is polyphyletic, Polyscias is paraphyletic with respect to six other genera, and thus it has been expanded to include all the species from Arthrophyllum, Cuphocarpus, Gastonia, Munroidendron, Reynoldsia, and

Tetraplasandra (Plunkett & Lowry 2010, Lowry & Plunkett 2010). By contrast, the polyphyly of

Schefflera will ultimately require the recognition of several separate genera. Only the species belonging to the small Schefflera § Schefflera clade (8 spp.) will retain that generic name, while the species belonging to the remaining four clades in Schefflera will require taxonomic transfers to several new or reinstated genera. The task of recircumscribing species belonging to Afro-

Malagasy Schefflera could proceed in one of two ways, either recognition of a single genus comprising all species in the clade, or two separate genera comprising species assigned to either

Neocussonia or Astropanax.

Future studies should focus on sampling from the 16 species left unsampled here and perhaps a morphometric approach to delimiting species in the Afro-Malagasy Schefflera clade, particularly for difficult species-complexes, including several in the Astropanax clade (e.g.,

Schefflera humblotiana, S. monophylla, and S. myriantha) and species belonging to the

“Palmate-Vantsilana” clade in Neocussonia. The present study provides the foundation for such future work, and makes a considerable contribution to the realignments necessary in both

Schefflera sensu lato and Araliaceae.

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TABLE 1: Species used for DNA samples used in this study, including geographic range and voucher information.

Taxon Geographic locality Source and accession no.

Outgroups Astrotricha pterocarpa Benth. Australia Queensland, Australia Plunkett 1527 (NY) Cussonia holstii Harms ex. Engl. Southern Africa Tanzania, Lowry 4986 (MO) Cussonia paniculata Eckl. & Zeyh. Southern Africa South Africa, Phillipson 5263 (MO) Cussonia thyrsiflora Thunb. Southern Africa South Africa, Phillipson 5110 (MO) Osmoxylon pectinatum (Merr.) Phillipson Taiwan Green Is., Taiwan Huang 756 (HAST) Schefflera digitata J.R. Forst. & G. Forst. New Zealand North Island, New Zealand Plunkett 2190 (NY) Seemannaralia gerrardii (Seem.) R. Vig. South Africa South Africa, Phillipson 5471 (MO)

Ingroup, Neocussonia subclade Schefflera “andohahelensis” Lowry ined. Madagascar Andohahela, Madagascar RBE 1505 (MO) Ivorona, Madagascar Lowry 7164 (MO) Ivorona, Madagascar Lowry 7169 (MO) Ivorona, Madagascar Lowry 7171 (MO) Schefflera “antoetrensis” Lowry ined. Madagascar Antoetra, Madagascar Lowry 7088 (MO) Antoetra, Madagascar Lowry 7091 (MO) Schefflera bojeri (Seem.) R. Vig. Madagascar Fianarantsoa, Madagascar Lowry 5818 (MO) Antsirabe, Madagascar Lowry 7071 (MO) Antsirabe, Madagascar Lowry 7109 (MO) Schefflera bracteolifera Frodin Madagascar Mahajanga, Madagascar Lowry 5327 (MO) Schefflera capuroniana (Bernardi) Bernardi Madagascar Antananarivo, Madagascar Bernardi 11118 (P) Ankazobe, Madagascar Plunkett 2328 (MO) Ambatovy, Madagascar Gostel 33 (MO) Schefflera favargeri Bernardi Madagascar Mahajanga, Madagascar Callmander 384 (MO) Mahajanga, Madagascar Callmander 444 (MO) Antsiranana, Madagascar Ratovoson 476 (MO) Mahajanga, Madagascar Lowry 5344A (MO) Schefflera “floretii” Lowry ined. Madagascar Ivorona, Madagascar Lowry 7160 (MO) Ivorona, Madagascar Lowry 7162 (MO)

Ivorona, Madagascar Lowry 7172 (MO) Andohahela, Madagascar RBE 1485 (MO) Andohahela, Madagascar RBE 1487 (MO) Ivorona, Madagascar RZK 4558 (MO) Ivorona, Madagascar RZK 4941 (MO) Schefflera fosbergiana (Bernardi) Bernardi Madagascar Antsiranana, Madagascar Schmidt 4322 (MO) Marojejy, Madagascar RD 3350 (MO) Makirovana, Madagascar RD 3337 (MO) Schefflera frodiniana Bernardi Madagascar Andohahela, Madagascar RBE 1486 (MO) Schefflera halleana Bernardi Madagascar Antsiranana, Madagascar Schmidt 4261 (MO) Marojejy, Madagascar RD 3339 (MO) Marojejy, Madagascar RD 3340 (MO) Marojejy, Madagascar RD 3343 (MO) Schefflera “humbertii” Lowry ined. Madagascar Antsiranana, Madagascar Lowry 5386 (MO) Schefflera longipedicellata (Lecomte) Bernardi Madagascar Toamasina, Madagascar Lowry 6153 (MO) Toamasina, Madagascar Lowry 6220 (MO) Antananarivo, Madagascar Lowry 6264 (MO) Antoetra, Madagascar Lowry 7098 (MO) Antoetra, Madagascar Lowry 7104 (MO) Ambatovy, Madagascar Gostel 21 (MO) Ambatovy, Madagascar Gostel 23 (MO) Ambatovy, Madagascar Gostel 33 (MO) Schefflera lukwangulensis (Tennant) Bernardi Tanzania Mabberly 1197 (MO) Schefflera macerosa Bernardi Madagascar Andohahela, Madagascar RBE 1499 (MO) Andohahela, Madagascar RBE 1509 (MO) Schefflera moratii Bernardi Madagascar Antsevabe, Madagascar Gostel 31 (MO) Antsevabe, Madagascar Gostel 34 (MO) Schefflera “purpuristyla” Lowry ined. Madagascar Mahajanga, Madagascar Callmander 432 (MO) Schefflera “rabenantoandroi” Lowry ined. Madagascar Tolagnaro, Madagascar Lowry 7148 (MO) Tolagnaro, Madagascar Lowry 7149 Schefflera rainaliana Bernardi Madagascar Tolagnaro, Madagascar McPherson 14791 (MO) Tolagnaro, Madagascar Lowry 4444 (MO)

Tolagnaro, Madagascar Lowry 7151 (MO) Schefflera staufferana Bernardi Madagascar Antsiranana, Madagascar McPherson 17219 (MO) Toamasina, Madagascar Ratovoson 170 (MO) Ambatovy, Madagascar Gostel 24 (MO) Ambatovy, Madagascar Gostel 25 (MO) Ambatovy, Madagascar Gostel 26 (MO) Antoetra, Madagascar Lowry 7082 (MO) Schefflera umbellifera (Sond.) Baillon Southern Africa S. Africa, Goldblatt 11950 (MO) Phillipson 5494 (MO) Schefflera vantsilana (Bak.) Bernardi Madagascar Toamasina, Madagascar Lowry 6225 (MO) Toamasina, Madagascar Randrianasolo 152 (MO) Ambatovy, Madagascar Gostel 32 (MO) Antoetra, Madagascar Lowry 7103 (MO) Schefflera “vohimenensis” Lowry ined. Madagascar Ivorona, Madagascar Plunkett 2378 (MO) Bemangedy, Madagascar Lowry 7156 (MO) Ivorona, Madagascar CR 4710 (MO) Ingroup, Astropanax subclade Schefflera barteri (Seem.) Harms Western sub-Saharan Africa and Cameroon, Etuge 5208 (P) São Tomé Gabon, McPherson 16998 (P) Cameroon, Thomas 6782 (NY) Schefflera goetzenii Harms Central and Eastern sub-Saharan Africa Zimbabwe, Lowry 4807 (MO) Rwanda, Auquier 3503 (BR) D.R. Congo, Cephas 358 (BR) Schefflera humblotiana Drake Madagascar Toamasina, Madagascar Schatz 3883 (P) Schefflera kivuensis P. Bamps Central sub-Saharan Africa Belgian Congo, Gutzwiller 2145 (BR) Schefflera mannii (Hook. f.) Harms Western sub-Saharan Africa Equitorial Guinea, Cavalho 4406 (BR) ?, s.n. ? Schefflera monophylla (Baker) Bernardi Madagascar Antsiranana, Madagascar Randrianasolo 409 (MO) Toamasina, Madagascar Randrianasolo 131 (MO)

Antoetra, Madagascar Lowry 7087 (MO) Antoetra, Madagascar Lowry 7097 (MO) Ivorona, Madagascar Lowry 7173 (MO) Andohahela, Madagascar RBE 1492 (MO) Schefflera myriantha (Baker) Drake Eastern sub-Saharan Africa, Comoro Tanzania, Lowry 4988 (MO) Islands, & Madagascar Antananarivo, Madagascar Lowry 5808 (MO) Antananarivo, Madagascar Lowry 5812 (MO) Antananarivo, Madagascar Lowry 5816 (MO) Mahajanga, Madagascar Lowry 5347 (MO) Mahajanga, Madagascar Lowry 5445 (MO) Anjavokely, Madagascar Lowry 5796 (P) Anjavokely, Madagascar Lowry 5798 (MO) Toamasina, Madagascar Lowry 6117 (MO) Tanzania, Mwangulango 501 (MO) Rwanda, P. Bamps 3050 (BR) Ethiopia, Danish-Ethiopian Exp. 1885 (BR) Malawi, Van der Linden L365 (BR) Antsiranana, Madagascar Ratovoson 470 (MO) Andohahela, Madagascar RBE 1498 (MO) Schefflera stolzii Harms Central-Eastern sub-Saharan Africa Gereau 3577 (P) Schefflera tessmannii Harms Western sub-Saharan Africa Gabon, Reitsma 2107 (P) Schefflera volkensii (Engl.) Harms Eastern sub-Saharan Africa Tanzania, Lowry 4987 (MO) Tanzania, Phillipson 5129 (P) Schefflera sp. Madagascar Makirovana, Madagascar RD 3332 (MO) Makirovana, Madagascar RD 3333 (MO) Makirovana, Madagascar RD 3334 (MO) Makirovana, Madagascar RD 3335 (MO) Makirovana, Madagascar RD 3336 (MO) Marojejy, Madagascar RD 3342 (MO) Marojejy, Madagascar RD 3344 (MO) Marojejy, Madagascar RD 3345 (MO) Marojejy, Madagascar RD 3347 (MO) Marojejy, RD 3348 (MO)

Marojejy, RD 3349 (MO) Marojejy, RD 3351 (MO) Marojejy, RD 3352 (MO)

TABLE 2: Oligonucleotide primers used for PCR amplification and DNA sequencing.

Primer Name Primer Region Primer Sequence (5’ – 3’) Primer Citation: Length Nuclear: ITS5-F ITS GGA AGT AAA AGT CGT AAC AAG G 22 bp White et al., 1990 C26A-R ITS TTT CTT TTC CTC CGC T 16 bp Wen and Zimmer, 1996 ETS-400F ETS GTT GGT CGG ATC CCT GCT TGT 21 bp Fiaschi & Plunkett (in press) 18S-2L (-r) ETS TGA CTA CTG GCA GGA TCA ACC AG 23 bp Linder et al., 2000

Plastid: ndhF-rpl32-F ndhF-rpl32 IGS GCA TAT TGA TAT GTC TGT TCC AT 23 bp Plunkett (unpubl.) ndhF-rpl32-R ndhF-rpl32 IGS AAG AGA TTT CCC TAA TGA CAA CGC 24 bp Plunkett (unpubl.) ndhF-rpl32-MF ndhF-rpl32 IGS GAG TAC TTG ATT CTG ATA TGA ATC 24 bp Gostel (unpubl.) ndhF-rpl32-MR ndhF-rpl32 IGS AGA ATC CGC CGT TAT GCC ATA G 22 bp Gostel (unpubl.) trnK-(rps16)_NF trnK-rps16 IGS AGC CGA GTA CTC TAC CGT TGA 21 bp Nicolas (unpubl.) (trnK)-rps16_NR trnK-rps16 IGS GAG CCG TCT ATC GAA TCG TTG CAA 24 bp Nicolas (unpubl.) trnK-rps16-MF trnK-rps16 IGS GCA CGG AAA TAA ATC GAT CCG C 22 bp Plunkett (unpubl.) trnK-rps16-MR trnK-rps16 IGS CGT TGG AAC TTT ACT AAC ACG 21 bp Plunkett (unpubl.) rpl32/trnL_F rpl32-trnL IGS GCG TTG TCA TTA GGG AAA TCT CTT 24 bp ?? rpl32/trnL_R rpl32-trnL IGS GCT TCC TAA GAG CAG CGT GTC T 22 bp ?? rpl32-trnL MF rpl32-trnL IGS CTA TCT CTA TAT CTA TAG AAG GGC AAA 27 bp Gostel (unpubl.) rpl32-trnL MR rpl32-trnL IGS TGA TTG GGT TGA AAG CAG GAG ATA TAT GGG AGA 38 bp Gostel (unpubl.) TTG AT

FIGURE LEGENDS

Fig. 1. The strict consensus of 51,600 trees resulting from the maximum parsimony (MP) analysis of 123 sequences from the nuclear ETS rDNA spacer. Tree length = 279 steps, CI =

Fig. 2. The strict consensus of 96,600 trees resulting from the maximum parsimony (MP) analysis of 123 sequences from the nuclear ITS rDNA spacer. Tree length = 271 steps, CI =

0.708, RI = 0.959. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. Parsimony bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as “<”.

Fig. 3. The strict consensus of 100,000 trees resulting from the maximum parsimony

(MP) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers. Tree length = 565 steps, CI = 0.665, and RI = 0.951. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. Parsimony bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as “<”.

Fig. 4. The strict consensus of 87,700 trees resulting from the maximum parsimony (MP) analysis of 123 sequences from the combined plastid markers, trnK-rps16, rpl32-trnL, and ndhF- rpl32. Tree length = 655 steps, CI = 0.612, RI = 0.918. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. Parsimony bootstrap percentages are

provided above the branches; bootstrap values less than 50% are recorded as “<”. Gray arrows indicate specimens whose placement has moved considerably from other analyses.

Fig. 5. The strict consensus of 94,200 trees resulting from the maximum parsimony (MP) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers and combined plastid markers, trnK-rps16, rpl32-trnL, and ndhF-rpl32. Tree length = 1,238 steps, CI = 0.608,

Fig. 6. The best tree (log likelihood = –2,279.056) based on maximum likelihood (ML) analysis of 123 sequences from the nuclear ETS rDNA spacer. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. ML bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as “<”.

Fig. 7. The best tree (log likelihood = –2,612.7117) based on maximum likelihood (ML) analysis of 123 sequences from the nuclear ITS rDNA spacer. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. ML bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as “<”.

Fig. 8. The best tree (log likelihood = –5,075.1868) based on maximum likelihood (ML) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. ML bootstrap

percentages are provided above the branches; bootstrap values less than 50% are recorded as

“<”.

Fig. 9. The best tree (log likelihood = –8,530.1115) based on maximum likelihood (ML) analysis of 123 sequences from the combined plastid markers, trnK-rps16, rpl32-trnL, and ndhF- rpl32. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. ML bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as “<”. Gray arrows indicate specimens whose placement has moved considerably from other analyses.

Fig. 10. The best tree (log likelihood = –14,328.1040) based on maximum likelihood

(ML) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers and combined plastid markers, trnK-rps16, rpl32-trnL, and ndhF-rpl32. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. ML bootstrap percentages are provided above the branches; bootstrap values less than 50% are recorded as “<”.

Fig. 11. The majority-rule tree based on the Bayesian-inference (BI) analysis of 123 sequences from the nuclear ETS rDNA spacer. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. BI posterior probabilities are provided above the branches.

Fig. 12. The majority-rule tree based on the Bayesian-inference (BI) analysis of 123 sequences from the nuclear ITS rDNA spacer. Brackets to the right of taxon labels correspond to

informal clade names discussed in the text. BI posterior probabilities are provided above the branches.

Fig. 13. The majority-rule tree based on the Bayesian-inference (BI) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. BI posterior probabilities are provided above the branches.

Fig. 14. The majority-rule tree based on the Bayesian-inference (BI) analysis of 123 sequences from the combined plastid markers, trnK-rps16, rpl32-trnL, and ndhF-rpl32. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. BI posterior probabilities are provided above the branches. Gray arrows indicate specimens whose placement has moved considerably from other analyses.

Fig. 15. The majority-rule tree based on the Bayesian-inference (BI) analysis of 123 sequences from the combined nuclear ITS + ETS rDNA spacers and combined plastid markers, trnK-rps16, rpl32-trnL, and ndhF-rpl32. Brackets to the right of taxon labels correspond to informal clade names discussed in the text. BI posterior probabilities are provided above the branches.

Fig. 16. The strict consensus tree resulting from maximum parsimony (MP) analysis of

123 sequences from the combined nuclear ITS + ETS rDNA spacers with geographic locality mapped onto branches using MacClade (version 4.08). DIVA (version 1.1a) output is provided

above branches to indicate ancestry for divergence nodes. “A” corresponds to continental Africa and “B” corresponds to Madagascar.

Fig. 17. The strict consensus tree resulting from maximum parsimony (MP) analysis of

123 sequences from the combined nuclear ITS + ETS rDNA spacers with carpel number states mapped onto branches.

Fig. 18. The strict consensus tree resulting from maximum parsimony (MP) analysis of

123 sequences from the combined nuclear ITS + ETS rDNA spacers with leaf composition character states mapped onto branches.

Fig. 19. The strict consensus tree resulting from maximum parsimony (MP) analysis of

123 sequences from the combined nuclear ITS + ETS rDNA spacers with inflorescence arrangement states mapped onto branches.

Fig. 20. The strict consensus tree resulting from maximum parsimony (MP) analysis of

123 sequences from the combined nuclear ITS + ETS rDNA spacers with pedicel lengths mapped onto branches.

Fig. 21. The strict consensus tree resulting from maximum parsimony (MP) analysis of

123 sequences from the combined nuclear ITS + ETS rDNA spacers with bract persistence mapped onto branches.

45 61 Schefflera favargeri 384 Schefflera favargeri 5344a Figure 1: MP, Schefflera favargeri 444 Schefflera favargeri 476 ETS Schefflera "humbertii" Schefflera halleana 4261 Schefflera cf. halleana 3339 65 Schefflera halleana 3343 Schefflera halleana 3340 Schefflera sp. 3332 Schefflera sp. 3333 Schefflera sp. 3334 Schefflera sp. 3342 Anticipes Schefflera sp. 3344 clade Schefflera sp. 3345 86 Schefflera bojeri 7071 Schefflera bojeri 7109 Schefflera bojeri 5818 64 Schefflera capuroniana 2328 Schefflera capuroniana 11118 < Schefflera “antoetrensis” 7088 Schefflera “antoetrensis” 7091 < Schefflera cf. capuroniana 28 Schefflera staufferana 24 Schefflera staufferana 26 61 Schefflera "vohimenensis" 2378 Schefflera "floretii" 4941 63 Schefflera "floretii" 7160 Schefflera "floretii" 7162 Schefflera "floretii" 7172 Schefflera "rabenantoandroi" 7148 Schefflera "rabenantoandroi" 7149 < Schefflera "vohimenensis" 7156 Schefflera "floretii" 4558 Palmate- Schefflera sp. 4710 Schefflera "floretii" 1485 Vantsilana < Schefflera "floretii" 1487 clade Schefflera longipedicellata 21 Schefflera longipedicellata 23 Neocussonia 65 Schefflera cf. longipedicellata 33 Schefflera vantsilana 6225 subclade Schefflera vantsilana 152 Schefflera vantsilana 32 Schefflera vantsilana 7103 62 Schefflera longipedicellata 6220 Schefflera longipedicellata 7098 < Schefflera longipedicellata 7104 < Schefflera longipedicellata 6153 Schefflera cf. staufferana 25 Schefflera sp. 3349 62 Schefflera sp. 3335 Schefflera sp. 3336 61 Schefflera sp. 3347 Schefflera sp. 3352 Schefflera sp. 3351 64 51 Schefflera fosbergiana 4322 Schefflera fosbergiana 3350 Schefflera cf. fosbergiana 3337 54 Schefflera rainaliana 4444 77 Schefflera rainaliana 14791 < Schefflera rainaliana 7151 78 Schefflera macerosa 1499 Schefflera macerosa 1509 84 Schefflera "purpuristyla" Schefflera bracteolifera Schefflera frodiniana Schefflera longipedicellata 6264 Schefflera staufferana 17219 100 Schefflera staufferana 170 Schefflera staufferana 7082 Schefflera moratii 31 Schefflera moratii 34 Schefflera sp. nov. 7164 Schefflera sp. nov. 7169 Schefflera sp. nov. 7171 Schefflera "andohahelensis" Schefflera lukwangulensis 98 Schefflera umbellifera 11950 Schefflera umbellifera 5494 Schefflera myriantha 5347 Schefflera myriantha 5445 Schefflera myriantha 470 87 Schefflera myriantha 5796 Schefflera myriantha 5798 Schefflera myriantha 5810 62 Schefflera myriantha 5808 Schefflera myriantha 5812 myriantha- Schefflera cf. myriantha 6117 Schefflera humblotiana monophylla 99 Schefflera monophylla 7087 clade 63 Schefflera monophylla 7097 Schefflera monophylla 7173 73 Schefflera cf. myriantha 5816 Schefflera monophylla 1492 51 Schefflera monophylla 409 Schefflera monophylla 131 71 Schefflera myriantha 1498 Astropanax Schefflera sp. 3348 Schefflera myriantha 501 subclade African 57 Schefflera myriantha 1885 Schefflera myriantha 3050 myriantha Schefflera myriantha L365 Schefflera myriantha 4988 clade < Schefflera goetzenii 4807 Schefflera goetzenii 398 88 Schefflera tessmannii Goetzenii Schefflera barteri 5208 Schefflera barteri 16998 clade Schefflera barteri 6782 Schefflera kivuensis 97 100 Schefflera volkensii 5129 Schefflera volkensii 4987 94 Schefflera mannii s.n. Schefflera mannii 4406 Schefflera goetzenii 3503 Schefflera stolzii 99 Cussonia thyrsiflora 83 97 Cussonia paniculata 68 Cussonia holstii 58 Seemannaralia gerrardii Outgroup Osmoxylon pectinatum Astrotricha pterocarpa Schefflera digitata

46 Schefflera bojeri 7071 60 Schefflera bojeri 7109 Figure 2: MP, Schefflera bojeri 5818 Schefflera cf. halleana 3339 ITS < Schefflera sp. 3332 Schefflera sp. 3334 Schefflera favargeri 384 Schefflera favargeri 476 Schefflera favargeri 5344a Schefflera favargeri 444 < Schefflera "humbertii" Anticipes Schefflera halleana 4261 Schefflera halleana 3343 clade Schefflera halleana 3340 Schefflera sp. 3333 Schefflera sp. 3342 Schefflera sp. 3344 Schefflera sp. 3345 < < Schefflera staufferana 17219 Schefflera staufferana 170 Schefflera staufferana 7082 Schefflera "antoetrensis" 7088 Schefflera "antoetrensis" 7091 56 Schefflera sp. 3335 53 Schefflera sp. 3336 Schefflera capuroniana 2328 Schefflera capuroniana 11118 89 Schefflera sp. nov. 7169 79 Schefflera sp. nov. 7171 Schefflera sp. nov. 7164 Schefflera "andohahelensis" 1505 67 Schefflera moratii 31 Schefflera moratii 34 66 Schefflera "vohimenensis" 7156 Schefflera "floretii" 4558 Schefflera sp. 4710 87 Schefflera "rabenantoandroi" 7148 Schefflera "rabenantoandroi" 7149 54 Schefflera "vohimenensis" 2378 Schefflera "floretii" 4941 Neocussonia Schefflera longipedicellata 21 Schefflera longipedicellata 23 Palmate- subclade Schefflera cf. longipedicellata 33 65 Schefflera vantsilana 6225 Vantsilana Schefflera vantsilana 152 clade Schefflera vantsilana 32 Schefflera vantsilana 7103 Schefflera "floretii" 7160 Schefflera "floretii" 7162 Schefflera "floretii" 7172 Schefflera "floretii" 1485 Schefflera "floretii" 1487 Schefflera longipedicellata 6220 63 Schefflera longipedicellata 7098 Schefflera longipedicellata 7104 73 Schefflera longipedicellata 6153 Schefflera sp. 3349 Schefflera longipedicellata 6264 88 Schefflera macerosa 1499 Schefflera macerosa 1509 82 Schefflera rainaliana 4444 86 Schefflera rainaliana 14791 < Schefflera rainaliana 7151 Schefflera frodiniana Schefflera cf. capuroniana 28 Schefflera bracteolifera Schefflera lukwangulensis Schefflera fosbergiana 4322 Schefflera cf. staufferana 25 Schefflera staufferana 24 100 Schefflera staufferana 26 Schefflera fosbergiana 3350 Schefflera cf. fosbergiana 3337 Schefflera sp. 3347 Schefflera sp. 3352 Schefflera sp. 3351 93 Schefflera umbellifera 11950 Schefflera umbellifera 5494 Schefflera "purpuristyla" Schefflera myriantha 5347 59 Schefflera myriantha 5445 Schefflera myriantha 470 Schefflera myriantha 5796 Schefflera myriantha 5808 90 Schefflera monophylla 409 Schefflera sp. 3348 Schefflera monophylla 7087 Schefflera monophylla 7097 Schefflera monophylla 7173 myriantha- Schefflera monophylla 131 monophylla 100 68 Schefflera cf. myriantha 5816 Schefflera cf. myriantha 6117 clade Schefflera myriantha 5798 Schefflera myriantha 5810 Schefflera myriantha 5812 Schefflera humblotiana Schefflera monophylla 1492 Astropanax 82 Schefflera myriantha 1498 64 Schefflera myriantha 501 subclade Schefflera myriantha 3050 African Schefflera myriantha 4988 myriantha Schefflera myriantha 1885 Schefflera myriantha L365 clade < 83 Schefflera volkensii 5129 Schefflera volkensii 4987 60 Schefflera mannii s.n. Schefflera mannii 4406 < Schefflera kivuensis Schefflera goetzenii 398 93 Schefflera tessmannii Schefflera barteri 5208 Goetzenii 100 66 Schefflera barteri 16998 Schefflera barteri 6782 clade 85 Schefflera goetzenii 4807 Schefflera goetzenii 3503 Schefflera stolzii 98 Cussonia thyrsiflora 74 Cussonia holstii Cussonia paniculata Seemannaralia gerrardii Outgroup < Astrotricha pterocarpa Osmoxylon pectinatum Schefflera digitata

47 65 Schefflera favargeri 384 Figure 3: MP, Schefflera favargeri 5344a Schefflera favargeri 444 Nuclear, combined 62 Schefflera cf. halleana 3339 Schefflera sp. 3332 Schefflera sp. 3334 Schefflera favargeri 476 Schefflera “humbertii” 60 Schefflera halleana 4261 Schefflera halleana 3343 Schefflera halleana 3340 Schefflera sp. 3333 58 Schefflera sp. 3342 Schefflera sp. 3344 Schefflera sp. 3345 94 Schefflera bojeri 7071 Schefflera bojeri 7109 Schefflera bojeri 5818 Anticipes 57 62 Schefflera staufferana 17219 Schefflera staufferana 170 clade 63 Schefflera “antoetrensis” 7088 Schefflera “antoetrensis” 7091 Schefflera staufferana 7082 62 Schefflera capuroniana 2328 Schefflera capuroniana 11118 88 Schefflera sp. 3335 Schefflera sp. 3336 63 Schefflera moratii 31 Schefflera moratii 34 Schefflera longipedicellata 6264 86 Schefflera sp. nov. 7169 83 Schefflera sp. nov. 7171 Schefflera sp. nov. 7164 Schefflera “andohahelensis” 63 Schefflera cf. capuroniana 28 Schefflera staufferana 24 Schefflera staufferana 26 Schefflera longipedicellata 21 Schefflera longipedicellata 23 Neocussonia 68 Schefflera cf. longipedicellata 33 Schefflera vantsilana 6225 subclade Schefflera vantsilana 152 Schefflera vantsilana 32 Schefflera vantsilana 7103 86 Schefflera “vohimenensis” 2378 Schefflera “floretii” 4941 62 Schefflera “floretii” 7160 Palmate- Schefflera “floretii” 7162 Vantsilana 65 Schefflera “floretii” 7172 62 Schefflera “vohimenensis” 7156 clade Schefflera “floretii” 4558 Schefflera sp. 4710 86 Schefflera “rabenantoandroi” 7148 Schefflera “rabenantoandroi” 7149 57 Schefflera “floretii” 1485 Schefflera “floretii” 1487 69 Schefflera longipedicellata 6220 Schefflera longipedicellata 7098 Schefflera longipedicellata 7104 Schefflera longipedicellata 6153 66 Schefflera cf. staufferana 25 Schefflera sp. 3349 98 Schefflera macerosa 1499 Schefflera macerosa 1509 61 Schefflera sp. 3347 Schefflera sp. 3352 72 Schefflera sp. 3351 78 Schefflera fosbergiana 4322 100 Schefflera fosbergiana 3350 Schefflera cf. fosbergiana 3337 97 Schefflera rainaliana 4444 70 Schefflera rainaliana 14791 Schefflera rainaliana 7151 Schefflera frodiniana 78 Schefflera “purpuristyla” Schefflera bracteolifera Schefflera lukwangulensis 100 Schefflera umbellifera 11950 Schefflera umbellifera 5494 Schefflera myriantha 5347 66 Schefflera myriantha 5445 Schefflera myriantha 470 67 Schefflera myriantha 5796 Schefflera myriantha 5808 66 Schefflera myriantha 5798 51 Schefflera myriantha 5810 Schefflera myriantha 5812 myriantha- Schefflera cf. myriantha 6117 100 Schefflera humblotiana monophylla 74 Schefflera monophylla 7087 Schefflera monophylla 7097 clade 63 Schefflera cf. myriantha 5816 95 Schefflera monophylla 7173 Schefflera monophylla 1492 84 Schefflera monophylla 409 Schefflera sp. 3348 96 Schefflera monophylla 131 Schefflera myriantha 1498 Astropanax 61 Schefflera myriantha 501 subclade Schefflera myriantha 3050 African 61 Schefflera myriantha 4988 myriantha Schefflera myriantha 1885 Schefflera myriantha L365 clade Schefflera goetzenii 398 90 Schefflera tessmannii Schefflera barteri 5208 Goetzenii Schefflera barteri 16998 66 Schefflera barteri 6782 clade Schefflera goetzenii 4807 100 Schefflera goetzenii 3503 100 Schefflera volkensii 5129 Schefflera volkensii 4987 98 Schefflera mannii s.n. Schefflera mannii 4406 Schefflera kivuensis Schefflera stolzii 99 99 Cussonia thyrsiflora 100 Cussonia paniculata Cussonia holstii 55 Seemannaralia gerrardii Outgroup 65 Astrotricha pterocarpa Osmoxylon pectinatum Schefflera digitata

48 Schefflera halleana 4261 Figure 4: MP, Schefflera cf. halleana 3339 61 Schefflera halleana 3343 Plastid, combined Schefflera halleana 3340 Schefflera sp. 3342 Schefflera sp. 3344 Schefflera sp. 3345 65 64 Schefflera sp. 3335 Schefflera sp. 3336 Schefflera sp. 3332 Schefflera sp. 3333 Schefflera sp. 3334 92 Schefflera fosbergiana 4322 Schefflera fosbergiana 3350 Schefflera sp. 3351 62 Schefflera moratii 31 Schefflera moratii 34 Schefflera umbellifera 11950 Schefflera umbellifera 5494 Schefflera longipedicellata 6220 Schefflera longipedicellata 6264 Schefflera longipedicellata 21 Schefflera longipedicellata 23 Schefflera longipedicellata 7098 Schefflera longipedicellata 7104 Schefflera longipedicellata 6153 Schefflera cf. longipedicellata 33 Schefflera “purpuristyla” Schefflera vantsilana 6225 Schefflera vantsilana 152 Schefflera vantsilana 32 Schefflera vantsilana 7103 Schefflera capuroniana 2328 Schefflera cf. capuroniana 28 Schefflera capuroniana 11118 Schefflera bracteolifera Schefflera "rabenantoandroi" 7148 Schefflera "rabenantoandroi" 7149 Schefflera lukwangulensis Schefflera rainaliana 4444 Neocussonia Schefflera rainaliana 14791 Schefflera rainaliana 7151 subclade Schefflera bojeri 7071 Schefflera bojeri 7109 Schefflera bojeri 5818 Schefflera favargeri 384 Schefflera favargeri 476 Schefflera favargeri 5344a Schefflera favargeri 444 Schefflera “humbertii” Schefflera “vohimenensis” 2378 Schefflera “vohimenensis” 7156 Schefflera "floretii" 4558 Schefflera sp. 4710 Schefflera "floretii" 4941 Schefflera staufferana 17219 Schefflera staufferana 170 Schefflera cf. staufferana 25 Schefflera staufferana 24 Schefflera staufferana 26 Schefflera staufferana 7082 Schefflera "antoetrensis" 7088 Schefflera "antoetrensis" 7091 Schefflera "floretii" 7160 Schefflera "floretii" 7162 Schefflera "floretii" 7172 Schefflera sp. nov. 7164 Schefflera sp. nov. 7169 Schefflera sp. nov. 7171 Schefflera "andohahelensis" Schefflera frodiniana Schefflera macerosa 1499 Schefflera macerosa 1509 Schefflera "floretii" 1485 Schefflera "floretii" 1487 Schefflera cf. fosbergiana 3337 Schefflera sp. 3347 Schefflera sp. 3352 Schefflera sp. 3349 65 Schefflera humblotiana Schefflera stolzii 100 98 Schefflera monophylla 7087 Schefflera monophylla 7097 Schefflera cf. myriantha 5816 86 Schefflera monophylla 7173 Schefflera monophylla 1492 Schefflera monophylla 409 Schefflera myriantha 5347 Schefflera myriantha 5445 myriantha- Schefflera myriantha 470 monophylla 74 Schefflera myriantha 5796 Schefflera cf. myriantha 6117 clade Schefflera myriantha 5798 Schefflera myriantha 5810 Schefflera myriantha 5808 61 Schefflera myriantha 5812 Schefflera myriantha 1498 Schefflera sp. 3348 Schefflera myriantha 1885 Astropanax 96 Schefflera myriantha 3050 African subclade 95 Schefflera myriantha L365 myriantha Schefflera myriantha 4988 Schefflera myriantha 501 clade 64 Schefflera tessmannii Schefflera barteri 5208 56 Schefflera barteri 6782 Schefflera monophylla 131 90 Schefflera goetzenii 4807 Schefflera goetzenii 398 67 65 Schefflera goetzenii 3503 Schefflera barteri 16998 99 Schefflera volkensii 5129 Schefflera volkensii 4987 Schefflera kivuensis 100 Schefflera mannii s.n. Schefflera mannii 4406 82 100 Cussonia thyrsiflora 69 Cussonia holstii Cussonia paniculata Seemannaralia gerrardii Outgroup Astrotricha pterocarpa Osmoxylon pectinatum Schefflera digitata

49 Schefflera halleana 4261 Figure 5: MP, Schefflera cf. halleana 3339 Schefflera halleana 3343 Nuclear + Plastid, combined Schefflera halleana 3340 60 Schefflera sp. 3332 Schefflera sp. 3333 Schefflera sp. 3334 Schefflera sp. 3342 76 Schefflera sp. 3344 Schefflera sp. 3345 65 Schefflera favargeri 384 Schefflera favargeri 5344A Schefflera favargeri 444 Schefflera favargeri 476 Schefflera “humbertii” 99 Schefflera bojeri 7071 Schefflera bojeri 7109 Anticipes Schefflera bojeri 5818 61 Schefflera “antoetrensis” 7088 clade 52 Schefflera “antoetrensis” 7091 Schefflera staufferana 7082 Schefflera staufferana 17219 Schefflera staufferana 170 65 Schefflera capuroniana 2328 Schefflera capuroniana 11118 98 Schefflera sp. 3335 Schefflera sp. 3336 Schefflera longipedicellata 6264 87 Schefflera moratii 31 Schefflera moratii 34 85 Schefflera sp. nov. 7169 63 Schefflera sp. nov. 7171 Schefflera sp. nov. 7164 Schefflera “andohahelensis” 75 Schefflera cf. capuroniana 28 Schefflera staufferana 24 Schefflera staufferana 26 62 Schefflera cf. longipedicellata 33 Schefflera vantsilana 32 Schefflera longipedicellata 21 Neocussonia < Schefflera longipedicellata 23 subclade Schefflera vantsilana 6225 Schefflera vantsilana 152 Schefflera vantsilana 7103 89 Schefflera “vohimenensis” 2378 Palmate- Schefflera “floretii” 4941 < Schefflera “floretii” 7160 Vantsilana Schefflera “floretii” 7162 clade < Schefflera “floretii” 7172 62 Schefflera “vohimenensis” 7156 Schefflera “floretii” 4558 Schefflera sp. 4710 85 Schefflera “rabenantoandroi” 7148 Schefflera “rabenantoandroi” 7149 60 Schefflera “floretii” 1485 Schefflera “floretii” 1487 54 Schefflera longipedicellata 6220 < Schefflera longipedicellata 7098 51 Schefflera longipedicellata 7104 < Schefflera longipedicellata 6153 Schefflera sp. 3349 Schefflera cf. staufferana 25 99 Schefflera macerosa 1499 Schefflera macerosa 1509 89 Schefflera fosbergiana 4322 Schefflera fosbergiana 3350 Schefflera sp. 3351 100 76 Schefflera cf. fosbergiana 3337 Schefflera sp. 3347 Schefflera sp. 3352 99 Schefflera rainaliana 4444 68 Schefflera rainaliana 14791 Schefflera rainaliana 7151 Schefflera frodiniana 89 Schefflera “purpuristyla” Schefflera bracteolifera Schefflera lukwangulensis 100 Schefflera umbellifera 11950 Schefflera umbellifera 5494 Schefflera myriantha 5347 Schefflera myriantha 5445 Schefflera myriantha 470 84 Schefflera myriantha 5796 Schefflera myriantha 5798 Schefflera myriantha 5810 Schefflera myriantha 5808 Schefflera myriantha 5812 myriantha- 100 Schefflera monophylla 7087 Schefflera monophylla 7097 71 monophylla 100 Schefflera cf. myriantha 5816 91 Schefflera monophylla 7173 clade 60 Schefflera monophylla 1492 85 Schefflera monophylla 409 Schefflera sp. 3348 62 Schefflera cf. myriantha 6117 Schefflera humblotiana 58 Schefflera monophylla 131 Astropanax Schefflera myriantha 1498 subclade 71 Schefflera myriantha 1885 African 84 Schefflera myriantha L365 99 Schefflera myriantha 3050 myriantha Schefflera myriantha 4988 Schefflera myriantha 501 clade 69 Schefflera tessmannii 80 Schefflera barteri 5208 89 Schefflera barteri 6782 Goetzenii < 69 Schefflera barteri 16998 96 Schefflera goetzenii 398 clade Schefflera goetzenii 4807 Schefflera goetzenii 3503 66 100 Schefflera volkensii 5129 Schefflera volkensii 4987 100 Schefflera kivuensis 100 Schefflera mannii s.n. Schefflera mannii 4406 Schefflera stolzii 100 Cussonia thyrsiflora 99 Cussonia holstii Cussonia paniculata 62 Seemannaralia gerrardii Outgroup 69 Astrotricha pterocarpa Osmoxylon pectinatum Schefflera digitata

50 Schefflera sp. 3334 Figure 6: ML, Schefflera favargeri 476 58 Schefflera sp. 3332 ETS Schefflera “humbertii” Schefflera halleana 3343 Schefflera halleana 3340 Schefflera sp. 3344 Schefflera favargeri 444 55 Schefflera favargeri 384 Schefflera favargeri 5344a Schefflera sp. 3342 Anticipes Schefflera halleana 4261 clade 69 Schefflera sp. 3345 Schefflera cf. halleana 3339 Schefflera sp. 3333 82 Schefflera “purpuristyla” Schefflera bracteolifera Schefflera "antoetrensis" 7088 Schefflera "antoetrensis" 7091 Schefflera staufferana 170 Schefflera sp. 3349 Schefflera longipedicellata 6153 Schefflera cf. staufferana 25 < 68Schefflera longipedicellata 7098 Schefflera longipedicellata 7104 Schefflera longipedicellata 6220 Schefflera staufferana 17219 Schefflera staufferana 7082 Schefflera sp. nov. 7171 Schefflera frodiniana 68 Schefflera sp. 3336 Schefflera sp. 3335 59 72Schefflera capuroniana 2328 Schefflera capuroniana 11118 Schefflera lukwangulensis Schefflera longipedicellata 6264 Schefflera rainaliana 14791 55 Schefflera rainaliana 4444 Schefflera rainaliana 7151 Neocussonia < 68 Schefflera macerosa 1499 subclade Schefflera macerosa 1509 66 Schefflera "floretii" 4941 Schefflera "vohimenensis" 2378 Schefflera "floretii" 7162 62 Schefflera "floretii" 7160 Schefflera "floretii" 7172 Schefflera "rabenantoandroi" 7149 Schefflera "vohimenensis" 7156 Schefflera sp. 4710 Palmate- < Schefflera "floretii" 4558 Schefflera "rabenantoandroi" 7148 Vantsilana Schefflera "floretii" 1487 Schefflera "floretii" 1485 clade Schefflera vantsilana 6225 < Schefflera vantsilana 7103 Schefflera vantsilana 32 Schefflera longipedicellata 23 Schefflera longipedicellata 21 Schefflera vantsilana 152 55 Schefflera cf. longipedicellata 33 Schefflera bojeri 7071 86 Schefflera bojeri 5818 Schefflera bojeri 7109 Schefflera sp. nov. 7169 Schefflera "moratii" 31 Schefflera "andohahelensis" Schefflera staufferana 24 51 Schefflera staufferana 26 Schefflera cf. capuroniana 28 Schefflera sp. nov. 7164 Schefflera "moratii" 34 62 Schefflera sp. 3351 65 Schefflera sp. 3347 56 Schefflera sp. 3352 96 Schefflera fosbergiana 3350 66 Schefflera fosbergiana 4322 Schefflera cf. fosbergiana 3337 97 Schefflera umbellifera 5494 Schefflera umbellifera 11950 Schefflera myriantha 5796 Schefflera myriantha 5347 Schefflera myriantha 470 Schefflera myriantha 5810 Schefflera myriantha 5812 89 Schefflera myriantha 5445 Schefflera myriantha 5798 70 Schefflera myriantha 5808 myriantha- Schefflera cf. myriantha 6117 Schefflera humblotiana 3883 monophylla Schefflera monophylla 7173 clade Schefflera monophylla 7097 56 Schefflera monophylla 7087 Schefflera monophylla 1492 97 76 Schefflera cf. myriantha 5816 Schefflera myriantha 1498 Schefflera sp. 3348 58 Schefflera monophylla 131 Astropanax Schefflera monophylla 409 Schefflera myriantha L365 subclade Schefflera myriantha 3050 African Schefflera myriantha 1885 myriantha 57 Schefflera myriantha 4988 Schefflera myriantha 501 clade < Schefflera kivuensis 65 Schefflera barteri 5208 < Schefflera goetzenii 4807 < 88 Schefflera barteri 6782 Goetzenii Schefflera goetzenii 398 Schefflera tessmannii clade Schefflera barteri 16998 Schefflera goetzenii 3503 99 94 Schefflera volkensii 4987 Schefflera volkensii 5129 Schefflera stolzii 90 Schefflera mannii 4406 Schefflera mannii s.n. 96 Cussonia thyrsiflora 73 Cussonia paniculata 88 Cussonia holstii Seemannaralia gerrardii Outgroup Schefflera digitata Osmoxylon pectinatum Astrotricha pterocarpa 0.02

51 Schefflera bojeri 5818 Figure 7: ML, 61 Schefflera bojeri 7071 Schefflera bojeri 7109 ITS Schefflera halleana 4261 Schefflera halleana 3343 Schefflera favargeri 476 Schefflera "humbertii" Schefflera favargeri 5344a Schefflera halleana 3340 Schefflera sp. 3333 Schefflera sp. 3344 Schefflera sp. 3345 < Schefflera sp. 3342 Schefflera favargeri 444 Schefflera favargeri 384 Schefflera cf. halleana 3339 < Schefflera sp. 3334 Schefflera sp. 3332 Schefflera staufferana 7082 Schefflera "antoetrensis" 7091 Anticipes Schefflera "antoetrensis" 7088 clade 64Schefflera staufferana 17219 Schefflera staufferana 170 51 Schefflera sp. 3336 < Schefflera sp. 3335 Schefflera capuroniana 2328 < Schefflera capuroniana 11118 Schefflera staufferana 24 Schefflera moratii 34 Schefflera moratii 31 65 78 Schefflera sp. nov. 7169 78 Schefflera sp. nov. 7171 Schefflera "andohahelensis" Schefflera sp. nov. 7164 Schefflera cf. capuroniana 28 Schefflera staufferana 26 Schefflera longipedicellata 6153 Schefflera longipedicellata 6220 Schefflera longipedicellata 7098 Neocussonia 68 Schefflera sp. 3349 subclade 66 Schefflera longipedicellata 7104 Schefflera longipedicellata 6264 54Schefflera "floretii" 1485 Schefflera "floretii" 1487 Schefflera "floretii" 7172 61Schefflera "floretii" 4941 Schefflera "vohimenensis" 2378 Schefflera vantsilana 32 Schefflera vantsilana 6225 Schefflera longipedicellata 23 Palmate- Schefflera vantsilana 7103 Schefflera "floretii" 7160 Vantsilana 88 Schefflera "rabenantoandroi" 7148 Schefflera "rabenantoandroi" 7149 clade Schefflera cf. longipedicellata 33 Schefflera "floretii" 7162 Schefflera vantsilana 152 68 Schefflera longipedicellata 21 Schefflera "vohimenensis" 7156 Schefflera "floretii" 4558 58 Schefflera sp. 4710 87 Schefflera macerosa 1509 Schefflera macerosa 1499 Schefflera fosbergiana 4322 Schefflera fosbergiana 3350 Schefflera sp. 3351 53Schefflera sp. 3347 < Schefflera bracteolifera < Schefflera sp. 3352 Schefflera cf. fosbergiana 3337 Schefflera rainaliana 14791 82 Schefflera rainaliana 7151 87 Schefflera rainaliana 4444 95 Schefflera frodiniana 1486 Schefflera lukwangulensis Schefflera cf. staufferana 25 92 Schefflera umbellifera 5494 Schefflera umbellifera 11950 Schefflera "purpuristyla" Schefflera cf. myriantha 5816 Schefflera monophylla 7087 Schefflera monophylla 7097 < Schefflera myriantha 5810 Schefflera myriantha 5798 < Schefflera myriantha 5812 Schefflera monophylla 1492 < Schefflera monophylla 7173 myriantha- Schefflera monophylla 131 Schefflera myriantha 1498 monophylla 82 Schefflera monophylla 409 clade Schefflera sp. 3348 74 Schefflera myriantha 5347 100 Schefflera myriantha 5796 Schefflera myriantha 5445 Schefflera myriantha 5808 Schefflera myriantha 470 80 Schefflera cf. myriantha 6117 Astropanax 66 Schefflera humblotiana Schefflera myriantha 3050 subclade Schefflera myriantha 4988 African Schefflera myriantha 501 myriantha Schefflera myriantha 1885 Schefflera myriantha L365 clade 87 Schefflera volkensii 5129 Schefflera volkensii 4987 Schefflera barteri 16998 Schefflera barteri 5208 86 Schefflera barteri 6782 Goetzenii Schefflera tessmannii 69 Schefflera goetzenii 398 clade 78 Schefflera goetzenii 3503 57 Schefflera goetzenii 4807 100 Schefflera stolzii 3577 61 Schefflera mannii 4406 Schefflera mannii s.n. Schefflera kivuensis Cussonia thyrsiflora 91 Cussonia holstii 71 Cussonia paniculata < Seemannaralia gerrardii Outgroup Osmoxylon pectinatum Schefflera digitata Astrotricha pterocarpa 0.02

52 Schefflera sp. 3344 Figure 8: ML, Schefflera "humbertii" Schefflera favargeri 476 Nuclear, combined Schefflera halleana 3340 Schefflera sp. 3333 Schefflera sp. 3342 Schefflera halleana 3343 62Schefflera sp. 3332 Schefflera sp. 3334 Schefflera cf. halleana 3339 Schefflera halleana 4261 69 Schefflera sp. 3345 64Schefflera favargeri 5344a 51 Schefflera favargeri 444 Schefflera favargeri 384 94Schefflera bojeri 7071 Schefflera bojeri 5818 Schefflera bojeri 7109 Anticipes Schefflera staufferana 7082 clade 61 57Schefflera staufferana 17219 Schefflera staufferana 170 66Schefflera "antoetrensis" 7088 Schefflera "antoetrensis" 7091 83 Schefflera sp. nov. 7169 Schefflera sp. nov. 7171 84 Schefflera sp. nov. 7164 Schefflera "andohahelensis" 1505 86 Schefflera sp. 3336 Schefflera sp. 3335 68 64Schefflera capuroniana 11118 Schefflera capuroniana 2328 Schefflera longipedicellata 6264 60Schefflera moratii 34 Schefflera moratii 31 56Schefflera staufferana 26 Schefflera cf. capuroniana 28 Schefflera staufferana 24 87 Schefflera “purpuristyla” Schefflera bracteolifera Neocussonia Schefflera rainaliana 4444 97 Schefflera rainaliana 14791 subclade 84 Schefflera rainaliana 7151 Schefflera frodiniana 1486 85 Schefflera "vohimenensis" 2378 Schefflera "floretii" 4941 81 Schefflera "floretii" 7162 Schefflera "floretii" 7160 Schefflera "floretii" 7172 60Schefflera "floretii" 1487 Schefflera "floretii" 1485 66Schefflera "floretii" 4558 Palmate- Schefflera sp. 4710 Schefflera "vohimenensis" 7156 Vantsilana 88 Schefflera "rabenantoandroi" 7148 Schefflera "rabenantoandroi" 7149 clade 74 Schefflera longipedicellata 21 Schefflera longipedicellata 23 Schefflera cf. longipedicellata 33 Schefflera vantsilana 32 Schefflera vantsilana 6225 Schefflera vantsilana 7103 67 Schefflera vantsilana 152 100 Schefflera macerosa 1499 Schefflera macerosa 1509 Schefflera longipedicellata 6220 65 Schefflera longipedicellata 7104 60 Schefflera longipedicellata 7098 51 Schefflera sp. 3349 Schefflera longipedicellata 6153 Schefflera cf. staufferana 25 Schefflera lukwangulensis 81 Schefflera fosbergiana 3350 100 Schefflera fosbergiana 4322 Schefflera cf. fosbergiana 3337 Schefflera sp. 3347 68 Schefflera sp. 3352 61 Schefflera sp. 3351 99 Schefflera umbellifera 11950 Schefflera umbellifera 5494 Schefflera myriantha 5808 Schefflera myriantha 5796 57 Schefflera myriantha 5347 Schefflera myriantha 5445 61 Schefflera myriantha 470 Schefflera myriantha 5812 51 Schefflera myriantha 5798 myriantha- 58 Schefflera myriantha 5810 Schefflera cf. myriantha 6117 monophylla Schefflera humblotiana Schefflera monophylla 7087 clade 100 98 57 Schefflera monophylla 7097 Schefflera cf. myriantha 5816 61 Schefflera monophylla 7173 Schefflera monophylla 1492 Schefflera myriantha 1498 95 Schefflera monophylla 131 Astropanax 81 Schefflera monophylla 409 Schefflera sp. 3348 subclade 66 Schefflera myriantha 3050 Schefflera myriantha 501 African 56 Schefflera myriantha 4988 myriantha Schefflera myriantha 1885 Schefflera myriantha L365 clade 60 Schefflera kivuensis 56 Schefflera barteri 5208 Schefflera barteri 6782 Schefflera barteri 16998 Goetzenii 76 Schefflera tessmannii 100 69 Schefflera goetzenii 398 clade Schefflera goetzenii 4807 Schefflera goetzenii 3503 100 Schefflera volkensii 4987 Schefflera volkensii 5129 Schefflera stolzii 94 Schefflera mannii 4406 Schefflera mannii s.n. 98 Cussonia thyrsiflora 97 Cussonia paniculata 98 Cussonia holstii Seemannaralia gerrardii Outgroup Schefflera digitata Osmoxylon pectinatum Astrotricha pterocarpa

0.02

53 Schefflera fosbergiana 3350 Figure 9: ML, Schefflera "floretii" 7172 Schefflera fosbergiana 4322 Plastid, combined Schefflera sp. 3351 Schefflera cf. fosbergiana 3337 77 Schefflera vohimenensis 2378 Schefflera "floretii" 1485 Schefflera sp. nov. 7169 Schefflera vantsilana 6225 Schefflera longipedicellata 7098 Schefflera "floretii" 1487 Schefflera sp. 3349 Schefflera "rabenantoandroi" 7149 Schefflera sp. 3347 Schefflera “vohimenensis” 7156 Schefflera rainaliana 14791 89 Schefflera rainaliana 4444 Schefflera rainaliana 7151 Schefflera cf. staufferana 25 Schefflera staufferana 24 61 Schefflera staufferana 26 Schefflera cf. capuroniana 28 Schefflera sp. nov. 7164 61 Schefflera “antoetrensis” 7091 Schefflera antoetrensis 7088 Schefflera staufferana 7082 Schefflera "floretii" 4558 Schefflera vantsilana 32 64 Schefflera cf. longipedicellata 33 < Schefflera longipedicellata 23 Schefflera "rabenantoandroi" 7148 Schefflera lukwangulensis Schefflera frodiniana Schefflera "floretii" 7162 Schefflera bojeri 7109 72 Schefflera bojeri 5818 Schefflera bojeri 7071 Schefflera "floretii" 4941 Schefflera longipedicellata 7104 Neocussonia Schefflera macerosa 1509 subclade Schefflera longipedicellata 6153 Schefflera sp. 4710 Schefflera longipedicellata 21 Schefflera vantsilana 7103 Schefflera sp. nov. 7171 Schefflera macerosa 1499 Schefflera "floretii" 7160 Schefflera staufferana 170 Schefflera sp. 3352 60Schefflera sp. 3344 Schefflera sp. 3345 Schefflera cf. halleana 3339 Schefflera halleana 3340 Schefflera halleana 3343 Schefflera sp. 3342 Schefflera halleana 4261 60 Schefflera sp. 3333 Schefflera sp. 3332 Schefflera sp. 3334 Schefflera sp. 3335 Schefflera sp. 3336 68 Schefflera capuroniana 11118 Schefflera favargeri 476 Schefflera favargeri 444 Schefflera moratii 34 Schefflera moratii 31 Schefflera capuroniana 2328 Schefflera "andohahelensis" 1505 Schefflera favargeri 384 Schefflera longipedicellata 6220 71Schefflera umbellifera 11950 55 Schefflera longipedicellata 6264 Schefflera umbellifera 5494 Schefflera vantsilana 152 Schefflera "humbertii" 5386 Schefflera favargeri 5344a Schefflera "purpuristyla" 432 Schefflera staufferana 17219 Schefflera bracteolifera Schefflera myriantha 5812 Schefflera myriantha 5798 58 Schefflera myriantha 5796 Schefflera myriantha 5810 Schefflera myriantha 5445 82 Schefflera myriantha 5347 Schefflera myriantha 5808 100 Schefflera monophylla 409 myriantha- Schefflera myriantha 470 monophylla Schefflera sp. 3348 Schefflera monophylla 7173 clade 73 Schefflera monophylla 1492 Schefflera cf. myriantha 5816 99 Schefflera monophylla 7097 Schefflera monophylla 7087 66 Schefflera cf. myriantha 6117 Schefflera myriantha 1498 Schefflera myriantha L365 Astropanax 91 Schefflera myriantha 4988 African 99 Schefflera myriantha 1885 myriantha subclade Schefflera myriantha 3050 93 Schefflera myriantha 501 clade 55 Schefflera barteri 16998 Schefflera monophylla 131 Schefflera goetzenii 4807 Schefflera goetzenii 398 Schefflera barteri 6782 92 67 Schefflera barteri 5208 Schefflera tessmannii Schefflera goetzenii 3503 93 Schefflera volkensii 5129 Schefflera volkensii 4987 99 Schefflera mannii s.n. 64 Schefflera mannii 4406 Schefflera kivuensis Schefflera stolzii Schefflera humblotiana Schefflera digitata 93 Cussonia holstii 100 Cussonia thyrsiflora 87 Cussonia paniculata Outgroup Seemannaralia gerrardii Astrotricha pterocarpa Osmoxylon pectinatum 0.0060

54 Schefflera sp. 3342 Figure 10: ML, 77 Schefflera sp. 3345 Schefflera halleana 3340 Nuclear + PLastid, combined Schefflera halleana 3343 Schefflera sp. 3344 Schefflera cf. halleana 3339 Schefflera halleana 4261 57 Schefflera sp. 3332 Schefflera sp. 3334 70 Schefflera sp. 3333 66Schefflera favargeri 5344a Schefflera favargeri 444 Schefflera favargeri 384 Schefflera "humbertii" 5386 Schefflera favargeri 476 93 Schefflera bojeri 7071 Schefflera bojeri 5818 Schefflera bojeri 7109 Schefflera staufferana 170 Anticipes Schefflera staufferana 17219 70Schefflera "antoetrensis" 7091 clade 56 Schefflera "antoetrensis" 7088 Schefflera staufferana 7082 Schefflera staufferana 26 84 Schefflera cf. capuroniana 28 Schefflera staufferana 24 91 Schefflera sp. nov. 7171 Schefflera sp. nov. 7169 56 Schefflera sp. nov. 7164 Schefflera "andohahelensis" 1505 83Schefflera capuroniana 2328 Schefflera capuroniana 11118 80 96Schefflera sp. 3336 Schefflera sp. 3335 85Schefflera moratii 34 Schefflera moratii 31 Schefflera longipedicellata 6264 90 Schefflera "vohimenensis" 2378 Neocussonia Schefflera "floretii" 4941 subclade Schefflera "floretii" 7160 61 Schefflera "floretii" 7172 Schefflera "floretii" 7162 Schefflera "floretii" 1487 Schefflera "floretii" 1485 Schefflera "vohimenensis" 7156 Schefflera "floretii" 4558 Palmate- Schefflera sp. 4710 Vantsilana 58Schefflera vantsilana 32 Schefflera cf. longipedicellata 33 clade Schefflera vantsilana 7103 Schefflera longipedicellata 21 58 Schefflera longipedicellata 23 Schefflera vantsilana 152 Schefflera vantsilana 6225 87Schefflera "rabenantoandroi" 7149 Schefflera "rabenantoandroi" 7148 Schefflera macerosa 1499 100 Schefflera macerosa 1509 Schefflera longipedicellata 7104 Schefflera longipedicellata 7098 Schefflera longipedicellata 6220 53Schefflera longipedicellata 6153 Schefflera sp. 3349 Schefflera cf. staufferana 25 Schefflera "bracteolifera" 77Schefflera "purpuristyla" Schefflera lukwangulensis Schefflera rainaliana 4444 100 Schefflera rainaliana 14791 99 75 Schefflera rainaliana 7151 Schefflera frodiniana 92 Schefflera fosbergiana 4322 Schefflera fosbergiana 3350 Schefflera sp. 3351 83 Schefflera sp. 3347 Schefflera sp. 3352 Schefflera cf. fosbergiana 3337 100 Schefflera umbellifera 5494 Schefflera umbellifera 11950 Schefflera myriantha 5812 59 Schefflera myriantha 5798 56 Schefflera myriantha 5810 53 Schefflera myriantha 5796 86 Schefflera myriantha 5347 85 Schefflera myriantha 5445 Schefflera myriantha 5808 Schefflera myriantha 470 myriantha- 82Schefflera cf. myriantha 6117 Schefflera humblotiana monophylla 57 Schefflera monophylla 7087 100 100 Schefflera cf. myriantha 5816 clade Schefflera monophylla 7097 85Schefflera monophylla 7173 60 Schefflera monophylla 1492 Schefflera sp. 3348 Schefflera monophylla 409 Schefflera myriantha 1498 Astropanax 65 Schefflera monophylla 131 subclade 75 Schefflera myriantha L365 90 Schefflera myriantha 1885 African Schefflera myriantha 3050 myriantha Schefflera myriantha 4988 99 68 Schefflera myriantha 501 clade 68 74 Schefflera barteri 5208 63 Schefflera barteri 6782 89 Schefflera tessmannii Goetzenii 83 Schefflera barteri 16998 93 Schefflera goetzenii 398 clade Schefflera goetzenii 4807 Schefflera goetzenii 3503 64 Schefflera kivuensis 100 Schefflera mannii s.n. 100 Schefflera mannii 4406 100 Schefflera volkensii 4987 Schefflera volkensii 5129 Schefflera stolzii Schefflera digitata Cussonia paniculata 100 Cussonia thyrsiflora 100 Cussonia holstii Outgroup Seemannaralia gerrardii Osmoxylon pectinatum Astrotricha pterocarpa

0.0070

55 .95 Schefflera favargeri 384 Schefflera favargeri 5344a Figure 11: Bayesian, Schefflera favargeri 444 Schefflera favargeri 476 ETS Schefflera “humbertii” Schefflera halleana 4261 Schefflera cf. halleana 3339 Anticipes .95 Schefflera halleana 3343 Schefflera halleana 3340 clade Schefflera sp. 3332 Schefflera sp. 3333 Schefflera sp. 3334 Schefflera sp. 3342 Schefflera sp. 3344 Schefflera sp. 3345 1 Schefflera bojeri 7071 Schefflera bojeri 7109 Schefflera bojeri 5818 .89 Schefflera “antoetrensis” 7088 Schefflera “antoetrensis” 7091 Schefflera staufferana 17219 Schefflera staufferana 170 Schefflera staufferana 7082 .98 Schefflera capuroniana 2328 .88 Schefflera capuroniana 11118 .99 Schefflera sp. 3335 Schefflera sp. 3336 Schefflera sp. nov. 7164 Schefflera sp. nov. 7169 Schefflera sp. nov. 7171 Schefflera ”andohahelensis” .99 Schefflera cf. capuroniana 28 Schefflera staufferana 24 Schefflera staufferana 26 Schefflera moratii 31 Schefflera moratii 34 Schefflera longipedicellata 6264 .99 Schefflera “vohimenensis” 2378 Schefflera ”floretii” 4941 .96 Schefflera “floretii” 7160 Neocussonia Schefflera ”floretii” 7162 Schefflera ”floretii” 7172 subclade Schefflera ”rabenantoandroi” 7148 Schefflera ”rabenantoandroi” 7149 .73 Schefflera ”vohimenensis” 7156 Schefflera ”floretii” 4558 Palmate- Schefflera sp. 4710 Vantsilana .71 Schefflera ”floretii” 1485 Schefflera ”floretii” 1487 clade Schefflera longipedicellata 21 Schefflera longipedicellata 23 .99 Schefflera cf. longipedicellata 33 Schefflera vantsilana 6225 Schefflera vantsilana 152 Schefflera vantsilana 32 Schefflera vantsilana 7103 1 Schefflera macerosa 1499 Schefflera macerosa 1509 .69 .99 Schefflera longipedicellata 6220 Schefflera longipedicellata 7098 Schefflera longipedicellata 7104 .91 Schefflera longipedicellata 6153 Schefflera cf. staufferana 25 Schefflera sp. 3349 .99 Schefflera sp. 3347 Schefflera sp. 3352 Schefflera sp. 3351 .96 .96 Schefflera fosbergiana 4322 1 Schefflera fosbergiana 3350 Schefflera cf. fosbergiana 3337 .96 Schefflera rainaliana 4444 Schefflera rainaliana 14791 Schefflera rainaliana 7151 Schefflera lukwangulensis 1 Schefflera “purpuristyla” Schefflera bracteolifera Schefflera frodiniana 1486 1 Schefflera umbellifera 11950 Schefflera umbellifera 5494 Schefflera myriantha 5347 Schefflera myriantha 5445 Schefflera myriantha 470 1 Schefflera myriantha 5796 Schefflera myriantha 5798 Schefflera myriantha 5810 .94 Schefflera myriantha 5808 Schefflera myriantha 5812 Schefflera cf. myriantha 6117 myriantha- Schefflera humblotiana monophylla 1 Schefflera monophylla 7087 .99 Schefflera monophylla 7097 clade Schefflera monophylla 7173 1 Schefflera cf. myriantha 5816 Schefflera monophylla 1492 .95 Schefflera monophylla 409 Schefflera monophylla 131 .98 Schefflera myriantha 1498 Schefflera sp. 3348 Astropanax Schefflera myriantha 501 .89 Schefflera myriantha 1885 African subclade Schefflera myriantha 3050 Schefflera myriantha L365 myriantha Schefflera myriantha 4988 clade .57 .96 .76 Schefflera goetzenii 4807 Schefflera barteri 5208 1 Schefflera goetzenii 398 .87 Schefflera tessmannii Schefflera barteri 16998 Schefflera barteri 6782 Schefflera kivuensis Schefflera goetzenii 3503 1 1 .69 Schefflera volkensii 5129 Schefflera volkensii 4987 Schefflera stolzii 1 Schefflera mannii s.n. Schefflera mannii 4406 1 .87 Cussonia thyrsiflora 1 Cussonia paniculata Cussonia holstii Seemannaralia gerrardii Outgroup Astrotricha pterocarpa Osmoxylon pectinatum Schefflera digitata

56 .97 Schefflera bojeri 7071 Figure 12: Bayesian, Schefflera bojeri 7109 ITS Schefflera bojeri 5818 .95 Schefflera cf. halleana 3339 Schefflera sp. 3332 Schefflera sp. 3334 Schefflera favargeri 384 Schefflera favargeri 476 Schefflera favargeri 5344a Schefflera favargeri 444 .91 Schefflera “humbertii” Anticipes Schefflera halleana 4261 Schefflera halleana 3343 clade Schefflera halleana 3340 Schefflera sp. 3333 Schefflera sp. 3342 Schefflera sp. 3344 Schefflera sp. 3345 .96 .99 Schefflera staufferana 17219 Schefflera staufferana 170 Schefflera staufferana 7082 Schefflera “antoetrensis” 7088 Schefflera “antoetrensis” 7091 Schefflera longipedicellata 6220 .98 Schefflera longipedicellata 7098 Schefflera longipedicellata 7104 1 Schefflera longipedicellata 6153 Schefflera sp. 3349 Schefflera longipedicellata 6264 .99 Schefflera sp. 3335 .97 Schefflera sp. 3336 Schefflera capuroniana 2328 Schefflera capuroniana 11118 .63 1 Schefflera sp. nov. 7169 1 Schefflera sp. nov. 7171 Schefflera sp. nov. 7164 Schefflera “andohahelensis” .98 Schefflera moratii 31 Schefflera moratii 34 Neocussonia Schefflera cf. capuroniana 28 Schefflera staufferana 24 subclade Schefflera staufferana 26 .97 Schefflera “vohimenensis” 7156 Schefflera “floretii” 4558 Schefflera sp. 4710 1 Schefflera “rabenantoandroi” 7148 Schefflera “rabenantoandroi” 7149 .99 Schefflera “vohimenensis” 2378 Schefflera “floretii” 4941 .92 Schefflera “floretii” 1485 Schefflera “floretii” 1487 Palmate- Schefflera longipedicellata 21 Vantsilana .89 Schefflera longipedicellata 23 Schefflera cf. longipedicellata 33 clade Schefflera vantsilana 6225 Schefflera vantsilana 152 Schefflera vantsilana 32 Schefflera vantsilana 7103 Schefflera “floretii” 7160 Schefflera “floretii” 7162 Schefflera “floretii” 7172 1 Schefflera rainaliana 4444 1 Schefflera rainaliana 14791 Schefflera rainaliana 7151 Schefflera frodiniana 1 Schefflera umbellifera 11950 Schefflera umbellifera 5494 1 .8 Schefflera fosbergiana 4322 Schefflera fosbergiana 3350 1 Schefflera macerosa 1499 Schefflera macerosa 1509 Schefflera “purpuristyla” Schefflera bracteolifera Schefflera lukwangulensis Schefflera cf. staufferana 25 Schefflera cf. fosbergiana 3337 Schefflera sp. 3347 Schefflera sp. 3352 Schefflera sp. 3351 .96 Schefflera myriantha 5798 Schefflera myriantha 5812 .56 Schefflera monophylla 7087 .65 Schefflera monophylla 7097 Schefflera cf. myriantha 5816 .98 Schefflera myriantha 5810 Schefflera monophylla 7173 .69 Schefflera monophylla 1492 myriantha- Schefflera monophylla 131 1 Schefflera myriantha 1498 monophylla Schefflera myriantha 5347 clade .99 Schefflera myriantha 5445 Schefflera myriantha 470 1 Schefflera myriantha 5796 Schefflera myriantha 5808 1 Schefflera monophylla 409 Schefflera sp. 3348 Schefflera cf. myriantha 6117 Astropanax Schefflera humblotiana 1 .98 Schefflera myriantha 501 subclade Schefflera myriantha 3050 African Schefflera myriantha 4988 myriantha Schefflera myriantha 1885 Schefflera myriantha L365 clade Schefflera goetzenii 398 1 Schefflera tessmannii Schefflera barteri 5208 Goetzenii .99 Schefflera barteri 16998 Schefflera barteri 6782 clade .64 .81 1 Schefflera goetzenii 4807 1 Schefflera goetzenii 3503 Schefflera stolzii 1 Schefflera volkensii 5129 Schefflera volkensii 4987 .86 Schefflera mannii s.n. Schefflera mannii 4406 Schefflera kivuensis 1 Cussonia thyrsiflora 1 Cussonia holstii Cussonia paniculata Seemannaralia gerrardii Outgroup Schefflera digitata Astrotricha pterocarpa Osmoxylon pectinatum

57 .97 Schefflera favargeri 384 Figure 13: Bayesian, Schefflera favargeri 5344a Nuclear, combined .51 Schefflera favargeri 444 .94 Schefflera sp. 3332 Schefflera sp. 3334 Schefflera cf. halleana 3339 Schefflera favargeri 476 Schefflera “humbertii” .96 Schefflera halleana 4261 Schefflera halleana 3343 Schefflera halleana 3340 Schefflera sp. 3333 .94 Schefflera sp. 3342 Schefflera sp. 3344 Schefflera sp. 3345 1 Schefflera bojeri 7071 Schefflera bojeri 7109 Schefflera bojeri 5818 Anticipes .96 .99 Schefflera staufferana 17219 clade Schefflera staufferana 170 .98 Schefflera “antoetrensis” 7088 Schefflera “antoetrensis” 7091 Schefflera staufferana 7082 .98 Schefflera capuroniana 2328 .89 Schefflera capuroniana 11118 1 Schefflera sp. 3335 Schefflera sp. 3336 1 Schefflera sp. nov. 7169 .71 1 Schefflera sp. nov. 7171 Schefflera sp. nov. 7164 Schefflera “andohahelensis” .99 Schefflera cf. capuroniana 28 Schefflera staufferana 24 Schefflera staufferana 26 .99 Schefflera moratii 31 Schefflera moratii 34 Schefflera longipedicellata 6264 Schefflera longipedicellata 21 Schefflera longipedicellata 23 Neocussonia .99 Schefflera cf. longipedicellata 33 Schefflera vantsilana 6225 subclade Schefflera vantsilana 152 Schefflera vantsilana 32 Schefflera vantsilana 7103 1 Schefflera “vohimenensis” 2378 Schefflera “floretii” 4941 Palmate- .97 Schefflera “floretii” 7160 Schefflera “floretii” 7162 Vantsilana 1 Schefflera “floretii” 7172 clade .99 Schefflera “vohimenensis” 7156 Schefflera “floretii” 4558 Schefflera sp. 4710 1 Schefflera “rabenantoandroi” 7148 Schefflera “rabenantoandroi” 7149 1 Schefflera “floretii” 1485 Schefflera “floretii” 1487 .52 1 Schefflera longipedicellata 6220 Schefflera longipedicellata 7098 .75 Schefflera longipedicellata 7104 .99 Schefflera longipedicellata 6153 Schefflera sp. 3349 Schefflera cf. staufferana 25 .81 1 Schefflera macerosa 1499 Schefflera macerosa 1509 .99 Schefflera sp. 3347 Schefflera sp. 3352 .99 Schefflera sp. 3351 1 Schefflera fosbergiana 4322 Schefflera fosbergiana 3350 1 Schefflera cf. fosbergiana 3337 1 Schefflera rainaliana 4444 1 Schefflera rainaliana 14791 Schefflera rainaliana 7151 Schefflera frodiniana 1 Schefflera “purpuristyla” Schefflera bracteolifera Schefflera lukwangulensis 1 Schefflera umbellifera 11950 Schefflera umbellifera 5494 Schefflera myriantha 5347 .95 Schefflera myriantha 5445 Schefflera myriantha 470 1 Schefflera myriantha 5796 Schefflera myriantha 5808 .99 Schefflera myriantha 5798 .96 Schefflera myriantha 5810 Schefflera myriantha 5812 Schefflera cf. myriantha 6117 myriantha- Schefflera humblotiana monophylla 1 Schefflera monophylla 7087 1 Schefflera monophylla 7097 clade 1 1 Schefflera cf. myriantha 5816 Schefflera monophylla 7173 .67 Schefflera monophylla 1492 Schefflera monophylla 131 Schefflera myriantha 1498 1 1 Schefflera monophylla 409 Schefflera sp. 3348 Astropanax .97 Schefflera myriantha 501 African subclade Schefflera myriantha 3050 .88 Schefflera myriantha 4988 myriantha Schefflera myriantha 1885 Schefflera myriantha L365 clade Schefflera goetzenii 398 1 1 Schefflera tessmannii 2107 Schefflera barteri 5208 Goetzenii .62 Schefflera barteri 16998 1 Schefflera barteri 6782 clade Schefflera goetzenii 4807 Schefflera goetzenii 3503 1 Schefflera kivuensis 2145 1 Schefflera volkensii 5129 Schefflera volkensii 4987 1 Schefflera mannii s.n. Schefflera mannii 4406 Schefflera stolzii 1 1 Cussonia thyrsiflora 1 Cussonia paniculata Cussonia holstii Seemannaralia gerrardii .61 Astrotricha pterocarpa Outgroup Osmoxylon pectinatum Schefflera digitata

58 Figure 14: Bayesian, Schefflera halleana 4261 Schefflera cf. halleana 3339 .99 Schefflera halleana 3343 Plastid Schefflera halleana 3340 Schefflera sp. 3342 Schefflera sp. 3344 Schefflera sp. 3345 .99 Schefflera sp. 3332 Schefflera sp. 3333 Schefflera sp. 3334 Schefflera sp. 3335 Schefflera sp. 3336 .76 Schefflera umbellifera 11950 .71 Schefflera longipedicellata 6220 .88 Schefflera umbellifera 5494 .71 Schefflera vantsilana 152 Schefflera longipedicellata 6264 Schefflera “humbertii” Schefflera cf. capuroniana 28 .95 Schefflera cf. staufferana 25 Schefflera staufferana 24 Schefflera staufferana 26 Schefflera staufferana 7082 .94 Schefflera “antoetrensis” 7088 Schefflera “antoetrensis” 7091 Schefflera sp. nov. 7164 1 Schefflera rainaliana 4444 Schefflera rainaliana 14791 Schefflera rainaliana 7151 .97 Schefflera bojeri 7071 Schefflera bojeri 7109 Schefflera bojeri 5818 .95 Schefflera cf. longipedicellata 33 Schefflera vantsilana 32 .98 Schefflera moratii 31 Schefflera moratii 34 Schefflera longipedicellata 21 Schefflera longipedicellata 23 Schefflera longipedicellata 7098 Schefflera longipedicellata 7104 Neocussonia Schefflera longipedicellata 6153 Schefflera “purpuristyla” subclade Schefflera vantsilana 6225 Schefflera vantsilana 7103 Schefflera capuroniana 2328 Schefflera capuroniana 11118 Schefflera bracteolifera Schefflera “rabenantoandroi” 7148 Schefflera “rabenantoandroi” 7149 Schefflera lukwangulensis Schefflera fosbergiana 4322 1 Schefflera favargeri 384 Schefflera favargeri 476 Schefflera favargeri 5344a Schefflera favargeri 444 Schefflera “vohimenensis” 2378 Schefflera “vohimenensis” 7156 Schefflera “floretii” 4558 Schefflera sp. 4710 Schefflera “floretii” 4941 Schefflera staufferana 17219 Schefflera staufferana 170 Schefflera “floretii” 7160 Schefflera “floretii” 7162 Schefflera “floretii” 7172 Schefflera sp. nov. 7169 Schefflera sp. nov. 7171 Schefflera “andohahelensis” Schefflera frodiniana Schefflera fosbergiana 3350 Schefflera macerosa 1499 Schefflera macerosa 1509 Schefflera “floretii” 1485 Schefflera “floretii” 1487 Schefflera cf. fosbergiana 3337 Schefflera sp. 3347 Schefflera sp. 3352 Schefflera sp. 3351 Schefflera sp. 3349 .52 Schefflera myriantha 5796 Schefflera myriantha 5798 1 Schefflera myriantha 5810 .6 Schefflera myriantha 5347 Schefflera myriantha 5445 .97 Schefflera monophylla 409 Schefflera myriantha 470 myriantha- Schefflera myriantha 5808 Schefflera myriantha 5812 monophylla Schefflera sp. 3348 1 Schefflera monophylla 7087 clade Schefflera monophylla 7097 1 Schefflera cf. myriantha 5816 Schefflera monophylla 7173 Schefflera cf. myriantha 6117 1 Schefflera monophylla 1492 Schefflera myriantha 1498 .6 .69 Schefflera myriantha 1885 African 1 Schefflera myriantha L365 Astropanax .54 1 Schefflera myriantha 4988 myriantha Schefflera myriantha 3050 subclade Schefflera myriantha 501 clade .74 .83 Schefflera monophylla 131 Schefflera barteri 16998 Schefflera goetzenii 4807 .9 .99 Schefflera tessmannii Schefflera barteri 5208 1 1 Schefflera barteri 6782 Schefflera goetzenii 398 Schefflera goetzenii 3503 1 Schefflera volkensii 5129 Schefflera volkensii 4987 1 .99 Schefflera mannii s.n. Schefflera mannii 4406 Schefflera kivuensis Schefflera humblotiana Schefflera stolzii 3577 Schefflera digitat 1 1 Cussonia thyrsiflora 1 Cussonia holstii Cussonia paniculata Outgroup Seemannaralia gerrardii Astrotricha pterocarpa Osmoxylonpectinatum

59 Schefflera halleana 4261 Figure 15: Bayesian, Schefflera cf. halleana 3339 .68 Schefflera halleana 3343 Nuclear + Plastid, combined Schefflera halleana 3340 Schefflera sp. 3342 Schefflera sp. 3344 1 Schefflera sp. 3345 .94 Schefflera sp. 3332 Schefflera sp. 3334 Schefflera sp. 3333 1 .99 Schefflera favargeri 384 Schefflera favargeri 5344a .87 Schefflera favargeri 444 Schefflera favargeri 476 1 Schefflera “humbertii” Schefflera bojeri .89 1 Schefflera “antoetrensis” 7088 .99 Schefflera “antoetrensis” 7091 Anticipes Schefflera staufferana 7082 clade 1 Schefflera staufferana 17219 Schefflera staufferana 170 .99 Schefflera capuroniana 2328 .74 Schefflera capuroniana 11118 .66 1 Schefflera sp. 3335 .77 Schefflera sp. 3336 .97 Schefflera longipedicellata 6264 1 Schefflera moratii 31 Schefflera moratii 34 1 Schefflera sp. nov. 7169 .99 Schefflera sp. nov. 7171 Schefflera sp. nov. 7164 Schefflera “andohahelensis” 1 Schefflera cf. capuroniana 28 Schefflera staufferana 24 Schefflera staufferana 26 1 Schefflera cf. longipedicellata 33 Neocussonia Schefflera vantsilana 32 Schefflera longipedicellata 21 subclade 1 Schefflera longipedicellata 23 Schefflera vantsilana 6225 Schefflera vantsilana 152 Schefflera vantsilana 7103 1 Schefflera “vohimenensis” 2378 Schefflera “floretii” 4941 Palmate- .98 .85 Schefflera “floretii” 7160 Schefflera “floretii” 7172 Vantsilana 1 Schefflera “floretii” 7162 clade .99 Schefflera “vohimenensis” 7156 Schefflera “floretii” 4558 Schefflera sp. 4710 1 Schefflera “rabenantoandroi” 7148 Schefflera “rabenantoandroi” 7149 1 Schefflera “floretii” 1485 Schefflera “floretii” 1487 .84 1 Schefflera longipedicellata 6220 Schefflera longipedicellata 7098 .99 Schefflera longipedicellata 7104 1 Schefflera longipedicellata 6153 Schefflera sp. 3349 .94 Schefflera cf. staufferana 25 1 Schefflera macerosa 1499 Schefflera macerosa 1509 .78 Schefflera fosbergiana 4322 .64 Schefflera fosbergiana 3350 Schefflera sp. 3351 1 .64 Schefflera sp. 3347 1 Schefflera sp. 3352 Schefflera cf. fosbergiana 3337 1 1 Schefflera rainaliana Schefflera frodiniana 1486 1 Schefflera “purpuristyla” Schefflera bracteolifera Schefflera lukwangulensis 1 Schefflera umbellifera 11950 Schefflera umbellifera 5494 .68 Schefflera myriantha 5347 .57 Schefflera myriantha 5445 .58 Schefflera myriantha 5808 Schefflera myriantha 470 1 Schefflera myriantha 5796 .98 Schefflera myriantha 5798 .51 Schefflera myriantha 5810 Schefflera myriantha 5812 myriantha- 1 Schefflera cf. myriantha 6117 Schefflera humblotiana monophylla 1 Schefflera monophylla 7087 clade Schefflera monophylla 7097 1 .73 1 Schefflera cf. myriantha 5816 Schefflera monophylla 7173 Schefflera monophylla 1492 1 1 Schefflera monophylla 409 Schefflera sp. 3348 Schefflera myriantha 1498 Astropanax 1 Schefflera monophylla 131 subclade .99 Schefflera myriantha 1885 1 Schefflera myriantha L365 African 1 Schefflera myriantha 3050 myriantha Schefflera myriantha 4988 Schefflera myriantha 501 clade .99 .92 .61 Schefflera barteri 5208 .99 Schefflera barteri 6782 1 Schefflera tessmannii Goetzenii 1 Schefflera goetzenii 398 .51 1 Schefflera barteri 16998 clade .86 Schefflera goetzenii 4807 Schefflera goetzenii 3503 1 Schefflera kivuensis 1 Schefflera mannii s.n. 1 Schefflera mannii 4406 1 Schefflera volkensii 5129 Schefflera volkensii 4987 Schefflera stolzii Schefflera digitata 52 1 Cussonia thyrsiflora 1 Cussonia holstii .72 Cussonia paniculata Outgroup Seemannaralia gerrardii Osmoxylon pectinatum Astrotricha pterocarpa

60 B Schefflera favargeri 384 Figure 16: Geography Schefflera favargeri 5344a Schefflera favargeri 444 B Schefflera cf. halleana 3339 Schefflera sp. 3332 Africa Schefflera sp. 3334 Schefflera favargeri 476 Schefflera “humbertii” Madagascar B Schefflera halleana 4261 Schefflera halleana 3343 Schefflera halleana 3340 Anticipes Schefflera sp. 3333 Equivocal B Schefflera sp. 3342 clade Schefflera sp. 3344 Schefflera sp. 3345 B Schefflera bojeri 7071 Other Schefflera bojeri 7109 Schefflera bojeri 5818 B B Schefflera staufferana 17219 Schefflera staufferana 170 B Schefflera “antoetrensis” 7088 Schefflera “antoetrensis” 7091 Schefflera staufferana 7082 B Schefflera capuroniana 2328 Schefflera capuroniana 11118 B Schefflera sp. 3335 Schefflera sp. 3336 B Schefflera moratii 31 Schefflera moratii 34 Schefflera longipedicellata 6264 B Schefflera sp. nov. 7169 B Schefflera sp. nov. 7171 Schefflera sp. nov. 7164 Schefflera “andohahelensis” B Schefflera cf. capuroniana 28 Schefflera staufferana 24 Schefflera staufferana 26 Schefflera longipedicellata 21 Schefflera longipedicellata 23 Neocussonia B Schefflera cf. longipedicellata 33 Schefflera vantsilana 6225 subclade Schefflera vantsilana 152 Schefflera vantsilana 32 Schefflera vantsilana 7103 B Schefflera “vohimenensis” 2378 Schefflera “floretii” 4941 B Schefflera “floretii” 7160 Palmate- Schefflera “floretii” 7162 Vantsilana B Schefflera “floretii” 7172 B Schefflera “vohimenensis” 7156 clade Schefflera “floretii” 4558 Schefflera sp. 4710 B Schefflera “rabenantoandroi” 7148 Schefflera “rabenantoandroi” 7149 B Schefflera “floretii” 1485 Schefflera “floretii” 1487 B Schefflera longipedicellata 6220 Schefflera longipedicellata 7098 Schefflera longipedicellata 7104 AB Schefflera longipedicellata 6153 Schefflera cf. staufferana 25 Schefflera sp. 3349 Schefflera macerosa 1499 Schefflera macerosa 1509 B Schefflera sp. 3347 Schefflera sp. 3352 B Schefflera sp. 3351 B Schefflera fosbergiana 4322 A Schefflera fosbergiana 3350 Schefflera cf. fosbergiana 3337 B Schefflera rainaliana 4444 B Schefflera rainaliana 14791 Schefflera rainaliana 7151 Schefflera frodiniana B Schefflera “purpuristyla” Schefflera bracteolifera Schefflera lukwangulensis Schefflera umbellifera 11950 Schefflera umbellifera 5494 Schefflera myriantha 5347 B Schefflera myriantha 5445 Schefflera myriantha 470 B Schefflera myriantha 5796 Schefflera myriantha 5808 B Schefflera myriantha 5798 B Schefflera myriantha 5810 Schefflera myriantha 5812 myriantha- Schefflera cf. myriantha 6117 A Schefflera humblotiana monophylla B Schefflera monophylla 7087 Schefflera monophylla 7097 clade B Schefflera cf. myriantha 5816 AB Schefflera monophylla 7173 Schefflera monophylla 1492 B Schefflera monophylla 409 Schefflera sp. 3348 A Schefflera monophylla 131 Schefflera myriantha 1498 Astropanax A Schefflera myriantha 501 subclade Schefflera myriantha 3050 African A Schefflera myriantha 4988 myriantha Schefflera myriantha 1885 Schefflera myriantha L365 clade Schefflera goetzenii 398 A Schefflera tessmannii Schefflera barteri 5208 Goetzenii A Schefflera barteri 16998 Schefflera barteri 6782 clade Schefflera goetzenii 4807 A Schefflera goetzenii 3503 A Schefflera volkensii 5129 Schefflera volkensii 4987 A Schefflera mannii s.n. Schefflera mannii 4406 Schefflera kivuensis Schefflera stolzii A A Cussonia thyrsiflora A Cussonia paniculata Cussonia holstii Seemannaralia gerrardii Outgroup Astrotricha pterocarpa Osmoxylon pectinatum Schefflera digitata

61 Schefflera favargeri 384 Figure 17: Carpel Number Schefflera favargeri 5344a Schefflera favargeri 444 Schefflera cf. halleana 3339 Schefflera sp. 3332 2–3 Carpellate Schefflera sp. 3334 Schefflera favargeri 476 Schefflera “humbertii” 4–5 Carpellate Schefflera halleana 4261 Schefflera halleana 3343 Schefflera halleana 3340 Anticipes Schefflera sp. 3333 6 + Carpellate Schefflera sp. 3342 clade Schefflera sp. 3344 Schefflera sp. 3345 Schefflera bojeri 7071 Missing Data Schefflera bojeri 7109 Schefflera bojeri 5818 Schefflera staufferana 17219 Schefflera staufferana 170 Schefflera “antoetrensis” 7088 Schefflera “antoetrensis” 7091 Schefflera staufferana 7082 Schefflera capuroniana 2328 Schefflera capuroniana 11118 Schefflera sp. 3335 Schefflera sp. 3336 Schefflera moratii 31 Schefflera moratii 34 Schefflera longipedicellata 6264 Schefflera sp. nov. 7169 Schefflera sp. nov. 7171 Schefflera sp. nov. 7164 Schefflera “andohahelensis” Schefflera cf. capuroniana 28 Schefflera staufferana 24 Schefflera staufferana 26 Schefflera longipedicellata 21 Schefflera longipedicellata 23 Schefflera cf. longipedicellata 33 Neocussonia Schefflera vantsilana 6225 subclade Schefflera vantsilana 152 Schefflera vantsilana 32 Schefflera vantsilana 7103 Schefflera “vohimenensis” 2378 Schefflera “floretii” 4941 Schefflera “floretii” 7160 Palmate- Schefflera “floretii” 7162 Vantsilana Schefflera “floretii” 7172 Schefflera “vohimenensis” 7156 clade Schefflera “floretii” 4558 Schefflera sp. 4710 Schefflera “rabenantoandroi” 7148 Schefflera “rabenantoandroi” 7149 Schefflera “floretii” 1485 Schefflera “floretii” 1487 Schefflera longipedicellata 6220 Schefflera longipedicellata 7098 Schefflera longipedicellata 7104 Schefflera longipedicellata 6153 Schefflera cf. staufferana 25 Schefflera sp. 3349 Schefflera macerosa 1499 Schefflera macerosa 1509 Schefflera sp. 3347 Schefflera sp. 3352 Schefflera sp. 3351 Schefflera fosbergiana 4322 Schefflera fosbergiana 3350 Schefflera cf. fosbergiana 3337 Schefflera rainaliana 4444 Schefflera rainaliana 14791 Schefflera rainaliana 7151 Schefflera frodiniana Schefflera “purpuristyla” Schefflera bracteolifera Schefflera lukwangulensis Schefflera umbellifera 11950 Schefflera umbellifera 5494 Schefflera myriantha 5347 Schefflera myriantha 5445 Schefflera myriantha 470 Schefflera myriantha 5796 Schefflera myriantha 5808 Schefflera myriantha 5798 Schefflera myriantha 5810 Schefflera myriantha 5812 myriantha- Schefflera cf. myriantha 6117 Schefflera humblotiana monophylla Schefflera monophylla 7087 Schefflera monophylla 7097 clade Schefflera cf. myriantha 5816 Schefflera monophylla 7173 Schefflera monophylla 1492 Schefflera monophylla 409 Schefflera sp. 3348 Schefflera monophylla 131 Schefflera myriantha 1498 Astropanax Schefflera myriantha 501 subclade Schefflera myriantha 3050 African Schefflera myriantha 4988 myriantha Schefflera myriantha 1885 Schefflera myriantha L365 clade Schefflera goetzenii 398 Schefflera tessmannii Schefflera barteri 5208 Goetzenii Schefflera barteri 16998 Schefflera barteri 6782 clade Schefflera goetzenii 4807 Schefflera goetzenii 3503 Schefflera volkensii 5129 Schefflera volkensii 4987 Schefflera mannii s.n. Schefflera mannii 4406 Schefflera kivuensis Schefflera stolzii Cussonia thyrsiflora Cussonia paniculata Cussonia holstii Seemannaralia gerrardii Outgroup Astrotricha pterocarpa Osmoxylon pectinatum Schefflera digitata

62 Schefflera favargeri 384 Figure 18: Leaf composition Schefflera favargeri 5344a Schefflera favargeri 444 Schefflera cf. halleana 3339 Schefflera sp. 3332 Palmately compound, 4+ leaflets Schefflera sp. 3334 Schefflera favargeri 476 Schefflera “humbertii” Palmately compound, 1-3 leaflets Schefflera halleana 4261 Schefflera halleana 3343 Schefflera halleana 3340 Anticipes Schefflera sp. 3333 Schefflera sp. 3342 clade Exclusively Unifoliolate Schefflera sp. 3344 Schefflera sp. 3345 Schefflera bojeri 7071 Missing Data Schefflera bojeri 7109 Schefflera bojeri 5818 Schefflera staufferana 17219 Schefflera staufferana 170 Schefflera “antoetrensis” 7088 Schefflera “antoetrensis” 7091 Schefflera staufferana 7082 Schefflera capuroniana 2328 Schefflera capuroniana 11118 Schefflera sp. 3335 Schefflera sp. 3336 Schefflera moratii 31 Schefflera moratii 34 Schefflera longipedicellata 6264 Schefflera sp. nov. 7169 Schefflera sp. nov. 7171 Schefflera sp. nov. 7164 Schefflera “andohahelensis” Schefflera cf. capuroniana 28 Schefflera staufferana 24 Schefflera staufferana 26 Schefflera longipedicellata 21 Schefflera longipedicellata 23 Schefflera cf. longipedicellata 33 Neocussonia Schefflera vantsilana 6225 subclade Schefflera vantsilana 152 Schefflera vantsilana 32 Schefflera vantsilana 7103 Schefflera “vohimenensis” 2378 Schefflera “floretii” 4941 Schefflera “floretii” 7160 Palmate- Schefflera “floretii” 7162 Vantsilana Schefflera “floretii” 7172 Schefflera “vohimenensis” 7156 clade Schefflera “floretii” 4558 Schefflera sp. 4710 Schefflera “rabenantoandroi” 7148 Schefflera “rabenantoandroi” 7149 Schefflera “floretii” 1485 Schefflera “floretii” 1487 Schefflera longipedicellata 6220 Schefflera longipedicellata 7098 Schefflera longipedicellata 7104 Schefflera longipedicellata 6153 Schefflera cf. staufferana 25 Schefflera sp. 3349 Schefflera macerosa 1499 Schefflera macerosa 1509 Schefflera sp. 3347 Schefflera sp. 3352 Schefflera sp. 3351 Schefflera fosbergiana 4322 Schefflera fosbergiana 3350 Schefflera cf. fosbergiana 3337 Schefflera rainaliana 4444 Schefflera rainaliana 14791 Schefflera rainaliana 7151 Schefflera frodiniana Schefflera “purpuristyla” Schefflera bracteolifera Schefflera lukwangulensis Schefflera umbellifera 11950 Schefflera umbellifera 5494 Schefflera myriantha 5347 Schefflera myriantha 5445 Schefflera myriantha 470 Schefflera myriantha 5796 Schefflera myriantha 5808 Schefflera myriantha 5798 Schefflera myriantha 5810 Schefflera myriantha 5812 myriantha- Schefflera cf. myriantha 6117 Schefflera humblotiana monophylla Schefflera monophylla 7087 Schefflera monophylla 7097 clade Schefflera cf. myriantha 5816 Schefflera monophylla 7173 Schefflera monophylla 1492 Schefflera monophylla 409 Schefflera sp. 3348 Schefflera monophylla 131 Schefflera myriantha 1498 Astropanax Schefflera myriantha 501 subclade Schefflera myriantha 3050 African Schefflera myriantha 4988 myriantha Schefflera myriantha 1885 Schefflera myriantha L365 clade Schefflera goetzenii 398 Schefflera tessmannii Schefflera barteri 5208 Goetzenii Schefflera barteri 16998 Schefflera barteri 6782 clade Schefflera goetzenii 4807 Schefflera goetzenii 3503 Schefflera volkensii 5129 Schefflera volkensii 4987 Schefflera mannii s.n. Schefflera mannii 4406 Schefflera kivuensis Schefflera stolzii Cussonia thyrsiflora Cussonia paniculata Cussonia holstii Seemannaralia gerrardii Outgroup Astrotricha pterocarpa Osmoxylon pectinatum Schefflera digitata

63 Schefflera favargeri 384 Figure 19: Inflorescence Arrangement Schefflera favargeri 5344a Schefflera favargeri 444 Schefflera cf. halleana 3339 Schefflera sp. 3332 Umbellate Schefflera sp. 3334 Schefflera favargeri 476 Schefflera “humbertii” Racemose Schefflera halleana 4261 Schefflera halleana 3343 Schefflera halleana 3340 Anticipes Schefflera sp. 3333 Missing Data Schefflera sp. 3342 clade Schefflera sp. 3344 Schefflera sp. 3345 Schefflera bojeri 7071 Schefflera bojeri 7109 Schefflera bojeri 5818 Schefflera staufferana 17219 Schefflera staufferana 170 Schefflera “antoetrensis” 7088 Schefflera “antoetrensis” 7091 Schefflera staufferana 7082 Schefflera capuroniana 2328 Schefflera capuroniana 11118 Schefflera sp. 3335 Schefflera sp. 3336 Schefflera moratii 31 Schefflera moratii 34 Schefflera longipedicellata 6264 Schefflera sp. nov. 7169 Schefflera sp. nov. 7171 Schefflera sp. nov. 7164 Schefflera “andohahelensis” Schefflera cf. capuroniana 28 Schefflera staufferana 24 Schefflera staufferana 26 Schefflera longipedicellata 21 Schefflera longipedicellata 23 Schefflera cf. longipedicellata 33 Neocussonia Schefflera vantsilana 6225 subclade Schefflera vantsilana 152 Schefflera vantsilana 32 Schefflera vantsilana 7103 Schefflera “vohimenensis” 2378 Schefflera “floretii” 4941 Schefflera “floretii” 7160 Palmate- Schefflera “floretii” 7162 Vantsilana Schefflera “floretii” 7172 Schefflera “vohimenensis” 7156 clade Schefflera “floretii” 4558 Schefflera sp. 4710 Schefflera “rabenantoandroi” 7148 Schefflera “rabenantoandroi” 7149 Schefflera “floretii” 1485 Schefflera “floretii” 1487 Schefflera longipedicellata 6220 Schefflera longipedicellata 7098 Schefflera longipedicellata 7104 Schefflera longipedicellata 6153 Schefflera cf. staufferana 25 Schefflera sp. 3349 Schefflera macerosa 1499 Schefflera macerosa 1509 Schefflera sp. 3347 Schefflera sp. 3352 Schefflera sp. 3351 Schefflera fosbergiana 4322 Schefflera fosbergiana 3350 Schefflera cf. fosbergiana 3337 Schefflera rainaliana 4444 Schefflera rainaliana 14791 Schefflera rainaliana 7151 Schefflera frodiniana Schefflera “purpuristyla” Schefflera bracteolifera Schefflera lukwangulensis Schefflera umbellifera 11950 Schefflera umbellifera 5494 Schefflera myriantha 5347 Schefflera myriantha 5445 Schefflera myriantha 470 Schefflera myriantha 5796 Schefflera myriantha 5808 Schefflera myriantha 5798 Schefflera myriantha 5810 Schefflera myriantha 5812 myriantha- Schefflera cf. myriantha 6117 Schefflera humblotiana monophylla Schefflera monophylla 7087 Schefflera monophylla 7097 clade Schefflera cf. myriantha 5816 Schefflera monophylla 7173 Schefflera monophylla 1492 Schefflera monophylla 409 Schefflera sp. 3348 Schefflera monophylla 131 Schefflera myriantha 1498 Astropanax Schefflera myriantha 501 subclade Schefflera myriantha 3050 African Schefflera myriantha 4988 myriantha Schefflera myriantha 1885 Schefflera myriantha L365 clade Schefflera goetzenii 398 Schefflera tessmannii Schefflera barteri 5208 Goetzenii Schefflera barteri 16998 Schefflera barteri 6782 clade Schefflera goetzenii 4807 Schefflera goetzenii 3503 Schefflera volkensii 5129 Schefflera volkensii 4987 Schefflera mannii s.n. Schefflera mannii 4406 Schefflera kivuensis Schefflera stolzii Cussonia thyrsiflora Cussonia paniculata Cussonia holstii Seemannaralia gerrardii Outgroup Astrotricha pterocarpa Osmoxylon pectinatum Schefflera digitata

64 Schefflera favargeri 384 Figure 20: Pedicel length Schefflera favargeri 5344a Schefflera favargeri 444 Schefflera cf. halleana 3339 Schefflera sp. 3332 > 5 mm Schefflera sp. 3334 Schefflera favargeri 476 Schefflera “humbertii” 0–5 mm Schefflera halleana 4261 Schefflera halleana 3343 Schefflera halleana 3340 Anticipes Schefflera sp. 3333 Absent Schefflera sp. 3342 clade Schefflera sp. 3344 Schefflera sp. 3345 Schefflera bojeri 7071 Missing Data Schefflera bojeri 7109 Schefflera bojeri 5818 Schefflera staufferana 17219 Schefflera staufferana 170 Schefflera “antoetrensis” 7088 Schefflera “antoetrensis” 7091 Schefflera staufferana 7082 Schefflera capuroniana 2328 Schefflera capuroniana 11118 Schefflera sp. 3335 Schefflera sp. 3336 Schefflera moratii 31 Schefflera moratii 34 Schefflera longipedicellata 6264 Schefflera sp. nov. 7169 Schefflera sp. nov. 7171 Schefflera sp. nov. 7164 Schefflera “andohahelensis” Schefflera cf. capuroniana 28 Schefflera staufferana 24 Schefflera staufferana 26 Schefflera longipedicellata 21 Schefflera longipedicellata 23 Schefflera cf. longipedicellata 33 Neocussonia Schefflera vantsilana 6225 subclade Schefflera vantsilana 152 Schefflera vantsilana 32 Schefflera vantsilana 7103 Schefflera “vohimenensis” 2378 Schefflera “floretii” 4941 Schefflera “floretii” 7160 Palmate- Schefflera “floretii” 7162 Vantsilana Schefflera “floretii” 7172 Schefflera “vohimenensis” 7156 clade Schefflera “floretii” 4558 Schefflera sp. 4710 Schefflera “rabenantoandroi” 7148 Schefflera “rabenantoandroi” 7149 Schefflera “floretii” 1485 Schefflera “floretii” 1487 Schefflera longipedicellata 6220 Schefflera longipedicellata 7098 Schefflera longipedicellata 7104 Schefflera longipedicellata 6153 Schefflera cf. staufferana 25 Schefflera sp. 3349 Schefflera macerosa 1499 Schefflera macerosa 1509 Schefflera sp. 3347 Schefflera sp. 3352 Schefflera sp. 3351 Schefflera fosbergiana 4322 Schefflera fosbergiana 3350 Schefflera cf. fosbergiana 3337 Schefflera rainaliana 4444 Schefflera rainaliana 14791 Schefflera rainaliana 7151 Schefflera frodiniana Schefflera “purpuristyla” Schefflera bracteolifera Schefflera lukwangulensis Schefflera umbellifera 11950 Schefflera umbellifera 5494 Schefflera myriantha 5347 Schefflera myriantha 5445 Schefflera myriantha 470 Schefflera myriantha 5796 Schefflera myriantha 5808 Schefflera myriantha 5798 Schefflera myriantha 5810 Schefflera myriantha 5812 myriantha- Schefflera cf. myriantha 6117 Schefflera humblotiana monophylla Schefflera monophylla 7087 Schefflera monophylla 7097 clade Schefflera cf. myriantha 5816 Schefflera monophylla 7173 Schefflera monophylla 1492 Schefflera monophylla 409 Schefflera sp. 3348 Schefflera monophylla 131 Schefflera myriantha 1498 Astropanax Schefflera myriantha 501 subclade Schefflera myriantha 3050 African Schefflera myriantha 4988 myriantha Schefflera myriantha 1885 Schefflera myriantha L365 clade Schefflera goetzenii 398 Schefflera tessmannii Schefflera barteri 5208 Goetzenii Schefflera barteri 16998 Schefflera barteri 6782 clade Schefflera goetzenii 4807 Schefflera goetzenii 3503 Schefflera volkensii 5129 Schefflera volkensii 4987 Schefflera mannii s.n. Schefflera mannii 4406 Schefflera kivuensis Schefflera stolzii Cussonia thyrsiflora Cussonia paniculata Cussonia holstii Seemannaralia gerrardii Outgroup Astrotricha pterocarpa Osmoxylon pectinatum Schefflera digitata

65 Schefflera favargeri 384 Figure 21: Bract morphology Schefflera favargeri 5344a Schefflera favargeri 444 Schefflera cf. halleana 3339 Schefflera sp. 3332 Bracts caducous Schefflera sp. 3334 Schefflera favargeri 476 Schefflera “humbertii” Bracts persistent Schefflera halleana 4261 Schefflera halleana 3343 Schefflera halleana 3340 Anticipes Schefflera sp. 3333 Data Missing Schefflera sp. 3342 clade Schefflera sp. 3344 Schefflera sp. 3345 Schefflera bojeri 7071 Schefflera bojeri 7109 Schefflera bojeri 5818 Schefflera staufferana 17219 Schefflera staufferana 170 Schefflera “antoetrensis” 7088 Schefflera “antoetrensis” 7091 Schefflera staufferana 7082 Schefflera capuroniana 2328 Schefflera capuroniana 11118 Schefflera sp. 3335 Schefflera sp. 3336 Schefflera moratii 31 Schefflera moratii 34 Schefflera longipedicellata 6264 Schefflera sp. nov. 7169 Schefflera sp. nov. 7171 Schefflera sp. nov. 7164 Schefflera “andohahelensis” Schefflera cf. capuroniana 28 Schefflera staufferana 24 Schefflera staufferana 26 Schefflera longipedicellata 21 Schefflera longipedicellata 23 Schefflera cf. longipedicellata 33 Neocussonia Schefflera vantsilana 6225 subclade Schefflera vantsilana 152 Schefflera vantsilana 32 Schefflera vantsilana 7103 Schefflera “vohimenensis” 2378 Schefflera “floretii” 4941 Schefflera “floretii” 7160 Palmate- Schefflera “floretii” 7162 Vantsilana Schefflera “floretii” 7172 Schefflera “vohimenensis” 7156 clade Schefflera “floretii” 4558 Schefflera sp. 4710 Schefflera “rabenantoandroi” 7148 Schefflera “rabenantoandroi” 7149 Schefflera “floretii” 1485 Schefflera “floretii” 1487 Schefflera longipedicellata 6220 Schefflera longipedicellata 7098 Schefflera longipedicellata 7104 Schefflera longipedicellata 6153 Schefflera cf. staufferana 25 Schefflera sp. 3349 Schefflera macerosa 1499 Schefflera macerosa 1509 Schefflera sp. 3347 Schefflera sp. 3352 Schefflera sp. 3351 Schefflera fosbergiana 4322 Schefflera fosbergiana 3350 Schefflera cf. fosbergiana 3337 Schefflera rainaliana 4444 Schefflera rainaliana 14791 Schefflera rainaliana 7151 Schefflera frodiniana Schefflera “purpuristyla” Schefflera bracteolifera Schefflera lukwangulensis Schefflera umbellifera 11950 Schefflera umbellifera 5494 Schefflera myriantha 5347 Schefflera myriantha 5445 Schefflera myriantha 470 Schefflera myriantha 5796 Schefflera myriantha 5808 Schefflera myriantha 5798 Schefflera myriantha 5810 Schefflera myriantha 5812 myriantha- Schefflera cf. myriantha 6117 Schefflera humblotiana monophylla Schefflera monophylla 7087 Schefflera monophylla 7097 clade Schefflera cf. myriantha 5816 Schefflera monophylla 7173 Schefflera monophylla 1492 Schefflera monophylla 409 Schefflera sp. 3348 Schefflera monophylla 131 Schefflera myriantha 1498 Astropanax Schefflera myriantha 501 subclade Schefflera myriantha 3050 African Schefflera myriantha 4988 myriantha Schefflera myriantha 1885 Schefflera myriantha L365 clade Schefflera goetzenii 398 Schefflera tessmannii Schefflera barteri 5208 Goetzenii Schefflera barteri 16998 Schefflera barteri 6782 clade Schefflera goetzenii 4807 Schefflera goetzenii 3503 Schefflera volkensii 5129 Schefflera volkensii 4987 Schefflera mannii s.n. Schefflera mannii 4406 Schefflera kivuensis Schefflera stolzii Cussonia thyrsiflora Cussonia paniculata Cussonia holstii Seemannaralia gerrardii Outgroup Astrotricha pterocarpa Osmoxylon pectinatum Schefflera digitata

Vita

Morgan Robert Gostel was born on January 30, 1985 in Henrico County, Virginia. He received his High School Diploma in 2003 from Douglas Southall Freeman High School in

Henrico Country, Virginia. Morgan attended Virginia Commonwealth University as an undergraduate student and received his Bachelor of Science in Biology in 2008.