TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Dividing and conquering the fastest-growing genus: Towards a natural sectional classification of the mega-diverse genus Begonia (Begoniaceae) Peter W. Moonlight,1,2 Wisnu H. Ardi,3 Luzmila Arroyo Padilla,4 Kuo-Fang Chung,5 Daniel Fuller,6 Deden Girmansyah,7 Ruth Hollands,1 Adolfo Jara-Muñoz,8 Ruth Kiew,9 Wai-Chao Leong,10 Yan Liu,11 Adi Mahardika,1 Lakmini D.K. Marasinghe,12 Meriel O’Connor,1 Ching-I Peng,5 Álvaro J. Pérez,13 Thamarat Phutthai,14 Martin Pullan,1 Sangeeta Rajbhandary,15 Carlos Reynel,16 Rosario R. Rubite,17 Julia Sang,18 David Scherberich,19 Yu-Min Shui,20 Mark C. Tebbitt,21 Daniel C. Thomas,22,23 Hannah P. Wilson,1,2 Nura H. Zaini24 & Mark Hughes1 1 Royal Botanic Garden Edinburgh, 20a Inverleith Row, Edinburgh EH3 5LR, U.K. 2 Institute of Biodiversity, Animal Health and Comparative Medicine, College of Medical, Veterinary and Life Sciences, University of Glasgow, Glasgow, G12 8QQ, U.K. 3 Bogor Botanic Gardens, Jl. Ir. Juanda No. 13, Bogor, Indonesia 4 Facultad de Ciencias Agrícolas, Universidad Autónoma Gabriel Rene Moreno, Km 9 al norte, El Vallecitio, Santa Cruz, Bolivia 5 Research Museum and Herbarium (HAST), Biodiversity Research Center, Academia Sinica, Taipei 11529, Taiwan 6 Department of Natural Sciences, University of Wisconsin – Superior, Superior, Wisconsin 54880, U.S.A. 7 Herbarium Bogoriense, Research Center for Biology, Indonesian Institute of Sciences, Jl. Raya Jakarta-Bogor Km 46, Cibinong, 16911, Bogor, West Java, Indonesia 8 Jardín Botánico de Bogotá “José Celestino Mutis”, Bogotá D.C., Colombia 9 Forest Research Institute Malaysia, Kepong, Selagnor, Malaysia 10 School of Forestry and Resource Conservation, National Taiwan University, No. 1, Sec. 4, Roosevelt Road, Taipei, 10617, Taiwan 11 Guangxi Institute of Botany, Guangxi Zhuangzu Autonomous Region and the Chinese Academy of Sciences, Guilin 541006, China 12 Department of Botany, Faculty of Natural Sciences, Open University of Sri Lanka, Nawala, Nugegoda, Sri Lanka 13 Herbario QCA, Escuela de Ciencas Biológicas, Pontifica Universidad Católica del Ecuador, Apartado 17-01-2184, Quito, Ecuador 14 Faculty of Environment and Resource studies, Mahidol University (Salaya campus), Nakhon Pathom 73170, Thailand 15 Central Department of Botany, Tribhuvan University, Kirtipur, Kathmandu, Nepal 16 Facultad de Ciencias Forestales, Universidad Nacional Agraria-La Molina, Avenida La Universidad, Apartado 456, Lima 12, Perú 17 University of the Philippines Manila, Department of Biology, College of Arts and Sciences, Padre Faura, Manila, Philippines 18 Botanical Research Centre, Sarawak Forestry, Kuching, Sarawak, Malaysia 19 Jardin Botanique de Lyon, Mairie de Lyon, 69205 Lyon cedex 01, France 20 Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, Yunnan, China 21 Department of Biological and Environmental Sciences, California University of Pennsylvania, California, Pennsylvania 15419-1394, U.S.A. 22 Singapore Botanic Gardens, Singapore 259569, Singapore 23 University of Hong Kong, Hong Kong, China 24 Leiden University, 2311 EZ Leiden, Netherlands Author for correspondence: Peter W. Moonlight, p.moonlight@rbge.org.uk DOI https://doi.org/10.12705/672.3 Abstract The pantropical genus Begonia is the sixth-largest genus of flowering plants, including 1870 species. The sections of Begonia are used frequently as analogues to genera in other families but, despite their taxonomic utility, few of the current sections have been examined in the light of molecular phylogenetic analyses. We present herein the largest, most representative phylogeny of Begonia published to date and a subsequent provisional sectional classification of the genus. We utilised three plastid markers for 574 species and 809 accessions of Begonia and used Hillebrandia as an outgroup to produce a dated phylogeny. The relationships between some species and sections are poorly resolved, but many sections and deeper nodes receive strong support. We recognise 70 sections of Begonia including 5 new sections: Astrothrix, Ephemera, Jackia, Kollmannia, and Stellandrae; 4 sections are reinstated from synonymy: Australes, Exalabegonia, Latistigma and Pereira; and 5 sections are newly synonymised. The new sectional classification is discussed with reference to identifying characters and previous classifications. Keywords Begonia; dating; phylogenetics; sectional classification Supplementary Material The Electronic Supplement (Table S1) and DNA sequence alignment are available from https://doi. org/10.12705/672.3.S1 and https://doi.org/10.12705/672.3.S2, respectively; an Excel table showing species names, voucher information, GenBank accession numbers and alignment name for each terminal is available from https://doi.org/10.12705/672.3.S3 Article history: Received: 20 Jun 2017 | returned for (first) revision: 21 Sep 2017 | (last) revision received: 18 Jan 2018 | accepted: 25 Jan 2018 | published: online fast track, n/a; in print and online issues, 9 May 2018 || Associate Editor: Javier Francisco-Ortega || © International Association for Plant Taxonomy (IAPT) 2018, all rights reserved Version of Record 267 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) INTRODUCTION Big genera remain the largest black holes in plant taxonomy (Frodin, 2004), which has a detrimental effect in related fields including evolutionary biology, biogeography, and conservation. Around 57 angiosperm genera have more than 200 species with the ten largest each containing more than 1250 species (Frodin, 2004). In the decade to 2015, 2064 species were published in these genera (Fig. 1), accounting for ca. 10% of species described in this timeframe (Bebber & al., 2010). The size of mega-diverse genera makes it unfeasible for a single taxonomist to monograph them within a career. Monographic work has instead tended to focus upon smaller plant groups (Wood & al., 2015), while taxonomists working on very large genera have largely focused on geographic or infrageneric groups (Atkins & al., 2013). The problems associated with geography-focussed taxonomy are well documented (Stace, 1991) thus we recommend putting future efforts into delimiting natural infrageneric groups through collaboration as advocated by Van Welzen & al. (2009). We suggest this will stimulate monographic work within large genera and help plug the taxonomic black hole. Begonia L. is one of the ten largest genera of plants (Frodin, 2004) and species number has increased faster than in any other genus from 2005 to 2015 (Fig. 1). The 341 new species published within this timeframe were not the result of a single work or group, with species published by 73 authors in 119 journal articles covering species from 29 countries across three continents. In 2015 alone, 105 Begonia species were published, more than in any other year since 1859 when Alphonse de Candolle published a treatment of the Begoniaceae C.Agardh. (Candolle, 1859). The number of accepted species of Begonia currently stands at 1870 species (Hughes & al., 2015–) and is likely to rise to well over 2000. The number of undescribed species from Borneo alone is estimated at over 300 (Julia & Kiew, 2014) and large parts of the taxonomic and geographic range of Begonia have not been covered in monographs or Floras since the 1850s (e.g., Brazil, India, B. sect. Knesebeckia (Klotzsch) A.DC., B. sect. Begonia). A stable, natural infra-generic classification of Begonia is required to aid systematic work within the genus. The author who progressed furthest towards a generic classification within the family was Klotzsch, who described 39 genera in two publications (Klotzsch, 1854, 1855). The majority of genera were relegated to sections of Begonia by Candolle (1859, 1864). In the context of a group whose species number was growing rapidly, in which the genera were poorly defined, and in which many of the species remained poorly known, Candolle concluded assigning new species to genera would remain problematic and result in considerable nomenclatural instability (Candolle, 1864). Candolle maintained only Casparya Klotzsch and Mezierea Gaudich. at the generic level. These genera were distinguished from Begonia by their apical fruit dehiscence and parietal placentation respectively, but these were later reduced to sections by Bentham and Hooker, who considered these characters weak (Bentham & Hooker, 1867). Symbegonia Warb. (Warburg, 1894) and Semibegoniella C.DC. (Candolle, 1908) were described to accommodate species with tepals fused into a lobed corolla tube. Semibegoniella was reduced to a section by Barkley & Baranov (1972) as species intermediate between the type and members of B. sect. Casparya had Fig. 1. Accumulation of species in the ten largest plant genera from 2005 to 2015. Data from the International Plant Names Index (IPNI, 2016). 268 TAXON 67 (2) • April 2018: 267–323 Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) by then been discovered and Forrest & Hollingsworth (2003) reduced Symbegonia to a section as molecular data showed it was highly nested within Begonia. Further sampling of both taxa and DNA regions (Forrest & al., 2005; Thomas & al., 2011; Moonlight & al., 2015) has shown that none of the previously recognised genera form a sister relationship with Begonia, with the exception of Hillebrandia Oliv. Subsequently, only Begonia and Hillebrandia are currently recognised at the generic level, with Hillebrandia being both phylogenetically and morphologically distinct from Begonia (Clement & al., 2004). Barkley & Baranov (1972) and Doorenbos & al. (1998) later classified the majority of known species of Begonia into sections and it is the latter work that is primarily followed today. Doorenbos & al. recognised 63 sections in 1998 and a further five have since been described (De Wilde & Plana, 2003; Forrest & Hollingsworth, 2003; Gu & al., 2007; Hughes & Takeuchi, 2015; Shui & al., 2002). All sections are restricted to a single continent with the notable exceptions of B. sect. Tetraphila A.DC. (De Wilde & al., 2011), which is present in both Africa and Asia, and B. sect. Doratometra (Klotzsch) A.DC., which is present in both South America and Asia (Radbouchoom & al., 2017), although the Asian range may be the result of an introduction in the latter case (Radbouchoom & al., 2017). The replacement of the generic classification of the Begoniaceae with one which is largely based on sections has downsides as well as benefits. It means the species binomials do not convey as much information about species relationships; a further disadvantange is that stating sectional placement is not a nomenclatural requirement, and hence species can be published “stateless” within the genus. However, a significant advantage of a sectional classification over a generic classification is that taxonomic instability does not produce nomenclatural instability. Species can be moved among sections with no impact upon their generic name or authority string (McNeill & al., 2012), a situation which is very welcome at present given that some of the sections are in a state of flux. The current morphological classification of Doorenbos & al. (1998) has served us well up to this point, but as our knowledge of Begonia phylogeny has improved and species become better known due to a resurgence of taxonomic activity in the genus, some improvements can be made. Doorenbos & al. were unable to assign ca. 50 poorly known species to section, only tentatively assigned a further ca. 80 species to a section, and suggested new sections would be required to encompass another 19 species. In addition a number of more recently published species have not been assigned to section by their authors (e.g., B. caparaoensis E.L.Jacques & L.Kollmann) or have been assigned to sections that do not follow the circumscriptions of Doorenbos & al. (1998) (e.g., B. acutitepala K.Y.Guan & D.K.Tian). Furthermore, the majority of the classification system remains untested by molecular phylogenetic methods and published phylogenies suggest a number of sections are paraphyletic (e.g., B. sect. Begonia [Moonlight & al., 2015]; B. sect. Platycentrum (Klotzsch) A.DC. [Thomas & al., 2011]) or polyphyletic (e.g., B. sect. Diploclinium (Lindl.) A.DC. [Thomas & al., 2011]; B. sect. Knesebeckia [Moonlight & al., 2015]). A phylogenetic update to the sectional classification of Begonia is required to work towards a natural sectional classification of Begonia. By natural classification, we mean one which is based on phylogeny, and also informed by morphology. We aim to delimit monophyletic sections where possible, and indeed this has proved feasible for most of the genus. There has been much debate about the acceptance of paraphyletic groups in classifications, and whether one should even attempt classifications based on monophyly. Paraphyletic groups are seen as either “bad taxonomy” (Ebach & al., 2006) or a fundamental consequence of evolution which need to be reflected in a practical taxonomy (Brummit, 2006). We are not fundamentally opposed to paraphyletic groups and have taken a pragmatic approach to sectional delimitation, based on the evidence at hand in terms of phylogeny, morphology and biogeography. We have left some sections as paraphyletic where this has either retained some utility for identification, or where it has not been possible to manufacture a taxonomy based on monophyly. We have based this updated classification on a chloroplast DNA phylogeny, for two reasons. Firstly, the taxon sampling available for the chloroplast DNA regions we have used is superior to that for nuclear DNA (specifically, ITS). The current sampling for ITS is heavily biased towards a small number of sections, and is particularly lacking in sampling in the Neotropics. Secondly, it is not possible to align ITS sequences with certainty across the genus as many base positions in the spacers are saturated with substitutions. Hybridisation and ensuing phylogenetic incongruence has been found in Begonia, but this is overwhelmingly limited to gene exchange between species within the same section (Hughes & al., 2015c). The results we present here are congruent with the only genus-wide phylogenetic study based on ITS and morphology (Forrest & al., 2005), but that lacks sufficient resolution to make a useful comparison. We consider the chloroplast DNA phylogeny presented here to be the best estimate of Begonia evolutionary history to date. While it is not the principle goal of our analysis, we recognise that a dated phylogeny will be of significant utility to authors studying the timing of evolutionary and biogeographic events in either Begonia or across multiple groups. The most comprehensive dated phylogeny of the Begoniaceae to date included 268 taxa (Moonlight & al., 2015); this study doubles this number and broadens the phylogenetic sampling considerably. While the divergence time estimation methods used in Moonlight & al. (2015) are relatively robust to incomplete sampling (Drummond & al., 2012), a sampling increase of this magnitude may reduce the variance on the inferred ages of some nodes. This study aims to: (1) Produce a well-sampled, well-resolved and dated phylogeny of the Begoniaceae; (2) Identify well-supported clades and assess how they relate to the sections of Begonia; (3) Recircumscribe natural sections where our sampling allows and identify sections or clades requiring further taxonomic and phylogenetic work to achieve a natural sectional classification of Begonia. Version of Record 269 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) TAXON 67 (2) • April 2018: 267–323 Fig. 2. Maximum-clade-credibility chronogram of BEAST analysis of the Begoniaceae. Node heights indicate mean ages. Rectangles beside taxon names indicate distributions: red, Hawaii; green, Americas; blue, Africa including Socotra; gold, Asia. Geological epochs are indicated by background colour: light grey, Miocene (23.0–5.3 Ma); mid-grey, Pliocene (5.3–2.6 Ma); dark grey, Holocene and Pleistocene (2.6–0 Ma). Asterisks indicate posterior clade probabilities < 0.85, crosses indicate ML bootstrap values < 75, double crosses indicate posterior clade probabilities < 0.85 and ML bootstrap values < 75. See the Results section for explanation of the clade acronyms. 270 Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) MATERIALS AND METHODS Plant material. — The dataset comprises 802 accessions of 574 species of the Begoniaceae including Hillebrandia sandwicensis, and 51 African (27%), 239 Asian (24%), and 283 Neotropical (40%) species of Begonia (see Appendix 1). All accessions were identified by experts in Begonia taxonomy, with reference to the most up to date taxonomy provided by Hughes & al. (2015–). Sampling focussed upon the inclusion of as many currently accepted sections of Begonia as possible with further emphasis placed upon the type of all published sections. We sampled species from 57 of the 68 currently accepted sections of Begonia. Types of section names were included from 66 of the 103 published sections of Begonia (see Electr. Suppl.: Table S1). In total, 1195 sequences were newly generated for this study. DNA extraction, amplification and sequencing. — Total genomic DNA was extracted from silica-dried or fresh material using Qiagen DNeasy Plant Mini Kits or QiaXtractor high-throughput purification (Qiagen, Germantown, Maryland, U.S.A.) in accordance with the manufacturer’s protocols. Three non-coding plastid DNA regions were chosen for amplification (ndhA intron, ndhF-rpl32 spacer, rpl32-trnL spacer) and the primers used were as in Thomas & al. (2011). The ndhA intron and ndhF-rpl32 spacer were amplified in 13.6 μl reactions containing 0.5 μl of template, 0.75 μl of both forward and reverse primers, 0.1 μl of Phusion polymerase (Thermo Fisher Scientific, Waltham, Massachusetts, U.S.A.), 8.125 μl of ddH 20, 2.5 μl of 10× reaction buffer, 1.25 μl of dNTPs, and 0.75 μl of DMSO. The rpl32-trnL spacer was amplified in 12.5 μl reactions containing 0.5 μl of template, 0.75 μl of both forward and reverse primers, 0.1 μl of Phusion polymerase (Thermo Fisher Scientific), 8.5 μl of ddH20, 2.5 μl of 10× reaction buffer, and 0.25 μl of dNTPs. The PCR temperature profile included: template denaturation at 95°C for 5 min; 34 cycles of denaturation at 98°C for 30 s, primer annealing at 50°C for 30 s, primer extension at 72°C for 30 s; and a final stage primer extension at 72°C for 10 min. PCR products were purified using ExoSAP-IT (Invitrogen/Life Technologies, Carlsbad, California, U.S.A.). Sequencing PCRs used the same primers as for amplification, in 10 μl reactions using BigDye sequencing kits (Invitrogen/Life Technologies). Samples were then sent to GenePool at the University of Edinburgh for sequencing. Phylogenetic analyses. — Sequences were edited with Geneious Pro v.7.1 (Biomatters, 2014) and aligned by hand in BioEdit v.7.2.5 (Hall, 1999). Phylogenetic and divergence-time estimations were carried out with an uncorrelated lognormal relaxed clock in BEAST v.1.8.3 (Drummond & al., 2012). No fossil data suitable for the calibration of divergence time estimation analyses exists within the Begoniaceae, thus we use a single secondary calibration point derived from a fossil-calibrated, Cucurbitales-Fagales wide analysis (Thomas & al., 2012). The age of the Begonia crown group was estimated at 24 Ma (± 3.57 Myr) with a normal distribution. Models of molecular evolution were determined upon a maximum-likelihood topology with jModelTest v.2 (Darriba & al., 2012), resulting in the selection of a GTR + I + Γ model. A tree prior of birth-death incomplete sampling was selected. Four sets of two searches each comprising two MCMC (Markov chain Monte Carlo) chains were run for between 3.2 × 107 and 3.4 × 107 generations on the Cipres Science Gateway (Miller & al., 2010) and sampled every 10,000 generations. Time-series plots were analysed in Tracer v.1.8.3 (Rambaut & Drummond, 2009) to ensure adequate sample size (> 200) for all parameters and convergence of parameters. Runs were combined in LogCombiner v.1.8.3 and the maximum clade credibility (MCC) chronogram was selected in TreeAnnotator v.1.8.3 (Drummond & al., 2012). Preliminary analyses were run with identical parameters on data for each DNA region. No well-supported incongruence was present among the three trees, so we proceeded with the combined analysis. Maximum likelihood (ML) phylogenetic analysis was carried out using RAxML (Stamatakis & al., 2008), with 100 rapid bootstrap inferences followed by a thorough ML search (10 replicates), using a gamma distribution of rates and a proportion of invariant sites. RESULTS Phylogenetic analyses. — The maximum-clade-credibility (MCC) chronogram of the Begoniaceae (Fig. 2) resolves broadly the same relationships as resolved in previous cpDNA analyses (Plana, 2003; Goodall-Copestake & al., 2010; Thomas & al., 2011; Moonlight & al., 2015). Selected clade-specific posterior probabilities and ages are presented in Table 1. African Begonia is resolved as an early-diverging grade with a single clade of Table 1. Bayesian posterior probabilities (PP), ML bootstrap support values (BS) and divergence time estimates (95% highest posterior density, HPD) for selected clades from a phylogeny of global Begonia (see Fig. 2). Clade PP BS Mean divergence age (95% HPD) [Ma] Begoniaceae – – 24.57 (33.03–16.76) Begonia 1.00 100 22.80 (30.41–16.14) YFAB 1.00 100 9.34 (14.59–4.92) FFAB 1.00 100 10.54 (15.45–6.73) MB 1.00 100 10.42 (15.40–5.34) SDAAB 1 & NC1 1.00 100 15.67 (21.78–10.40) SDAAB 1 1.00 100 7.11 (11.58–3.71) NC1 1.00 100 13.18 (18.30–8.76) Asian Begonia 1.00 100 14.32 (14.9–9.23) Asian Clade C 0.98 72 12.15 (16.66–7.79) Asian Clade D 1.00 94 11.10 (15.39–7.28) SDAAB 2 & NC2 1.00 100 16.42 (21.89–10.66) SDAAB 2 1.00 100 5.05 (9.55–2.13) NC2 1.00 92 15.41 (20.92–10.43) NC2-i 1.00 85 12.47 (17.87–7.98) NC2-ii 1.00 100 13.98 (18.70–9.00) NC2-iii 1.00 96 11.20 (16.00–7.53) Version of Record 271 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Asian Begonia (including Socotran Begonia [SB]) and two clades of Neotropical Begonia (each as sister to a clade of Seasonally Dry Adapted African Begonia [SDAAB]) nested within. We resolve Yellow-flowered African Begonia (YFAB) as sister to the remainder of Begonia (PP: 1.0; BS: 100). The relationships among early-diverging clades of African Begonia were unresolved in earlier analyses (Plana, 2003; GoodallCopestake & al., 2010; Moonlight & al., 2015). The second dichotomy within Begonia resolves African Begonia (excluding SDAAB and SB) as sister to the remainder of the genus, although there is poor support for both this clade (PP: 0.74; BS: 56) and the relationships among its major subclades (PP: 0.48; BS: 19). This relationship was unresolved in earlier analyses (Plana, 2003; Goodall-Copestake & al., 2010). Within this clade, we resolve B. sect. Exalabegonia Warb. as sister to the reciprocally monophyletic fleshy-fruited African Begonia (FFAB) and Malagasy Begonia (MB). The majority of Asian Begonia is resolved within two clades. Asian Clade C and Asian Clade D as identified by Thomas & al. (2011). We resolve Asian Clade C as nested within a grade we informally name as Early Diverging Asian Begonia (EDAB), which also includes SB. The relationships within EDAB, and among major Asian clades are poorly supported. We replicate the results of Moonlight & al. (2015) in resolving Neotropical Clade 1 (NC1) and Neotropical Clade 2 (NC2) as sister to SDAAB 1 and SDAAB 2 respectively (both PP: 1.0; BS: 100). Within NC2, we resolve the same three major subclades as Moonlight & al. (2015) and informally name them: NC2-i, containing Central American and Mexican species; NC2-ii, cane-like Andean and Caribbean species nested within a diverse South American grade; NC2-iii, primarily western South American tuberous, rhizomatous, and scandent species. We resolve NC2-i as sister to NC2-ii, with both sister to NC2-iii. Divergence time estimates. — We estimate a much younger date (24.6 [33.0–16.8] Ma) for the divergence of Begonia and Hillebrandia than that found by Clement & al. (2004) (50.9 [62.8–39.0] to 64.7 [80.1–49.3]). However, the dates we estimate within the genus Begonia (see our Table 1) are similar to those of previous analyses (see table 1 in Moonlight & al., 2015 and table 1 in Thomas & al., 2012). We estimate ages for all major clades of 1–3 Ma older than previous estimates, with the exception of NC1, which is estimated as 13.2 (18.3–8.8) Ma compared to 14.5 (20.1–9.0) Ma in Moonlight & al. (2015). DISCUSSION With few exceptions, species in the Begoniaceae are easily recognisable as such. The majority of species share a more or less asymmetric leaf; and dry, trilocular fruits while all share an inferior ovary and a ring of collar cells on the seed (De Lange & Bouman, 1999). Debate regarding the classification of species within the family has historically focused upon the delimitation of genera and, more recently, sections. All previous sectional and generic classifications of Begonia and the Begoniaceae have their merits with respects to particular sections or clades of Begonia, but we see little value in their 272 TAXON 67 (2) • April 2018: 267–323 comparative assessment. We limit our discussion to the currently accepted sectional classification of Begonia by Doorenbos & al. (1998) and its subsequent updates (Forrest & Hollingsworth, 2003; De Wilde & Plana, 2003; Rubite & al., 2013; Hughes & Takeuchi, 2015; Moonlight & Jara-Muñoz, 2017; Moonlight & al., 2017b). We accept previously accepted sections where the rules of nomenclature require and refer only to previous classifications where they best fit the results of our phylogenetic analyses. In updating the sectional classification of Begonia to take into account information from molecular phylogenetic analyses we have reached an often-uneasy compromise among conflicting criteria: monophyly, stability and utility. It is unlikely that if we were to design a sectional classification without the burden of history, we would have settled upon that which we present. However, even given our often incomplete knowledge, particularly with regards to phylogenetic taxon sampling, the classification we present represents a significant advance from previous sectional classifications of Begonia. In this discussion, we reference both sampled and unsampled species and sections, highlighting clades and sections we consider in need of further work to achieve a completed sectional classification of Begonia. Divergence time estimates The divergence of Begonia and Hillebrandia. — (Fig. 2). The sister relationship between Hillebrandia and Begonia is well established (Clement & al., 2004), and our use of the former as an outgroup confirms the most early-diverging lineages of Begonia are African. Our analyses suggest the two genera diverged 24.6 (33.0–16.8) Ma, much more recently than the range of 50.9 to 64.7 Ma found by Clement & al. (2004). Our divergence time estimates are congruent with a hypothesis of Begonia and Hillebrandia diverging during the Oligocene when suitable habitat was available for Begoniaceae species in the Hawaiian archipelago. The Hawaiian distribution of Hillebrandia may therefore be best explained by a single dispersal from Africa to Hawaii and not a boreotropical or MalaysianPacific origin as suggested by Clement & al. (2004). However, our divergence time analyses are based upon a single, secondary calibration point at the base of Begonia and therefore of limited utility outside the genus. Divergence time estimates within Begonia. — (Table 1). Divergence time estimates for all major clades within Begonia are largely in line, and certainly within the margin of error, of the previous most densely sampled and dated phylogeny of Begonia (Moonlight & al., 2015) (Table 1). In general, we find dates for major clades 1–3 Ma older than those in Moonlight & al. (2015). This is likely because taxon sampling, both in terms of species number and the phylogenetic diversity of those species, is significantly higher in this paper. Indeed, the clades where we see greatest increase in divergence time estimates are those where sampling density has increased most significantly. For example, we now sample almost three times as many species in NC2 as Moonlight & al. (2015), and the mean divergence time estimation for this group has increased from 12.5 (17.2–7.97) Ma to 15.4 (20.9–10.4) Ma. Conversely, the number of species in NC1 has less than doubled and this is the Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) only major clade whose divergence time estimate has decreased (from 14.5 [20.1–8.9] Ma to 13.2 [18.3–8.8] Ma). It is possible that the divergence time estimates of large, undersampled clades (such as B. sect. Petermannia (Klotzsch) A.DC.) may increase given further species-level sampling while relatively well-sampled clades will remain stable. Phylogenetic relationships and sectional monophyly African Begonia (including Malagasy and Socotran Begonia) (Figs. 3, 4). — Sectional delimitation within African Begonia has remained relatively stable compared to that of Asian and American Begonia. This is due to a combination of factors. Most notably fewer species occur in Africa and the morphological gaps between clades are larger (presumably because extinction rates have been higher here due to a combination of the relatively long time that Begonia has existed in Africa coupled with large-scale climatic changes). African Begonia systematics has also benefitted greatly from having a particularly large group of researchers (mostly based in the Netherlands) concentrate their efforts in this geographical area (e.g., De Wilde, 1985a). Other geographical regions are yet to receive the same degree of attention by monographers. The majority of the continent’s sections have been included in recent revisions (B. sect. Augustia (Klotzsch) A.DC. [Irmscher, 1961]; B. sect. Baccabegonia J.M.Reitsma [Reitsma, 1985]; B. sect. Chasmophila J.J.de Wilde & Plana [De Wilde & Plana, 2003]; B. sect. Cristasemen J.J.de Wilde [De Wilde, 1985b]; B. sect. Loasibegonia A.DC. [Sosef, 1994]; B. sect. Mezierea (Gaudich.) Warb. [Klazenga & al., 1994]; B. sect. Peltaugustia (Warb.) F.A.Barkley [Hughes & Miller, 2002]; B. sect. Rostrobegonia Warb., [Irmscher, 1961]; B. sect. Scutobegonia Warb. [Sosef, 1994]; B. sect. Squamibegonia Warb. [De Wilde & Arends, 1980]; B. sect. Tetraphila [De Wilde, 2002]) or floristic treatments (Malagasy Begonia [Keraudren, 1983]) and members of all currently accepted sections except B. sect. Muscibegonia A.DC. have further been included in molecular phylogenetic studies (Plana, 2003; Plana & al., 2004). Our sampling includes some species not included in these studies and vice versa but the sectional relationships and major clades resolved in the studies are identical, thus we discuss the monophyly of sections of African Begonia with reference to all three phylogenies. ● Yellow-flowered African Begonia (YFAB) (Fig. 3). – We resolve a well-supported clade containing all sampled YFAB species as sister to the rest of Begonia. Included are all sampled members of B. sect. Filicibegonia A.DC., B. sect. Loasibegonia and B. sect. Scutobegonia as reciprocally monophyletic groups with B. sect. Scutobegonia resolved as sister to the two other sections. This mirrors the results of Plana (2003) and Plana & al. (2004) despite little overlap in sampled species between theirs and our study. We, therefore, recognise existing circumscriptions of these three sections. We further note the monotypic B. sect. Chasmophila was resolved as sister to this clade and that B. sect. Cristasemen was resolved as sister to the wider clade by Plana (2003) and Plana & al. (2004). Species in both of these sections have yellow flowers and are recognised as currently circumscribed. ● Begonia sect. Exalabegonia (Fig. 3). – Begonia sect. Mezierea was recognised as polyphyletic by Plana (2003) and Plana & al. (2004) with Malagasy species resolved within a clade of Malagasy sections and the two mainland African species (B. meyeri-johannis Engl., B. oxyloba Welw. ex Hook.f.) resolved as a separate clade at the unresolved base of their tree. We achieve greater resolution and resolve the two mainland species, treated here as B. sect. Exalabegonia, as sister to a clade including all Malagasy sections as sister to FFAB. ● Malagasy Begonia (MB) (Fig. 3). – Both our and previous studies resolve a clade containing all sampled species of MB, with B. sect. Erminea A.DC. as sister to the remainder of MB. We resolve as sister to B. sect. Erminea a clade containing Malagasy species of B. sect. Mezierea and all sampled species of B. sect. Nerviplacentaria A.DC. and B. sect. Quadrilobaria A.DC. From both our sampling and Plana (2003) it is clear that B. sect. Quadrilobaria is not monophyletic: B. goudotii A.DC. as sister to B. sect. Mezierea and B. nossibea as nested within B. sect. Nerviplacentaria. ● Fleshy-fruited African Begonia (FFAB) (Fig. 3). – We resolve a clade including all sampled species of B. sect. Tetraphila, B. sect. Squamibegonia, and B. sect. Baccabegonia, which together include all of FFAB except B. sect. Mezierea. The relationship between these sections is poorly resolved, and like Plana (2003) and Plana & al. (2004), we resolve B. sect. Squamibegonia and B. sect. Baccabegonia as sister clades nested within B. sect. Tetraphila. ● Seasonally Dry Adapted African Begonia (SDAAB) (Fig. 3). – The three sections of SDAAB are resolved into two well-supported clades. The first of these (SDAAB 1) contains all sampled members of B. sect. Sexalaria A.DC. and B. sect. Rostrobegonia resolved as sister groups, with both sister to NC1. The second (SDAAB 2) contains all sampled members of B. sect. Augustia, as sister to NC2. ● Socotran Begonia (Fig. 4). – Begonia sect. Peltaugustia is resolved as a monophyletic group within early-diverging Asian Begonia as shown by previous studies (Thomas & al., 2011, 2012; Moonlight & al., 2015). Early-diverging Asian Begonia (Fig. 4). — Like previous studies, we resolve Asian Begonia as monophyletic with Socotran Begonia nested within. The majority of Asian species are resolved in two major clades: clades C and D as defined by Thomas & al. (2011). The remainder of Asian and Socotran Begonia are resolved within a number of smaller clades as a grade with Asian Begonia clade C nested within. The relationships within this clade are poorly resolved. ● Begonia sect. Haagea (Fig. 4). – The monotypic B. sect. Haagea (Klotzsch) A.DC. represented by B. dipetala is resolved as sister to the type of the polyphyletic B. sect. Reichenheimia (Klotzsch) A.DC. (see below). Collectively these two sections are resolved as sister to Socotran Begonia. ● Tuberous Begonia sect. Reichenheimia (Fig. 4). – We resolve the section as polyphyletic. Begonia thwaitesii Hook. was previously considered as a subspecies of another Sri Lankan endemic (B. tenera Dryand.) as B. tenera var. thwaitesii (Hook.) Jayasuriya. Following field observations of both taxa by one of the authors, we consider them as distinct at species level. Version of Record 273 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) TAXON 67 (2) • April 2018: 267–323 Fig. 3. Maximum-clade-credibility chronogram of BEAST analysis of the Begoniaceae focusing on African taxa. Node heights indicate mean ages. Geological epochs are indicated by background colour: light grey, Miocene (23.0–5.3 Ma); mid-grey, Pliocene (5.3–2.6 Ma); dark grey, Holocene and Pleistocene (2.6–0 Ma). Asterisks indicate posterior clade probabilities < 0.85, crosses indicate ML bootstrap values < 75, double crosses indicate posterior clade probabilities < 0.85 and ML bootstrap values < 75. Circles indicate species placed in sections different to their previous placement. 274 Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Fig. 4. Maximum-clade-credibility chronogram of BEAST analysis of the Begoniaceae focusing on Early-diverging Asian Begonia and Asian Begonia Clade C. Node heights indicate mean ages. Geological epochs are indicated by background colour: light grey, Miocene (23.0–5.3 Ma); mid-grey, Pliocene (5.3–2.6 Ma); dark grey, Holocene and Pleistocene (2.6–0 Ma). Asterisks indicate posterior clade probabilities < 0.85, crosses indicate ML bootstrap values < 75, double crosses indicate posterior clade probabilities < 0.85 and ML bootstrap values < 75. Circles indicate species placed in sections different to their previous placement. Version of Record 275 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Begonia thwaitesii is the type of B. sect. Reichenheimia. The section as circumscribed in Doorenbos & al. (1998) includes both rhizomatous species, which are most diverse in Malesia, and tuberous species, which are most diverse on mainland Asia. We create B. sect. Jackia sect. nov. to encompass the rhizomatous species (see below) but our sampling of tuberous species is poor and those species we do sample are resolved as polyphyletic. Begonia albococcinea Hook. and B. floccifera Bedd. are resolved with the unassigned B. malabarica Lam. and B. hymenophylla Gagnep. in a clade with members of B. sect. Parvibegonia A.DC. and B. brandisiana Kurz in the “Diploclinium grade”. The placement of the remainder of the section remains unclear. ● Begonia dioica (Fig. 4). – We resolve B. dioica Buch.Ham. ex D.Don within early-diverging Asian Begonia, although its position as sister to three Indo–Sri Lankan species is poorly supported (PP: 0.22; BS: 59). Asian Begonia (Clade C) (Fig. 4). — Clade C contains primarily mainland Asian species including the “Diploclinium grade” and the “Platycentrum-Sphenanthera clade”. ● Begonia pteridiformis (Fig. 4). – We resolve B. pteridiformis Phutthai as sister to a clade containing B. sect. Parvibegonia, the “Diploclinium grade”, and the “PlatycentrumSphenanthera clade”. ● Begonia sect. Parvibegonia (Fig. 4). – We resolve a clade containing all sampled species of B. sect. Parvibegonia and B. smithiae Geddes as sister to the “Diploclinium grade”. ● The “Diploclinium grade” (Fig. 4). – The majority of species of B sect. Diploclinium were sampled; B. demissa Craib; B. boisiana Gagnep.; B. brandisiana; and B. alicida C.B.Clarke, the type of B. sect. Alicida C.B.Clarke, are resolved as a grade with the “Platycentrum-Sphenanthera clade” (see below) nested within. This is largely consistent with previous studies (Rajbhandary & al., 2011: Thomas & al., 2011, 2012), and the group was informally named “Diploclinium grade” by Rajbhandary & al. (2011). We follow this notation. We resolve the type of B. sect. Diploclinium, B. grandis Dryand., as sister to the rest of the clade. Our placement of B. sect. Alicida within the “Diploclinium grade” is poorly supported and does not match that observed by the ITS phylogeny presented in Rajbhandary & al. (2011), who resolved the section as sister to B. sect. Parvibegonia and B. smithiae. ● The “Platycentrum-Sphenanthera clade” (Fig. 4). – That B sect. Platycentrum and B. sect. Sphenanthera (Hassk.) Warb. form a clade in which both sections are polyphyletic has been demonstrated repeatedly (Tebbitt & al., 2006; Rajbhandary & al., 2011; Thomas & al., 2011, 2012). We confirm both this and the findings of Rajbhandary & al. (2011) that both species of B. sect. Monopteron (A.DC.) Warb., B. griffithiana (A.DC.) Warb. and B. nepalensis (A.DC.) Warb., are nested within this clade. We synonymise B. sect. Sphenanthera and B. sect. Monopteron with B. sect. Platycentrum while recognising a “Sphenanthera group” within B. sect. Platycentrum. Asian Begonia (Clade D) (Figs. 5, 6). — Clade D is most diverse in Malesia but also includes the predominantly Chinese B. sect. Coelocentrum Irmsch. ● Begonia sect. Coelocentrum (Fig. 5). – Begonia sect. Coelocentrum in our analysis forms a clade including B. leprosa 276 TAXON 67 (2) • April 2018: 267–323 Hance, the type of B. sect. Leprosae (T.C.Ku) Y.M.Shui. This is consistent with the section as recircumscribed by Chung & al. (2014) and we follow their delimitation of B. sect. Coelocentrum with B. sect. Leprosae treated as a synonym. Chung & al. further noted that B. bataiensis Kiew and B. longicarpa K.Y.Guan & D.K.Tian, included within B. sect. Leprosae by Shui & al. (2002), do not fall within their B. sect. Coelocentrum. Here we treat B. bataiensis as unassigned to section due to its isolated phylogenetic position in Chung & al. (2014), and B. longicarpa as belonging to B. sect. Platycentrum based on its phylogenetic placement in Rajbhandary & al. (2011) and Chung & al. (2014). We confirm the multiple origin of fleshy fruits with Begonia (Thomas & al., 2011), which is especially prevalent in B. sect. Coelocentrum and sect- Platycentrum. ● Begonia peltatifolia (Fig. 5). – Begonia peltatifolia H.L.Li was treated as a doubtful member of B. sect. Reichenheimia I by Doorenbos & al. (1998) and B. sect. Diploclinium by Gu & al. (2007). It differs considerably from both sections as circumscribed here, in being a relatively large rhizomatous species with very succulent leaves. Given its isolated phylogenetic position here and in the phylogeny of Chung & al. (2014), we treat B. peltatifolia as unassigned to section. ● The “Begonia amphioxus clade” (Fig. 5). – A number of Bornean and mainland Asian members of B. sect. Petermannia are resolved in a well-supported clade as sister to most of the remaining Malesian Begonia in Clade D. We resolve Vietnamese B. cucphuongensis H.Q.Nguyen & Tebbitt in this clade, a species that was unassigned to section upon its description and considered most similar to the unplaced B. boisiana. This hypothesis is unsupported by our analysis (B. boisiana is placed in the “Diploclinium grade”). We also include B. amphioxus Sands within this group, which as originally assigned to B. sect. Platycentrum (Sands, 1990). Doorenbos & al. (1998) later treated this species as a possible new section, while Tebbitt & Maciver (1999) noted morphological and phylogenetic similarities between it and B. sect. Petermannia. We informally name species within this clade the “Begonia amphioxus clade” and treat them within B. sect. Petermannia. ● Begonia sect. Ridleyella (Fig. 5). – Begonia kingiana Irmsch. and B. speluncae Ridl. are resolved as sister to the remainder of Asian Begonia. Begonia kingiana is our only sampled species of B. sect. Ridleyella Irmsch. sensu Doorenbos & al. (1998). Although we could not include the extinct B. eriomischia Ridl., the type of the section, we are confident it would fall within this group given its peltate leaves and retuse stamens. We transfer B. speluncae to B. sect. Ridleyella. ● Begonia sect. Baryandra (Fig. 5). – Begonia sect. Baryandra A.DC. was recently recircumscribed to include a group consisting of largely Philippines species previously in B. sect. Diploclinium (Rubite & al., 2013). This was based upon nrITS data, which resolved the recircumscribed section as a monophyletic group far removed from the type of B. sect. Diploclinium. We resolve the same clade and follow the circumscription of Rubite & al. (2013). ● Begonia sect. Jackia sect. nov. (Fig. 6). – Begonia sect. Reichenheimia sensu Doorenbos & al. (1998) is resolved in our analysis as polyphyletic (Fig. 2). A large number of rhizom- Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Fig. 5. Maximum-clade-credibility chronogram of BEAST analysis of the Begoniaceae focusing on Asian Begonia Clade D (part 1). Node heights indicate mean ages. Geological epochs are indicated by background colour: light grey, Miocene (23.0–5.3 Ma); mid-grey, Pliocene (5.3–2.6 Ma); dark grey, Holocene and Pleistocene (2.6–0 Ma). Asterisks indicate posterior clade probabilities < 0.85, crosses indicate ML bootstrap values < 75, double crosses indicate posterior clade probabilities < 0.85 and ML bootstrap values < 75. Circles indicate species placed in sections different to their previous placement. Version of Record 277 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) atous Malaysian and Sumatran species previously assigned to this section are resolved in a clade sister to the remainder of B. sect. Petermannia and all sampled species of B. sect. Bracteibegonia A.DC. and B. sect. Symbegonia. (Warb.) L.L.Forrest & Hollingsw. This rhizomatous clade is distant both geographically and phylogenetically from the type of B. sect. Reichenheimia, B. thwaitesii from Sri Lanka. Species within the Malaysian and Sumatran clade further differ from the remainder of B. sect. Reichenheimia in their rhizomatous (vs. tuberous) habit and we include them here in B. sect. Jackia sect. nov. A single species of B. sect. Jackia, B. olivacea Ardi, is resolved as sister to B. sect. Bracteibegonia. Begonia olivacea is a Sumatran endemic which morphologically clearly fits in B. sect. Jackia and not B. sect. Bracteibegonia. We suggest chloroplast capture is the most likely cause of the unexpected placement of B. olivacea and we include it here within B. sect. Jackia. ● Begonia sect. Bracteibegonia (Fig. 6). – All sampled species currently classified within B sect. Bracteibegonia are resolved as a monophyletic group sister to the remainder of B. sect. Petermannia and B. sect. Symbegonia. However, further sampling is needed of the potentially dozens of species which are currently in B. sect. Petermannia, but which may fit better into B. sect. Bracteibegonia following a better understanding of how to define these sections morphologically. Our concept of this section follows Hughes & al. (2015b) and includes species with hairy stems, a low-growing spreading habit, short petioles, leaf lamina base often cuneate, short peduncles, a lax and fasciculate androecium, fruits which often have bristles or hairs and pistillate flowers with persistent tepals. ● The “Petermannia-Symbegonia clade” (Fig. 6). – The majority of B. sect. Petermannia is resolved as a grade with B. sect. Symbegonia nested within. It has been clear since it was transferred from a genus that B. sect. Symbegonia is nested within B. sect. Petermannia (Forrest & Hollingsworth, 2003). The clade of B. sect. Petermannia containing B. sect. Symbegonia includes species from Malesia both west and east of Wallace’s line. We informally name this group the “Petermannia-Symbegonia clade” and suggest further study is required to determine whether this group should be split up to allow the continued recognition of B. sect. Symbegonia. ● Unsampled Asian Begonia. – We have been unable to sample members of four sections of Asian Begonia: B. sect. Apterobegonia Warb., B. sect. Monophyllon A.DC., B. sect. Oligandrae M.Hughes & W.N.Takeuchi, and B. sect. Putzeysia (Klotzsch) A.DC. The affinities of two of these sections appear clear and are discussed above (B. sect. Monophyllon, B. sect. Putzeysia) but those of the remainder are less apparent. Begonia sect. Apterobegonia is a monotypic section of unknown affinity and we retain its sectional status. The affinity of B. sect. Oligandrae may lie with B. sect. Symbegonia as discussed by Hughes & Takeuchi (2015). We also did not sample any members of the six species of the “calcarea group”, as defined by Kiew & al. (2016), and they have not been included in previous phylogenetic analyses. We follow Kiew & al. (2016) and do not assign these species to section. 278 TAXON 67 (2) • April 2018: 267–323 Neotropical Clade 1 (Fig. 7). — Neotropical Clade 1 (NC1) as defined by Moonlight & al. (2015) contains the majority of American species from east of Amazonia and a small number of widespread groups found throughout the Americas. The major clades and their branching order replicates those described by Moonlight & al., but we double the number of sampled species within this clade. NC1 can be split into three major clades: two clades of “cane-like” Brazilian Begonia; a clade we informally name the “core Pritzelia clade”; and B. sect. Wageneria (Klotzsch) A.DC. and B. sect. Donaldia (Klotzsch) A.DC. nested within a grade of small groups of phylogenetically and morphologically distinct species we describe as B. sect. Kollmannia sect. nov. and B. sect. Stellandrae sect. nov. ● Begonia sect. Gaerdtia (Fig. 7). – The majority of species in B. sect. Gaerdtia (Klotzsch) A.DC. including the type are resolved in a monophyletic group including B. coccinea Hook., which was previously classified in B. sect. Pritzelia (Klotzsch) A.DC. ● Begonia sect. Latistigma (Fig. 7). – All sampled Brazilian species of B. sect. Knesebeckia, as circumscribed by Doorenbos & al. (1998), are only distantly related to the type of the section (B. incarnata Link & Otto). We resolve a group of species including B. aconitifolia A.DC., the type of B. sect. Latistigma A.DC., which we reinstate, as sister to B. sect. Gaerdtia. ● Begonia sect. Tetrachia (Fig. 7). – We resolve B. sect. Tetrachia Brade as sister to the remainder of NC1, although this placement is poorly supported. Relationships within B. sect. Tetrachia are also poorly supported. ● The “Begonia acetosa clade” (Fig. 7). – We resolve as sister to B. sect. Kollmannia sect. nov. a well-supported clade containing two rhizomatous species currently classified in B. sect. Pritzelia: B. acetosa Vell. and B. subacida Irmsch. This relationship is well supported. ● Begonia sect. Kollmannia sect. nov. (Fig. 7). – We resolve B. thelmae L.B.Sm. & Wassh. and B. jaguarensis L.Kollmann & al. as sister to the “Begonia acetosa clade”. ● Begonia sect. Donaldia (Fig. 7). – Begonia sect. Donaldia is resolved as sister to a clade containing B. sect. Stellandrae sect. nov., and the “Wageneria clade”. Both this clade and its relationships to related sections are well supported, and we also resolve B. dichotoma Jacq. and B. scabrida A.DC. within the clade. ● Begonia sect. Stellandrae sect. nov. (Fig. 7). – We resolve B. hoehneana as sister to the “Wageneria clade”, although this position is poorly supported. ● The “Wageneria clade” (Fig. 7). – All sampled species of B. sect. Wageneria are resolved within a clade sister to B. sect. Stellandrae sect. nov. This clade is remarkable within Begonia for the lack of sequence divergence among its species, indicated in Fig. 7 by the short branch lengths and poor support for the majority of relationships within the clade. A number of species of B. sect. Pritzelia are also resolved within this clade, which is highly supported. ● The “core Pritzelia clade” (Fig. 7). – The majority of species in B. sect. Pritzelia are resolved in a clade containing B. dietrichiana Irmsch., the type of B. sect. Pritzelia. We further resolve B. sect. Scheidweilaria (Klotzsch) A.DC.; the monotypic B. sect. Trendelenbergia (Klotzsch) A.DC.; and B. venosa Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Fig. 6. Maximum-clade-credibility chronogram of BEAST analysis of the Begoniaceae focusing on Asian Begonia Clade D (part 2). Node heights indicate mean ages. Geological epochs are indicated by background colour: light grey, Miocene (23.0–5.3 Ma); mid-grey, Pliocene (5.3–2.6 Ma); dark grey, Holocene and Pleistocene (2.6–0 Ma). Asterisks indicate posterior clade probabilities < 0.85, crosses indicate ML bootstrap values < 75, double crosses indicate posterior clade probabilities < 0.85 and ML bootstrap values < 75. Circles indicate species placed in sections different to their previous placement. Version of Record 279 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) TAXON 67 (2) • April 2018: 267–323 Fig. 7. Maximum-clade-credibility chronogram of BEAST analysis of the Begoniaceae focusing on Neotropical Clade 1. Node heights indicate mean ages. Geological epochs are indicated by background colour: light grey, Miocene (23.0–5.3 Ma); mid-grey, Pliocene (5.3–2.6 Ma); dark grey, Holocene and Pleistocene (2.6–0 Ma). Asterisks indicate posterior clade probabilities < 0.85, crosses indicate ML bootstrap values < 75, double crosses indicate posterior clade probabilities < 0.85 and ML bootstrap values < 75. Circles indicate species placed in sections different to their previous placement. 280 Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Skan ex Hook.f. within this clade, and we transfer all to B. sect. Pritzelia. This clade is well supported and informally named the “core Pritzelia clade”. Neotropical Clade 2 (Figs. 8–10). — Neotropical Clade 2 (NC2) as defined by Moonlight & al. (2015) contains a small number of species and sections from eastern South America and the vast majority of species and sections from the remainder of the Americas. The relationships we observe within NC2 replicate those found by Moonlight & al. but our markedly improved sampling has allowed us to recircumscribe and describe a number of sections within the clade. The clade, NC2 can be split into three main clades: a largely Central American and Mexican clade (NC2-i); an eastern South American, Caribbean, and western South American clade of primarily cane-like species (NC2-ii); and a western South American clade of primarily rhizomatous and tuberous species (NC2-iii). NC2-i (Fig. 8). — This clade is well supported, as is the sub-clade containing our much-expanded sampling of B. sect. Gireoudia. ● Tuberous Central American and Mexican Begonia (Fig. 8). – Our sampling of northern Central American and Mexican tuberous B. sect. Knesebeckia and B. sect. Quadriperigonia Ziesenh. is poor, and we sample only a single species from each. Begonia gracilis Kunth of B. sect. Quadriperigonia is resolved as sister to rhizomatous Central American and Mexican Begonia. This section is morphologically homogenous and it is likely monophyletic. We recognise the section as circumscribed by Doorenbos & al. (1998). From Central American and Mexican B. sect. Knesebeckia we sample only the type of the section (B. incarnata), which is nested within rhizomatous Central American and Mexican Begonia. This placement is surprising, given the many morphological differences between these two sections and we suggest it is unlikely to be representative of the species’ true affinities. ● Fleshy-fruited Central American and Mexican Begonia (Fig. 8). – We resolve B. oaxacana A.DC. and B. heydei C.DC., the type of B. sect. Parietoplacentalia Ziesenh. and the monotypic B. sect. Urniformia Ziesenh. respectively, as sister to the remainder of NC2. The sister relationship of these two sections is well supported but their placement is poorly supported. ● Rhizomatous Central American and Mexican Begonia (Fig. 8). – All sampled species of B. sect. Gireoudia (Klotzsch) A.DC. and B. sect. Weilbachia (Klotzsch & Oersted ex Klotzsch) A.DC. are resolved in a well-supported clade. We resolve B. sect. Weilbachia as polyphyletic within a paraphyletic B. sect. Gireoudia and newly synonymise B. sect. Weilbachia with B. sect. Gireoudia. NC2-ii (Fig. 9). — The branching order among the early-diverging clades of NC2-ii is well-resolved and all major clades are well supported. We resolve a clade containing five well-supported, early-diverging clades from eastern South America of which three correspond to currently recognised sections: B. sect. Solananthera A.DC; B. sect. Trachelocarpus (Müll.Berol.) A.DC.; and B. sect. Microtuberosa Moonlight & Tebbitt. We recognise B. sect. Solananthera and B. sect. Trachelocarpus as delimited by Doorenbos & al. (1998) and B. sect. Microtuberosa as delimited by Moonlight & al. (2017b). We also resolve a clade of Brazilian members of B. sect. Knesebeckia with thyrsoid inflorescences, which we describe as the new section Astrothrix sect. nov., and a clade of members of B. sect. Gaerdtia, for which we reinstate B. sect. Pereira Brade. These five sections are sister to a large clade of cane-like species most diverse in the Caribbean and Andes but with representatives throughout the Americas. This group includes B. sect. Ruizopavonia A.DC.; B. sect. Casparya (Klotzsch) Warb.; B. sect. Semibegoniella (C.DC.) Baranov & Barkley; B. sect. Doratometra; B. sect. Begonia; B. sect. Hydristyles A.DC. and B. sect. Cyathocnemis (Klotzsch) A.DC. and is exceptional among Begonia in the large levels of sequence divergence among clades and species, and appears to have an elevated rate of sequence evolution. ● Begonia sect. Astrothrix sect. nov. (Fig. 9). – We resolve a clade containing two species (B. santos-limae Brade, B. bullatifolia L.Kollmann) of B. sect. Knesebeckia. Together with four unsampled species, these species have been recognised as forming a morphologically and geographically distinct group within B. sect. Knesebeckia (Kollmann, 2009; Kollmann & Peixoto, 2012). ● Begonia sect. Pereira (Fig. 9). – A fourth eastern South American clade contains B. edmundoi Brade, B. lubbersii E.Morren, and B. pseudolubbersii Brade, all recognised by Doorenbos & al. (1998) in B. sect. Gaerdtia. We reinstate B. sect. Pereira, typified by B. edmundoi, to include these species. ● Begonia sect. Pilderia and sect. Rossmannia (Fig. 9). – We resolve a clade containing B. sect. Pilderia (Klotzsch) A.DC. as recircumscribed by Moonlight and Jara (Moonlight & Jara-Muñoz, 2017) as sister to the monotypic B. sect. Rossmannia A.DC. These sections are together resolved as sister to the remainder of NC2-ii. ● Begonia sect. Ephemera sect. nov. (Fig. 9). – A well-supported clade containing the majority of sampled South American members of B. sect. Begonia is resolved as sister to the remainder of NC2-ii. Begonia hirtella Link, treated by Doorenbos & al. (1998) in B. sect. Doratometra, is included in this clade. ● Begonia sect. Ruizopavonia (Fig. 9). – The relationships between the next three early-diverging, well-supported clades within NC2-ii remain unresolved. The first of these contains primarily species treated by Doorenbos & al. within B. sect. Ruizopavonia; however, this section is polyphyletic. Species resolved within this clade share raised veins on the underside of the leaves, elliptic or oblong anthers longer than the filaments, and fewer than five tepals in the female flower (except B. tiliifolia and B. harlingii). We therefore recircumscribe B. sect. Ruizopavonia to include those species with these characters. ● Horn-fruited Andean Begonia (Fig. 9). – Begonia sect. Casparya and B. sect. Semibegoniella together form a distinctive group of Andean species with horn-like fruits that dehisce along the back of the locules. The two sections were distinguished by Doorenbos & al. (1998) by the connate tepals of B. sect. Semibegoniella, its less numerous stamens, and its simple placentae (usually bifid in B. sect. Casparya). While we note that our results suggest the two sections may not be reciprocally monophyletic, we refrain from making formal changes to their circumscriptions while we await more detailed morphological and molecular data (A. Jara, unpub. data). Version of Record 281 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) TAXON 67 (2) • April 2018: 267–323 Fig. 8. Maximum-clade-credibility chronogram of BEAST analysis of the Begoniaceae focusing on Neotropical Clade 2-i. Node heights indicate mean ages. Geological epochs are indicated by background colour: light grey, Miocene (23.0–5.3 Ma); mid-grey, Pliocene (5.3–2.6 Ma); dark grey, Holocene and Pleistocene (2.6–0 Ma). Asterisks indicate posterior clade probabilities < 0.85, crosses indicate ML bootstrap values < 75, double crosses indicate posterior clade probabilities < 0.85 and ML bootstrap values < 75. Circles indicate species placed in sections different to their previous placement. 282 Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) ● Begonia sect. Lepsia (Fig. 9). – The remainder of Begonia sect. Ruizopavonia is resolved in a clade including B. holtonis A.DC., the type of B. sect. Meionanthera A.DC., and B. foliosa Kunth, the type of B. sect. Lepsia (Klotzsch) A.DC. ● Begonia sect. Doratometra (Fig. 9). – Three species of Begonia sect. Doratometra are resolved as sister to the remainder of NC2-ii. We do not sample B. wallichiana Lehm., the type of the section, but reproduce the circumscription of the section of Doorenbos & al. (1998) with the exception of B. hirtella Link, which was treated by Doorenbos & al. in B. sect. Doratometra and we include in B. sect. Ephemera sect. nov. (see discussion above). ● Caribbean Begonia sect. Begonia (Fig. 9). – All sampled Caribbean species of B. sect. Begonia are resolved as a grade with the remainder of NC2-ii nested within. The first dichotomy within this grade separates all sampled species of Caribbean B. sect. Begonia (except B. acutifolia Jacq.) from the remainder of NC2-ii (including B. acutifolia). The second dichotomy resolves B. acutifolia as sister to Bolivian Begonia and B. sect. Cyathocnemis. ● Bolivian Begonia (Fig. 9). – The Bolivian species of Begonia are among the most poorly known of the genus, with most known species described with incomplete descriptions, few having been covered in floristic accounts, and several other species awaiting description (P.W. Moonlight, unpub. data; M.C. Tebbitt, unpub. data). The members of B. sect. Hydristyles (which is almost restricted to Bolivia), monotypic B. sect. Warburgina O.Kuntze, and the Bolivian members of B. sect. Ruizopavonia are particularly problematic. The only species of these groups we sample (B. fissistyla Irmsch.) is not nested within any other sections and as sister to B. rubriflora L.Kollmann, which was described as a member of B. sect. Begonia from cultivated material of uncertain origins presumed to be from Argentina. We cannot speculate upon the placement of the remainder of the species of Bolivian B. sect. Hydristyles, B. sect. Ruizopavonia or B. sect. Warburgina. ● Begonia sect. Cyathocnemnis (Fig. 9). – The final clade of NC2-ii contains primarily species of B. sect. Cyathocnemis. We also resolve B. obtecticaulis Irmsch. within this clade. NC2-iii (Fig. 10). — We increase the sampling within NC2-iii from seven species in Moonlight & al. (2015) to over 50 in our current analysis. Much of the clade remains poorly resolved but we resolve a number of clear, reciprocally monophyletic clades within it. All species included are Andean and the majority are rhizomatous or tuberous members of B. sect. Knesebeckia and B. sect. Eupetalum (Lindl.) A.DC., although the scandent B. sect. Gobenia A.DC. is also nested within this clade. ● Begonia bifurcata (Fig. 10). – Northern Andean B. bifurcata L.B.Sm. & B.G.Schub. is resolved as sister to a primarily northern Andean clade including B. sect. Barya (Klotzsch) A.DC., the “cremnophila group”, B. sect. Eupetalum and B. sect. Knesebeckia group I. This relationship is well supported. ● Northern Andean Begonia sect. Eupetalum (Fig. 10). – We resolve a well-supported clade including all sampled species with multifid styles as sister to a clade containing the simple-styled “cremnophila group”, B. sect. Knesebeckia I and B. sect. Barya. Resolved as the most early-diverging clade of this sister group are B. geraniifolia Hook. and B. weberbaueri Irmsch., which also have simple styles, but the placement of this clade is poorly supported. Our result is not inconsistent with B. geraniifolia, B. weberbaueri, and B. sect. Eupetalum forming a clade but we note our analyses do not strictly support this hypothesis. ● The “cremnophila clade” (Fig. 10). – We resolve a moderately well-supported clade containing B. cremnophila Tebbitt, B. speculum Moonlight & Tebbitt and B. urubambensis Tebbitt as sister to B. sect. Knesebeckia group I. The position of this clade is poorly supported and its affinity to B. sect. Knesebeckia group I and B. sect. Eupetalum remains unclear. These species are superficially very different, but share characters including their asymmetric, cymose inflorescences; four, unequal male tepals; anther morphology; five subequal female tepals; and stigmatic papillae morphology. We informally name this group of species the “cremnophila group”. ● Begonia sect. Knesebeckia group I and B. sect. Barya (Fig. 10). – We resolve a clade containing a number of species of the “acerifolia group” as defined by Tebbitt (2016); and B. monadelpha Ruiz & Pav. ex A.DC., the type of B. sect. Barya. The clade is well-supported but there is no support for relationships among subclades. ● Begonia sect. Australes (Fig. 10). – We resolve all sampled South American tuberous and caulescent members of B. sect. Eupetalum (sensu Doorenbos & al., 1998) with bifid styles within a well-supported clade. This clade corresponds to and includes the type of former B. sect. Australes. Begonia boliviensis A.DC. of B. sect. Barya is also resolved within this clade. ● Begonia sect. Knesebeckia group III (Fig. 10). – The remainder of Andean B. sect. Knesebeckia are resolved in two clades, the “maynensis group” as defined by Moonlight & al. (2017a) and part of the “acerifolia group” as defined by Tebbitt (2016): B. arrogans Irmsch., B. ludwigii Irmsch., B. serotina A.DC. and B. wollnyi Herzog, the tuberous species B. piurensis L.B.Sm. & B.G.Schub. and B. parcifolia C.DC. Together with B. froebelii A.DC., these two clades form a grade within which a clade containing B. lutea L.B.Sm. & B.G.Schub., B. sect. Knesebeckia group II, and B. sect. Gobenia is nested. The relationship between these clades and their close relatives are poorly supported and results are not inconsistent with B. sect. Knesebeckia III forming a monophyletic group, although we find no particular support for this hypothesis. ● Begonia lutea (Fig. 10). – Begonia lutea is resolved as sister to a clade containing B. sect. Knesebeckia group II and B. sect. Gobenia A.DC. as reciprocally monophyletic groups. This placement is poorly supported. ● Begonia sect. Gobenia (Fig. 10). – Begonia sect. Gobenia forms a well-supported clade including all sampled species of the section resolved as sister to B. sect. Knesebeckia group II. Relationships within B. sect. Gobenia are poorly supported. ● Begonia sect. Knesebeckia group II (Fig. 10). – Resolved as sister to B. sect. Gobenia are two undescribed limestone endemics from the Alto Mayo region of northern Perú. These species best fit Doorenbos & al.’s (1998) circumscription of B. sect. Knesebeckia and we informally name this clade B. sect. Knesebeckia group II. Version of Record 283 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) TAXON 67 (2) • April 2018: 267–323 Fig. 9. Maximum-clade-credibility chronogram of BEAST analysis of the Begoniaceae focusing on Neotropical Clade 2-ii. Node heights indicate mean ages. Geological epochs are indicated by background colour: light grey, Miocene (23.0–5.3 Ma); mid-grey, Pliocene (5.3–2.6 Ma); dark grey, Holocene and Pleistocene (2.6–0 Ma). Asterisks indicate posterior clade probabilities < 0.85, crosses indicate ML bootstrap values < 75, double crosses indicate posterior clade probabilities < 0.85 and ML bootstrap values < 75. Circles indicate species placed in sections different to their previous placement. 284 Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Fig. 10. Maximum-clade-credibility chronogram of BEAST analysis of the Begoniaceae focusing on Neotropical Clade 2-iii. Node heights indicate mean ages. Geological epochs are indicated by background colour: light grey, Miocene (23.0–5.3 Ma); mid-grey, Pliocene (5.3–2.6 Ma); dark grey, Holocene and Pleistocene (2.6–0 Ma). Asterisks indicate posterior clade probabilities < 0.85, crosses indicate ML bootstrap values < 75, double crosses indicate posterior clade probabilities < 0.85 and ML bootstrap values < 75. Circles indicate species placed in sections different to their previous placement. Version of Record 285 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) TAXONOMIC TREATMENT Begonia sect. Alicida C.B.Clarke in Hooker, Fl. Brit. India 2: 637. 1879 ≡ Begonia subg. Alicida (C.B.Clarke) C.B.Clarke in Bot. J. Linn. Soc. 18: 116. 1881 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 2. 1972): Begonia alicida C.B.Clarke. — [Fig. 13A]1 Species list. – 4 species: B. alicida C.B.Clarke, B. tricuspidata C.B.Clarke, B. triradiata C.B.Clarke, B. vagans Craib. Distribution. – ASIA: Myanmar, Thailand. Taxonomic notes. – Our placement of B. sect. Alicida within the “Diploclinium grade” does not match that observed by Rajbhandary & al. (2011), who resolved B. alicida as sister to a clade of B. sect Parvibegonia. Given the conflict between datasets, we refer to the description in Doorenbos & al. (1998: 64) and follow the circumscription therein. Begonia sect. Apterobegonia Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 143. 1894 – Type: Begonia deliculata Parish ex C.B.Clarke. — [Fig. 13B] Species list. – 2 species: B. delicatula Parish ex C.B.Clarke, B. phutthaii M.Hughes. Distribution. – ASIA: Myanmar, Thailand. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 65) and follow the circumscription therein. Begonia sect. Astrothrix Moonlight, sect. nov. – Type: Begonia bullatifolia L.Kollmann. — [Fig. 15A] Description. – Acaulescent, rhizomatous herbs, perennial, with an indumentum of stellate hairs. Stipules persistent. Leaves alternate, basifixed or peltate, blade subsymmetrical or asymmetrical, veins palmate. Inflorescence axillary, thyrsoid, protandrous, bracts deciduous or persistent. Male flowers: with 4 free perianth segments; stamens 20 to 50, filaments fused for less than half their length or free to the base (B. espiritosantensis), anthers obovate, dehiscing via lateral slits, connective not projecting or slightly projecting. Female flowers: bracteoles absent or 2 at the base of the ovary, with 5 to 6 free perianth segments; ovary and fruit with 3 wings, wings unequal, 3-locular, placentas axile, bifid, bearing ovules on both surfaces; styles 3, free to base, bifid, stigmatic papillae in a once spirally twisted band. Fruit a capsule. Species list. – 5 species: B. bullatifolia L.Kollmann, B. espiritosantensis E.L.Jacques & Mamede, B. fragae L.Kollmann & Peixoto, B. leopoldinensis L.Kollmann, B. santos-limae Brade. Distribution. – AMERICAS: Brazil (Espírito Santo, Rio de Janeiro). Etymology. – The epithet is from the Greek astro- (starry) and thrix (hairs), referring to the stellate indumentum shared by all species in the section. Taxonomic notes. – Begonia sect. Astrothrix represents a geographically and morphologically distinct group of species previously classified within B. sect. Knesebeckia. The section 1 286 Colour plates are ordered geographically and collected at the end of the Taxonomic Treatment (p. 308 ff.). TAXON 67 (2) • April 2018: 267–323 is distinguished from other current and former members of B. sect. Knesebeckia by their unique combination of a rhizomatous habit, stellate indumentum, and thyrsoid inflorescence. The section may be most easily confused with B. sect. Tetrachia as both share a stellate indumentum, basifixed or peltate leaves, and both usually have 5 free female and 4 male perianth segments. These two sections are best distinguished by their inflorescence structure (cymose in B. sect. Tetrachia; thyrsoid in B. sect. Astrothrix). Begonia sect. Augustia (Klotzsch) A.DC., Prodr. 15(1): 384. 1864 ≡ Augustia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 124. 1854 – Type (designated by Barkley & Baranov, in Buxtonian 1(Suppl. 1): 2. 1972): Augustia dregei (Otto & Dietr.) Klotzsch ≡ Begonia dregei Otto & Dietr. — [Fig. 11A] Species list. – 12 species: B. angolensis Irmsch., B. brevibracteata Kupicha, B. dregei Otto & A.Dietr., B. geranioides Hook.f., B. homonyma Steud., B. princeae Gilg, B. pygmea Irmsch., B. riparia Irmsch., B. stolzii Irmsch., B. sutherlandii Hook.f., B. tayloriana Irmsch., B. wakefieldii Gilg ex Engl. Distribution. – AFRICA: Angola, Democratic Republic of the Congo, Kenya, Malawi, Mozambique, South Africa, Swaziland, Tanzania, Zaire, Zimbabwe. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 66) and follow the circumscription therein. Begonia sect. Australes L.B.Sm. & B.G.Schub. in Darwiniana 5: 80. 1941 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 2. 1972): Begonia micranthera Griseb. — [Fig. 15B] Species list. – 20 species: B. barborkae Halda, B. baumannii Lem., B. boliviensis A.DC., B. chrysantha Tebbitt, B. cinnabarina Hook., B. clarkei Hook.f., B. crinita Oliv. ex Hook.f., B. fulgens Lem., B. germaineana Tebbitt, B. heliantha Tebbitt, B. herrerae L.B.Sm. & B.G.Schub., B. krystofii Halda, B. micranthera Griseb., B. pearcei Hook.f., B. phantasma Tebbitt, B. sleumeri L.B.Sm. & B.G.Schub., B. tafiensis Lillo, B. tominana Golding, B. veitchii Hook.f., B. weddelliana A.DC. Description. – Caulescent or rarely acaulescent (in some populations of B. clarkei), tuberous herbs, perennial. Stipules usually persistent, rarely deciduous (in some populations of B. micranthera). Leaves alternate, basifixed, blade usually asymmetrical, rarely symmetrical to subsymmetrical, veins palmate or palmate-pinnate. Inflorescence axillary, cymose, protandrous, bracts persistent. Male flowers: with 4 free perianth segments; stamens 15 to 150, filaments on a rounded receptacle or arranged along a 0.1–2.5 cm long column, anthers ellipsoid, cuboid, or obovoid, dehiscing via lateral slits, connective not projecting or slightly projecting. Female flowers: bracteoles usually absent or rarely 2 at the base of the ovary, with 5 or rarely 6 free perianth segments; ovary and fruit usually with 3 wings, wings unequal, rarely wings reduced to ribs, usually 3-locular, rarely 4-locular (in some populations of B. micranthera), placentas axile, bifid, bearing ovules on both surfaces; styles usually 3, rarely 4 (in some populations of B. micranthera), free or slightly fused at base, usually bifid Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) or rarely 2-lobed (in some populations of B. clarkei), stigmatic papillae usually in a spirally twisted band. Fruit a capsule. Distribution. – AMERICAS: Argentina (Catamarca, Jujuy, Salta, Tucumán), Bolivia, Ecuador, Peru. Taxonomic notes. – Begonia sect. Australes as circumscribed here includes all tuberous and caulescent members of B. sect. Eupetalum sensu Doorenbos & al. (1998) with bifid styles. We also include B. boliviensis in this section, which was previously treated in B. sect. Barya. This species possesses the characteristic features of B. sect. Australes but differs in its columnar androecium and red, forward projecting tepals. These characters appear to be adaptations to hummingbird pollination independent of those of B. sect. Barya. We transfer B. boliviensis into B. sect. Australes. We note two collections of B. clarkei from Cuzco, Perú are resolved within B. sect. Eupetalum allied to B. pleiopetala A.DC. These collections are from near the northern end of the range of B. clarkei and B. sect. Australes where the species occurs sympatrically with B. pleiopetala (which is herein recognised in a more narrowly defined B. sect. Eupetalum). We propose recent chloroplast capture as a possible explanation for the observed disagreement between molecules and morphology and continue to classify B. clarkei within B. sect. Australes since this species is caulescent and has bifid styles. Begonia sect. Baccabegonia Reitsma in Agric. Univ. Wageningen Pap. 84(3): 97. 1985 – Type: Begonia baccata Hook.f. — [Fig. 11B] Species list. – 2 species: B. baccata Hook.f., B. crateris Exell. Distribution. – AFRICA: São Tomé and Príncipe (São Tomé). Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 68) and follow the circumscription therein. Begonia sect. Barya (Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 122. 1859 ≡ Barya Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 122. 1854 – Type: Barya monadelpha (Ruiz ex Klotzsch) Klotzsch ≡ Begonia monadelpha Ruiz & Pav. ex A.DC. — [Fig. 15C] Species list. – 2 species: B. monadelpha Ruiz & Pav. ex A.DC., B. soror Irmsch. Distribution. – AMERICAS: Ecuador, Peru. Taxonomic notes. – We follow the circumscription of Doorenbos & al. (1998: 70), although B. boliviensis is transferred to B. sect. Australes (see above). We refer to the description therein but note the section no longer includes tuberous species. Begonia sect. Baryandra A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 122. 1859 – Type: Begonia oxysperma A.DC. — [Fig. 13C] Species list. – 64 species: B. acclivis Coyle, B. acuminatissima Merr., B. alba Merr., B. alvarezii Merr., B. angilogensis Merr., B. anisoptera Merr., B. bernadusii V.S. Guanih, B. biliranensis Merr., B. blancii M.Hughes & C.I Peng, B. calcicola Merr., B. camiguinensis Elmer, B. castilloi Merr., B. chingipengii Rubite, B. chloroneura P. Wilkie & Sands, B. cleopatrae Coyle, B. collisiae Merr., B. colorata Warb., B. copelandii Merr., B. coronensis Merr., B. culasiensis C.I Peng, Rubite, C.W.Lin & K.F.Chung, B. diwolii Kiew, B. doloisii Rimi, B. elmeri Merr., B. fenicis Merr., B. gitingensis Elmer, B. gueritziana Gibbs, B. gutierrezii Coyle, B. hernandioides Merr., B. hughesii Rubite & C.I Peng, B. isabelensis Quisumb. & Merr., B. klemmei Merr., B. lancilimba Merr., B. longinoda Merr., B. longiscapa Warb., B. luzonensis Warb., B. manillensis A.DC., B. merrilliana C.I Peng, Rubite, C.W.Lin & K.F.Chung, B. mindorensis Merr., B. moneta C.I Peng, Rimi & C.W.Lin, B. neopurpurea L.B.Sm. & Wassh., B. nigritarum (Kamel) Steud., B. obtusifolia Merr., B. oxysperma A.DC., B. parva Merr., B. rubiteae M.Hughes, B. rubrifolia Merr., B. rufipila Merr., B. serpens Merr., B. sharpeana F.Muell., B. simunii Rimi, B. subnummularifolia Merr., B. suborbiculata Merr., B. sykakiengii Rubite, C.I Peng, C.W.Lin & K.F.Chung, B. tagbanua M.Hughes, C.I Peng & Rubite, B. tandangii C.I Peng & Rubite, B. taraw M.Hughes, C.I Peng & Rubite, B. tayabensis Merr., B. titoevangelistae D. Tandang & Rubite, B. trichocheila Warb., B. vanoverberghii Merr., B. vespipropinqua F.Y.Chong, B. wadei Merr. & Quisumb., B. wilkiei Coyle, B. woodii Merr. Distribution. – ASIA: Indonesia (Kalimantan, Moluccas), Japan (Nansei-Shoto), Malaysia (Sabah, Sarawak), Philippines, Papua New Guinea, Taiwan. Taxonomic notes. – We refer to the description in Rubite & al. (2013: 4) and follow the circumscription therein. Begonia sect. Begonia – Type: Begonia obliqua L. — [Fig. 15D] = Begonia sect. Podandra A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 122. 1859 – Type: B. decandra Pav. ex A.DC. – Begonia “sect. Begoniastrum” A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 123. 1859, not validly published (Art. 22.2). – Begonia (“sect. Begoniastrum”) “subsect. Eubegonia” Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 149. 1894, not validly published (Art. 21.3 & 22.2). Species list. – 41 species: B. abbottii Urb., B. acutifolia Jacq., B. alcarrasica J.Sierra, B. azuensis Urb. & Ekman, B. banaoensis J.Sierra, B. bissei J.Sierra, B. bolleana Urb. & Ekman, B. brachyclada Urb. & Ekman, B. brachypoda O.E.Schulz, B. bullata Urb. & Ekman, B. cowellii Nash, B. cubensis Hassk., B. decandra Pav. ex A.DC., B. domingensis A.DC., B. eciliata O.E.Schulz, B. exilis O.E.Schulz, B. glaberrima Urb. & Ekman, B. jamaicensis A.DC., B. leivae J.Sierra, B. leuconeura Urb. & Ekman, B. libanensis Urb., B. linearifolia J.Sierra, B. lomensis Britton & P.Wilson, B. maestrensis Urb., B. minor Jacq., B. notiophila Urb., B. obliqua L., B. odorata Willd., B. pensilis L.B.Sm. & Wassh., B. platyptera Urb., B. plumieri A.DC., B. purdieana A.DC., B. purpurea Sw., B. pycnantha Urb. & Ekman, B. repens Lam., B. retusa O.E.Schulz, B. rotundifolia Lam., B. schulziana Urb. & Ekman, B. stipulacea Willd., B. vincentina O.E.Schulz, B. wrightiana A.DC. Species whose membership is doubtful. – 5 species: B. admirabilis Brade, B. konderreisiana L.B.Sm. & R.C.Sm., B. lindmanii Brade, B. lanstyakii Brade, B. lucidissima Golding & Kareg. Version of Record 287 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Distribution. – AMERICAS: Cuba, Dominica, Dominican Republic, Grenada, Guadeloupe, Haiti, Jamaica, Martinique, Montserrat, Puerto Rico, Saint Kitts & Nevis, Saint Lucia, Saint Vincent, Trinidad-Tobago. — Doubtful species: Brazil (Bahia, Espírito Santo, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Rio de Janeiro, Santa Catarina, São Paulo), Paraguay. Taxonomic notes. – All sampled non-Caribbean species of B. sect. Begonia are only distantly related to the type of the section and are transferred to other sections (see B. sect. Ephemera sect. nov.; the “core Pritzelia clade”) with the exception of an unsampled group of many-flowered eastern South American species. We assign these species doubtfully to B. sect. Begonia, recognising a much-diminished section restricted to the Caribbean. The placement of B. acutifolia as sister to the remainder of NC2-ii resolves B. sect. Begonia as paraphyletic with the remainder of NC2-ii nested within. Begonia acutifolia is a Cuban species most similar to a number of other Cuban species that differ from the remainder of Caribbean B. sect. Begonia in their relatively straight, lanceolate leaves (e.g., B. banaoensis J.Sierra, B. cubensis Haask., B. maestrensis Urb.). We sample only B. acutifolia of this poorly known group and assign all species to B. sect. Begonia. We refer to the description in Doorenbos & al. (1998: 74), which fits B. sect. Begonia despite its significantly reduced circumscription. Begonia sect. Bracteibegonia A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 127. 1859 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 2. 1972): Begonia bracteata Jack. – [Fig. 13D] Species list. – 13 species: B. aberrans Irmsch., B. barbellata Ridl., B. beludruvenea M.Hughes, B. bracteata Jack, B. fasciculata Jack, B. flexula Ridl., B. horsfieldii Miq. ex A.DC., B. jackiana M.Hughes, B. lepida Blume, B. lepidella Ridl., B. pilosa Jack, B. triginticollium Girm., B. verecunda M.Hughes Distribution. – ASIA: Indonesia (Java, Sumatra), Malaysia (Peninsular Malaysia). Taxonomic notes. – Our concept of this section follows Hughes & al. (2015b) and includes species with hairy stems, a low-growing spreading habit, leaves with short petioles, short peduncles, a lax and fasciculate androecium and fruits which often have bristles or hairs and pistillate flowers with persistent tepals. Many species from Borneo currently classified in B. sect. Petermannia have this suite of characters and may need to be transferred to B. sect. Bracteibegonia following further research. However, given the flood of new taxa in B. sect. Petermannia published during the writing of this paper (e.g., Ardhaka & Ardi, 2016; Julia & Kiew, 2016; Julia & al., 2016), we thought it premature to attempt a recircumscription of these two sections. Begonia sect. Casparya (Klotzsch) Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 146. 1894 ≡ Casparya Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 127. 1854 ≡ Begonia subg. Casparya (Klotzsch) C.B.Clarke in Bot. J. Linn. Soc. 18: 115. 1881 – Type 288 TAXON 67 (2) • April 2018: 267–323 (designated by Doorenbos & al. in Wageningen Agric. Univ. Pap. 98(2): 80, 1998): Casparya coccinea Klotzsch = Begonia urticae L.f. — [Fig. 15E] = Stibadotheca Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 127. 1854 (“Stiradotheca”) ≡ Casparya sect. Stibadotheca (Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 116. 1859 ≡ Begonia (sect. Casparya) subsect. Stibadotheca (Klotzsch) Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 146. 1894 – Type (designated by Baranov & Barkley in Buxtonian 1(Suppl. 1): 2. 1972): Stibadotheca ferruginea (L.f.) Klotzsch ≡ Begonia ferruginea L.f. = Sassea Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 128. 1854 ≡ Begonia (sect. Casparya) subsect. Sassea (Klotzsch) Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 146. 1894 – Type (designated by Baranov & Barkley in Buxtonian 1(Suppl. 1): 2. 1972): Sassea urticae (L.f.) Klotzsch = Begonia urticae L.f. = Isopteryx Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 127. 1854 (“Isopteris”) ≡ Casparya sect. Isopteryx (Klotzsch) A.DC., Prodr. 15(1): 270. 1864 – Lectotype (designated Swart, ING Card 10021. 1960): Isopteryx umbellata (Kunth) Klotzsch ≡ Begonia umbellata Kunth. = Casparya sect. Aetheopteryx A.DC., Prodr. 15(1): 271. 1864 ≡ Begonia (sect. Casparya) subsect. Astheopteryx (A.DC.) Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 146. 1894 – Type: Casparya trispathulata A.DC. ≡ Begonia trispathulata (A.DC.) Warb. = Casparya sect. Andiphila A.DC., Prodr. 15(1): 271. 1864 ≡ Begonia (sect. Casparya) subsect. Andiphila (A.DC.) Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 146. 1894 – Type (designated by Baranov & Barkley in Buxtonian 1(Suppl. 1): 3. 1972): Casparya trianae A.DC. ≡ Begonia trianae (A.DC.) Warb. Species list. – 28 species: B. antioquensis (A.DC.) Warb., B. brevipetala (A.DC.) Warb., B. chlorolepis L.B.Sm. & B.G. Schub., B. colombiana L.B.Sm. & B.G.Schub., B. cornuta L.B.Sm. & B.G.Schub., B. diffusa L.B.Sm. & B.G.Schub., B. diversistipulata Irmsch., B. ferruginea L.f., B. formosissima Sandwith, B. fuchsiiflora (A.DC.) F.A.Barkley & Baranov, B. gamolepis L.B.Sm. & B.G.Schub., B. hirta (Klotzsch) L.B. Sm. & B.G.Schub., B. liesneri Burt-Utley & Utley, B. lipolepis L.B.Sm., B. mariae L.B.Sm., B. montana (A.DC.) Warb., B. nevadensis Dorr, B. raimondii Irmsch., B. solaniflora Jara, B. toledana L.B.Sm. & B.G.Schub., B. trianae (A.DC.) Warb., B. trispathulata (A.DC.) Warb., B. trujillensis L.B.Sm., B. umbellata Kunth, B. ursina L.B.Sm. & B.G.Schub., B. urticae L.f., B. vareschii Irmsch., B. wilburi Burt-Utley & Utley Distribution. – AMERICAS: Colombia, Costa Rica, Ecuador, Panamá, Perú, Venezuela. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 80) and follow the circumscription therein. Begonia sect. Chasmophila J.J.de Wilde & Plana in Edinburgh J. Bot. 60(2): 122. 2003 – Type: Begonia iucunda Irmsch. Species list. – 1 species: B. iucunda Irmsch. Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Distribution. – AFRICA: Democratic Republic of the Congo, Republic of the Congo. Taxonomic notes. – We refer to the description in De Wilde & Plana (2003: 122) and follow the circumscription therein. Begonia sect. Coelocentrum Irmsch. in Mitt. Inst. Allg. Bot. Hamburg 10: 553. 1939 – Type (designated by Baranov & Barkley in Buxtonian 1(Suppl. 1): 3. 1972): Begonia porteri H.Lév. & Vaniot. — [Fig. 13E] = Begonia sect. Leprosae (T.C.Ku) Y.M.Shui in Bot. Bull. Acad. Sin. 43(4): 321. 2002 – Type: B. leprosa Hance. Species list. – 69 species: B. arachnoidea C.I Peng, Yan Liu & S.M.Ku, B. asteropyrifolia Y.M.Shui & W.H.Chen, B. aurantiflora C.I Peng, Yan Liu & S.M.Ku, B. auritistipula Y.M.Shui & W.H.Chen, B. austroguangxiensis Y.M.Shui & W.H.Chen, B. babeana Aver. & H.Q.Nguyen, B. bamaensis Yan Liu & C.I Peng, B. biflora T.C.Ku, B. ×breviscapa C.I Peng, Yan Liu & S.M. Ku, B. calciphila C.I Peng, B. chongzuoensis Yan Liu, S.M.Ku & C.I Peng, B. circularis C.I Peng & C.W.Lin, B. cirrosa L.B.Sm. & Wassh., B. crystallina Y.M.Shui & W.H.Chen, B. curvicarpa S.M.Ku, C.I Peng & Yan Liu, B. cylindrica D.R.Liang & X.X.Chen, B. daxinensis T.C.Ku, B. debaoensis C.I Peng, Yan Liu & S.M.Ku, B. fangii Y.M.Shui & C.I Peng, B. ferox C.I Peng & Yan Liu, B. filiformis Irmsch., B. fimbribracteata Y.M.Shui & W.H.Chen, B. guangxiensis C.Y.Wu, B. guixiensis Yan Liu, S.M.Ku & C.I Peng, B. huangii Y.M.Shui & W.H.Chen, B. jingxiensis D.Fang & Y.G.Wei, B. kui C.I Peng, B. langsonensis C.I Peng & C.W.Lin, B. lanternaria Irmsch., B. leipingensis D.K.Tian, L.H.Yang & C.Li, B. leprosa Hance, B. liuyanii C.I Peng, S.M.Ku & W.C. Leong, B. locii C.I Peng, C.W.Lin & N.H. Nguyen, B. longa C.I Peng & W.C. Leong, B. longgangensis C.I Peng & Yan Liu, B. longistyla Y.M.Shui & W.H.Chen, B. luochengensis S.M.Ku, C.I Peng & Yan Liu, B. luzhaiensis T.C.Ku, B. masoniana Irmsch., B. melanobullata C.I Peng & C.W.Lin, B. montaniformis C.I Peng, C.W.Lin & N.H. Nguyen, B. morsei Irmsch., B. nahangensis Aver. & H.Q.Nguyen, B. ningmingensis D.Fang, Y.G.Wei & C.I Peng, B. obliquefolia S.H.Huang & Y.M.Shui, B. ornithophylla Irmsch., B. pengii S.M.Ku & Yan Liu, B. phuthoensis H.Q.Nguyen, B. picturata Yan Liu, S.M.Ku & C.I Peng, B. platycarpa Y.M.Shui & W.H.Chen, B. porteri H.Lév. & Vaniot, B. pseudodaxinensis S.M.Ku, Yan Liu & C.I Peng, B. pseudodryadis C.Y.Wu, B. pseudoleprosa C.I Peng, Yan Liu & S.M.Ku, B. pulvinifera C.I Peng & S.M.Ku, B. retinervia D.Fang, D.H.Qin & C.I Peng, B. rhynchocarpa Y.M.Shui & W.H.Chen, B. rugosula Aver., B. semiparietalis Yan Liu, S.M.Ku & C.I Peng, B. setulosopeltata C.Y.Wu, B. sinofloribunda Dorr, B. sonlaensis Aver., B. subcoriacea C.I Peng, Yan Liu & S.M.Ku, B. ufoides C.I Peng, Y.H.Qin & C.W.Lin, B. umbraculifolia Y.Wan & B.N.Chang, B. variegata Y.M.Shui & W.H.Chen, B. variifolia Y.M.Shui & W.H.Chen, B. wangii T.T.Yu, B. zhengyiana Y.M.Shui Distribution. – ASIA: China (Guangdong, Guangxi, Guizhou, Yunnan), Vietnam. Taxonomic notes. – We refer to the description in Chung & al. (2014) and follow the circumscription therein. Begonia sect. Cristasemen J.J.de Wilde in Agric. Univ. Wageningen Pap. 84(3): 115. 1984 – Type: Begonia thomeana C.DC. – [Fig. 11D] Species list. – 1 species: B. thomeana C.DC. Distribution. – AFRICA: Gabón, São Tomé and Príncipe (São Tomé). Taxonomic notes. – We refer to the description in De Wilde (1985a: 115) and follow the circumscription therein. Begonia sect. Cyathocnemis (Klotzsch) A.DC., Prodr. 15(1): 332. 1864 ≡ Cyathocnemis Klotzsch in Abh. Königl. Akad. Wiss. Berlin 1854: 220. 1855 – Type: Cyathocnemis obliqua Klotzsch ≡ Begonia cyathophora Poepp. & Endl. — [Fig. 15F] = Moschkowitzia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 127. 1854 – Type: Moschkowitzia fagopyroides (Kunth) Klotzsch = Begonia denticulata Kunth. Species list. – 18 species: B. altoperuviana A.DC., B. bracteosa A.DC., B. brevicordata L.B.Sm. & B.G.Schub., B. cryptocarpa L.B.Sm. & B.G.Schub., B. cyathophora Poepp. & Endl., B. falcifolia Hook.f., B. laxa L.B.Sm. & B.G.Schub., B. lophoptera Rolfe, B. lucifuga Irmsch., B. magdalenae L.B.Sm. & B.G.Schub., B. obtecticaulis Irmsch., B. pseudoglauca Irmsch., B. roezlii Regel, B. stenotepala L.B.Sm. & B.G.Schub., B. subciliata A.DC., B. subspinulosa Irmsch., B. suprafastigiata Irmsch., B. tribracteata Irmsch. Species whose membership is doubtful. – 4 species: B. denticulata Kunth, B. machrisiana L.B.Sm. & B.G.Schub., B. nubicola L.B.Sm. & B.G.Schub., B. wageneriana (Klotzsch) Hook. Distribution. – AMERICAS:Bolivia, Colombia, Peru. — Doubtful species: Brazil (Amazonas, Goiás), Venezuela. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 88) and follow the circumscription therein with the exclusion of B. viridiflora A.DC. (see B. sect. Ruizopavonia) and the addition of B. obtecticaulis. The female flowers of this species were unknown in 1998 but match those of B. sect. Cyathocnemis (P.W. Moonlight, unpub. data). Our sampling lacks northern Andean species (e.g., B. denticulata Kunth., B. nubicola L.B.Sm. & B.G.Schub.) and the single Brazilian species (B. machrisiana L.B.Sm. & B.G.Schub.) so we follow Doorenbos & al. (1998) in doubtfully assigning these species to the section. Begonia sect. Diploclinium (Lindl.) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 129. 1859 ≡ Diploclinium Lindl., Veg. Kingd., ed. 2: 319. 1847 – Type: Diploclinium evansianum (Andrews) Lindl. = Begonia grandis Dryand. — [Fig. 13F] = Begonia sect. Trilobaria A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 131. 1859 – Type: Begonia ovatifolia A.DC. – Begonia “subg. Eubegonia” C.B.Clarke in Bot. J. Linn. Soc. 18: 117. 1881, not validly published. Species list. – 85 species: B. acutitepala K.Y.Guan & D.K.Tian, B. adscendens C.B.Clarke, B. alveolata T.T.Yu, B. anceps Irmsch., B. arboreta Y.M.Shui, B. arnottiana (Wight) A.DC., B. asperifolia Irmsch., B. brandisiana Kurz, B. burmensis L.B.Sm. & Wassh., B. cavaleriei H.Lév., B. cehengensis Version of Record 289 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) T.C.Ku, B. concanensis A.DC., B. coptidi-montana C.Y.Wu, B. cordifolia (Wight) Thwaites, B. demissa Craib, B. dentatobracteata C.Y.Wu, B. dioica Buch.-Ham. ex D.Don, B. discreta Craib, B. exposita Phutthai & M.Hughes, B. fengii T.C.Ku, B. festiva Craib, B. fimbristipula Hance, B. flagellaris H.Hara, B. fordii Irmsch., B. fulgurata C.I Peng, C.W.Lin & Phutthai, B. garrettii Craib, B. gigabracteata H-Z.Li & H.Ma, B. glechomifolia C.M.Hu ex C.Y.Wu & T.C.Ku, B. grandis Dryand., B. guaniana H.Ma & Z.D.Wei, B. guishanensis S.H.Huang & Y.M.Shui, B. howii Merr. & Chun, B. hymenocarpa C.Y.Wu, B. imitans Irmsch., B. incerta Craib, B. incondita Craib, B. jayaensis Kiew, B. josephii A.DC., B. kanburiensis Phutthai, B. labordei H.Lév., B. leptoptera H.Hara, B. lithophila C.Y.Wu, B. lowiana King, B. lushaiensis C.E.C.Fischer, B. malipoensis S.H.Huang & Y.M.Shui, B. mashanica D.Fang & D.H.Qin, B. minicarpa H.Hara, B. morifolia T.T.Yu, B. modestiflora Kurz, B. muliensis T.T.Yu, B. murina Craib, B. notata Craib, B. obsolescens Irmsch., B. ovatifolia A.DC., B. parvuliflora A.DC., B. pedunculosa Wall., B. pengchingii Phutthai & M.Hughes, B. picta Sm., B. poilanei Kiew, B. pseudosubperfoliata Phutthai & M.Hughes, B. putii Craib, B. rabilii Craib, B. ravenii C.I Peng & Y.K.Chen, B. rhodophylla C.Y.Wu, B. rong jiangensis T.C.Ku, B. rotundilimba S.H.Huang & Y.M.Shui, B. rubella Buch.Ham. ex D.Don, B. ruboides C.M.Hu ex C.Y.Wu & T.C.Ku, B. satrapis C.B.Clarke, B. saxifragifolia Craib, B. scintillans Dunn, B. sino-vietnamica C.Y.Wu, B. soluta Craib, B. suboblata D.Fang & D.H.Qin, B. subperfoliata Parish ex Kurz, B. subviridis Craib, B. summoglabra T.T.Yu, B. surculigera Kurz, B. taliensis Gagnep., B. tonkinensis Gagnep., B. tribenensis C.R.Rao, B. wengeri C.E.C.Fischer, B. wenshanensis C.M.Hu ex C.Y.Wu, B. wilsonii Gagnep., B. yui Irmsch. Species whose membership is doubtful. – 16 species: B. acaulis Merr. & L.M.Perry, B. alta Aver., B. bartlettiana Merr. & L.M.Perry, B. clavicaulis Irmsch., B. havilandii Ridl., B. hymenophylloides Kingdon-Ward ex L.B.Sm. & Wassh., B. kaniensis Irmsch., B. longovillosa A.DC., B. minjemensis Irmsch., B. miranda Irmsch., B. pyrrha Ridl., B. subcyclophylla Irmsch., B. vitiensis A.C.Sm., B. xingyiensis T.C.Ku, B. xinyiensis T.C.Ku, B. xishuensis T.C.Ku Distribution. – ASIA: Bhutan, China (Guangdong, Guangxi, Guizhou, Hainan, Sichuan, Yunnan), India (Arunachal Pradesh, Assam, Darjeeling, Kerala, Manipur, Meghalaya, Mizoram, Nagaland, Sikkim, Tamil Nadu, Tripura), Laos, Malaysia (Peninsular Malaysia), Myanmar, Nepal, Pakistan, Papua New Guinea, Sri Lanka, Taiwan, Thailand, Vietnam, West Papua. — Doubtful species: Fiji (Santo Island, Vanua Levu), Indonesia (West Papua), Malaysia (Sarawak), Papua New Guinea, Philippines. Taxonomic notes. – Merging B. sect. Diploclinium, B. sect. Platycentrum and B. sect. Sphenanthera to produce a monophyletic section is impractical and would result in a highly polymorphic section distinguished by no obvious morphological characters. It is more feasible to split B. sect. Diploclinium into its constituent clades and search for morphological characters distinguishing those clades. Our current sampling is too low to permit this thus we recognise a paraphyletic B. sect. Diploclinium and acknowledge that further morphological and 290 TAXON 67 (2) • April 2018: 267–323 phylogenetic work is required to recircumscribe the section into natural groups. Begonia boisiana is also phylogenetically placed within the “Diploclinium grade”. This is a morphologically aberrant species for the grade (e.g., it differs by its entire placentae and shrubby habit) and has a conflicting placement in a phylogeny composed of both nuclear and chloroplast data, where it appears in a clade with B. sect. Coelocentrum and sect. Petermannia (Chung & al., 2014). This species was unplaced to section by Doorenbos & al. (1998), which we follow here. The similarity between B. brandisiana, B. burmensis L.B.Sm. & Wassh. and B. demissa was noted by Hughes (2008), but the three species have been placed in different sections by Doorenbos & al. (1998): B. brandisiana in B. sect. Reichenheimia, B. burmensis in B. sect. Lauchea, and B. demissa in B. sect. Parvibegonia. The three species are not only morphologically similar but also do not fit easily into any of the three sections currently used to accommodate them. We transfer B. brandisiana, B. demissa, and B. burmensis into B. sect. Diploclinium, based on the position of the first two taxa in the “Diploclinium grade”. Begonia dioica was treated by Doorenbos & al. (1998) within B. sect. Diploclinium and was resolved within the “Diploclinium grade” (see below) in an ITS phylogeny by Rajbhandary & al. (2011) as sister to B. ravenii C.I Peng & Y.K.Chen. Begonia dioica fits well morphologically in with the remainder of B. sect. Diploclinium in being tuberous with almost symmetric leaves. We consider it likely that the early-diverging position of B. dioica within Asian Begonia is correct and that the slightly incongruent position in the ITS phylogeny could be due to later hybridisation. Until further data becomes available on its phylogenetic placement, and further species in B. sect. Diploclinium are sampled, we leave B. dioica in B. sect. Diploclinium. Begonia sect. Donaldia (Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 141. 1859 ≡ Donaldia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 127. 1854 – Lectotype (designated Swart, ING Card 09630. 1959): Donaldia ulmifolia (Willd.) Klotzsch ≡ Begonia ulmifolia Willd. — [Fig. 15G] Species list. – 3 species: B. gesnerioides L.B.Sm. & B.G.Schub., B. saxicola A.DC., B. ulmifolia Willd. Distribution. – AMERICAS: Brazil (Acre, Alagoas, Bahia, Ceará, Espírito Santo, Minas Gerais, Pará, Paraíba, Pernambuco, Rio de Janeiro, Sergipe), Colombia, French Guiana, Guyana, Peru, Suriname, Trinidad-Tobago, Venezuela. Taxonomic notes. – The number of species in B. sect. Donaldia has fallen greatly since 1998, all other species having been synonymised with either B. saxicola A.DC. or B. ulmifolia Willd. (Jacques & Mamede, 2005). We recognise this diminished section with the addition of B. gesnerioides L.B.Sm. & B.G.Schub., which we resolve within the section. Begonia gesnerioides was tentatively assigned to B. sect. Ruizopavonia A.DC. by Doorenbos & al. (1998) but both type collection and specimens collected at its type locality appear indistinguishable from B. ulmifolia and are likely the same species (P.W. Moonlight, unpub. data). Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Also resolved within B. sect. Donaldia are B. dichotoma Jacq., including both a Colombian and a Venezuelan collection, and B. scabrida A.DC. Neither species can be morphologically grouped with B. sect. Donaldia to the exclusion of the members of B. sect. Pritzelia resolved within the “Wageneria clade”. These two species are the only Colombian and Venezuelan members of this group we sampled. We consider it highly likely that an ancestor of B. scabrida and B. dichotoma hybridised with a member of B. sect. Donaldia resulting in chloroplast capture. This may extend to the other three Venezuelan close relatives of these species: B. boucheana (Klotzsch) A.DC., B. otophylla L.B.Sm. & B.G.Schub., and B. verruculosa L.B.Sm. Given the clear morphological similarities between these species and the “Wageneria clade”, which are not shared by B. sect. Donaldia, we retain all as members of B. sect. Pritezlia. We refer to the description in Doorenbos & al. (1998: 98) and follow the circumscription therein. Begonia sect. Doratometra (Klotzsch) A.DC., Prodr. 15(1): 383. 1864 ≡ Doratometra Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 126. 1854 – Type: Doratometra wallichiana (Lehm.) Klotzsch ≡ Begonia wallichiana Lehm. — [Fig. 15H] = Begonia sect. Poecilia A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 142. 1859 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 3. 1972): Begonia pavoniana A.DC. = Begonia humilis Dryand. = Hoffmannella Klotzsch ex. A.DC., Prodr. 15(1): 299. 1864 – Type: Hoffmannella rosea Klotzsch ex. A.DC. = Begonia semiovata Liebm. Species list. – 8 species: B. bernicei Aymard & G.A.Romero, B. hirsuta Aubl., B. humilis Dryand, B. prieurii A.DC., B. semiovata Liebm., B. steyermarkii L.B.Sm. & B.G.Schub., B. subcostata Rusby, B. wallichiana Lehm. Distribution. – AMERICAS: Brazil (Acre, Bahia, Ceará, Espírito Santo, Minas Gerais, Pará, Parana, Pernambuco, Rio de Janeiro, Roraima, Santa Catarina, São Paulo), Colombia, Costa Rica, Ecuador, French Guiana, Guadeloupe, Guatemala, Guyana, Honduras, Jamaica, Martinique, Mexico (Hidalgo, Oaxaca, Querétaro, San Luis Potosi, Tamaulipas, Veracruz), Nicaragua, Panama, Peru, Saint Vincent, Suriname, Trinidad and Tobago, Venezuela. ASIA: Vietnam (B. wallichiana). Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 99) and follow the circumscription therein with the exception of B. hirtella (see below). We note the unusual South American and Vietnamese distribution of B. wallichiana Lehm. Given the weedy nature of this species its occurance in Asia is likely the result of human introduction rather than a natural, long-distance dispersal event. Begonia sect. Ephemera Moonlight, sect. nov. – Type: Begonia exigua Irmsch. — [Fig. 15I] Description. – Caulescent, rhizomatous (B. delicata Gregório & J.A.S.Costa) or erect succulent herbs, annual. Stipules persistent. Leaves alternate, basifixed, blade subsymmetrical to asymmetrical, veins palmate to palmate-pinnate. Inflorescence axillary, few flowered, cymose, bracts persistent. Male flowers: with 2 or 4 free perianth segments; stamens 3 to numerous, free or filaments fused at the base or united in a column, anthers oblong, dehiscing via lateral slits, connective produced. Female flowers: bracteoles 2 or 3 at the base of the ovary, with 5 free perianth segments; ovary and fruit with 3 wings, wings subequal or unequal, 3-locular, placentas axile, bifid, bearing ovules on all surfaces; styles 3, free, bifid, stigmatic papillae in a spirally twisted band. Fruit a capsule. Species list. – 14 Species: B. alchemilloides Meisn. ex A.DC., B. balansae C.DC., B. cacauicola L.B.Sm. ex S.F. Smith & Wassh., B. cucullata Willd., B. delicata Gregório & J.A.S.Costa, B. descoleana L.B.Sm. & B.G.Schub., B. exigua Irmsch., B. fischeri Schrank, B. hassleri C.DC., B. hirtella Link, B. mollicaulis Irmsch., B. obovatistipula C.DC., B. per-dusenii Brade, B. subvillosa Klotzsch. Distribution. – AMERICAS: Argentina (Buenos Aires, Chaco, Corrientes, Entre Ríos, Formosa, Jujuy, Misiones, Salta, Santa Fe, Santa Cruz, Tucumán), Bolivia, Brazil (Acre, Bahia, Distrito Federal, Durango, Espírito Santo, Goiás, Minas Gerais, Mato Grosso, Mato Grosso do Sul, Paraná, Pernambuco, Rio de Janeiro, Rio Grande do Sul, Roraima, Santa Catarina, São Paulo, Tocantins), Colombia, Costa Rica, Cuba, Ecuador, El Salvador, Guatemala, Guyana, Honduras, Jamaica, México (Chiapas, Hidalgo, Oaxaca, San Luis Potosí, Tabasco, Veracruz), Nicaragua, Panama, Paraguay, Peru, Uruguay, Venezuela. Etymology. – The epithet is from the Greek ephḗmeros (fleeting), referring to the annual life cycle of all species in this section, which is highly unusual in Begonia. Taxonomic notes. – The species we treat in B. sect. Ephemera were treated in B. sect. Begonia or sect. Doratometra (B. hirtella Link) by Doorenbos & al. (1998) and share an annual life history; a succulent, upright habit (1 rhizomatous species); persistent stipules, bracts, and bracteoles; and few-flowered cymose inflorescences. Despite the high support for this clade, it has proven difficult to find characters that distinguish B. sect. Ephemera from B. sect. Doratometra but the majority of species are larger and more robust than those of the latter section. Those species most similar to those of B. sect. Doratometra (e.g., B. hirtella and B. fischeri Schrank) can be distinguished by their combination of 4 male tepals, 5 female tepals, and bifid placentae. These characters are all found within B. sect. Doratometra but not in combination. We note that many of the species of B. sect. Doratometra are poorly known and highly variable. A revision of this group is long overdue, and may help further distinguish these two sections. Some species we treat within B. sect. Doratometra may later prove to be members of B. sect. Ephemera. Begonia sect. Erminea A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 143. 1859 – Type: Begonia erminea L’Hér. — [Fig. 11C] Species list. – 14 species: B. androrangensis Humb. ex Keraudren & Bosser, B. antongilensis Humb. ex Keraudren & Bosser, B. betsimisaraka Humb. ex Keraudren & Bosser, B. bogneri Ziesenh., B. erminea L’Hér., B. keraudrenae Bosser, B. lemurica Keraudren, B. marojejyensis Humb., B. monicae Version of Record 291 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Aymonin & Bosser, B. nana L’Hér., B. nosymangabensis Scherber. & Duruiss., B. pteridoides Scherber. & Duruiss., B. razafinjohanyi Aymonin & Bosser, B. tsimihety Humb. ex Keraudren & Bosser. Species whose membership is doubtful. – 1 species: B. bosseri Keraudren. Distribution. – AFRICA: Madagascar. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 102) and follow the circumscription therein. We likewise doubtfully assign B. bosseri Keraudren to this section (see Doorenbos & al., 1998 for discussion). Begonia sect. Eupetalum (Lindl.) A.DC., Prodr. 15(1): 280. 1864 ≡ Eupetalum Lindl., Intr. Nat. Syst. Bot., ed. 2: 440. 1836 – Type: Eupetalum petalodes Lindl. = Begonia geraniifolia Hook. — [Fig. 15J] = Begonia sect. Begoniella A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 120. 1859 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 3. 1972): Begonia gracillima A.DC. = Begonia pleiopetala A.DC. = Huszia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 121. 1854 ≡ Begonia sect. Huszia (Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser 4, 11: 121. 1859 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 4. 1972): Huszia octopetala (L’Hér.) Klotzsch ≡ Begonia octopetala L’Hér. Species list. – 16 species: B. aequatorialis L.B.Sm. & B.G.Schub.*, B. anemoniflora Irmsch.*, B. anemonoides Azara, B. geraniifolia Hook., B. macra A.DC.*, B. marinae Tebbitt*, B. neoharlingii L.B.Sm. & Wassh., B. novogranatae A.DC.*, B. octopetala L’Hér.*, B. pleiopetala A.DC.*, B. polypetala A.DC.*, B. pseudopleiopetala Tebbitt*, B. rosacea Putz.*, B. rubricaulis Hook.*, B. tumbezensis Irmsch.*, B. weberbaueri Irmsch. — [*The “octopetala group”] Distribution. – AMERICAS: Argentina (Jujuy, Salta, Tucumán), Bolivia, Colombia, Ecuador, Peru, Venezuela. Taxonomic notes. – The circumscription of B. sect. Eupetalum adopted by Doorenbos & al. (1998) was an uneasy alliance of geophytic Andean species. The majority were previously included as members of two synonymised sections. Smith & Schubert (1941) described B. sect. Australes to encompass caulescent, southern Andean species with bifid styles whereas B. sect. Huszia included acaulescent, northern Andean species with multifid styles. Doorenbos & al. (1998) united both groups with geophytic Andean species fitting neither category, including B. weberbaueri Irmsch. and B. geraniifolia, the type of B. sect. Eupetalum, the latter of which possesses both multifid styles and an upright stem. We proceed by recognising a reduced circumscription of B. sect. Eupetalum including only B. sect. Huszia, B. geraniifolia, and B. weberbaueri but recognise that further work is required to resolve the relationships among these taxa. Begonia lutea was doubtfully included in B. sect. Eupetalum by Doorenbos & al. (1998), differing from his circumscription of the section in its entire placentae. The species further differs from our circumscription in its bifid styles and yellow tepals, and is resolved as distantly related to all species included in the section. We treat B. lutea as unassigned to section. 292 TAXON 67 (2) • April 2018: 267–323 Begonia sect. Exalabegonia Warb. in Gartenflora 49: 2. 1900 – Type: Begonia heddei Warb. = Begonia oxyloba Welw. ex Hook.f. — [Fig. 11E] Description. – Caulescent herbs, erect or scandent, perennial. Stipules deciduous. Leaves alternate, basifixed, blade subsymmetric, veins palmate. Inflorescence axillary, a symmetrical cyme, unisexual (B. meyeri-johannis) or bisexual (B. oxyloba), bracts persistent. Male flowers: with 2 or 4 free perianth segments; stamens 7–100, filaments fused for < 1 mm their length or free to base, anthers obovate to oblong or elliptical, dehiscing via lateral slits, connective not projecting or slightly projecting (B. oxyloba). Female flowers: bracteoles absent, with 2–4 free perianth segments; ovary and fruit indehiscent, wingless, 3to 5-locular, placentas parietal for > 60% of the ovary, bifid, bearing ovules on all surfaces; styles 3–5, free to base, bifid, stigmatic papillae in a spirally twisted band. Fruit a berry. Species list. – 2 species: B. meyeri-johannis Engl., B. oxyloba Welw. ex Hook.f. Distribution. – AFRICA: Angola, Burundi, Cameroon, Democratic Republic of the Congo, Equatorial Guinea (Bioko, Equatorial Guinea), Gabón, Ghana, Guinea, Ivory Coast, Kenya, Liberia, Madagascar, Malawi, Mozambique, Nigeria, Republic of the Congo, Rwanda, Sierra Leone, Togo, Uganda. Taxonomic notes. – Plana (2003) discussed at length the possibility of creating a section to include mainland B. oxyloba and B. meyeri-johannis, but was unable to find characters that link the two species to the exclusion of Malagasy B. sect. Mezierea. However, given the addition of further molecular evidence supporting the polyphyly of this group, we consider it prudent to separate Malagasy and mainland species into two sections. Begonia oxyloba is the type of B. sect. Exalabegonia thus we reinstate this section to include B. oxyloba and B. meyeri-johannis. Begonia sect. Filicibegonia A.DC., Prodr. 15(1): 392. 1864 – Type: Begonia aspleniifolia Hook.f. ex A.DC. — [Fig. 11F] = Begonia (sect. Scutobegonia) ser. Longicaules Engl. in Engler & Drude, Veg. Erde 9(3, 2): 616. 1921 – Type (designated by Doorenbos & al. in Wageningen Agric. Univ. Pap. 98(2): 107. 1998): Begonia elatostemmoides Hook.f. Species list. – 8 species: B. aspleniifolia Hook.f. ex A.DC., B. auriculata Hook.f., B. elatostemmoides Hook.f., B. gossweileri Irmsch., B. macrocarpa Warb., B. minutifolia N.Hallé, B. sciaphila Gilg ex Engl., B. sessilifolia Hook.f. Distribution. – AFRICA: Angola, Cameroon, Democratic Republic of the Congo, Equatorial Guinea (Bioko), Gabón, Republic of the Congo. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 107) and follow the circumscription therein. Begonia sect. Gaerdtia (Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 128. 1859 ≡ Gaerdtia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 123. 1854 – Type (designated by Barkey & Baranov in Buxtonian 1(Suppl. 1): 4. 1972): Gaerdtia maculata (Raddi) Klotzsch ≡ Begonia maculata Raddi. — [Fig. 15K] Species list. – 8 species: B. coccinea Hook., B. corallina Carrière, B. dichroa Sprague, B. kunthiana Walp., B. lunaris Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) E.L.Jacques, B. macduffieana L.B.Sm. & B.G.Schub., B. maculata Raddi, B. undulata Schott Distribution. – AMERICAS: Brazil (Espírito Santo, Minas Gerais, Pará, Rio de Janeiro, São Paulo). Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 80) and recognise a reduced circumscription of the section (as discussed under Begonia sect. Pereira below). Begonia coccinea is resolved within the B. sect. Gaerdtia and differs from the remainder of the section only in having entire placentae and thus we transfer it into the section. Begonia sect. Gireoudia (Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 133. 1859 ≡ Gireoudia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 125. 1854 – Type (designated by Barkey & Baranov in Buxtonian 1(Suppl. 1): 4. 1972): Gireoudia involucrata (Liebm.) Klotzsch ≡ Begonia involucrata Liebm. — [Fig. 15L] = Rachia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 124. 1854 ≡ Begonia sect. Rachia (Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 131. 1859 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 4. 1972): Rachia incana (Lindl.) Klotzsch = Begonia peltata Otto & A.Dietr. = Weilbachia Klotzsch & Oerst. ex Klotzsch in Abh. Königl. Akad. Wiss. Berlin 1854: 239. 1855 ≡ Begonia sect. Weilbachia (Klotzsch & Oerst. ex Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 134. 1859, syn. nov. – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 8. 1972): Weilbachia reptans Klotzsch = Begonia ludicra A.DC. = Begonia sect. Auriformia Ziesenh. in Begonian 41: 13. 1974 – Type: Begonia bakeri C.DC. = Begonia cardiocarpa Liebm. = Begonia sect. Liebmannia Ziesenh. in Begonian 43: 64. 1976 – Type: Begonia alice-clarkiae Ziesenh. = Begonia sect. Psathuron A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 131. 1859 – Type: Begonia pinetorum A.DC. Species list. – 110 species: B. acutiloba Liebm., B. aguabuenensis Burt-Utley & Utley, B. alice-clarkiae Ziesenh., B. almedana Burt-Utley & Utley, B. aridicaulis Ziesenh., B. assurgens Irmsch., B. barkeri Knowles & Wescott, B. barsalouxiae Standl. & L.H.J.Williams, B. bettinae Ziesenh., B. boqueronensis Burt-Utley & Utley, B. bowerae Ziesenh., B. breedlovei Burt-Utley, B. broussonetiifolia A.DC., B. buseyi Burt-Utley, B. calderonii Standl., B. calzadae Burt-Utley & Utley, B. campanensis Burt-Utley & Utley, B. cardiocarpa Liebm., B. carletonii Standl., B. carolineifolia Regel, B. carrieae Ziesenh., B. chiapensis Burt-Utley, B. chivatoa Ziesenh., B. conchifolia A.Dietr., B. confusa L.B.Sm. & B.G.Schub., B. copeyana C.DC., B. corredorana C.DC., B. corzoensis Ziesenh., B. crassicaulis Lindl., B. cristobalensis Ziesenh., B. croatii Burt-Utley, B. dressleri Burt-Utley, B. erythrotricha C.DC., B. faustinoi Burt-Utley & Utley, B. fimbriata Liebm., B. fonsecae Standl., B. fortunensis Burt-Utley & Utley, B. fusca Liebm., B. garagarana C.DC., B. gentryi Burt-Utley & Utley, B. glandulosa Hook., B. heracleifolia Schltdl. & Cham., B. hispidavillosa Ziesenh., B. hondurensis Burt-Utley & Utley, B. hubertii Ziesenh., B. hydrocotylifolia Otto ex Hook., B. imperialis Lem., B. involucrata Liebm., B. jaliscana Burt-Utley, B. jarmilae Halda, B. karwinskyana A.DC., B. kenworthyae Ziesenh., B. knoopii Ziesenh., B. kortsiae Ziesenh., B. lindleyana Walp., B. louis-williamsii Burt-Utley, B. ludicra A.DC., B. lyman-smithii Burt-Utley & Utley, B. lyniceorum Burt-Utley, B. makrinii Burt-Utley & Utley, B. manicata Brongn. ex Cels, B. mariti Burt-Utley, B. matudae Burt-Utley & Utley, B. mazae Ziesenh., B. mcphersonii Burt-Utley & Utley, B. mexicana G.Karst ex Fotsch, B. militaris L.B.Sm. & B.G.Schub., B. morii Burt-Utley, B. morrisiorum Rekah Morris & P.D.McMillan ex Rekah Morris, B. motozintlensis Burt-Utley & Utley, B. mucronistipula C.DC., B. multinervia Liebm., B. multistaminea Burt-Utley, B. nelumbiifolia Schltdl. & Cham., B. panamensis Burt-Utley & Utley, B. peltata Otto & A.Dietr., B. philodendroides Ziesenh., B. pinetorum A.DC., B. plantaginea L.B.Sm. & B.G.Schub., B. plebeja Liebm., B. polygonata Liebm., B. popenoei Standl., B. pringlei S. Watson, B. pruinata (Klotzsch) A.DC., B. pseudodaedalea P.D.McMillan & Rekah Morris, B. pseudopeltata Burt-Utley & Utley, B. pudica L.B.Sm. & B.G.Schub., B. purpusii Houghton ex Ziesenh., B. pustulata Liebm., B. quaternata L.B.Sm. & B.G.Schub., B. rafael-torresii Burt-Utley, B. reptans Benth., B. rhizocaulis (klotzsch) A.DC., B. roseibractea Ziesenh., B. sartorii Liebm., B. sericoneura Liebm., B. skutchii Burt-Utley & Utley, B. sousae Burt-Utley, B. sparsipila Baker f., B. squarrosa Liebm., B. stigmosa Lindl., B. strigillosa A.Dietr., B. sukutensis Burt-Utley & Utley, B. thiemei C.DC., B. trichosepala C.DC., B. turrialbae Burt-Utley & Utley, B. urophylla Hook., B. vestita C.DC., B. violifolia A.DC., B. xilitlensis Burt-Utley. Species whose membership is doubtful. – 1 Species: B. setulosa Bertol. Distribution. – AMERICAS: Belize, Colombia, Costa Rica, Ecuador, El Salvador, Guatemala, Honduras, México (Chiapas, Colima, Durango, Guerrero, Hidalgo, Jalisco, Distrito Federal, México State, Michoacán, Morelos, Nayarit, Nuevo León, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosí, Sinaloa, Tabasco, Tamaulipas, Veracruz, Yucatán), Nicaragua, Panama, Venezuela. Taxonomic notes. – All sampled species of B. sect. Gireoudia and B. sect. Weilbachia (Klotzsch & Oersted ex Klotzsch) A.DC. are resolved in a well-supported clade. We resolve B. sect. Weilbachia as polyphyletic within a paraphyletic B. sect. Gireoudia. Begonia sect. Gireoudia and B. sect. Weilbachia encompass mainly rhizomatous but also upright species, and differ primarily in locule number (3 in B. sect. Gireoudia; 2 in B. sect. Weilbachia). Species in B. sect. Weilbachia also tend to be smaller and often have thin, elongated rhizomes although others (e.g., B. popenoei Standl.) have large rhizomes more typical of B. sect. Gireoudia. Given the morphological similarity of these two sections, and the polyphyly of B. sect. Weilbachia within B. sect. Gireoudia, we conclude bilocular fruits have evolved multiple times within the clade and newly synonymise B. sect. Weilbachia with B. sect. Gireoudia. We refer to the descriptions in Doorenbos & al. (1998) of B. sect. Gireoudia (p. 113) and B. sect. Weilbachia (p. 214). Version of Record 293 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Begonia sect. Gobenia A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 119. 1859 – Type (designated by Barkley & Baranov, in Buxtonian 1(Suppl. 1): 4. 1972): B. maurandiae A.DC. — [Fig. 16A] Species list. – 16 species: B. aeranthos L.B.Sm. & B.G.Schub., B. dodsonii L.B.Sm. & Wassh., B. geminiflora L.B.Sm. & Wassh., B. hederacea A.DC., B. hitchcockii Irmsch., B. maurandiae A.DC., B. pululahuana C.DC., B. rubrotincta L.B.Sm. & B.G.Schub., B. secunda L.B.Sm. & Wassh., B. segregata L.B.Sm. & B.G.Schub., B. sodiroi C.DC., B. spadiciflora L.B.Sm. & B.G.Schub., B. tropaeolifolia A.DC., B. truncicola Sodiro ex C.DC., B. wurdackii L.B.Sm. & B.G.Schub., B. ynesiae L.B.Sm. & Wassh. Distribution. – AMERICAS: Colombia, Ecuador, Peru. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 117) and follow the circumscription therein with the addition of B. aeranthos L.B.Sm. & B.G.Schub. This species was treated by Doorenbos & al. (1998) in B. sect. Wageneria but differs only from the traditional members of B. sect. Gobenia in minor characters. Begonia sect. Haagea (Klotzsch) A.DC., Prodr. 15(1): 391. 1864 ≡ Haagea Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 125. 1854 – Type: Haagea dipetala (Graham) Klotzsch ≡ Begonia dipetala Graham. — [Fig. 13G] Species list. – 1 species: B. dipetala Graham. Distribution. – ASIA: India (Karnataka, Kerala, Tamil Nadu), Sri Lanka. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 119) and follow the circumscription therein. Begonia sect. Hydristyles A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 132. 1859 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 4. 1972): Begonia bridgesii A.DC. — [Fig. 16B] Species list. – 10 species: B. andina Rusby, B. bridgesii A.DC., B. fissistyla Irmsch., B. juntasensis Kuntze, B. peltigera Irmsch., B. rubriflora L.Kollmann, B. santarosensis Kuntze, B. subcaudata Rusby ex L.B.Sm. & B.G.Schub., B. unduavensis Rusby, B. unilateralis Rusby. Species whose membership is doubtful. – 1 species: B. ophiogyna L.B.Sm. & B.G.Schub. Distribution. – AMERICAS: Argentina, Bolivia, Peru. — Doubtful species: Colombia. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 119) and follow the circumscription therein with the addition of the previously-unplaced Argentinian B. rubriflora L.Kollmann. This species is phylogenetically placed within the section. Begonia sect. Jackia M.Hughes, sect. nov. – Type: Begonia sublobata Jack. – [Fig. 13H] Description. – Acaulescent rhizomatous herbs, perennial, glabrous or hairy. Stipules persistent. Leaves basifixed or peltate, asymmetric to subsymmetric, veins palmate. Inflorescence axillary, a dichasial cyme, bracts deciduous. Male flowers: with 294 TAXON 67 (2) • April 2018: 267–323 2 or 4 free perianth segments; stamens numerous, filaments fused into a column, anthers obovate, dehiscing via lateral slits, connective retuse. Female flowers: with 2, 3 or 4 free perianth segments, ovary and fruit with 3 wings, wings subequal, 3-locular, placentas axile, entire, bearing ovules on both surfaces; styles 3, free, bifid, stigmatic papillae in a once-spirally twisted band. Fruit a capsule, usually recurved at maturity. Species list. – 53 species: B. andersonii Kiew & S.Julia, B. bimaensis Undaharta & Ardhaka, B. bonii Gagnep., B. caespitosa Jack, B. cladotricha M.Hughes, B. coriacea Hassk., B. corneri Kiew, B. droopiae Ardi, B. fluvialis M.Hughes, B. forbesii King, B. foxworthyi Burkill ex Ridl., B. gesneriifolia Aver., B. goegoensis N.E.Br., B. halabanensis M.Hughes, B. hasskarliana (Miq.) A.DC., B. hosensis C.W.Lin & C.I Peng, B. ignorata Irmsch., B. inversa Irmsch., B. ionophylla Irmsch., B. karangputihensis Girm., B. kemumuensis M.Hughes, B. korthalsiana Miq. ex M.Hughes, B. kudoensis Girm., B. lengguanii Kiew, B. lilliputana M.Hughes, B. lobbii A.DC., B. longipedunculata Golding & Kareg., B. lugrae Ardhaka & Undaharta, B. minuscula Aver., B. mollis A.DC., B. muricata Blume, B. natunaensis C.W.Lin & C.I Peng, B. nurii Irmsch., B. olivacea Ardi, B. orbiculata Jack, B. pasamanensis M.Hughes, B. pseudomuricata Girm., B. puspitae Ardi, B. rajah Ridl., B. raoensis M.Hughes, B. reginula Kiew, B. sendangensis Ardi, B. simolapensis Ardi, B. stictopoda (Miq.) A.DC., B. sublobata Jack, B. sudjanae C.-A.Jansson, B. sumbawaensis Girm., B. sychnantha L.B.Sm. & Wassh., B. tambelanensis (Irmsch.) Kiew, B. tigrina Kiew, B. trichopoda Miq., B. vietnamensis H.Q.Nguyen & C.I Peng, B. yappii Ridl. Distribution. – ASIA: China (Guangxi, Jiangxi), Indonesia (Bali, Java, Kalimantan, Lesser Sunda Islands, Sumatra), Malaysia (Peninsular Malaysia, Sarawak), Vietnam. Etymology. – In honour of William Jack (1795–1822), a pioneering botanist from Scotland who collected in Penang, Singapore and Sumatra. Taxonomic notes. – Created to hold Southeast Asian species previously in B. sect. Reichenheimia, the type of which (B. thwaitesii) is a tuberous species endemic to Sri Lanka in early-diverging Asian Begonia and not associated with the rhizomatous species we move here to B. sect. Jackia. The only species phylogenetically sampled here from B. sect Reichenheimia (B. albo-coccinea, B. floccifera, B. tenera, B. thwaitesii) are placed in early-diverging Asian Begonia. The non-Malesian species we include in B. sect Jackia from continental Southeast Asia and China fit here reasonably well morphologically, but given their geographic location they may not be closely related and further phylogenetic data is needed to confirm this. Begonia sect. Kollmannia Moonlight, sect. nov. – Type: B. jaguarensis L.Kollmann, R.S.Lopes & Peixoto. — [Fig. 16C] Description. – Caulescent, semi-erect to repent herbs, perennial. Stipules persistent. Leaves alternate, basifixed, blade asymmetrical, veins palmate-pinnate. Inflorescence axillary, racemose, bracts persistent. Male flowers: with four free perianth segments; stamens 6 to 8, filaments fused into a column for around half their length, anthers oblong, dehiscing via lateral Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) slits, connective projecting. Female flowers: bracteoles 2 at the base of the ovary, with 5 free perianth segments; ovary and fruit with 3 wings, wings subequal, 3-locular, placentas axile, entire, bearing ovules on all surfaces; styles 3, united at the base, bifid, stigmatic papillae in a spirally twisted band. Fruit a capsule. Species list. – 2 species: B. jaguarensis L.Kollmann, R.S.Lopes & Peixoto, B. thelmae L.B.Sm. & Wassh. Distribution. – AMERICAS: Brazil (Espírito Santo). Etymology. – The epithet honours Ludovic Jean Charles Kollmann, who has described more than 80 species of Brazilian plants including the type of B. sect. Kollmannia and 14 other species of Begonia. Taxonomic notes. – Upon its description, Smith & Wasshausen (1981) were only able to classify B. thelmae L.B.Sm. & Wassh. to section by extending the boundaries of B. sect. Begoniastrum A.DC. (a superfluous name for B. sect. Begonia). Doorenbos & al. (1998) were unable to assign the species to any of their sections. The recently described B. jaguarensis L.Kollmann & al. shares with B. thelmae its creeping habit; fimbriate stipules; few-flowered, racemose inflorescences; and androecium united on a column. We create B. sect. Kollmannia to include these two species. Begonia sect. Kollmannia is most similar to B. hoehneana of the monotypic B. sect. Stellandrae sect. nov., which differs in its reniform leaf blades, and cymose inflorescences. We describe the placentae of B. sect. Kollmannia as entire but note the discussion in Kollmann & al. (2015) regarding the placentae of B. thelmae. Like Kollmann & al. (2015), we only observe entire placentae in cultivated specimens but are aware that this character may vary in wild populations. Begonia sect. Knesebeckia (Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 125. 1859 ≡ Knesebeckia Klotzsch, Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 122. 1854 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 4. 1972): Knesebeckia incarnata (Link & Otto) Klotzsch ≡ Begonia incarnata Link & Otto. — [Fig. 16D, core Knesebeckia; Fig. 16E, Knesebeckia group I; Fig. 16F, Knesebeckia group II; Fig. 16G, Knesebeckia group III] = Begonia sect. Apteron C.DC. in Bull. Herb. Boissier 2(8): 326. 1908 – Type: Begonia exalata C.DC. = Begonia sect. Cylindribegonia L.B.Sm. & B.G.Schub. in Contr. Gray Herb. 127: 25. 1939 – Type: Begonia cylindrata L.B.Sm. & B.G.Schub. = Begonia sect. Dissepbegonia Ziesenh. in Begonian 15: 20. 1948 – Type: Begonia cavum Ziesenh. Species list (core Knesebeckia). – 23 species: B. cavum Ziesenh., B. cebadillensis Houghton ex L.B.Sm. & B.G.Schub., B. cuernavacensis Ziesenh., B. cylindrata L.B.Sm. & B.G. Schub., B. extranea L.B.Sm. & B.G.Schub., B. falciloba Liebm., B. fernaldiana L.B.Sm. & B.G.Schub., B. gracilioides BurtUtley & Utley, B. hintoniana L.B.Sm. & B.G.Schub., B. ignea (Klotzsch) Warsz. ex A.DC., B. incarnata Link & Otto, B. lachaoensis Ziesenh., B. macvaughii Burt-Utley & Utley, B. martinezii Burt-Utley & Utley, B. michoacana L.B.Sm. & B.G.Schub., B. monophylla Pav. ex A.DC., B. nemoralis L.B.Sm. & B.G.Schub., B. relicta L.B.Sm. & B.G.Schub., B. tenuis BurtUtley & Utley, B. tlapensis Burt-Utley & Utley, B. uniflora S.Watson, B. viscida Ziesenh., B. weberlingii Irmsch. Species whose membership is doubtful. – 4 species: B. barkleyana L.B.Sm., B. molinana Burt-Utley, B. mysteriosa L.Kollmann & A.Fontana, B. stenocardia L.B.Sm. & B.G.Schub. Species list (Knesebeckia group I). – 2 species: B. acerifolia Kunth*, B. velata L.B.Sm. & B.G.Schub.* Species list (Knesebeckia group II). – 3 species: B. exalata C.DC., B. microcarpa A.DC., B. pastoensis L.B.Sm. & B.G.Schub. Species list (Knesebeckia group III). – 20 species: B. albomaculata C.DC.**, B. arrogans Irmsch.*, B. brandbygeana L.B.Sm. & B.G.Schub., B. chemillenensis Moonlight & Tebbitt**, B. gorgonea Tebbitt, B. hydrophylloides L.B.Sm. & B.G.Schub., B. leathermaniae O’Reilly & Kareg.*, B. ludwigii Irmsch.*, B. lugonis Wassh., B. maynensis A.DC.**, B. oellgaardii L.B.Sm. & Wassh., B. parcifolia C.DC., B. parodiana L.B.Sm. & B.G.Schub., B. piurensis L.B.Sm. & B.G.Schub., B. scorpiocaulis Moonlight & Tebbitt**, B. serotina A.DC.*, B. sparreana L.B.Sm. & Wassh., B. thyrsoidea Irmsch., B. triramosa Irmsch., B. wollnyi Herzog* [*The “acerifolia group”; **the “maynensis group”] Distribution (core Knesebeckia). – AMERICAS: Costa Rica, El Salvador, Guatemala, Mexico (Chiapas, Colima, Guerrero, Jalisco, México Distrito Federal, México State, Michoacán, Nayarit, Nuevo León, Oaxaca, Veracruz), Panamá. — Doubtful species: AMERICAS: Bolivia, Brazil (Espírito Santo, Paraná, Santa Catarina), Colombia; Honduras, Venezuela. Distribution (Knesebeckia group I). – AMERICAS: Bolivia, Ecuador, Peru. Distribution (Knesebeckia group II). – AMERICAS: Colombia, Ecuador, Peru. Distribution (Knesebeckia group III). – AMERICAS: Argentina (Salta), Bolivia, Brazil (Acre), Ecuador, Peru. Taxonomic notes. – Central American and Mexican B. sect. Knesebeckia is similar to B. sect. Quadriperigonia in its tuberous, upright habit but differs in its axillary, cymose inflorescences. We sample only a single species of this group (B. incarnata, the type of the section name), which is resolved within B. sect. Gireoudia. This placement is surprising and we continue to recognise Central American and Mexican B. sect. Knesebeckia and suggest further sampling of the 22 other species in B. sect. Knesebeckia to confirm the placement of the group. All sampled South American members of B. sect. Knesebeckia are distantly related to B. incarnata. The section has been used as a “taxonomic dustbin”, including species with vastly different habits lacking synapomorphies. We recognise a much-reduced section including only Central American and Mexican species; reinstate the synonymised B. sect. Latistigma (see below); describe a new section (see B. sect. Astrothrix); and transfer species into existing sections (see B. sect. Tetrachia below). We further identify three numbered groups of B. sect. Knesebeckia. Informally naming numbered groups within sections of Begonia has precedence. Doorenbos & al. (1998) identified three morphological groups within both B. sect. Version of Record 295 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Diploclinium and B. sect. Reichenheimia for “ease of survey”. We justify our inclusion of informal numbered groups upon similar grounds, although our groups are phylogenetic rather than morphological and we note that these groups are unlikely to remain assigned to B. sect. Knesebeckia in the future. Begonia sect. Knesebeckia group I includes B. acerifolia Kunth and B. velata L.B.Sm. & B.G.Schub., which form part of the informally named “acerifolia group” of B. sect. Knesebeckia (Tebbitt, 2016). Begonia monadelpha, the type of B. sect. Barya, differs from these two species primarily in its red, tubular flowers and columnar androecium, an apparent adaptation to hummingbird pollination. Together, these three species form a clade but we have been unable to identify synapomorphies that distinguish these species from the remainder of the “acerifolia group” (see B. sect. Knesebeckia group III), and are accordingly reluctant to recircumscribe B. sect. Barya. We compromise by recognising B. sect. Barya and informally naming the remainder of the clade B. sect. Knesebeckia group I. Further morphological and phylogenetic work on the B. acerifolia group is required before formal changes are proposed. Our sampling of B. sect. Knesebeckia group II is limited to two undescribed species from northern Peru. These species share their upright stems with lower portions trailing and rooting at the nodes; simple, glandular indumentum; cymose inflorescences; paired bracteoles; four male and five female flower tepals; columnar androecia; equal-winged ovaries and fruits; and lobed stigmas. These characters are shared with three Ecuadorian and Colombian species: B. exalata C.DC., B. microcarpa A.DC., and B. pastoensis A.DC. Begonia exalata is the type of B. sect. Apteron C.DC., but we refrain from reinstating the section to include these species while we await the description of the two undescribed Peruvian species and phylogenetic confirmation of their proposed relationship with B. exalata. We have been unable to identify synapomorphies that unite Begonia sect. Knesebeckia group III to the exclusion of the members of B. sect. Knesebeckia group I. In particular, the polyphyly of the morphologically well-defined “acerifolia group” is unexpected and may be the result of chloroplast capture. We refrain from describing this group as a section and await further work to identify synapomorphies for the group and to confirm its monophyly. Five unsampled South American species do not fall within any of these groups and are doubtfully assigned to B. sect. Knesebeckia. Begonia molinana is doubtfully assigned because its fragmentary type collection lacks the characters required to assign it to section. Begonia bifurcata is resolved as sister to a clade containing B. sect. Eupetalum, B. sect. Cremnophila, and B. sect. Knesebeckia group I. This species is most similar to B. froebelii (see B. sect. Australes) and its placement in our phylogeny is surprising. We refrain from describing a small section for B. bifurcata and treat it as unplaced to section while we await further phylogenetic data. We refer to the description in Doorenbos & al. (1998: 123), which fits B. sect. Knesebeckia despite its significantly reduced circumscription. 296 TAXON 67 (2) • April 2018: 267–323 Begonia sect. Latistigma A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 119. 1859 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 5. 1972): Begonia aconitifolia A.DC. — [Fig. 16H] Description. – Caulescent, erect suffrutescent herbs, perennial. Stipules deciduous. Leaves alternate, basifixed, blade asymmetrical, palmatifid, veins palmate. Inflorescence axillary, cymose, bracts deciduous. Male flowers: with 2–4 free perianth segments; stamens ca. 50, free, anthers obovate, dehiscing via lateral slits, connective not projecting. Female flowers: bracteoles absent, with 5 free perianth segments; ovary and fruit with 3 wings, wings subequal or unequal, 3-locular, placentas axile, bifid, bearing ovules on all surfaces; styles 3, free, bifid, stigmatic papillae in a spirally twisted band. Fruit a capsule. Species list. – 5 species: B. aconitifolia A.DC., B. olbia Kerch., B. pachypoda L.Kollmann & Peixoto, B. platanifolia Schott, B. teuscheri Linden ex André. Distribution. – Brazil (Espírito Santo, Minas Gerais, Rio de Janeiro, São Paulo). Taxonomic notes. – We treat a number of species previously included within B. sect. Knesebeckia as belonging to the reinstated B. sect. Latistigma. These species are recognised by their upright habit with swollen stem bases, palmately dissected leaves, and large flowers. All are restricted to the Mata Atlântica of Brazil. Begonia sect. Lauchea (Klotzsch) A.DC., Prodr. 15(1): 353. 1864 ≡ Lauchea Klotzsch in Abh. Akad. Berlin 1854: 241. 1855 – Type: Lauchea verticillata (Hook.) Klotzsch ≡ Begonia adenopoda Lem. — [Fig. 13I] = Begonia sect. Dysmorphia A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 136. 1859 – Type: Begonia crenata Dryand. = Begonia sect. Papyracea C.B.Clarke in Hooker, Fl. Brit. India, 2: 649. 1879 – Type (designated here): Begonia adenopoda Lem. Species list. – 5 species: B. adenopoda Lem., B. crenata Dryand., B. namkadingensis C.J.Yang, Souladeth & Tagane, B. pteridiformis Phutthai, B. tenasserimensis Phutthai & M.Hughes Distribution. – ASIA: Myanmar, Thailand. Taxonomic notes. – Begonia pteridiformis and B. tenasserimensis were placed originally in B. sect. Parvibegonia on account of their tuberous habits and bilocular fruit with one enlarged wing and bifid placentae (Phutthai & Sridith, 2010; Phutthai & Hughes, 2017). These characters are shared with B. namkadingensis C.J.Yang & al., which was described as unplaced to section (Yang & al., 2018), and B. adenopoda, the type of B. sect. Lauchea. All these species further share an unusual fruit morphology (covered in glandular hairs, with the largest wing being fleshy; Fig. 13I) and symmetric lanceolate leaves. We recognise an expanded B. sect. Lauchea including B. namkadingensis, B. pteridiformis and B. tenasserimensis but excluding B. burmensis (see B. sect. Diploclinium). Begonia sect. Lepsia (Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 141. 1859 ≡ Lepsia Klotzsch in Ber. Bekanntm. Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 123. 1854 – Type: Lepsia microphylla Klotzsch = Begonia foliosa Kunth. — [Fig. 16I] = Tittelbachia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 126. 1854 ≡ Begonia sect. Tittelbachia (Klotzsch) A.DC., Prodr. 15(1): 291. 1864 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 8. 1972): Tittelbachia fuchsioides (Hook.) Klotzsch = Begonia foliosa Kunth. = Begonia sect. Meionanthera A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 141. 1859 – Type: Begonia holtonis A.DC. Species list. – 8 species: B. barrigae L.B.Sm. & B.G.Schub., B. confinis L.B.Sm. & Wassh., B. foliosa Kunth, B. guaduensis Kunth, B. holtonis A.DC., B. meridensis A.DC., B. opuliflora Putz., B. praerupta Irmsch. Distribution. – AMERICAS: Brazil (Amazonas, Mato Grosso, Para, Rondônia), Colombia, Ecuador, Guyana, Panamam Peru, Suriname, Venezuela. Taxonomic notes. – Begonia sect. Lepsia sensu Doorenbos & al. (1998) was a small section of shrub-like northern Andean species. We transfer a number of former members of B. sect. Ruizopavonia into this section. The redefined B. sect. Lepsia differs from B. sect. Ruizopavonia in having smooth leaf undersurfaces with indistinct primary and secondary veins, globose or subglobose anthers shorter than the filaments (except B. foliosa), and five tepals in the female flower. Included in the species we transfer is B. holtonis, the type of B. sect. Meionanthera. This section was described in the same publication as B. sect. Lepsia (Candolle, 1859) so neither name has priority. We treat B. sect. Lepsia as the accepted name to maintain nomenclatural stability as B. sect. Lepsia was previously accepted. Begonia sect. Mezierea (Gaudich.) Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 139. 1894 ≡ Mezierea Gaudich., Voy. Bonite, Bot. 3: t. 32. 1841 – Type: Mezierea salaziensis Gaudich. ≡ Begonia salaziensis (Gaudich.) Warb. — [Fig. 11H] = Mezierea sect. Aneupteron A.DC., Prodr. 15(1): 407. 1864 – Type: Mezierea salaziensis Gaudich. ≡ Begonia salaziensis (Gaudich.) Warb. Species list. – 4 species: B. comorensis Warb., B. humbertii Keraudren, B. salaziensis (Gaudich.) Warb., B. seychellensis Hemsl. Distribution. – AFRICA: Comoros, Madagascar, Mauritius, Réunion, Seychelles. Taxonomic notes. – Malagasy and Indian Ocean B. sect. Mezierea appears monophyletic and we recircumscribe the section to include only these species, which represent all fleshyfruited Malagasy species excluding B. masoalaensis M.Hughes. This species differs from B. sect. Mezierea in its tuberous habit and axile placentation, and Hughes (2011) suggest it may be more closely related to other Malagasy sections with these characters (e.g., B. sect. Erminea or B. sect. Nerviplacentaria). We follow Hughes (2011) in not assigning this species to section while we await further molecular data. Begonia sect. Loasibegonia A.DC., Prodr. 15(1): 389. 1864 – Type: Begonia prismatocarpa Hook. — [Fig. 11G] = Begonia (sect. Scutobegonia) ser. Cordifolia Engl. in Engler & Drude, Veg. Erde 9(3, 2): 616. 1921 – Type: Begonia pseudoviola Gilg. = Begonia (sect. Scutobegonia) ser. Euscutatae Engl. in Engler & Drude, Veg. Erde 9(3, 2): 617. 1921 – Type (designated by Sosef in Wageningen Agric. Univ. Pap. 94(1): 136. 1994): Begonia quadrialata Warb. Species list. – 21 species: B. adpressa Sosef, B. aequatoguineensis Sosef & Nguema, B. atroglandulosa Sosef, B. duncanthomasii Sosef, B. gentilii De Wild., B. heterochroma Sosef, B. letouzeyi Sosef, B. microsperma Warb., B. minuta Sosef, B. potamophila Gilg, B. prismatocarpa Hook., B. pseudoviola Gilg, B. pulcherrima Sosef, B. quadrialata Warb., B. salisburyana Irmsch., B. scapigera Hook.f., B. schaeferi Engl., B. scutifolia Hook.f., B. sosefiana J.J.de Wilde & Valk., B. staudtii Gilg, B. stellata Sosef. Distribution. – AFRICA: Angola, Burundi, Cameroon, Democratic Republic of the Congo, Equatorial Guinea, Gabón, Ghana, Guinea, Ivory Coast, Liberia, Nigeria, Republic of the Congo, Rwanda, Sierra Leone, Togo. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 131) and follow the circumscription therein. Begonia sect. Monophyllon A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 135. 1859 – Type: Begonia prolifera A.DC. — [Fig. 13J] Species list. – 2 species: B. paleacea Kurz, B. prolifera A.DC. Distribution. – ASIA: Malaysia (Peninsular Malaysia), Myanmar, Singapore, Thailand. Taxonomic notes. – Begonia sect. Monophyllon A.DC. includes two species, B. paleacea and B. prolifera, which are small, tuberous plants with 2-locular fruits similar to those in B. sect. Parvibegonia. It is differentiated from B. sect. Parvibegonia by inflorescences arising from the base of the leaves whereas those of B. sect. Parvibegonia are terminal. We retain B. sect. Monophyllon as a separate section awaiting further molecular evidence, refer to the description in Doorenbos & al. (1998: 136), and follow the circumscription therein. Begonia sect. Microtuberosa Moonlight & Tebbitt in Eur. J. Taxon. 281: 4. 2017 – Type: Begonia elachista Moonlight & Tebbitt. — [Fig. 16J] Species list. – 1 species: B. elachista Moonlight & Tebbitt. Distribution. – AMERICAS: Peru. Taxonomic notes. – We refer to the description in Moonlight & al. (2017: 4) and follow the circumscription therein. Begonia sect. Muscibegonia A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 143. 1859 – Type: Begonia perpusilla A.DC. Species list. – 2 species: B. kalabenonensis Humb. ex Keraudren & Bosser, B. perpusilla A.DC. Distribution. – AFRICA: Madagascar. Taxonomic notes. – We note the similarity of this unsampled section to B. sect. Erminea, which is separated only by its seed morphology (Doorenbos & al., 1998). Further phylogenetic work is required to determine whether the two sections should Version of Record 297 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) remain distinct. We refer to the description in Doorenbos & al. (1998: 138) and follow the circumscription therein. Begonia sect. Nerviplacentaria A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 144. 1859 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 5. 1972): Begonia lyallii A.DC. — [Fig. 11I] Species list. – 10 species: B. baronii Baker, B. cladocarpoides Humb. ex Aymonin & Bosser, B. coursii Humb. ex Keraudren, B. henrilaportei Scherber. & Duruiss., B. lyallii A.DC., B. madecassa Keraudren, B. majungaensis Guillaumin, B. mangorensis Humb. ex Keraudren & Bosser, B. marnieri Keraudren, B. perrieri Bois Distribution. – AFRICA: Madagascar. Taxonomic notes. – The similarity between B. sect. Nerviplacentaria and B. sect. Quadrilobaria has long been noted, with the former only differing in its basally woody stems (Doorenbos & al., 1998). We refer to the description in Doorenbos & al. (1998: 139), reproduce the existing circumscriptions of these sections, and echo Plana (2003) in advocating for further work to determine the phylogenetic and morphological boundaries of these sections. We further assign B. perrieri Bois to B. sect. Nerviplacentaria (unassigned by Doorenbos & al., 1998) because of its affinity to caulescent members of the section (e.g., B. madecassa Keraudren and B. majungaensis Guillamin). Begonia sect. Oligandrae M.Hughes & W.N.Takeuchi in Phytotaxa 197(1): 38. 2015 – Type: Begonia oligandra Merr. & L.M.Perry. — [Fig. 13K] Species list. – 5 species: B. brassii Merr. & L.M.Perry, B. chambersiae W.N.Takeuchi, B. oligandra Merr. & L.M.Perry, B. pentandra W.N.Takeuchi, B. sandsiana W.N.Takeuchi. Distribution. – ASIA: Indonesia (West Papua), Papua New Guinea. Taxonomic notes. – We refer to the description in Hughes & Takeuchi (2015: 38) and follow the circumscription therein. Begonia sect. Parietoplacentalia Ziesenh. in Begonian 36: 36. 1969 – Type: Begonia candollei Ziesenh. = B. oaxacana A.DC. — [Fig. 16K] = Begonia sect. Hexaptera Ziesenh. in Begonian 41: 13. 1974 – Type: Begonia serrulatoala C.DC. = Begonia oaxacana A.DC. Species list. – 3 species: B. oaxacana A.DC., B. udisilvestris C.DC., B. yunckeri Standl. Distribution. – AMERICAS: Costa Rica, El Salvador, Guatemala, Honduras, México (Chiapas, Guerrero, Hidalgo, Jalisco, Puebla, Veracruz), Nicaragua, Panama. Taxonomic notes. – Begonia sect. Parietoplacentalia and B. sect. Urniformia share an erect to semi-scandent habit; an inflorescence structure with male flowers basal and female flowers distal; male flowers with four tepals; female flowers with three tepals; and fleshy fruit. The fruit of B. sect. Parietoplacentalia differ in their beaked rather than horned fruit with bipartite rather than entire placentation. We refer to the description in Doorenbos & al. (1998: 143) and follow the circumscription therein. 298 TAXON 67 (2) • April 2018: 267–323 Begonia sect. Parvibegonia A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 136. 1859 ≡ Begonia subg. Parvibegonia (A.DC.) C.B.Clarke in Bot. J. Linn. Soc. 18: 115. 1881 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 5. 1972): Begonia martabanica A.DC. — [Fig. 13L] = Begonia sect. Heeringia Irmsch. in Mitt. Inst. Allg. Bot. Hamburg 8: 158. 1929, syn. nov. – Type: Begonia sibthorpioides Ridl. Species list. – 28 species: B. andamensis Parish ex C.B.Clarke, B. bella Phutthai, B. brevicaulis A.DC., B. canarana Miq., B. carnosula Ridl., B. elisabethae Kiew, B. erosa Blume, B. flaccidissima Kurz, B. geoffrayi Gagnep., B. grantiana Craib, B. grata Geddes ex Craib, B. integrifolia Dalzell, B. martabanica A.DC., B. parishii C.B.Clarke, B. phamiana Kiew, B. phoeniogramma Ridl., B. procridifolia Wall. ex A.DC., B. pumila Craib., B. rimarum Craib, B. rupicola Miq., B. sibthorpioides Ridl., B. sinuata Wall. ex Meisn., B. socia Craib, B. tenuifolia Dryand., B. thaipingensis King, B. variabilis Ridl., B. wattii C.B.Clarke, B. zollingeriana A.DC. Distribution. – ASIA: Cambodia, India (Andaman Islands, Karnataka, Manipur, Meghalaya, Mizoram, Nagaland), Indonesia (Bali, Java, Sumatra), Laos, Malaysia (Peninsular Malaysia), Myanmar, Nepal, Thailand, Vietnam. Taxonomic notes. – The placement of B. smithiae in this section in both our phylogeny and Rajbhandary & al. (2011) is unusual as it is rhizomatous (not tuberous) and has 3-locular fruit (not 2-locular). We treat this species as unplaced to section, rather than assigned to B. sect. Platycentrum as in Doorenbos & al. (1998). We also resolve B. hymenophylla of B. sect. Reichenheimia in this clade, which fits morphologically with the exception of the fruits having 3 locules. Begonia hymenophylla is perhaps most similar to B. sibthorpioides Ridl., the type of B. sect. Heeringia Irmsch., which is also resolved within B. sect. Parvibegonia. Both differ slightly from core B. sect. Parvibegonia in their diminutive size and monochasial inflorescence structure (as opposed to dichasia ending in monochasia), but otherwise the small, tuberous habit fits well. We reclassify both species in B. sect. Parvibegonia and newly synonymise B. sect. Heeringia with B. sect. Parvibegonia. Begonia sect. Peltaugustia (Warb.) Barkley in Phytologia 24: 156. 1972 ≡ Begonia (sect. Augustia) subsect. Peltaugustia Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 140. 1894 – Type: Begonia socotrana Hook.f. — [Fig. 11J] Species list. – 2 species: B. samhaensis M.Hughes & A.G.Mill., B. socotrana Hook.f. Distribution. – AFRICA: Yemen (Socotra). Taxonomic notes. – We refer to the description in Hughes & Miller (2002: 277) and follow the circumscription therein. Begonia sect. Pereira Brade in Rodriguésia 18: 33. 1945 – Type: Begonia edmundoi Brade. — [Fig. 16L] Description. – Caulescent, erect suffrutescent herbs, one species with a subterranean tubercule-like structure (B. caparaoensis), perennial. Stipules deciduous or persistant. Leaves alternate, peltate or rarely basifixed (B. pseudolubbersii), blade subsymmetrical to asymmetrical, veins pinnate or Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) palmate-pinnate. Inflorescence axillary, cymose, bracts deciduous. Male flowers: with 4 free perianth segments; stamens 30–100, free or fused for less than half their length, anthers obovate, dehiscing via lateral slits, connective not extended or slightly-projecting. Female flowers: bracteoles 1–2 or absent, with 5 free perianth segments; ovary and fruit with 3 wings, wings subequal or unequal, 3-locular, placentas axile, bifid, bearing ovules on all surfaces or only the outside surfaces; styles 3, free, one- or two-lobed, the lobes flattened, stigmatic papillae in a twisted band. Fruit a capsule. Species list. – 5 species: B. caparaoensis E.L.Jacques & L.Kollmann, B. edmundoi Brade, B. jureiensis S.J.Gomes & Mamede, B. lubbersii E. Morren, B. pseudolubbersii Brade. Distribution. – AMERICAS: Brazil (Minas Gerais, Rio de Janeiro, São Paulo). Taxonomic notes. – The species we treat in B. sect. Pereira published by 1998 were all treated within B. sect. Gaerdtia by Doorenbos & al. (1998) while those published subsequently were left unassigned by their respective authors (Gomes da Silva & Mamede, 2000; Jacques & Kollmann, 2009). They differ from the species we treat in B. sect. Gaerdtia in their usually peltate leaves (basifixed in B. pseudolubbersii) with palmate-pinnate venation (rather than pinnate); and their flattened, reniform stigmas (rather than bifid). Begonia edmundoi Brade is the type of B. sect. Pereira thus we reinstate this section to include these species. Begonia sect. Petermannia (Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 128. 1859 ≡ Petermannia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 124. 1854 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 6. 1972): Petermannia cumingiana Klotzsch ≡ B. cumingiana (Klotzsch) A.DC. — [Fig. 14A] Species list. – 416 species: B. abbreviata C.I Peng, B. acidulenta S.Julia & Kiew, B. addrinii S.Julia & Kiew, B. adenodes Irmsch., B. adliniana Rimi, B. aequata A.Gray, B. affinis Merr., B. agusanensis Merr., B. aiensis C.W.Lin & C.I Peng, B. aketajawensis Ardi & D.C.Thomas, B. alabensis Kiew, B. albobracteata Ridl., B. amidalae C.W.Lin & C.I Peng, B. amphioxus Sands, B. androturba Coyle, B. angustilimba Merr., B. anseriana C.W.Lin & C.I Peng, B. anthonyi Kiew, B. apayaoensis Merr., B. apiensis Kiew & S.Julia, B. armykapii S.Julia & C.Y.Ling, B. articulata Irmsch., B. artior Irmsch., B. atricha (Miq.) A.DC., B. augustae Irmsch., B. awongii Sands, B. axillaris Ridl., B. axillipara Ridl., B. bahakensis Sands, B. baik C.W.Lin & C.I Peng, B. baikoides S.Julia & C.Y.Ling, B. bakunensis S.Julia, B. baramensis Merr., B. basintaliana Rimi, B. baturongensis Kiew, B. bayae S.Julia & Kiew, B. beccarii Warb., B. belagaensis S.Julia, B. benaratensis S.Julia, B. bengohensis S.Julia, B. berhamanii Kiew, B. beryllae Ridl., B. bintang Rimi, B. binuangensis Merr., B. bipinnatifida J.J.Sm., B. bolsteri Merr., B. bonthainensis Hemsl., B. borneensis A.DC., B. bosuangiana S.Julia, B. brachybotrys Merr. & L.M.Perry, B. brangbosangensis Girm., B. brevipes Merr., B. brevirimosa Irmsch., B. bruneiana Sands, B. burbidgei Stapf, B. calliantha Merr. & L.M.Perry, B. capituliformis Irmsch., B. carnosa Teijsm. & Binn., B. casiguranensis Quisumb. & Merr., B. caudata Merr., B. cauliflora Sands, B. celata S.Julia & Kiew, B. celebica Irmsch., B. chaiana Kiew & S.Julia, B. chiasmogyna M.Hughes, B. chlorandra Sands, B. chlorosticta Sands, B. chongii Sands, B. ciliifera Merr., B. cincinnifera Irmsch., B. clemensiae Merr. & L.M.Perry, B. cognata Irmsch., B. comestibilis D.C.Thomas & Ardi, B. compacta S.Julia & Kiew, B. congesta Ridl., B. conipila Irmsch. ex Kiew, B. conniegeriae S.Julia & Kiew, B. consanguinea Merr., B. contracta Warb., B. corrugata Kiew & S.Julia, B. crassa S.Julia & Kiew, B. crispipila Elmer, B. crockerensis S.Julia, B. cucphuongensis H.Q.Nguyen & Tebbitt, B. cumingiana (Klotzsch) A.DC., B. cumingii A.Gray, B. cuneatifolia Irmsch., B. cyanescens Sands, B. danumensis F.Y.Chong, B. darthvaderiana C.W.Lin & C.I Peng, B. densiretis Irmsch., B. devexa S.Julia & Kiew, B. didyma D.C.Thomas & Ardi, B. diffusiflora Merr. & L.M.Perry, B. dimorpha S.Julia, B. dinosauria C.W.Lin & C.I Peng, B. divaricata Irmsch., B. divergens Kiew & S.Julia, B. djamuensis Irmsch., B. dolichobracteata Girm., B. dolichocarpa Girm., B. dolichotricha Merr., B. dosedlae Gilli, B. duhungensis Girm., B. eberhardtii Gagnep., B. edanoi Merr., B. edgariana S.Julia & Kiew, B. elatostematoides Merr., B. elatostemma Ridl., B. eliasii Warb., B. erythrogyna Sands, B. esculenta Merr., B. eutricha Sands, B. everettii Merr., B. fasciculiflora Merr., B. felis C.W.Lin & C.I Peng, B. filibracteosa Irmsch., B. flacca Irmsch., B. flammea Rimi, B. flavovirens Kiew & S.Julia, B. flexicaulis Ridl., B. fruticella Ridl., B. fuscisetosa Sands, B. galeolepis Ardi & D.C.Thomas, B. gambutensis Ardi & D.C.Thomas, B. gelasensis Rimi & Kimahim, B. gemella Warb. ex L.B.Sm. & Wassh., B. geniculata Jack, B. georgei Coyle, B. gibbsiae Irmsch. ex Sands, B. gilgiana Irmsch., B. glabricaulis Irmsch., B. gomantongensis Kiew, B. gracilipes Merr., B. grandipetala Irmsch., B. gusilii Rimi, B. guttapilla D.C.Thomas & Ardi, B. hainanensis Chun & F.Chun, B. harauensis M.Hughes, B. hekensis D.C.Thomas, B. heliostrophe Kiew, B. heteroclinis Miq. ex Koord., B. hexaptera Sands, B. hidirii Tawan, Ipor & Meekiong, B. hirsuticarpa C.W.Lin & C.I Peng, B. hirsuticaulis Irmsch., B. hirtitepala S.Julia & Kiew, B. hispidissima Zipp. ex Koord., B. holosericea (Teijsm. & Binn.) Teijsm. & Binn., B. holosericeoides Ardi & D.C.Thomas, B. holttumii Irmsch., B. hooveriana Wiriad., B. hullettii Ridl., B. humboldtiana Gibbs, B. humericola Sands, B. humilicaulis Irmsch., B. ignita C.W.Lin & C.I Peng, B. imbricata Sands, B. imperfecta Irmsch., B. incisa A.DC., B. incompta Kiew, B. inobongensis Kiew, B. inopinata V.S.Guanih, B. inostegia Stapf, B. inostegioides F.Y.Chong, B. insueta D.C.Thomas & Ardi, B. insularum Irmsch., B. iridifolia C.W.Lin & C.I Peng, B. isoptera Dryand. ex Sm., B. isopteroidea King, B. jagorii Warb., B. jamilahana Y.W. Low, Joffre & Ariffin, B. jamilahanuiana S.Julia & Kiew, B. jamiliana Rimi, B. jaranpusangensis Girm., B. jenginensis S.Julia & Kiew, B. jiewhoei Kiew, B. joffrei S.Julia, B. johariana S.Julia & C.Y.Ling, B. jugamensis S.Julia & Kiew, B. julaihiana S.Julia & C.Y.Ling, B. juliasangii Kiew, B. kachak K.G.Pearce, B. kasutensis K.G.Pearce, B. keeana Kiew, B. keithii Kiew, B. kelliana Irmsch., B. kerstingii Irmsch., B. kiamfeei Kiew & S.Julia, B. kibambangensis V.S.Guanih & F.Y.Chong, B. kinabaluensis Sands, B. kinahimiae Rimi, B. kipandiensis S.Julia, Version of Record 299 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) B. koordersii Warb. ex L.B.Sm. & Wassh., B. kuchingensis C.W.Lin & C.I Peng, B. kurakura Tawan, Ipor & Meekiong, B. labiensis (Sands) S.Julia, B. lacera Merr., B. lacustris Irmsch. ex Peekel, B. lagunensis Elmer, B. lailana Kiew & Geri, B. lambii Kiew, B. lambirensis Kiew & S.Julia, B. lamriana Rimi, B. lamxayiana Souvann., B. lancifolia Merr., B. laruei M.Hughes, B. lasioura D.C.Thomas & Ardi, B. latistipula Merr., B. lauterbachii Warb., B. lawii C.W.Lin & C.I Peng, B. layang-layang Kiew, B. lazat Kiew & Reza Azmi, B. ledermannii Irmsch., B. leptantha C.B.Rob., B. leucochlora Sands, B. leucosticta Warb., B. leucotricha Sands, B. lichenora C.W.Lin & C.I Peng, B. linauensis S.Julia, B. littleri Merr., B. loheri Merr., B. lombokensis Girm., B. longibractea Merr., B. longiseta Irmsch., B. longistipula Merr., B. ltahensis Girm., B. lucychongiana S.Julia & Kiew, B. lunatistyla Irmsch., B. macgregorii Merr., B. macintyreana M.Hughes, B. madaiensis Kiew, B. magentifolia Kiew & S.Julia, B. magnicarpa C.W.Lin & C.I Peng, B. malachosticta Sands, B. malindangensis Merr., B. malmquistiana Irmsch., B. mamutensis Sands, B. manuselaensis Ardhaka & Ardi, B. mariaensis Rimi & Simun, B. masarangensis Irmsch., B. matangensis S.Julia & Kiew, B. mearnsii Merr., B. media Merr. & L.M.Perry, B. megacarpa Merr., B. megalantha Merr., B. mekongensis Girm. & Wiriad., B. melanosticta F.Y.Chong & V.S.Guanih, B. melikopia Kiew, B. melinauensis S.Julia & Kiew, B. mendumae M.Hughes, B. mentewangensis Girm., B. meriraiensis Kiew & S.Julia, B. merrittii Merr., B. metallicolor C.W.Lin & C.I Peng, B. mindanaensis Warb., B. minutiflora Sands, B. monantha Warb., B. montis-bismarckii Warb., B. moszkowskii Irmsch., B. multidentata Warb., B. multijugata M.Hughes, B. murudensis Merr., B. murumensis S.Julia & C.Y.Ling, B. mystacina L.B.Sm. & Wassh., B. nagaensis Kiew & S.Julia, B. naumoniensis Irmsch., B. negrosensis Elmer, B. nephrophylla Undaharta & Ardi, B. niahensis K.G.Pearce, B. nix C.W.Lin & C.I Peng, B. nobmanniae D.C.Thomas & Ardi, B. nothobaramensis Joffre, B. novoguineensis Merr. & L.M.Perry, B. oblongata Merr., B. oblongifolia Stapf, B. oligantha Merr., B. otophora Merr. & L.M.Perry, B. oxyura Merr. & L.M.Perry, B. ozotothrix D.C.Thomas, B. padangensis Irmsch., B. palawanensis Merr., B. panayensis Merr., B. paoana Kiew & S.Julia, B. papuana Warb., B. papulifolia S.Julia & C.Y.Ling, B. papyraptera Sands, B. paracauliflora Rimi, C.I Peng & S.M.Ku, B. parvilimba Merr., B. pediophylla Merr. & L.M.Perry, B. peekelii Irmsch., B. pellionoides Y.M.Shui & W.H.Chen, B. pendula Ridl., B. penrissenensis Kiew & S.Julia, B. pentaphragmifolia Ridl., B. peridoticolia Rimi, C.I Peng & C.W.Lin, B. perryae L.B.Sm. & Wassh., B. pinnatifida Merr. & L.M.Perry, B. piring Kiew & S.Julia, B. platyphylla Merr., B. pleioclada Irmsch., B. plieranensis S.Julia & C.Y.Ling, B. polilloensis Tebbitt, B. polyclada C.I Peng, C.W.Lin & Rubite, B. postarii Kiew, B. prasinimarginata S.Julia, B. prionota D.C.Thomas & Ardi, B. promethea Ridl., B. propinqua Ridl., B. pryeriana Ridl., B. pseudolateralis Warb., B. pubescens Ridl., B. punbatuensis Kiew, B. punchak Kiew & S.Julia, B. quercifolia A.DC., B. racemosa Jack, B. rachmatii Tebbitt, B. rambaiensis Kiew, B. rambutan Rimi, B. ramlanii Rimi & Handry, B. ramosii Merr., B. ramosissima Kiew & S.Julia, B. ranaiensis Girm., 300 TAXON 67 (2) • April 2018: 267–323 B. randiana Merr. & L.M.Perry, B. rantemarioensis D.C.Thomas & Ardi, B. renek Rimi, B. repanda Blume, B. retakensis (Sands) Joffre, B. rhodantha Ridl., B. rhodochaeta Kiew & S.Julia, B. rhodoneura S.Julia, B. rhodotricha S.Julia & C.Y.Ling, B. rhombipetala S.Julia & C.Y.Ling, B. rieckei Warb., B. rizalensis Merr., B. roseopunctata Kiew, B. rotundibracteata Kiew, B. rubida Ridl., B. rubrobracteolata S.Julia & C.Y.Ling, B. rubrosetosa Aver., B. ruthiae S.Julia, B. sageaensis Wiriad., B. salomonensis Merr. & L.M.Perry, B. samarensis Merr., B. sanguineopilosa D.C.Thomas & Ardi, B. sarangica Kiew & S.Julia, B. sarasinorum Irmsch., B. sarawakensis Ridl., B. sarmentosa L.B.Sm. & Wassh., B. saxatilis Blume, B. serapatensis Kiew & S.Julia, B. serraticauda Merr. & L.M.Perry, B. serratipetala Irmsch., B. setiamensis S.Julia & Kiew, B. sibutensis Sands, B. siccacaudata J.Door., B. simulans Merr. & L.M.Perry, B. siregarii Ardi & D.C.Thomas, B. sirukitii S.Julia & C.Y.Ling, B. sogerensis Ridl., B. somervillei Hemsl., B. sphenocarpa Irmsch., B. spilotophylla F.Muell., B. stenogyna Sands, B. stevei M.Hughes, B. stichochaete K.G.Pearce, B. stilandra Merr. & L.M.Perry, B. strachwitzii Warb. ex Irmsch., B. strictinervis Irmsch., B. strictipetiolaris Irmsch., B. subelliptica Merr. & L.M.Perry, B. subisensis K.G.Pearce, B. sublongipes Y.M.Shui, B. subprostrata Merr., B. subtruncata Merr., B. suffrutescens Merr. & L.M.Perry, B. superciliaris C.W.Lin & C.I Peng, B. sympodialis Irmsch., B. tafaensis Merr. & L.M.Perry, B. taniana V.S.Guanih, B. tawaensis Merr., B. tebiang S.Julia & Kiew, B. temburongensis Sands, B. tenericaulis Ridl., B. tenuissima S.Julia & C.Y.Ling, B. timorensis (Miq.) Golding & Kareg., B. tindan Rimi & Kimahim, B. tinjanii S.Julia, B. tomaniensis Rimi, B. torajana D.C.Thomas & Ardi, B. torricellensis Warb., B. triangularis Kiew & C.Y.Ling, B. ubahribuensis S.Julia & Kiew, B. umbratica S.Julia, B. urdanetensis Elmer, B. urunensis Kiew, B. vaccinioides Sands, B. vanderentii Rossiti, B. vanderwateri Ridl., B. varipeltata D.C.Thomas, B. vermeulenii D.C.Thomas, B. vuijckii Koord., B. vulgare S.Julia & Kiew, B. wallacei C.W.Lin & C.I Peng, B. walteriana Irmsch., B. warburgii K.Schum. & Lauterb., B. wariana Irmsch., B. watuwilensis Girm., B. weberi Merr., B. weigallii Hemsl., B. wenzelii Merr., B. wrayi Hemsl., B. xiphophylla Irmsch., B. xiphophylloides Kiew, B. yiii Kiew & S.Julia, B. zamboangensis Merr. Distribution. – ASIA: Brunei, China (Hainan, Yunnan), East Timor, Indonesia (Java, Kalimantan, Lesser Sunda Islands, Maluku, Sulawesi, Sumatra, West Papua), Malaysia (Peninsular Malaysia, Sabah, Sarawak), Papua New Guinea, Philippines, Solomon Islands, Thailand, Vietnam. Taxonomic notes. – We recognise a circumscription of B. sect. Petermannia based on Doorenbos & al. (1998) and Kiew & al. (2015) with some modifications. We recommend further research to determine which species belong in B. sect. Bracteibegonia, and the relationships between the “Begonia amphioxus clade” and the “Petermannia-Symbegonia clade”. The range of morphology present in B. sect. Petermannia is much more apparent following recent research on Borneo by Kiew & al. (2015), and it may be that the section can be divided in more manageable units in the future. However for the time being we recognise a large and paraphyletic concept of the section. Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Doorenbos & al. (1998) noted the unusual peltate, lanceolate leaves of B. amphioxus and its dichasial inflorescences with solitary female flowers. These characters are not shared by the other species within the “Begonia amphioxus clade” and we are unable to separate this group from the “PetermanniaSymbegonia clade” by morphology. Species in both clades share a characteristic inflorescence morphology being protandrous and with basal female flowers; and perforate anther endothecium cell wall thickening. Furthermore, species in both clades are resolved together (although with poor support) in nrDNA and mtDNA phylogenies (see review in Thomas & al., 2011). Thomas & al. (2011) discussed a scenario whereby transfer of a foreign plastid into Bornean B. sect. Petermannia followed by diversification within Borneo would account for the observed conflict among phylogenies and with morphology. We agree this scenario is likely, but note the presence of non-Bornean B. cucphuongensis (Vietnam) and B. hainanensis Chun & F.Chun (China: Hainan) in the “Begonia amphioxus clade”. These species may represent a back-colonisation from Borneo or be relatives of the plants from which chloroplast capture occurred. Begonia saxatilis is moved here, although initially described in the genus Diploclinium, it has a typical B. sect. Petermannia inflorescence with female flowers at the base. Begonia warburgii was treated by Doorenbos & al. (1998) as only a tentative member of B. sect. Petermannia. We assign this species to the section with confidence because of the similarity between this species and B. bipinnatifida. The Bornean B. piring has paired female flowers at the base of a male-flower cyme and so fits very comfortably in B. sect. Petermannia, and it is moved here from B. sect. Diploclinium. Begonia rubrosetosa is also moved from B. sect. Diploclinium, as it lacks tubers and is likely allied to other caulescent species from Vietnam in B. sect. Petermannia. Begonia beccarii was previously unplaced (Hughes, 2008), and is moved to B. sect. Petermannia as it is very close to, if not synonymous with, B. promethea Ridl., which is in the same section. Begonia sect. Pilderia (Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 141. 1859 ≡ Pilderia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 127. 1854 – Type (designated by Swart, ING Card 10693. 1960): Pilderia urticifolia Klotzsch = Begonia buddleiifolia A.DC. — [Fig. 17A] Species list. – 6 species: B. buddleiifolia A.DC., B. glandulifera Griseb., B. humillima L.B.Sm. & Wassh., B. jenmanii Tutin, B. mariannensis Wassh. & T.McClellan, B. tepuiensis Moonlight & Jara Distribution. – AMERICAS: Colombia, Ecuador, Guyana, Peru, Trinidad-Tobago, Venezuela. Taxonomic notes. – We refer to the description in Moonlight & Jara-Muñoz (2017) and follow the circumscription therein. Begonia sect. Platycentrum (Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 134. 1859 ≡ Platycentrum Klotzsch in Abh. Königl. Akad. Wiss. Berlin 1854: 243. 1855 ≡ Begonia subg. Platycentrum (Klotzsch) C.B.Clarke in Bot. J. Linn. Soc. 18: 118. 1881 – Type: Platycentrum xanthiana (Hook.) Klotzsch ≡ B. xanthinum Hook. — [Fig. 14B] = Begonia sect. Elasticae C.B.Clarke in Bot. J. Linn. Soc. 18: 115. 1881 – Type (designated here): B. episcopalis C.B. Clarke = Begonia griffithiana (A.DC.) Warb. = Begonia sect. Euplatycentrum C.B.Clarke in Bot. J. Linn. Soc. 18: 115. 1881, nom. illeg. – Type: Begonia xanthina Hook. = Begonia sect. Monolobium T.C.Ku in Gu & al., Fl. China 13: 207. 2007 – Type: Begonia wutaiana C.I Peng & Y.K.Chen. = Begonia sect. Sphenanthera (Hassk.) Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 140. 1894 ≡ Sphenanthera Hassk. in Verslagen Meded. Afd. Natuurk. Kon. Akad. Wetensch. 4: 139. 1855 ≡ Casparya sect. Sphenanthera (Hassk.) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 118. 1859, syn. nov. – Type: Sphenanthera robusta (Blume) Hassk. ex Klotzsch ≡ Begonia robusta Blume. = Casparya sect. Holoclinium A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 118. 1859 ≡ Begonia sect. Holoclinium (A.DC.) Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 142. 1894, syn. nov. – Type: Casparya trisulcata A.DC. = Begonia longifolia Blume. = Casparya sect. Polyschisma A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 119. 1859 ≡ Begonia sect. Polyschisma (A.DC.) Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 149. 1894, syn. nov. – Type: Casparya crassicaulis A.DC. = Begonia multangula Blume. = Mezierea sect. Monopteron A.DC., Prodr. 15(1): 406. 1864 ≡ Begonia sect. Monopteron (A.DC.) Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 142. 1894, syn. nov. – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 5. 1972): Mezierea nepalensis A.DC. ≡ Begonia nelapensis (A.DC.) Warb. Species list. – 171 species: B. ×chungii C.I Peng & S.M.Ku, B. ×taipeiensis C.I Peng, B. abdullahpieei Kiew, B. aborensis Dunn*, B. acetosella Craib*, B. adenostegia Stapf, B. aequilateralis Irmsch., B. algaia L.B.Sm. & Wassh., B. alpina L.B.Sm. & Wassh., B. altissima Ridl., B. annulata K.Koch, B. aptera Blume*, B. areolata Miq., B. augustinei Hemsl., B. austrotaiwanensis Y.Chen & C.I Peng, B. balansana Gagnep.*, B. baliensis Girm.*, B. baviensis Gagnep., B. beddomei Hook., B. bouffordii C.I Peng, B. brevipedunculata Y.M.Shui, B. brevisetulosa C.Y.Wu, B. burkillii Dunn*, B. caobangensis C.I Peng & C.W.Lin, B. cathayana Hemsl., B. cathcartii Hook.f., B. ceratocarpa S.H.Huang & Y.M.Shui*, B. chishuiensis T.C.Ku, B. chitoensis T.S.Lui & M.J.Lai, B. chuyunshanensis C.I Peng & Y.K.Chen, B. chlorocarpa Irmsch. ex Sands*, B. circumlobata Hance, B. coelocentroides Y.M.Shui & Z.D.Wei, B. coptidifolia H.G.Ye, F.G.Wang, Y.S.Ye & C.I Peng, B. crocea C.I Peng, B. cucurbitifolia C.Y.Wu, B. daweishanensis S.H.Huang & Y.M.Shui, B. decora Stapf, B. deliciosa Linden ex Fotsch, B. diadema Linden ex Rodigas, B. dielseiana E.Pritz. ex Diels, B. difformis (Irmsch.) W.C. Leong, C.I Peng & K-F.Chung, B. digyna Irmsch., B. discrepans Irmsch., B. dryadis Irmsch., B. duclouxii Gagnep., B. edulis H.Lév., B. emeiensis C.M.Hu ex C.Y.Wu & T.C.Ku, B. flaviflora H.Hara, B. formosana (Hayata) Masam., B. forrestii Irmsch., B. foveolata Irmsch., B. fraseri Kiew, B. gagnepainiana Irmsch., B. goniotis C.B.Clarke, Version of Record 301 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) B. griffithiana (A.DC.) Warb., B. gulinqingensis S.H.Huang & Y.M.Shui, B. hahiepiana H.Q.Nguyen & Tebbitt*, B. halconensis Merr.*, B. handelii Irmsch.*, B. hatacoa Buch.-Ham. Ex D.Don, B. hayamiana Nob.Tanaka*, B. hayatae Gagnep.*, B. hekouensis S.H.Huang, B. hemsleyana Hook.f., B. herveyana King, B. hongkongensis F.W.Xing, B. houttuynioides T.T.Yu & W.P.Fang, B. iridescens Dunn, B. jinyunensis C.I Peng, B.Ding & Q.Wang, B. kachinensis Nob.Tanaka*, B. khaophanomensis Phutthai & M.Hughes, B. kingdon-wardii Tebbitt*, B. klossii Ridl., B. koelzii R.Camfield, B. koksunii Kiew, B. lacerata Irmsch., B. laminariae Irmsch., B. lancangensis S.H.Huang*, B. lecomtei Gagnep., B. lempuyangensis Girm.*, B. leuserensis M.Hughes, B. limprichtii Irmsch., B. lipingensis Irmsch., B. longanensis C.Y.Wu, B. longialata K.Y.Guan & D.K. Tian, B. longicarpa K.Y.Guan & D.K.Tian*, B. longicaulis Ridl., B. longiciliata C.Y.Wu, B. longifolia Blume*, B. lukuana Y.C.Liu & C.H.Ou*, B. macrotoma Irmsch., B. maguanensis S.H.Huang & Y.M.Shui, B. manhaoensis S.H.Huang & Y.M.Shui, B. maxwelliana King, B. megalophyllaria C.Y.Wu, B. megaptera A.DC., B. menglianensis Y.Y.Qian, B. mengtzeana Irmsch., B. multangula Blume*, B. multibracteata Girm.*, B. myanmarica C.I Peng & Y.D.Kim, B. nantoensis M.J.Lai & N.J.Chung, B. nepalensis (A.DC.) Warb., B. nuwakotensis S.Rajbhandary, B. obovoidea Craib*, B. obversa C.B.Clarke, B. oligocarpa A.DC. ex Koord., B. oreodoxa Chun & F.Chun ex C.Y.Wu & T.C.Ku, B. oreophila Kiew, B. palmata D.Don, B. paucilobata C.Y.Wu, B. paupercula King, B. pavonina Ridl., B. pedatifida H.Lév., B. peii C.Y.Wu, B. perakensis King, B. pinglinensis C.I Peng, B. praetermissa Kiew, B. prolixa Craib, B. pseudoscottii Girm.*, B. psilophylla Irmsch., B. pulchrifolia D.K.Tian & C.H.Li, B. purpureofolia S.H.Huang & Y.M.Shui, B. reflexisquamosa C.Y.Wu, B. repenticaulis Irmsch., B. rex Putz., B. rheifolia Irmsch., B. rhoephila Ridl., B. rhyacophila Kiew, B. robusta Blume*, B. rockii Irmsch., B. roxburghii (Miq.) A.DC.*, B. rubinea H-Z.Li, B. rubropunctata S.H.Huang & Y.M.Shui, B. sandalifolia C.B.Clarke, B. scitifolia Irmsch., B. scortechinii King, B. scottii Tebbitt*, B. shilendrae Rekha Morris & P.D.Mcillan, B. siamensis Gagnep., B. sikkimensis A.DC., B. silletensis (A.DC.) C.B.Clarke*, B. sizemoreae Kiew, B. smithiana T.T.Yu, B. sphenantheroides C.I Peng, B. subhowii S.H.Huang, B. taiwaniana Hayata, B. taligera S.Rajbhandary, B. tamdaoensis C.I Peng, B. tampinica Burkill ex Irmsch., B. tarokoensis M.J.Lai, B. tengchiana C.I Peng & Y.K.Chen, B. tessaricarpa C.B.Clarke*, B. tetralobata Y.M.Shui, B. teysmanniana (Miq.) Tebbitt, B. thomsonii A.DC., B. togashii Nob.Tanaka & C.I Peng, B. truncatiloba Irmsch., B. tsaii Irmsch., B. tsoongii C.Y.Wu, B. tuberculosa Girm., B. vallicola Kiew, B. venusta King, B. versicolor Irmsch., B. villifolia Irmsch., B. wui-senioris C.I Peng, B. wutaiana C.I Peng & Y.K.Chen, B. wyepingiana Kiew, B. xanthina Hook., B. yingjiangensis S.H.Huang. — [*The “Sphenanthera group”] Species whose membership is doubtful. – 6 species: B. amabilis Linden, B. dux C.B.Clarke, B. gungshanensis C.Y.Wu, B. langbianensis Baker f., B. polytricha C.Y.Wu, B. setifolia Irmsch. Distribution. – ASIA: Bhután, China (Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Heilongjiang, Hong 302 TAXON 67 (2) • April 2018: 267–323 Kong, Hubei, Jiangsi, Sichuan, Tibet, Yunnan), India (Arunachal Pradesh, Assam, Darjeeling, Manipur, Meghalaya, Mizoram, Nagaland, Sikkim, Tripura), Indonesia (Bali, Java, Lesser Sunda Islands, Maluku, Sulawesi, Sumatra), Laos, Malaysia (Peninsular Malaysia, Sabah), Myanmar, Nepal, Philippines, Taiwan, Thailand, Vietnam. Taxonomic notes. – Maintaining B. sect. Platycentrum separate from B. sect. Sphenanthera (Hughes & Girmansyah, 2011) is no longer tenable in the face of molecular data from both chloroplast and nuclear data showing the latter to be polyphyletic (Tebbitt & al., 2006; Thomas & al., 2011; this study). Although the fruit type of the vast majority of species in B. sect. Platycentrum (2-locular splash-cup, dehiscent, with one enlarged wing) is instantly diagnostic, the fruits of species in B. sect. Sphenanthera vary in locule number (3–8) and shape, being united only by having a fleshy fruit wall. The type of B. sect. Sphenanthera, B. robusta Blume, has a fruit which is intermediate between the two sections, having one wing enlarged, but with fleshy walls and three locules; Doorenbos & al. (1998) state “if it did not have a 3-locular ovary, one would not hesitate to put B. robusta in Platycentrum”. The newly enlarged B. sect. Platycentrum is characterised by a rhizomatous habit (with a few exceptions; e.g., B. longifolia Blume and allies, which are caulescent, but have a rhizomatous stem base), highly convolute stigmas, stamens on a column, and anthers dehisching through long lateral slits with an extended connective. We distinguish the members of the section with fleshy fruit walls in the informally named “Sphenanthera group” of B. sect. Platycentrum. Begonia sect. Monopteron shares with B. sect. Platycentrum its two locular ovaries with bifid placentae and we replicate Rajbhandary & al. (2011) in placing the section within the B. sect. Platycentrum. The two sections were differentiated by the presence or absence of two lateral wings on the fruits, which otherwise share an enlarged dorsal wing. We synonymise B. sect. Monopteron with B. sect. Platycentrum herein. Begonia halconensis Merr. from Mindoro in the Philippines is moved here from B. sect. Petermannia based on its very close resemblance to the widespread B. longifolia, and phylogenetic evidence in Tebbitt & al. (2006). We refer to the descriptions in Doorenbos & al. (1998) of B. sect. Platycentrum (p. 155) and B. sect. Sphenanthera (p. 194). Begonia sect. Pritzelia (Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 137. 1859 ≡ Pritzelia Klotzsch in Ber. Bekanntm. Verh. Königl.Preuss. Akad. Wiss. Berlin 1854: 126. 1854 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 6. 1972): Pritzelia fischeri (Otto & A.Dietr.) Klotzsch ≡ Begonia dietrichiana Irmsch. — [Fig. 17B] = Ewaldia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 123. 1854 ≡ Begonia sect. Ewaldia (Klotzsch) A.DC., Prodr. 15(1): 372. 1864 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 4. 1972): Ewaldia lobata (Schott) Klotzsch = Begonia rufa Thunb. = Gurltia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 123. 1854 – Type: Gurltia meyeri (Otto & A.Dietr.) Klotzsch = Begonia tomentosa Schott. Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) = Scheidweileria Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 123. 1854 ≡ Begonia sect. Scheidweileria (Klotzsch) A.DC., Prodr. 15(1): 370. 1864, syn. nov. – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 7. 1972): Scheidweilaria muricata (Scheidw.) Klotzsch = Begonia pentaphylla Walp. = Saueria Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 122. 1854 ≡ Begonia sect. Saueria A.DC., Prodr. 15(1): 288. 1864 – Type: Saueria sulcata (Scheidw.) Klotzsch = Begonia dichotoma Jacq. = Trendelenburgia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 123. 1854 ≡ Begonia sect. Trendelenburgia (Klotzsch) A.DC. in Martius, Fl. Bras. 4(1): 377. 1861, syn. nov. – Type: Trendelenburgia fruticosa Klotzsch ≡ Begonia fruticosa (Klotzsch) A.DC. = Steineria Klotzsch in Abh. Königl. Akad. Wiss. Berlin 1854: 243. 1855 ≡ Begonia sect. Steineria A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 288. 1859 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 6. 1972): Steineria ferruginea Klotzsch = Begonia hookeriana Gardner. = Nephromischus Klotzsch in Braun & al., Append. Gen. Sp. Hort. Berol. 1855: 1. 1855 vel 1856 – Type: Nephromischus rutilans (Van Houtte) Klotzsch ≡ Begonia rutilans Van Houtte. = Begonia sect. Philippomartia A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 139. 1859 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 2. 1972): Begonia neglecta A.DC. = Begonia sect. Plurilobarica A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 136. 1859 – Type: Begonia schlumbergeriana A.DC. = Begonia sect. Dasystyles A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 143. 1859 – Type: Begonia frigida hort. ex A.DC. = Begonia sect. Bradea Toledo in Arq. Bot. Estado Sao Paulo 2(3): 61. 1946 – Type: Begonia rufoserica Toledo. = Begonia sect. Gehrtia Toledo in Arq. Bot. Estado Sao Paulo 2(3): 62. 1946 – Type: Begonia adiantiformis Toledo = Begonia itatiensis Irmsch. ex Brade. Species list. – 147 species: B. acetosa Vell., B. acida Vell., B. altamiroi Brade, B. angularis Raddi, B. angulata Vell., B. apparicioi Brade, B. ×antonietae Brade, B. arborescens Raddi, B. bahiensis A.DC., B. besleriifolia Schott, B. bidentata Raddi, B. biguassuensis Brade, B. bonitoensis Brade, B. boraceiensis Handro, B. boucheana (Klotzsch) A.DC., B. bradei Irmsch., B. brevilobata Irmsch., B. bufoderma L.B.Sm. & Wassh., B. callosa L.Kollmann, B. calvescens (Brade ex L.B.Sm. & R.C.Sm.) E.L.Jacques & Mamede, B. camposportoana Brade, B. capanemae Brade, B. caraguatatubensis Brade, B. cariocana Brade ex L.B.Sm. & Wassh., B. catharinensis Brade, B. collaris Brade, B. concinna Schott, B. cordata Vell., B. cornitepala Irmsch., B. crispula Brade, B. curtii L.B.Sm. & B.G.Schub., B. declinata Vell., B. densifolia Irmsch., B. dentatiloba A.DC., B. dichotoma Jacq., B. dietrichiana Irmsch., B. digitata Raddi, B. echinosepala Regel, B. elianeae Gregório & J.A.S. Costa, B. epipsila Brade, B. fabulosa L.B.Sm. & Wassh., B. fernando-costae Irmsch., B. fiebrigii C.DC., B. fluminensis Brade, B. forgetiana Hemsl., B. friburgensis Brade, B. frigida hort. ex A.DC., B. fruticosa (Klotzsch) A.DC., B. fuscocaulis Brade, B. gardneri A.DC., B. garuvae L.B.Sm. & R.C.Sm., B. gehrtii Irmsch., B. hammoniae Irmsch., B. handroi Brade, B. heringeri Brade, B. hilariana A.DC., B. hispida Schott, B. hookeriana Gardner, B. hugelii (Klotzsch) A.DC., B. incisoserrata A.DC., B. inculta Irmsch., B. insularis Brade, B. isopterocarpa Irmsch., B. itaguassuensis Brade, B. itatiaiensis Brade, B. itatinensis Irmsch. ex Brade, B. itupavensis Brade, B. jocelinoi Brade, B. juliana Loefgr. ex Irmsch., B. larorum L.B.Sm. & Wassh., B. lineolata Brade, B. listada L.B.Sm. & Wassh., B. longibarbata Brade, B. lossiae L.Kollmann, B. luxurians Scheidw., B. magdalenensis Brade, B. matogrossensis L.B.Sm. ex S.F.Smith & Wassh., B. mattos-silvae L.B.Sm. ex S.F.Smith & Wassh., B. membranacea A.DC., B. metallica W.G.Sm., B. moysesii Brade, B. neglecta A.DC., B. neocomensium A.DC., B. nuda Irmsch., B. obscura Brade, B. occhionii Brade, B. odeteiantha Handro, B. olsoniae L.B.Sm. & B.G.Schub., B. organensis Brade, B. otophylla L.B.Sm. & B.G.Schub., B. paganuccii Gregório & J.A.S.Costa, B. paleata Schott ex A.DC., B. paranaensis Brade, B. parviflora Poepp. & Endl., B. parvifolia Schott, B. parvistipulata Irmsch., B. paulensis A.DC., B. peltifolia Schott, B. pentaphylla Walp., B. peristegia Stapf, B. pickelii Irmsch., B. pilgeriana Irmsch., B. pinheironis L.B.Sm. ex S.F.Smith & Wassh., B. piresiana Handro, B. pluvialis L.B.Sm. ex S.F.Smith & Wassh., B. polyandra Irmsch., B. princeps A.DC., B. pulchella Raddi, B. ramentacea Paxton, B. reniformis Dryand., B. riedelii A.DC., B. rigida Linden ex Regel, B. rotunda Vell., B. rubropilosa A.DC., B. rufa Thunb., B. rufoserica Toledo, B. rupium Irmsch., B. russelliana L.B.Sm. ex S.F.Smith & Wassh., B. rutilans hort.Van-Houtte ex A.DC., B. salesopolensis S.J.Gomes & Mamede, B. sanguinea Raddi, B. sarmentacea hort. ex Brilmayer, B. saxifraga A.DC., B. scabrida A.DC., B. scabridoidea L.B.Sm. & Wassh., B. scharffiana Regel, B. scharffii Hook.f., B. schlumbergeriana Lem., B. semidigitata Brade, B. serranegrae L.B.Sm. ex S.F.Smith & Wassh., B. solimutata L.B.Sm. & Wassh., B. solitudinis Brade, B. spinibarbis Irmsch., B. squamipes Irmsch., B. stenolepis L.B.Sm. & R.C.Sm., B. stenophylla A.DC., B. subacida Irmsch., B. sylvatica Meisn. ex A.DC., B. sylvestris A.DC., B. toledoana Handro, B. tomentosa Schott, B. valdensium A.DC., B. valida Goebel, B. venosa Skan ex Hook.f., B. verruculosa L.B.Sm., B. vicina Irmsch., B. windischii L.B.Sm. ex S.F.Smith & Wassh. Species whose membership is doubtful. – 1: B. erecta Vell. Distribution. – AMERICAS: Argentina (Misiones), Bolivia, Brazil (Alagoas; Bahia, Espírito Santo, Minas Gerais, Mato Grosso, Paraná, Pernambuco, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo), Colombia, Costa Rica, Ecuador, Panamá, Paraguay, Peru, Venezuela. Taxonomic notes. – We place all sampled rhizomatous species of B. sect. Pritzelia within the “core Pritzelia clade” (e.g., B. olsoniae, B. fluminensis, B. moysesii) except B. acetosa and B. subacida. We have been unable to identify morphological characters that separate these two species from B. sect. Pritzelia to our satisfaction. We retain all rhizomatous members of B. sect. Pritzelia within the section and await further study of these species. Version of Record 303 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) We further resolve B. sect. Scheidweilaria and the monotypic B. sect. Trendelenbergia within this clade, both of which we reduce to synonyms of B. sect Pritzelia. The latter was maintained as separate from B. sect. Pritzelia by Doorenbos & al. (1998) due to the types’ supposed lack of cystoliths. Herbarium work (P.W. Moonlight, unpub. data) confirms the presence of cystoliths in B. fruticosa. The similarity between B. sect. Scheidweilaria and B. sect. Pritzelia was noted upon by Doorenbos & al. (1998), the former differing primarily in their compound leaves (except B. parviflora) and anthers that are shorter than the filaments. Begonia venosa is resolved in the “core Pritzelia clade” as sister to the very similar Begonia curtii L.B.Sm. & B.G.Schub. These two species share the cystoliths and bifid placentae (Moonlight, unpublished data) characteristic of B. sect. Pritzelia and we reclassify B. venosa in this section. Begonia matogrossensis was tentatively described by its authors in B. sect. Ruizopavonia on account of its superficial similarity to B. holtonis A.DC. (Smith & Wasshausen, 1999), and B. besleriifolia and B. elianeae were both described in the same section. The female flowers of B. matogrossensis are unknown, but the habit and staminate flower morphology of all three species are consistent with B. sect. Pritzelia. The pistillate flower morphology and fruit morphology of B. besleriifolia and B. elianeae are also both consistent with the section. Given their Brazilian distribution (B. sect. Ruizopavonia is an Andean section) and the presence of cystoliths in the leaf laminae of B. matogrossensis and B. elianeae (characteristic of B. sect. Pritzelia but absent in B. sect. Ruizopavonia), we assign all three species to B. sect. Pritzelia. We have been unable to confirm the presence of cystoliths in B. elianeae but they are expected to be present. We refer to the description in Doorenbos & al. (1998: 159), which fits our new amended circumscription. Like Doorenbos & al. (1998), we treat B. erecta Vell as doubtfully assigned to section as it is only known from an illustration. Begonia sect. Putzeysia (Klotzsch) A.DC., Prodr. 15(1): 314. 1864 ≡ Putzeysia Klotzsch in Abh. Königl. Akad. Wiss. Berlin 1854: 254. 1855 – Type: Putzeysia gemmipara (Hook.f. & Thomson) Klotzsch ≡ Begonia gemmipara Hook.f. & Thomson. — [Fig. 14C] Species list. – 1 species: B. gemmipara Hook.f. & Thomson. Distribution. – ASIA: Bhutan, India (Darjeeling, Sikkim), Nepal. Taxonomic notes. – We agree with Doorenbos & al. (1998) in noting the similarity of this species to those of B. sect. Diploclinium, although it differs in its seeds having adpressed papillae, and in having unique clusters of axillary tubercles. We retain B. sect. Putzeysia as a monotypic section and refer to the description in Doorenbos & al. (1998: 164). Begonia sect. Quadrilobaria A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 130. 1859 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 6. 1972): Begonia nossibea A.DC. — [Fig. 11K] Species list. – 20 species: B. anjuanensis Humb. ex Keraudren & Bosser, B. ankaranensis Humb. ex Keraudren 304 TAXON 67 (2) • April 2018: 267–323 & Bosser, B. antsingyensis Humb. ex Keraudren & Bosser, B. antsiranensis Aymonin & Bosser, B. bagotiana Humb. ex Keraudren & Bosser, B. bekopakensis Aymonin & Bosser, B. bernieri A.DC., B. boiviniana A.DC., B. decaryana Humb. ex Keraudren & Bosser, B. francoisii Guillaumin, B. goudotii A.DC., B. heteropoda Baker, B. isalensis Humb. ex Keraudren & Bosser, B. leandrii Humb. ex Keraudren & Bosser, B. mananjebensis Humb. ex Keraudren & Bosser, B. neoperrieri Humb. ex Keraudren & Bosser, B. nossibea A.DC., B. sambiranensis Humb. ex Keraudren & Bosser, B. trullifolia Guillaumin, B. tsaratananensis Aymonin & Bosser. Distribution. – AFRICA: Comoros, Madagascar. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 165) and follow the circumscription therein with the addition of B. trullifolia (unassigned by Doorenbos & al., 1998) because of its similarity to B. goudotii. Begonia sect. Quadriperigonia Ziesenh. in Begonian 35: 257. 1968 – Type: Begonia abaculoides Ziesenh. = Begonia boisseri A.DC. — [Fig. 17C] = Begonia (sect. Begoniastrum) subsect. Euknesebeckia Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 149. 1894 – Type: Begonia monoptera Link & Otto = Begonia balmisiana Bálmis. Species list. – 19 species: B. angustiloba A.DC., B. anodifolia A.DC., B. balmisiana Bálmis, B. biserrata Lindl., B. boissieri A.DC., B. bulbillifera Link & Otto, B. dealbata Liebm., B. fusibulba C.DC., B. gracilior Burt-Utley & McVaugh, B. gracilis Kunth, B. macdanielsii Standl., B. ornithocarpa Standl., B. palmeri S.Watson, B. pedata Liebm., B. portillana S.Watson, B. rhodochlamys L.B.Sm. & B.G.Schub., B. sandtii Houghton ex Ziesenh., B. tapatia Burt-Utley & McVaugh, B. uruapensis Sessé & Moç. Species whose membership is doubtful. – 1 species: B. racemiflora Ortiges ex C.Chev. Distribution. – AMERICAS: El Salvador, Guatemala, Honduras, Mexico (Aguascalientes, Baja California, Chiapas, Chihuahua, Colima, Durango, Guanajuato, Guerrero, Hidalgo, Jalisco, México Distrito Federal, México State, Michoacán, Morelos, Nayarit, Nuevo León, Oaxaca, Puebla, Querétaro, San Luis Potosí, Sinaloa, Sonora, Tabasco, Tamaulipas, Yucatán, Zacatecas). Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 167) and follow the circumscription therein. Begonia sect. Reichenheimia (Klotzsch) A.DC. Prodr. 15(1): 385. 1864 ≡ Reichenheimia Klotzsch in Abh. Königl. Akad. Wiss. Berlin 1854: 174. 1855 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 7. 1972): Reichenheimia thwaitesii (Hook.) Klotzsch ≡ Begonia thwaitesii Hook. — [Fig. 14D] = Mitscherlichia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 124. 1854 ≡ Begonia sect. Mitscherlichia (Klotzsch) A.DC., Prodr. 15(1): 389. 1864 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 7. 1972): Begonia albo-coccinea Hook. = Begonia subg. Aschisma C.B.Clarke in Bot. J. Linn. Soc. Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) 18: 117. 1881 – Type (designated here): Begonia fibrosa C.B.Clarke. = Begonia sect. Uniplacentales C.B.Clarke in Hooker, Fl. Brit. India 2: 652. 1879 – Type (designated here): Begonia fibrosa C.B.Clarke. Species list. – 20 species: B. albococcinea Hook., B. cardiophora Irmsch., B. chingii Irmsch., B. fibrosa C.B.Clarke, B. floccifera Bedd., B. harmandii Gagnep., B. henryi Hemsl., B. hymenophylla Gagnep., B. intermixta Irmsch., B. morelii Irmsch. ex Kareg., B. nivea Parish ex Kurz, B. nymphaeifolia T.T.Yu, B. parvula H.Lév. & Vaniot, B. phrixophylla Blatt. & McCann, B. pierrei Gagnep., B. pumilio Irmsch., B. subpeltata Wight, B. tenera Dryand., B. thwaitesii Hook., B. trichocarpa Dalzell. Distribution. – ASIA: Cambodia, China (Guangxi, Guizhou, Hainan, Yunnan), India (Karnataka, Nagaland), Laos, Myanmar, Sri Lanka, Thailand, Vietnam. Taxonomic notes. – This section is much reduced from its earlier concept, following the description of B. sect Jackia which now contains many of the species hitherto included in it. It likely represents a phylogenetically disparate assemblage of species united by having a tuberous habit and entire placentae. We await further study of the affinities of individual species and anticipate further phylogenetic work may result in the continued reduction of B. sect. Reichenheimia. Begonia sect. Ridleyella Irmsch. in Mitt. Inst. Allg. Bot. Hamburg 8: 105. 1929 – Type: Begonia eiromischa Ridl. — [Fig. 14E] Description. – Acaulescent, rhizomatous herbs, perennial. Stipules persistent. Leaves alternate, peltate or basifixed (B. padawanensis), blade subsymmetrical, veins peltate to palmate-pinnate, raised above. Inflorescence axillary, few flowered, cymose, bracts deciduous or persistent. Male flowers: with 3 or 4 free perianth segments; androecium zygomorphic, stamens 25–35, free, anthers obovate, dehiscing via lateral slits, connective retuse. Female flowers: bracteoles absent or present at the base of the ovary, with 3 to 5 free perianth segments; ovary and fruit with 3 wings, wings equal or subequal, 3-locular, placentas axile, entire, bearing ovules on both surfaces; styles 3, free or united at the base, bifid, stigmatic papillae in a spirally twisted band. Fruit a capsule. Species list. – 7 species: B. burttii Kiew & S.Julia, B. eiromischa Ridl., B. kingiana Irmsch., B. padawanensis C.W.Lin & C.I Peng, B. payung Kiew & S.Julia, B. serianensis C.W.Lin & C.I Peng, B. speluncae Ridl. Distribution. – ASIA: Malaysia (Peninsular Malaysia, Sarawak). Taxonomic notes. – In the phylogeny we resolve B. speluncae as sister to B. kingiana, and the two species share a creeping, rhizomatous habit; peltate leaves; prominent venation on the leaf upper surfaces; four tepals in the male flower and five tepals in the female flower; asymmetric androecium; and stamens with a retuse apex. These characteristics are shared by B. burttii, B. padawanensis, B. payung and B. serianensis (although B. padawanensis has basifixed leaves) from Sarawak. These species differ however from B. kingiana in their tri- rather than bi-locular ovaries. We transfer these four species to B. sect. Ridleyella. The recircumscribed section is morphologically most similar to B. sect. Jackia, sharing the rhizomatous habit of that section, but distinguished by five tepals in the female flower; asymmetric androecium and retuse stamens. All species assigned to B. sect. Ridleyella share a preference for limestone habitats. We agree with Doorenbos & al. (1998) in concluding the assignment of B. pumila to the section is doubtful, and we move that species to B. sect. Parvibegonia. Begonia sect. Rossmannia A.DC., Prodr. 15(1): 333. 1864 ≡ Rossmannia Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 125. 1854 – Type: Rossmannia repens Klotzsch = Begonia rossmanniae A.DC. — [Fig. 17D] Species list. – 1 species: B. rossmanniae A.DC. Distribution. – AMERICAS: Brazil (Acre), Colombia, Ecuador, Peru. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 174), follow the circumscription therein, and refer to the discussion in Moonlight & Jara-Muñoz (2017) for comparisons with the closely related B. sect. Pilderia. Begonia sect. Rostrobegonia Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 141. 1894 – Type: Begonia rostrata Welw. ex Hook.f. — [Fig. 11L] Species list. – 9 species: B. bequaertii Robyns & Lawalrée, B. engleri Gilg, B. johnstonii Oliv. ex Hook.f., B. nyassensis Irmsch., B. rostrata Welw. ex Hook.f., B. rumpiensis Kupicha, B. schliebenii Irmsch., B. sonderiana Irmsch., B. wollastonii Baker f. Distribution. – AFRICA: Angola, Democratic Republic of the Congo, Ghana, Ivory Coast, Kenya, Liberia, Malawi, Mali, Mozambique, Nigeria, Sierra Leone, South Africa, Tanzania, Togo, Uganda, Zimbabwe. Taxonomic notes. – Begonia sect. Rostrobegonia and B. sect. Sexalaria differ in their style morphology and chromosome number thus we retain them as separate sections herein. We refer to the description in Doorenbos & al. (1998: 176) and follow the circumscription therein. Begonia sect. Ruizopavonia A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 139. 1859 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 7. 1972): Begonia alnifolia A.DC. — [Fig. 17E] Species list. – 26 species: B. alnifolia A.DC., B. boreoharlingii Tebbitt & Moonlight, B. botryoides Moonlight & Tebbitt*, B. carpinifolia Liebm., B. consobrina Irmsch., B. convallariodora C.DC., B. cooperi C.DC., B. cuatrecasana L.B.Sm. & B.G.Schub., B. cymbalifera L.B.Sm. & B.G.Schub., B. dugandiana L.B.Sm. & B.G.Schub., B. estrellensis C.DC., B. extensa L.B.Sm. & B.G.Schub., B. glauca (Klotzsch) Ruiz & Pav. ex A.DC., B. glaucoides Irmsch., B. harlingii L.B.Sm. & Wassh.*, B. juninensis Irmsch., B. lignescens C.V.Morton, B. peruviana A.DC., B. pilosella Irmsch., B. prionophylla Irmsch., B. rubiginosipes Irmsch., B. seemanniana A.DC., B. tiliifolia C.DC.*, Version of Record 305 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) B. tonduzii C.DC., B. viridiflora A.DC., B. xylopoda L.B.Sm. & B.G.Schub. — [*The “tiliifolia group”] Species whose membership is doubtful. – 6 species: B. bangii Kuntze, B. buchtienii Irmsch., B. chaetocarpa Kuntze, B. leptostyla Irmsch., B. oblanceolata Rusby, B. varistyla Irmsch. Distribution. – AMERICAS: Bolivia, Colombia, Costa Rica, Ecuador, Guatemala, Honduras, México (Chiapas), Nicaragua, Panama, Peru, Venezuela. Taxonomic notes. – We recognise a much reduced circumscription of B. sect. Ruizopavonia relative to Doorenbos & al. (1998). We resolve a clade with species that share raised veins on the underside of the leaves, elliptic or oblong anthers longer than the filaments, and fewer than five female tepals (except B. boreoharlingii, B. harlingii and B. tiliifolia). These characters are shared by the type B. alnifolia thus we recircumscribe B. sect. Ruizopavonia to include these species. The remainder of species are transferred to B. sect. Cyathocnemis, B. sect. Donaldia, B. sect. Lepsia and B. sect. Pritzelia as discussed under those sections. We place B. viridiflora in B. sect. Ruizopavonia as it resolves within the section and shares the section’s characteristic scandent habit and pinnate leaves. We only tentatively assign the Bolivian members of B. sect. Ruizopavonia with multifid styles to the section. We sample no species of this problematic group and note the difficulty separating these poorly-known species from B. sect. Hydristyles (see discussion in Doorenbos & al., 1998). We consider it more likely that their true affinity is with that section, but we are reluctant to change their section without phylogenetic data. Begonia sect. Scutobegonia Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 140. 1894 – Type: Begonia lacunosa Warb. — [Fig. 12A] Species list. – 25 species: B. aggeloptera N.Hallé, B. anisosepala Hook.f., B. ciliobracteata Warb., B. clypeifolia Hook.f., B. dewildei Sosef, B. erectocaulis Sosef, B. erectotricha Sosef, B. ferramica N.Hallé, B. hirsutula Hook.f., B. lacunosa Warb., B. laporteifolia Warb., B. lopensis Sosef & M.Leal, B. mbangensis Sosef, B. mildbraedii Gilg, B. monte-alenensi Sosef, B. montis-elephantis J.J.de Wilde, B. peperomioides Hook.f., B. puberula Sosef, B. scutulum Hook.f., B. susaniae Sosef, B. vankerckhovenii De Wild., B. vittarifolia N.Hallé, B. wilksii Sosef, B. zairensis Sosef, B. zenkeriana L.B.Sm. & Wassh. Distribution. – AFRICA: Angola, Cameroon, Democratic Republic of the Congo, Equatorial Guinea, Gabón, Ghana, Ivory Coast, Nigeria, Republic of the Congo. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 184) and follow the circumscription therein. Begonia sect. Semibegoniella (C.DC.) F.A.Barkley & Baranov in Buxtonian 1 (Suppl. 1): 7. 1972 ≡ Semibegoniella C.DC. in Bull. Herb. Boissier 2(7): 307. 1908 – Type: Semibegoniella sodiroi C.DC. = Begonia longirostris Benth. — [Fig. 17F] = Begoniella Oliv. in Trans. Linn. Soc. Bot. 28: 513. 1873 – Type: Begonia whitei Oliv. ≡ Begonia oliveri L.B.Sm. & B.G.Schub. Species list. – 15 species: B. hexandra Irmsch., B. holmnielseniana L.B.Sm. & Wassh., B. irmscheri L.B.Sm. & B.G. 306 TAXON 67 (2) • April 2018: 267–323 Schub., B. kalbreyeri (Oliv.) L.B.Sm. & B.G.Schub., B. killipiana L.B.Sm. & B.G.Schub., B. lehmannii (Irmsch.) L.B.Sm. & B.G.Schub., B. libera (L.B.Sm. & B.G.Schub.) L.B.Sm. & B.G.Schub., B. longirostris Benth., B. napoensis L.B.Sm. & B.G.Schub., B. oliveri L.B.Sm. & B.G.Schub., B. pectennervia L.B.Sm. & Wassh., B. silverstonii Jara, B. suaviola Jara, B. tetrandra Irmsch., B. valvata L.B.Sm. & B.G.Schub. Distribution. – AMERICAS: Colombia, Ecuador. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 187) and follow the circumscription therein. Begonia sect. Sexalaria A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 127. 1859 – Type: Begonia annobonensis A.DC. — [Fig. 12B] Species list. – 1 species: B. annobonensis A.DC. Distribution. – AFRICA: Cameroon, Equatorial Guinea (Annobón), São Tomé and Príncipe (Príncipe, São Tomé). Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 189) and follow the circumscription therein. Begonia sect. Solananthera A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 128. 1859 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 7. 1972): Begonia solananthera A.DC. — [Fig. 17G] Species list. – 3 species: B. integerrima Spreng., B. radicans Vell., B. solananthera A.DC. Distribution. – AMERICAS: Brazil (Bahia, Espírito Santo, Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo). Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 192) and follow the circumscription therein. Begonia sect. Squamibegonia Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 139. 1894 – Type: Begonia ampla Hook.f. — [Fig. 12C] Species list. – 3 species: B. ampla Hook.f., B. bonushenricus J.J.de Wilde, B. poculifera Hook.f. Distribution. – AFRICA: Angola, Burundi, Cameroon, Democratic Republic of the Congo, Equatorial Guinea (Annobón, Bioko), Gabón, Nigeria, Republic of the Congo, Rwanda, São Tomé and Príncipe (Príncipe, São Tomé), Tanzania, Uganda. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 197) and follow the circumscription therein. Begonia sect. Stellandrae Moonlight, sect. nov. – Type: Begonia hoehneana Irmsch. — [Fig. 17H] Description. – Caulescent, repent herbs, perennial. Stipules persistent. Leaves alternate, basifixed, blade subsymmetrical, veins palmate. Inflorescence axillary, a dichasial cyme, bracts deciduous. Male flowers: with 4 free perianth segments; stamens 4 or 8, free, spreading from the apex of the column, anthers linear, dehiscing via lateral slits, connective projecting. Female flowers: bracteoles 2 at the base of the ovary, with 5 free perianth segments; ovary and fruit with 3 wings, wings subequal, 3-locular, placentas axile, entire, bearing ovules on both surfaces; styles 3, free, bifid, stigmatic papillae in a twice-spirally twisted band. Fruit a capsule. Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Species list. – 1 species: B. hoehneana Irmsch. Distribution. – AMERICAS: Brazil (São Paulo). Etymology. – The epithet is from the Latin stella (star) and andro (male), referring to the type’s stamens, which point outwards in a star-like fashion from the apex of a column. Taxonomic notes. – The position of B. hoehneana has been uncertain since its description. Irmscher (1953) placed it within B. sect. Pritzelia but noted the species’ unusual habit and androecium. Doorenbos & al. (1998) did not classify the species to section. Given its phylogenetically isolated position and the large morphological differences between B. hoehneana and the “Wageneria clade”, we create B. sect. Stellandrae to encompass this species. Begonia sect. Stellandrae is easily distinguished from all other American sections of Begonia by its combination of a creeping habit and anthers spreading from the apex of a long column. Begonia sect. Symbegonia (Warb.) L.L.Forrest & Hollingsw. in Pl. Syst. Evol 241: 208. 2003 ≡ Symbegonia Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 149. 1894 – Type: Symbegonia fulvo-villosa (Warb.) Warb. ≡ Begonia fulvovillosa Warb. — [Fig. 14F] Species list. – 13 species: B. arfakensis (Gibbs) L.L.Forrest & Hollingsw., B. argenteomarginata Tebbitt, B. fulvo-villosa Warb., B. mooreana (Irmsch.) L.L.Forrest & Hollingsw., B. pulchra (Ridl.) L.L.Forrest & Hollingsw., B. strigosa (Warb.) L.L.Forrest & Hollingsw., B. symbracteosa L.L.Forrest & Hollingsw., B. symgeraniifolia L.L.Forrest & Hollingsw., B. symhirta L.L.Forrest & Hollingsw., B. sympapuana L.L. Forrest & Hollingsw., B. symparvifolia L.L.Forrest & Hollingsw., B. symsanguinea L.L.Forrest & Hollingsw., B. yapenensis M.Hughes Distribution. – ASIA: Indonesia (West Papua), Papua New Guinea. Taxonomic notes. – Begonia sect. Symbegonia is nested within B. sect. Petermannia but is highly distinctive, with all species displaying fused tepals and elongate stigmas in the female flowers and many also displaying fused tepals in the male flower and columnar androecia (Hughes & al., 2015a). Following Forrest & Hollingsworth (2003), who transferred the genus Symbegonia into Begonia, we are reluctant to sink the section into B. sect. Petermannia and lose the utility of such a recognisable group, despite the fact it renders B. sect. Petermannia paraphyletic. The position of B. arfakensis in the phylogeny sister to an unidentified member of B. sect. Petermannia from Papua also renders B. sect. Symbegonia polyphyletic; this is a marked incongruence between morphological and molecular data and is potentially the result of chloroplast capture. We refer to the description in Forrest & Hollingsworth & al. (2003) and follow the circumscription therein. Begonia sect. Tetrachia Brade in Rodriguesia 18: 21. 1945 – Type: Begonia quadrilocularis Brade = Begonia egregia N.E.Br. — [Fig. 17I] Description. – Caulescent, erect succulent herbs, perennial. Stipules deciduous. Leaves alternate, peltate or basifixed, blade subsymmetrical or asymmetrical, veins peltate-pinnate to palmate-pinnate. Inflorescence axillary, many flowered, cymose, bracts deciduous. Male flowers: with 2 or 4 free perianth segments; stamens 25 to numerous, free or filaments united in a column, anthers oblong or obovate, dehiscing via lateral slits, connective produced. Female flowers: bracteoles 2 at the base of the ovary, with 5 or 6 (B. egregia) free perianth segments; ovary and fruit with 3 wings, wings subequal, 3- or 4-locular (B. egregia), placentas axile, entire or bifid, bearing ovules on both surfaces; styles 3, free or united at the base, bifid, stigmatic papillae in a spirally twisted band. Fruit a capsule. Species list. – 16 species: B. aguiabrancensis L.Kollmann, B. albidula Brade, B. egregia N.E.Br., B. fellereriana Irmsch., B. goldingiana L.Kollmann & A.Fontana, B. grisea A.DC., B. ibitiocensis E.L.Jacques & Mamede, B. kuhlmannii Brade, B. lealii Brade, B. novalombardiensis L.Kollmann, B. obdeltata Gregório & E.L.Jacques, B. pernambucensis Brade, B. ruhlandiana Irmsch., B. ruschii L.Kollmann, B. umbraculifera Hook.f., B. wasshauseniana L.Kollmann & Peixoto Distribution. – AMERICAS: Brazil (Alagoas, Bahia, Espírito Santo, Minas Gerais, Paraíba, Pernambuco, Rio de Janeiro) Taxonomic notes. – Begonia sect. Tetrachia was previously classified as monotypic, containing only B. egregia (Doorenbos & al., 1998). Our analyses resolve this species as nested within a clade containing three species of Brazilian Begonia currently classified within B. sect. Pritzelia and B. sect. Knesebeckia. Almost all of these species have stellate or peltate, scale-like hairs (B. umbraculifera is glabrous); frequently peltate leaves; axile, usually simple placentation; and cymose inflorescences. These characteristics are shared with ten unsampled Brazilian dry-adapted species of B. sect. Begonia, B. sect. Knesebeckia and B. sect. Pritzelia. We expand B. sect. Tetrachia to include these species but note that B. egregia remains unique within the section in its female flowers with 6 tepals and four locules (rather than 5 tepals and three locules). Begonia sect. Tetraphila A.DC., Prodr. 15(1): 517. 1864 – Type: Begonia mannii Hook.f. — [Fig. 12D] = Begonia sect. Fuscibegonia Warb. in Engler & Prantl, Nat. Pflanzenfam. 3(6a): 141. 1894 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 8. 1972): Begonia eminii Warb. = Begonia (sect. Fuscibegonia) ser. Subaequilaterales Engl. in Engler. & Drude, Veg. Erde 9(3, 2): 618. 1921 – Type (designated by Doorenbos & al. in Wageningen Agric. Univ. Pap. 98(2): 201. 1998): Begonia polygonoides Hook.f. = Begonia (sect. Fuscibegonia) ser. Inaequilaterales Engl. in Engler & Drude, Veg. Erde 9(3, 2): 620. 1921 – Type (designated by Doorenbos & al. in Wageningen Agric. Univ. Pap. 98(2): 201. 1998): Begonia eminii Warb. = Begonia sect. Irmschera Ziesenh. in Begonian 38(7): 161. 1971 – Type: Begonia mauricei Ziesenh. = Begonia loranthoides Hook.f. Species list. – 31 species: B. afromigrata J.J.de Wilde, B. capillipes Gilg, B. cavallyensis A.Chev., B. ebolowensis Version of Record 307 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) TAXON 67 (2) • April 2018: 267–323 Fig. 11. Sections of African Begonia. A, B. sect. Augustia, B. dregei, RBGE living accession number 20000906, South Africa, McLellan, T. 223 (E); B, B. sect. Baccabegonia, B. baccata, Sao Tomé, Berthold, C. & Gardner, M.F. 5 (E); C, B. sect. Erminea, B. bogneri, RBGE living accession number 19860844; D, B. sect. Cristasemen, B. thomeana, Sao Tomé, Berthold, C. & Gardner, M.F. 13 (E); E, B. sect. Exalabegonia, B. oxyloba, Guinea, Bidault, E. & Diabaté, M. 665 (MO); F, B. sect. Filicibegonia, B. aspleniifolia, Gabon, cultivated in private collection; G, B. sect. Loasibegonia, B. scutifolia, Cameroon, Droisart, V. & al. 1214 (MO); H, B. sect. Mezierea, B. salaziensis, Mauritius, cultivated in St. Andrews BG; I, B. sect. Nerviplacentaria, B. henrilaportei, Madagascar, Scherberich, D. 969 (LYJB); J, B. sect. Peltaugustia, B. samhaensis, RBGE living accession number 19990398, Socotra, Hughes, M. H6 (E); K, B. sect. Quadrilobaria, B. nossibea, Madagascar; L, B. sect. Rostrobegonia, B. johnstonii, RBGE living accession number 20131209, Barber, S. & al. 14 (E). — Photographs by P.W. Moonlight (A, C, J, L); M. Gardner (B, D); E. Bidault (E), J. Duruisseau (F, K), V. Droisart (G), M. Hughes (H), D. Scherberich (I, K inset). 308 Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Engl., B. elaeagnifolia Hook.f., B. eminii Warb., B. furfuracea Hook.f., B. fusialata Warb., B. fusicarpa Irmsch., B. gabonensis J.J.de Wilde, B. horticola Irmsch., B. karperi J.C.Arends, B. kisuluana Büttner, B. komoensis Irmsch., B. letestui J.J.de Wilde, B. longipetiolata Gilg, B. loranthoides Hook.f., B. mannii Hook.f., B. molleri (C.DC.) Warb., B. oxyanthera Warb., B. pelargoniiflora J.J.de Wilde & J.C.Arends, B. polygonoides Hook.f., B. preussii Warb., B. rubromarginata Gilg, B. rubronervata De Wild., B. rwandensis J.C.Arends, B. squamulosa Hook.f., B. subalpestris A.Chev., B. subscutata De Wild., B. tatoniana R.Wilczek, B. zimmermannii Peter ex Irmsch. Distribution. – AFRICA: Angola, Burundi, Cameroon, Central African Republic, Democratic Republic of the Congo, Equatorial Guinea (Equatorial Guinea, Bioko), Gabón, Ghana, Guinea, Ivory Coast, Kenya, Liberia, Nigeria, Republic of the Congo, Rwanda, Sierra Leone, São Tomé and Príncipe (Príncipe, São Tomé), South Sudan, Tanzania, Uganda. ASIA: Laos; Thailand (B. afromigrata). Taxonomic notes. – We resolve B. sect. Baccabegonia and B. sect. Squamibegonia as nested within B. sect. Tetraphila. These sections differ in a number of characters (e.g., male tepal number, fruit dehiscence, see Doorenbos & al., 1998 for more details) and are thus useful for diagnostic purposes. If B. sect. Squamibegonia and B. sect. Baccabegonia are to be retained, B. sect. Tetraphila will require recircumscription if it is to be a monophyletic section. This section is the largest in Africa and contains considerable morphological diversity, and it is likely morphological characters will be found to aid recircumscription. We refer to the description in Doorenbos & al. (1998: 207) and follow the circumscription therein. Begonia sect. Trachelocarpus (Müll.Berol.) A.DC. in Martius, Fl. Bras. 4(1): 381. 1861 ≡ Trachelocarpus Müll.Berol. in Ann. Bot. Syst. 4: 909. 1857 – Type: Trachelocarpus rhizocarpus Müll.Berol. = Begonia depauperata Schott. — [Fig. 17J] = Trachelanthus Klotzsch in Abh. Königl. Akad. Wiss. Berlin 1854: 240. 1855, non Trachelanthus G.Kunze – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 8. 1972): Trachelanthus rhizocarpus Klotzsch = Begonia depauperata Schott. Species list. – 5 species: B. depauperata Schott, B. fulvosetulosa Brade, B. herbacea Vell., B. lanceolata Vell., B. velloziana Walp. Distribution. – AMERICAS: Brazil (Paraná, Rio de Janeiro, Santa Catarina, São Paulo). Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 204) and follow the circumscription therein. Begonia sect. Urniformia Ziesenh. in Begonian 41: 12. 1974 – Type: Begonia pitteri C.DC. = Begonia heydei C.DC. — [Fig. 17K] Species list. – 1 species: B. heydei C.DC. Fig. 12. Sections of African Begonia. A, B. sect. Scutobegonia, B. mildbradei, Bidault, E. & Akouangou, A. 1511 (MO); B, B. sect. Sexalaria, B. annobonensis, Sao Tomé, Berthold, C. & Gardner, M.F. 50 (E); C, B. sect. Squamibegonia, B. ampla, RBGE living accession number 19990258, Sao Tomé, Gascoigne 15 (E); D, B. sect. Tetraphila, B. gabonensis, RBGE living accession number 20101692. — Photographs by E. Bidault (A), M. Gardner (B), P.W. Moonlight (C, D). Version of Record 309 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) TAXON 67 (2) • April 2018: 267–323 Fig. 13. Sections of Asian Begonia. A, B. sect. Alicida, B. alicida, Thailand; B, B. sect. Apterobegonia, B. phutthaii, Thailand, Middleton, D.J. & al. 5822, 5827 (E); C, B. sect. Baryandra, B. oxysperma, Mount Banahaw, Luzon, the Philippines; D, B. sect. Bracteibegonia, B. bracteata, Sumatra, Girmansyah, D. & Hughes, M. DEDEN1495 (E); E, B. sect. Coelocentrum, B. porteri, Guangxi, Peng, C.-I 19840 (HAST); F, B. sect. Diploclinium, B. grandis, Yunnan, China; G, B. sect. Haagea, B. dipetala, Sri Lanka, Peng, C.-I 21011 (HAST); H, B. sect. Jackia, B. sublobata, Sumatra, Girmansyah, D. DEDEN1486 (E); I, B. sect. Lauchea, B. adenopoda, Thailand, Middleton, D.J. & al. PK78473 (E); J, B. sect. Monophyllon, B. prolifera, Thailand, Peng, C.-I 24195 (HAST); K, B. sect. Oligandrae, B. pentandra, Papua New Guinea, Takeuchi, W.N. & al. 23126 (A); L, B. sect. Parvibegonia, B. martabanica, Kanchanaburi, Thailand. — Photographs by C.I Peng (A, C, D [inset], E, G, J, L), D. Middleton (B, I), M. Hughes (D, F, H), W. Takeuchi (K). 310 Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Distribution. – AMERICAS: Costa Rica, Guatemala, Panama. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 209) and follow the circumscription therein. Begonia sect. Wageneria (Klotzsch) A.DC. in Ann. Sci. Nat., Bot., ser. 4, 11: 137. 1859 ≡ Wageneria Klotzsch in Ber. Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1854: 126. 1854 – Type (designated by Barkley & Baranov in Buxtonian 1(Suppl. 1): 3. 1972): Wageneria fagifolia (Fisch. ex Otto & A.Dietr.) Klotzsch ≡ Begonia fagifolia Fisch. ex Otto & A.Dietr. — [Fig. 17L] = Begonia sect. Enita Brade in Rodriguésia 32: 160. 1957 – Type: Wageneria convolvulacea Klotzsch ≡ Begonia convolvulacea (Klotzsch) A.DC. Species list. – 7 species: B. convolvulacea (Klotzsch) A.DC., B. epibaterium Mart. ex A.DC., B. fagifolia Fisch. ex Otto & A.Dietr., B. glabra Aubl., B. inconspicua Brade, B. polygonifolia A.DC., B. smilacina A.DC. Distribution. – AMERICAS: Belize, Brazil (Acre, Alagoas, Amapá, Amazonas, Bahia, Ceará, Distrito Federal, Espírito Santo, Goiás, Minas Gerais, Paraíba, Paraná, Pernambuco, Rio de Janeiro, Rondônia, Santa Catarina, São Paulo, Sergipe), Bolivia, Colombia, Costa Rica, Cuba, Ecuador, French Guiana, Guatemala, Guyana, Honduras, Jamaica, México (Chiapas, Guerrero, Hidalgo, Oaxaca, Puebla, Querétaro, Tabasco, Tamaulipas, Veracruz), Nicaragua, Panama, Peru, Suriname, Trinidad-Tobago, Venezuela (Venezuela, Venezuelan Antilles). Taxonomic notes. – Begonia sect. Wageneria was separated from B. sect. Pritzelia by Doorenbos & al. (1998) due to its distinct scandent habit. We resolve all members of the section within a well-supported clade with a number of species of B. sect. Pritzelia characterised by large, palmately veined and slightly bullate leaves and lax, cymose inflorescences. These species include B. reniformis, which was previously placed by Klotzsch in his genus Wageneria. However, a number of species treated within Wageneria by Klotzsch are resolved in the “core Pritzelia clade” (e.g., B. hispida and B. hugelii) The difference between these groups remains unclear. We refer to the description in Doorenbos & al. (1998: 210), follow the circumscription therein. and informally name the wider clade the “Wageneria clade”. Further morphological and phylogenetic work is required to determine which other similar members of B. sect. Pritzelia resolve within the “Wageneria clade” (e.g., Fig. 14. Sections of Asian Begonia. A, B. sect. Petermannia, B. cummingiana, Philippines; B, B. sect. Platycentrum, B. xanthina, Yunnan, China; C, B. sect. Putzeysia, B. gemmipara, West Bengal, India; D, B. sect. Reichenheimia, B. tenera, Sri Lanka; E, B. sect. Ridleyella, B. kingiana, Peninsula Malaysia, Peng, C.-I 21226 (HAST); F, B. sect. Symbegonia, B. arfakensis, RBGE living accession number 20090808, West Papua, Argent, G. & al. 305. — Photographs by C.I Peng (A, B, E), P. Blanc (C), M.Hughes (D, F). Version of Record 311 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) TAXON 67 (2) • April 2018: 267–323 Fig. 15. Sections of American Begonia. A, B. sect. Astrothrix, B. fragae, Brasil, Fraga C.N. 2323 (UPCB); B, B. sect. Australes, B. micranthera, Bolivia, Sarkinen & al. 2144 (FHO); C, B. sect. Barya, B. monadelpha, Peru, Moonlight, P. & Daza, A. 148 (E); D, B. sect. Begonia, B. odorata, RBGE living accession number 20082086; E, B. sect. Casparya, B. urticae, Peru, Moonlight, P. & Daza, A. 154 (E); F, B. sect. Cyathocnemis, B. altoperuviana, Peru, Moonlight, P. & Daza, A. 83 (E); G, B. sect. Donaldia, B. saxicola, Bahia, Cardoso, D. 1305 (HUEFS); H, B. sect. Doratometra, B. semiovata, Peru, Moonlight, P. & Daza, A. 172 (E); I, B. sect. Ephemera, B. fischeri, Peru, Moonlight, P. & Daza, A. 124 (E); J, B. sect. Eupetalum, B. geraniifolia, Peru, Moonlight, P. & Daza, A. 116 (E); K, B. sect. Gaerdtia, B. maculata, RBGE living accession number 19891579; L, B. sect. Gireoudia, B. carolineifolia, RBGE living accession number 20100417, Twyford, A. 584 (E). — Photographs by L. Kollmann (A), T. Sarkinen (B), P.W. Moonlight (C–F, H–L), D. Cardoso (G). 312 Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Fig. 16. Sections of American Begonia. A, B. sect. Gobenia, B. tropaeolifolia, Ecuador, Tebbitt, M. & Gutierrez, D. 850 (QCA); B, B. sect. Hydristyles, B. fissistyla, A. Orejuela 2830 (USM); C, B. sect. Kollmannia, B. jaguarensis, Espírito Santo, Kollmann, L. & Lopes, R.S. 11418 (MBML); D, Core B. sect. Knesebeckia, B. monophylla, Nuevo León; E, B. sect. Knesebeckia group I, B. velata, Peru, Tebbitt, M. & Daza, A. 838 (MOL); F, B. sect. Knesebeckia group II, B. aff. microcarpa, Peru, Moonlight, P. & Daza, A. 156 (E); G, B. sect. Knesebeckia group III, B. chemillenensis, Peru, Moonlight, P. & Daza, A. 314 (E); H, B. sect. Latistigma, B. aconitifolia, RBGE living accession number 20140767; I, B. sect. Lepsia, B. foliosa, RBGE living accession number 19480286; J, B. sect. Microtuberosa, B. elachista, Moonlight, P. & Daza, A. 318 (E); K, B. sect. Parietoplacentalia, B. oaxacana, Knapp, S. & A. Munro 9984 (BM); L, B. sect. Pereira, B. lubbersii, Berlin Botanic Garden. — Photographs by M.C. Tebbitt (A, E), A. Orejuela (B), L. Kollmann (C), S. Eduardo (D), P.W. Moonlight (F–J), A. Munro (K), J.-P. Bachelot Pierre (L). Version of Record 313 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) TAXON 67 (2) • April 2018: 267–323 Fig. 17. Sections of American Begonia. A, B. sect. Pilderia, B. buddleiifolia, RBGE living accession number 20160126, Peru, Moonlight, P. & Daza, A. 165 (E); B, B. sect. Pritzelia, B. parviflora, Peru, Moonlight, P. & Daza, A. 171 (E); C, B. sect. Quadriperigonia, B. gracilis, Chihuahua; D, B. sect. Rossmannia, B. rossmanniae, Peru, Moonlight, P.W. & Daza, A. 315 (E); E, B. sect. Ruizopavonia, B. peruviana, Peru, Moonlight, P. & Daza, A. 82 (E); F, B. sect. Semibegoniella, B. longirostris, Ecuador; G, B. sect. Solananthera, B. radicans, São Paulo; H, B. sect. Stellandrae, B. hoehneana, RBGE living accession number 20141394; I, B. sect. Tetrachia, B. grisea, Bahia, Moonlight, P.W. & Fernandes, M. 750 (HUEFS); J, B. sect. Trachelocarpus, B. herbacea, RBGE living accession number 19731859; K, B. sect. Urniformia, B. heydei, RBGE living accession number 20131992; L, B. sect. Wagneria, B. glabra, Peru, P. Moonlight & A. Daza 240 (E). — Photographs by P.W. Moonlight (A, B, D, E, H–J, L), A. McDonnell (C), A. Jara-Muñoz (F), A. Benedito (G), C.I Peng (K). 314 Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) B. caraguatatubensis Brade, B. collaris Brade). A revision of these sections is overdue, and may clarify the boundaries between B. sect. Pritzelia and B. sect. Wageneria. Begonia sect. Warburgina Kuntze, Revis. Gen. Pl. 3(2): 105. 1898 – Type: Begonia comata Kuntze. Species list. – 1 species: B. comata Kuntze. Distribution. – AMERICAS: Bolivia. Taxonomic notes. – We refer to the description in Doorenbos & al. (1998: 213) and follow the circumscription therein. Species Unassigned to Section (do not match current sections): 32 species: Begonia antaisaka Humb. ex Keraudren & Bosser, B. archboldiana Merr. & L.M.Perry, B. bataiensis Kiew, B. bifurcata L.B.Sm. & B.G.Schub., B. boisiana Gagnep., B. calcarea Ridl.*, B. crassula Aver., B. cremnophila Tebbitt**, B. erythrothrix Tebbitt & Moonlight, B. fractiflexa S.Julia & Kiew, B. froebelii A.DC., B. gracilicyma Irmsch. ex M.Hughes, B. herteri Irmsch., B. hohuanensis S.S.Ying, B. kanaensis Kiew & C.Y.Ling*, B. laccophora Sands, B. lingiae S.Julia*, B. lutea L.B.Sm. & B.G.Schub., B. malabarica Lam., B. masoalaensis M.Hughes, B. peltatifolia H.L.Li, B. physandra Merr. & L.M.Perry, B. rigidifolia Aver., B. rubrotepala S.Julia*, B. sabahensis Kiew & J.H.Tan*, B. sadirensis Kiew & S.Julia*, B. smithiae Geddes, B. speculum Moonlight & Tebbitt**, B. tanala Humb. ex Keraudren & Bosser, B. thyrsoidea Irmsch., B. urubambensis Tebbitt**, B. viscosa Aver. & H.Q.Nguyen. — [*The “calcarea group”; **the “cremnophila group”] Species unassigned to section (imperfectly known): 50 species: B. anaimalaiensis Bedd., B. brongniartiana Lem., B. brongniartii Lem., B. burkei hort., B. capensis L.f., B. castaneifolia hort. ex Otto & A.Dietr., B. cerasiphylla L.B.Sm. & Wassh., B. chakensis S.Julia & C.Y.Ling, B. complicata (Hassk.) A.DC., B. diptera Dryand., B. donkelaariana Lem., B. eriocaulon Neumann, B. gemmirhiza H.Lév., B. groenewegensis hort. ex K. Koch & G.A. Fintelmann, B. herteri Irmsch., B. hochbaumii hort. ex E.Otto, B. intermedia Veitch ex Van Houtte, B. khasiana C.B.Clarke, B. lansbergeae Linden & Rodigas, B. lemaoutii hort. ex Vallerand, B. macrotis Vis., B. maracuyensis Parodi, B. martynia hort., B. medusae Linden, B. mexera hort., B. meysselliana Linden, B. microptera Hook., B. muroptera hort., B. mutabilis Harland, B. nigricans L.H.Bailey, B. nobilis hort., B. northiana hort. ex Gentil, B. nummularifolia Putz., B. opulifolia Loudon, B. orchidiflora Griff., B. palmatiloba Linden & André, B. paniculata D.Parodi, B. pantherina Putz. ex Linden, B. papillaris Herb. ex Cels, B. paraguayensis D.Parodi, B. phyllomaniaca Mart., B. piperoides Linden, B. porteana Van Geert, B. preseriana hort., B. putzeysii hort., B. stipularis Spreng., B. strigulosa (Hassk.) A.DC., B. sunorchis C.Chev., B. tascellezii hort., B. villifera Galeotti. AUTHOR CONTRIBUTIONS PWM and MH designed the research; PWM, WHA, LAP, K-FC, DG, AJ-M, RK, W-CL, YL, LDKM, C-IP, AJP, TP, SR, CR, RRR, DS, MCT, DCT, NHZ and MH collected the samples; PWM, DF, RH, AJ-M, AM, LDKM, MO’C, TP, DCT, NHZ and MH provided the molecular data; PWM and MH performed the analyses; MP provided database support; PWM, WA, K-FC, AJ-M, RK, C-IP, TP, SR, RRR, SJ, DS, Y-MS, MCT, DCT and MH interpreted the results; PWM and MH wrote the manuscript; and all authors were involved in editing the manuscript. — K-FC, https://orcid.org/0000-0003-3628-2567, bochung@ gate.sinica.edu.tw; DG, deden_bo@yahoo.com; MH, https://orcid. org/0000-0002-2168-0514, mhughes@rbge.ac.uk; RK, ruth@frim. gov.my; PWM, https://orcid.org/0000-0003-4342-2089; C-IP, https:// orcid.org/0000-0002-9175-1146; DCT, https://orcid.org/0000-00021307-6042, Daniel_THOMAS@nparks.gov.sg; HPW , https://orcid. org/0000-0002-5231-3987 ACKNOWLEDGEMENTS We are grateful to the curators of the following herbaria for access to specimens and images: A, AA, ABD, AMAZ, ANDA, B, BISH, BK, BKF, BKL, BM, BO, BRIT, C, CAS, CEB, CHAPA, CLEMS, COL, CPUN, CR, CUVC, E, EAC, F, FHO, FI, FMB, FRIM, G, GB, GH, HAST, HBG, HLDG, HOXA, HUA, HUEFS, HUSA, HUT, IZTA, K, KATH, KEP, KUN, K-W, L, LAE, LPB, LY, MEX, MEXU, MICH, MO, MOL, NY, OXF, P, PDA, PNH, PSU, QCA, QCNE, QPLS, RB, S, SING, TI, TUCH, U, UC, UPNG, US, USM, VEN, W, WAG and Z. We thank M.L. Hart and L.L. Forrest (RBGE) for technical support; Pablo Hendrigo Alves de Melo, Toby Pennington, Flávia Pezzini and Tiina Sarkinen for silica gel material; the Royal Botanic Garden Edinburgh, Glasgow Botanic Gardens, HAST Institute Taipei, and Lyon Botanic Garden for access to living material; and the many people who provided images of living plants (see figure captions 11–17). This research arose out of projects supported by the M.L. MacIntyre Trust, The Sibbald Trust, The Royal Society of Edinburgh Bilateral Program, and The Davis Expedition Fund. The Royal Botanic Garden Edinburgh is supported by the Scottish Goverments’ Rural and Environment Science and Analytical Services Division. We are indebted to two anonymous reviewers and the Taxon editorial team, in particular Franz Stadler, for their assistance in correcting errors and omissions. LITERATURE CITED Ardhaka, I.M. & Ardi, W.H. 2016. A new species of Begonia (Begoniaceae) from Mansuela National Park, Seram. 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PhytoKeys 51: 1–282. https:// doi.org/10.3897/phytokeys.51.7104 TAXON 67 (2) • April 2018: 267–323 Yang, C.-J., Tagane, S., Souladeth, P., Okabe, N., Hu, J.-M. & Yahara, T. 2018. Flora of Nam Kading National Protected Area III: Begonia namkadingensis (Begoniaceae), a new species in limestone area. Phytotaxa 334: 195–199. https://doi.org/10.11646/ phytotaxa.334.2.8 Appendix 1. GenBank accession numbers and voucher specimen information. Taxon, collector plus collection number (herbarium code), ndhA intron, ndhF-rpl32 spacer, rpl32-trnL spacer (“–” indicates missing sequence; accessions starting with “MH” are newly obtained for this study) Begonia abdullahpieei, C.I Peng 22727 (HAST), MH207018, MH207427, MH207848; B. aberrans, M. Hughes 786 (E), KP712927, KP713097, KP713247; B. aborensis, J. Durisseau 100493 (LBG), MH207019, MH207428, MH207849; B. acerifolia, P.W. Moonlight & A. Daza 96 (E), –, MH207429, –; B. acerifolia, M. Tebbitt 768 (QCA), MH207020, MH207430, MH207850; B. acerifolia, M. Tebbitt 788 (QCA), MH207021, MH207431, MH207851; B. acerifolia, M. Tebbitt & A. Daza 797 (E), MH207022, MH207433, –; B. acerifolia, M. Tebbitt & A. Daza 810 (E), MH207023, MH207434, MH207852; B. acerifolia, M. Tebbitt & A. Daza 831 (E), MH207024, MH207435, –; B. acerifolia, M. Tebbitt & A. Daza 843 (E), MH207025, MH207436, MH207853; B. acerifolia, A. Parada 3342 (USZ), –, MH207432, –; B. acerifolia, P.W. Moonlight & A. Daza 120 (E), MH207026, MH207437, MH207854; B. acerifolia, T. Sarkinen 2058 (E), KP713031, KP713140, –; B. acerifolia, C.-I Peng 20869 (HAST), KP713069, KP713076, –; B. aceroides, T. Phutthai 243 (E), JF756385, JF756469, JF756553; B. acetosa, M. Tebbitt, S.M. Swensen & J. Yeadon 15 (BKL), KP712965, KP713154, KP713324; B. acetosella, Glasgow B.G. 003-152-95 (E), JF756367, JF756451, JF756535; B. acida, C.-I Peng 21294 (HAST), KP712957, KP713148, KP713330; B. aconitifolia, C.-I Peng 22224 (HAST), KP712925, –, –; B. aconitifolia, Tebbitt s.n., MH235394, MH235397, MH235432; B. acuminatissima, R.R. Rubite R321 (PNH), KR186445, KR186532, KR186706; B. acutifolia, M. Tebbitt 127 (BKL), KP713047, KP713166, KP713248; B. acutifolia, L.L. Forrest 160 (E), KP713055, KP713165, KP713249; B. alabensis, F. Lens 072 (SBG), MH207030, MH207441, MH207858; B. alabensis, F. Lens 078 (SBG), MH207031, MH207442, MH207859; B. albo-coccinea, C.-I Peng P23302 (HAST), KR186447, KR186534, KR186708; B. albomaculata, P.W. Moonlight & A. Daza 126 (E), MH207032, MH207443, MH207860; B. albomaculata, P.W. Moonlight & A. Daza 198 (E), MH207033, MH207444, MH207861; B. albomaculata, P.W. Moonlight & A. Daza 213 (E), MH207034, MH207445, –; B. alice-clarkiae, E20110203, KP713007, KP713200, KP713233; B. alicida, T. Phutthai 139 (E), JF756388, JF756472, JF756556; B. almedana, A. Twyford 264 (E), MH207035, MH207446, MH207862; B. altoperuviana, P. W. Moonlight & A. Daza 84 (E), MH207036, MH207447, MH207863; B. altoperuviana, M. Tebbitt & A. Daza 823 (E), MH207037, MH207448, MH207864; B. amphioxus, D.C. Thomas 08116 (E), MH207038, MH207449, MH207865; B. ampla, Suksuwan 45 E00198091 (E), KP712979, –, –; B. anemoniflora, P.W. Moonlight & A. Daza 222 (E), MH207039, MH207450, MH207866; B. anemoniflora, P.W. Moonlight & A. Daza 230 (E), MH207040, MH207451, MH207867; B. angularis, M. Tebbitt 78 (BKL), KP712948, KP713217, KP713279; B. angustifolia, A. JaraMuñoz 2765 (COL), MH207041, MH207452, –; B. anisoptera, R.R. Rubite R479 (PNH), KR186448, KR186535, KR186709; B. annobonensis, M.F. Gardner, C.E. Berthold 49 (E), MH207042, –, MH207868; B. aptera, Smith & Galloway 67 (E), JF756369, JF756453, JF756537; B. arborescens var. confertiflora, M. Tebbitt 119 (BKL), KP712953, KP713162, KP713334; B. areolata, SUBOE 81 (E), MH207043, MH207453, MH207869; B. areolata, Gao & Hollingsworth 24141 (E), JF756366, JF756450, JF756534; B. arfakensis, ABEG 305 (E), MH207044, MH207454, MH207870; B. argenteomarginata, Glasgow B.G. 008-038-87 (E), JF756417, JF756501, JF756585; B. arrogans, P.W. Moonlight & A. Daza 27 (E), MH207045, MH207455, –; B. arrogans, P.W. Moonlight & A. Daza 285 (E), MH207046, MH207456, MH207871; B. atricha, SUBOE 11 (E), MH207047, MH207457, MH207872; B. augustae, B. Sénéchal 070409 (LBG), MH207048, MH207458, MH207873; B. baccata, M.E. Gardner & C.E. Berthold 5 (E), MH207049, MH207459, MH207874; B. baik, C.I Peng 24235 (HAST), MH207050, –, MH207875; B. balansana, J. Durisseau 120338 (LBG), MH207051, MH207460, MH207876; B. baramensis, P34 S09 (SBG), –, MH207461, MH207877; B. barbellata, P33 SBG s.n. (SBG), MH207052, MH207462, MH207878; B. barkeri, HAST RM08 304 A, KP713034, KP713138, –; B. barkeri, A. Twyford 530 E00668768 (E), KP713061, KP713081, KP713245; B. barkleyana, M. Peixoto 041650. (LBG), MH207053, MH207463, MH207879; B. baviensis, HNE 388 (E), MH207054, MH207464, MH207880; B. beludruvenea, M. Hughes & Taufiq 1541 (E), –, MH207465, MH207881; B. beryllae, F. Lens 071 (SBG), MH207055, MH207466, MH207882; B. cf. beryllae, F. Lens 074 (SBG), MH207091, MH207501, MH207917; B. bifurcata, M. Tebbitt 782 (QCA), –, MH207467, MH207883; B. bifurcata, P.W Moonlight & A. Daza 105 (E), KX756296, KX756307, MH207884; B. bifurcata, P.W Moonlight & A. Daza 117 (E), KX756278, KX756308, MH207885; B. biliranensis, R.R. Rubite R311 (PNH), KR186449, KR186536, KR186710; B. bipinnatifida, Kreb s.n. (SBG), MH207058, MH207470, MH207886; B. bissei, W.G.C. 655 (E), MH207059, –, MH207887; B. blancii, C.-I Peng P22545 (HAST), KR186450, KR186537, KR186711; B. bogneri, L.L. Forrest 200 E00171239 (E), KP712977, KP713185, –; B. boisiana, no voucher, MH207060, MH207471, MH207888; B. boliviensis, no voucher, MH207062, MH207473, MH207890; B. boliviensis, M. Tebbitt 704 (USZ), MH207063, MH207474, MH207891; B. boliviensis, M. Tebbitt s.n. (USZ), MH207064, MH207475, MH207892; B. boliviensis, M. Tebbitt s.n. (USZ), MH207065, MH207476, MH207893; B. boliviensis, L.L. Forrest 182 (E), JF756346, JF756430, JF756514; B. bonthainensis, D.C. Thomas 09-63 (E), KP712932, KP713108, KP713327; B. bonthainensis, C.-I Peng P22531 (HAST), KR186451, KR186538, KR186712; B. boquetensis, B. Sénéchal 60681 (LBG), MH207066, MH207477, MH207894; B. botryoides, M. Tebbitt 777 (QCA), MH207067, MH207478, MH207895; B. bracteata, W.H. Ardi 25 (E), KP712991, KP713110, KP713323; B. bracteosa, M. Tebbitt & A. Daza 820 (E), MH207068, MH207479, MH207896; B. bradei, M. Tebbitt 121 (BKL), KP712960, KP713225, KP713287; B. brandisiana, Chamchamroon 3614 (E), JF756379, JF756463, JF756547; B. breedlovei, 08301 (HAST) , KP713036, KP713109, –; B. brevicordata, M. Tebbitt & A. Daza 822 (E), MH207069, MH207480, MH207897; B. brevirimosa, Reeve 142 (E), JF756414, JF756498, JF756582; B. broussonetifolia, B. Sénéchal 110685 (LBG), MH207070, MH207481, MH207898; B. bruneiana, NHZ34 (SBG), MH207071, MH207482, MH207899; B. bryophila ined., Chien-I Huang 6100 (E), MH207072, MH207483, MH207900; B. buddleiifolia, A. JaraMuñoz 2599 (E), KX756286, KX756327, MH207901; B. buddleiifolia, P.W. Moonlight & A. Daza 165 (E), MH207074, MH207485, MH207902; B. buddleiifolia, P.W. Moonlight & A. Daza 316 (E), MH207075, MH207486, MH207903; B. bufoderma, Lyon B.G. 80415 (LBG), MH207076, MH207487, MH207904; B. bullatifolia, C.-I Peng 21323 (HAST), KP713073, KP713128, –; B. bullatifolia, J. Duruisseau 120348 (LBG), KX756303, –, MH207905; B. burbidgei, no voucher, MH207079, MH207489, MH207906; B. burbidgei, F. Lens 040 (SBG), MH207080, MH207490, MH207907; B. calcicola, C.-I Peng P20761 (HAST), KR186452, KR186539, KR186713; B. calderonii, A. Twyford 455 (E), MH207081, MH207491, MH207908; B. callosa, Lyon B.G. 90451 (LBG), MH207082, MH207492, MH207909; B. calzadae, A. Twyford 445 (E), MH207083, MH207493, MH207910; B. camiguinensis, R.R. Rubite R506 (PNH), KR186453, KR186540, KR186714; B. capanemae, M. Tebbitt, Swensen & Yeadon 7 (BKL), KP712949, KP713160, KP713280; B. capensis, Salamone, C. - LBG 20504, MH235383, MH235414, MH235434; B. capituliformis, Kinho & A.D. Poulson 169 (E), JN133309, JN133384, JN133413; B. cardiocarpa, Glasgow B.G. 006-006-92 (E), KP713018, KP713184, KP713301; B. carolinefolia, A. Twyford 457 (E), MH207085, MH207495, MH207912; B. carolineifolia, M. Tebbitt 80 (BKL), KP713033, KP713103, KP713244; B. carrieae, Tebbitt 151 (BKL), MH207086, MH207496, MH207913; B. carrieae, Lyon B.G. s.n. (LBG), MH207087, MH207497, –; B. castilloi, R.R. Rubite R98 (PNH), KR186454, KR186541, KR186715; B. catharinensis, B. Sénéchal 50723 (LBG), MH207088, MH207498, MH207914; B. cathcartii, B. Sénéchal 130528 (LBG), MH207089, MH207499, MH207915; B. aff. cauliflora, F. Lens 065 (SBG), MH207027, MH207438, MH207855; B. ceratocarpa, Y.M. Shui & al. B2015207 (KUN), MH207090, MH207500, MH207916; B. chemillenensis, P.W. Moonlight & A. Daza 278 (E), MH207096, MH207505, MH207922; B. chemillenensis, P.W. Moonlight & A. Daza 287 (E), –, MH207506, MH207923; B. chemillenensis, P.W. Moonlight & A. Daza 292 (E), MH207097, MH207507, MH207924; B. chemillenensis, P.W. Moonlight & A. Daza 294 (E), –, MH207508, –; B. chemillenensis, P.W. Moonlight & A. Daza 313 (E), MH207098, MH207509, MH207925; B. chemillenensis, P.W. Moonlight & A. Daza 314 (E), MH207099, MH207510, MH207926; B. chemillenensis, P.W. Moonlight & A. Daza 317 (E), MH207100, –, –; B. chiasmogyna, D.C. Thomas 07-29 (E), JN133310, JN133385, 318 Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Appendix 1. Continued. JN133414; B. chingipengii, C.-I Peng P23368 (HAST), KR186455, KR186542, KR186716; B. chlorolepis, A. JaraMuñoz 2327 (COL), MH207101, MH207511, MH207927; B. chloroneura, L.L. Forrest 128 (E), JF756394, JF756478, JF756562; B. chloroneura, P. Wilkie & al. 29015 (E), KR186456, KR186543, KR186717; B. chlorosticta, L.L. Forrest 41 (E), JF756402, JF756486, JF756570; B. chrysantha, M. Tebbitt 748 (USZ), MH207102, MH207512, MH207928; B. cinnabarina, M. Tebbitt 705 (USZ), MH207103, MH207513, MH207929; B. cinnabarina, M. Tebbitt 745 (USZ), MH207104, MH207514, MH207930; B. clarkei, M. Tebbitt & A. Daza 798 (E), –, MH207515, –; B. clarkei, M. Tebbitt & A. Daza 802 (E), MH207105, MH207516, –; B. clarkei, M. Tebbitt & A. Daza 804 (E), MH207106, MH207517, MH207931; B. clarkei, M. Tebbitt & A. Daza 806 (E), MH207107, MH207518, MH207932; B. clarkei, M. Tebbitt & A. Daza 811 (E), MH207108, MH207519, MH207933; B. clarkei, M. Tebbitt & A. Daza 815 (E), MH207109, MH207520, MH207934; B. clarkei, M. Tebbitt & A. Daza 818 (E), MH207110, MH207521, MH207935; B. clarkei, M. Tebbitt & A. Daza 819 (E), MH207111, MH207522, MH207936; B. clarkei, M. Tebbitt 824 (E), KX756293, KX756304, MH207937; B. clarkei, M. Tebbitt 738 (USZ), –, MH207524, –; B. cleopatrae, RBGE - Philippine National Herbarium Expedition 1998-25416 (E), JF756390, JF756474, JF756558; B. cleopatrae, P. Wilkie & al. 25373 (E), KR186457, KR186544, KR186718; B. coccinea, Glasgow B.G. 001-119-85, KP712929, KP713137, KP713275; B. colombiana, D.C. 43980 (COL), MH207113, MH207525, MH207938; B. comestibilis, D.C. Thomas & W.H. Ardi 09-62 (E), JN133311, JN133386, JN133415; B. conchifolia, M. Tebbitt 89 (BKL), KP713021, KP713083, KP713305; B. aff. congesta, D.C. Thomas 09-05 (E), JN133306, JN133381, JN133410; B. aff. congesta, D.C. Thomas 09-05 (E), JN133324, JN133399, JN133428; B. consobrina, Lyon B.G. 140666 (LBG), MH207114, MH207526, MH207939; B. convallariodora, B. Sénéchal CRBS/EB201117 (LBG), MH207115, MH207527, MH207940; B. convolvulacea, L.L. Forrest 168 E0020588 (E), KP712951, KP713146, KP713290; B. copelandii, R.R. Rubite R238 (PNH), KR186514, KR186601, KR186774; B. copeyana, B. Sénéchal CRBS/EB201139 (LBG), MH207116, MH207528, MH207941; B. corallina, Lyon B.G. 880164 (LBG), MH207117, –, –; B. cornitepala, B. Sénéchal 130507 (LBG), MH207118, MH207529, MH207942; B. corredorana, RBGE 20071055, MH207119, –, –; B. corrugata, D.C. Thomas 09-02 (E), JF756401, JF756485, JF756569; B. coursii, 20132227 (E), MH207120, MH207530, MH207943; B. crassicaulis, M. Tebbitt 84 (BKL), KP713022, KP713084, KP713300; B. crassipes, Faden 86/67 C.-I Peng 20867 (HAST), KP712967, KP713180, –; B. cremnophila, M. Tebbitt 753 (USZ), KX756282, KX756309, MH207944; B. croatii, D. Gould 100344 (LBG), MH207122, MH207532, MH207945; B. cubensis, M. Tebbitt 66 (BKL), KP713049, KP713170, KP713257; B. cucphuongensis, C.I Peng 20227 (HAST), MH207123, MH207533, MH207946; B. cucphuongensis, CK040 (KUN), MH207124, MH207534, MH207947; B. cucullata, J. P. Grienay 110707 (LBG), MH207125, MH207535, MH207948; B. culasiensis, R.R. Rubite R234 (PNH), KR186459, KR186546, KR186719; B. culasiensis, C.-I Peng P23793 (HAST), KR186458, KR186545, no data; B. curtii, S. Follin 70392 (LBG), MH207126, MH207536, MH207949; B. cyanescens, S06 (SBG), MH207127, MH207537, MH207950; B. cyathophora, no voucher, KP713075, KP713171, KP713255; B. cyathophora, no voucher, KP713075, KP713171, KP713255; B. decora, S. Neale 8C (E), JF756355, JF756439, JF756523; B. demissa, T. Phutthai 221 (E), JF756384, JF756468, JF756552; B. dentatiloba, M. Tebbitt 126 (BKL), KP712983, KP713218, KP713299; B. descoleana, Lyon B.G. 120349 (LBG), MH207128, MH207538, MH207951; B. diadema, D. Permingeat 50584 (LBG), MH207129, MH207539, MH207952; B. dichotoma, Glasgow B.G. 03714766 (E), MH207130, MH207540, MH207953; B. dichotoma, Goldsmith U163 (E), MH207131, MH207541, MH207954; B. dichotoma, Glasgow B.G. 037-147-66 (E), KP713056, KP713177, KP713283; B. dichroa, Glasgow B.G. 001-096-95, KP712931, KP713192, KP713343; B. didyma, D.C. Thomas 08-77 (E), KP712936, KP713129, KP713226; B. dietrichiana, M. Tebbitt 147 (BKL), KP712976, KP713191, KP713271; B. dioica, Chien-I Huang 13651 (E), MH207132, MH207542, MH207955; B. dipetala, D.C. Thomas 100468 (E), JF756341, JF756425, JF756509; B. dipetala, C.-I Peng P22520 (HAST), KR186460, KR186547, KR186720; B. dolichocarpa, P. Wilkie 1014 (E), MH207133, MH207543, MH207956; B. doloisii, PH 115 (SBG), MH207134, –, MH207957; B. domingensis, Lyon B.G. 140495 (LBG), MH207135, MH207544, MH207958; B. dominicalis, Glasgow B.G. 008-139-69 (E), KP713052, KP713169, KP713267; B. dregei, T. McLellan 415 (E), JF756338, JF756422, JF756506; B. dregei, T. McLellan 212 E00198090 (E), KP713014, KP713174, KP713309; B. dregei, C.-I Peng P20868 (HAST), KR186461, KR186548, KR186721; B. dregei, No voucher (AS balcony), MH235380, MH235408, MH235418; B. droopiae, C.P. Puglisi 123 (E), MH207136, MH207545, MH207959; B. dux, C.I Peng 23656 (HAST), MH207137, MH207546, MH207960; B. echinosepala, M. Tebbitt, Swensen & Yeadon 5 (BKL), KP712944, KP713220, KP713278; B. edmundoi, L.L. Forrest 196 (E), KP712994, KP713216, KP713261; B. egregia, L.L. Forrest 173 E00205197 (E), KP712969, KP713153, –; B. elachista, Moonlight 318 (E), KX756285, KX756324, MH207961; B. elachista, Moonlight 318 (E), KX756297, KX756324, MH207962; B. elaegnifolia, Glasgow B.G. 010-079-97 (E), –, MH207549, MH207963; B. elisabethae, T. Phutthai 239 (E), JF756381, JF756465, JF756549; B. aff. elisbethae, D.C. Thomas 08-149 (E), KP712937, KP713123, KP713240; B. elmeri, R.R. Rubite R319 (PNH), KR186462, KR186549, KR186722; B. eminii, C.-I Peng 22515 (HAST), KP712962, KP713193, KP713311; B. engleri, no voucher, KP712984, KP713133, KP713342; B. epipsila, C.-I Peng 21292 (HAST), KP712946, KP713187, KP713318; B. erythrogyna, S. Follin 90517 (LBG), MH207140, MH207550, MH207964; B. aff. erythrogyna, F. Lens 002 (SBG), MH207028, MH207439, MH207856; B. exigua, F.F. Pezzini 146 (E), MH207141, MH207551, MH207965; B. exigua, P.H. Alves de Melo 4993 (HRCB), MH207142, MH207552, MH207966; B. fagifolia, Glasgow B.G. 010-075-71 (E), KP712954, KP713142, KP713293; B. faustinoi, L.L. Forrest 187 (E), KP713008, KP713088, KP713228; B. fenicis, D.C. Thomas 08-119 (E), JF756392, JF756476, JF756560; B. fenicis, C.-I Peng P10794 (HAST), KR186464, KR186551, KR186724; B. fenicis, NK 11979 (HAST), KR186465, KR186552, KR186725; B. ferruginea, A. JaraMuñoz 2739 (COL), MH207143, MH207553, MH207967; B. fischeri, D.C. 41113 (COL), MH207146, MH207557, MH207971; B. fischeri, A. JaraMuñoz 2560 (COL), MH207144, MH207555, MH207969; B. fischeri, P.W. Moonlight & A. Daza 74 (E), –, MH207554, MH207968; B. fischeri, P.W. Moonlight & A. Daza 124 (E), MH207145, MH207556, MH207970; B. fissistyla, L.L. Forrest 157 E00205201 (E), KP713051, KP713173, KP713250; B. fissistyla, Tebbitt, Swensen & Yeadon 8 (BKL), MH235391, MH235415, MH235423; B. cf. fissistyla, Lyon B.G. 890537 (LBG), MH207092, MH207502, MH207918; B. flacca, D.C. Thomas & W.H. Ardi 09-133 (E), JN133312, JN133387, JN133416; B. flagellaris, Chien-I Huang 6010 (E), MH207147, MH207558, MH207972; B. flagellaris, Chien-I Huang 6010 (E), MH207148, MH207559, MH207973; B. flagellaris, S. Rajbhandary & S. Bista S33 (E), JF756353, JF756437, JF756521; B. floccifera, L.L. Forrest 238 (E), JF756343, JF756427, JF756511; B. fluminensis, C.-I Peng 22328 (HAST), KP712939, KP713221, KP713314; B. fluvialis, M. Hughes 1489 (E), MH207150, MH207561, MH207975; B. foliosa, Unknown s.n. E19480286 (E) , KP713060, KP713176, KP713310; B. forbesii, C.-I Peng P22685 (HAST), KR186467, KR186554, KR186727; B. foxworthyii, C.-I Peng P22721 (HAST), KR186468, KR186555, KR186728; B. friburgensis, Lyon B.G. 31318 (LBG), MH207151, MH207562, MH207976; B. froebelii, M. Tebbitt 786 (QCA), KX756288, KX756305, MH207977; B. fruticosa, Lyon B.G. 070728 (LBG), MH207153, MH207564, MH207978; B. fruticosa, F. Pautz 20532 (LBG), –, MH207565, –; B. fulvosetulosa, B. Sénéchal 70728 (LBG), KX756279, KX756331, –; B. furfuracea, B. Sénéchal 31610 (LBG), MH207154, MH207566, MH207979; B. aff. fusca, A. Twyford 677 (E), MH207029, MH207440, MH207857; B. fuscisetosa, 20120800 (E), MH207155, –, MH207980; B. gabaldonensis, C.-I Peng P23356 (HAST), KR186469, KR186556, KR186729; B. gabonensis, L.L. Forrest 230 E00205158 (E), KP712985, KP713215, KP713321; B. galeolepis, Kreb s.n. (SBG), MH207156, –, MH207981; B. gamolepis, A. JaraMuñoz 2616 (COL), MH207157, MH207567, MH207982; B. garagarana, B. Sénéchal CRBS/EB201120 (LBG), MH207158, MH207568, MH207983; B. gardneri, D. Permingeat 50722 (LBG), MH207159, MH207569, MH207984; B. gehrtii, M. Tebbitt 85 (BKL), MH207160, MH207570, MH207985; B. gehrtii, C.-I Peng 21202 (HAST), KP712941, KP713222, KP713313; B. geraniifolia, M. Tebbitt & A. Daza 840 (E), MH207162, MH207572, MH207987; B. geraniifolia, P.W Moonlight & A. Daza 116 (E), KX756283, KX756311, MH207988; B. germaniana, M. Tebbitt 709 (USZ), MH207164, MH207574, MH207989; B. gesnerioides, P.W. Moonlight & A. Daza 183 (E), –, MH207575, MH207990; B. gesnerioides, P.W. Moonlight & A. Daza 184 (E), –, MH207576, –; B. gigabracteata, C.I Peng 22174 (HAST), MH207165, MH207577, MH207991; B. gitingensis, R.R. Rubite R255 (PNH), KR186470, KR186557, KR186730; B. glabra, Z. Badcock 7 (E), MH207166, –, MH207992; B. glabra, s.n. (HAST), KP712943, KP713145, KP713291; B. glabra, P.W. Moonlight & A. Daza 38 (E), MH207168, MH207579, MH207994; B. glabra, P.W. Moonlight & A. Daza 26 (E), MH207169, –, MH207995; B. glabra, P.W. Moonlight & A. Daza 153 (E), MH207170, MH207580, MH207996; B. glabra, No voucher, MH235379, –, –; B. glandulifera, M. Gardner 6608 (E), KX756294, KX756328, MH207997; B. goegoensis, D.C. Thomas 08-107 (E), JF756376, JF756460, JF756544; B. goudotii, V. Plana 120 (E), JF756347, JF756431, JF756515; B. gracilis, Z. Badcock 9 (E), KP713004, KP713078, KP713260; B. grandis, D.C. Thomas 08-145 (E), JF756351, JF756435, JF756519; B. griffithiana, C.I Peng 20851 (HAST), MH207172, MH207582, MH207998; B. grisea, Lyon B.G. 11433 (LBG), MH207173, MH207583, MH207999; Version of Record 319 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) TAXON 67 (2) • April 2018: 267–323 Appendix 1. Continued. B. guaduensis, C.-I Peng 22518 (HAST), KP713054, KP713178, KP713294; B. cf. guaduensis, DC 43489 (COL), MH207093, MH207503, MH207919; B. gueritziana, C.-I Peng P22311 (HAST), KR186471, KR186558, KR186731; B. gueritziana, C.-I Peng P22342 (HAST), KR186472, KR186559, KR186732; B. aff. gueritziana, P22344 (HAST), MH235385, MH235401, MH235426; B. gutierrezii, P. Blanc s.n. (E), KR186473, KR186560, KR186733; B. guttapila, D.C. Thomas 08-81 (E), JF756405, JF756489, JF756573; B. hainanensis, C.I Peng 19543 (HAST), MH207174, MH207584, MH208000; B. hainanensis, Y.M. Shui & al. B2011044 (KUN), MH207175, MH207585, MH208001; B. handelii, J. Durisseau 130285 (LBG), MH207176, MH207586, MH208002; B. harauensis, D.C. Thomas 09-134 (E), JN133330, JN133405, JN133434; B. hatacoa, D.C. Thomas 08-110 (E), JF756354, JF756438, JF756522; B. hekensis, D.C. Thomas & W.H. Ardi 08-43 (E), JN133313, JN133388, JN133417; B. heliantha, R.T. Pennington & A.Daza 1113 (E), –, MH207587, –; B. henrilaportei, J. Duruisseau 060667 (LBG), MH207177, MH207588, MH208003; B. heracleifolia, A. Twyford 597 (E), MH207178, MH207589, MH208004; B. cf. heracleifolia, B. Sénéchal 50849 (LBG), MH207094, –, MH207920; B. herbacea, L.L. Forrest 163 E00205153 (E), KP713015, KP713202, MH208005; B. hernandioides, D.C. Thomas 08-118 (E), JF756393, JF756477, JF756561; B. hernandioides, C.-I Peng P21006 (HAST), KR186474, KR186561, KR186734; B. hernandioides, R.R. Rubite R106 (PNH), KR186475, KR186562, KR186735; B. heydei, P22624 (HAST), MH207180, MH207591, –; B. heydei, Lyon B.G. 140136 (LBG), MH207181, MH207592, –; B. heydei, C.-I Peng 22624 (HAST), KP713030, KP713114, –; B. hirtella, P.W. Moonlight & A. Daza 163 (E), MH207182, MH207593, MH208006; B. hirtella, P.W. Moonlight & A. Daza 286 (E), MH207183, MH207594, MH208007; B. hispida var. cucullifera, M. Tebbitt 122 (BKL), KP713064, KP713159, KP713281; B. hispidissima, Kinho & A.D. Poulson 168 (E), JN133314, JN133389, JN133418; B. hitchcockii, A. JaraMuñoz 2758 (COL), KX756291, KX756313, MH208008; B. hitchcockii, EQJD04 (LBG), KX756292, KX756316, MH208009; B. hitchcockii, P.W Moonlight & A. Daza 123 (E), KX756290, KX756315, MH208010; B. hitchcockii, P.W. Moonlight & A. Daza 151 (E), MH207187, MH207598, MH208011; B. hoehneana, 20131494 (E), MH207188, MH207599, MH208012; B. holtonis, Forrest 152 (E), KP713062, KP713175, KP713307; B. holtonis, Glasgow B.G. 01112984 (E), MH207190, MH207601, MH208014; B. holtonis, A. JaraMuñoz 2382 (COL), MH207191, MH207603, MH208015; B. holttumii, J. Duruisseau 100498 (LBG), MH207193, MH207602, MH208017; B. holttumii, C.I Peng 22736 (HAST), MH207192, –, MH208016; B. hookeriana, Lyon B.G. 880089 (LBG), MH207194, MH207604, MH208018; B. hugelii, B. Sénéchal 20121 (LBG), MH207195, MH207605, MH208019; B. hughesii, C.-I Peng P23466 (HAST), KR186476, KR186563, KR186736; B. hughesii, C.-I Peng P23475 (HAST), KR186477, KR186564, KR186737; B. humbertii, 20132230 (E), MH207196, –, MH208020; B. humilis, D. Scherberich 1077 (LBG), –, MH207606, MH208021; B. hydrocotylifolia, M. Tebbitt 86 (BKL), KP713039, KP713085, KP713246; B. hymenophylla, T. Phutthai 232 (E), JF756382, JF756466, JF756550; B. ignorata, C.-I Peng P22725 (HAST), KR186478, KR186565, KR186738; B. imbricata, F. Lens 075 (SBG), MH207197, MH207607, MH208022; B. imperialis, Caen 8979 (E), MH207198, MH207608, MH208023; B. imperialis, Glasgow B.G. 0028979 (E), MH207199, MH207609, –; B. incarnata, A. Twyford 587 (E), KP713065, KP713090, KP713232; B. aff. incarnata, A. Twyford 587 E00619704 (E), KP713065, KP713090, KP713232; B. inobangensis, F. Lens 062 (SBG), MH207200, MH207610, MH208024; B. inostegia, F. Lens 021 (SBG), MH207201, MH207611, MH208025; B. insularis, M. Peixoto 041658 (LBG), MH207202, MH207612, MH208026; B. integerrima, M. Tebbitt 69 (BKL), KP713000, KP713099, KP713242; B. involucrata, Tebbitt, Swensen & Yeadon 23 (BKL), KP712995, KP713082, KP713332; B. involucrata, 20070426 (E), MH207204, MH207614, MH208028; B. isoptera, SUBOE 74 (E), MH207205, MH207615, MH208029; B. itaguassuensis, Lyon B.G. 10570 (LBG), MH207206, MH207616, MH208030; B. itatiensis, Lyon B.G. 140316 (LBG), MH207207, MH207617, MH208031; B. jaguarensis, Lyon B.G. 120352 (LBG), MH207208, MH207618, MH208032; B. jamilahana, NHZ14 (SBG), MH207209, MH207619, MH208033; B. jocelinoi, C.-I Peng 23331 (HAST), KP712982, KP713143, KP713315; B. joffrei, S15 (SBG), MH207210, MH207620, MH208034; B. johnstonii, no voucher, KP712996, KP713134, KP713339; B. juliana, M. Tebbitt 132 (BKL), KP712950, KP713158, KP713285; B. karangputihensis, C. Puglesi & al. 53 (E), MH207211, –, –; B. karwinskyana, RM-04-16/P20880 (HAST), KP713037, KP713086, KP713316; B. kemumuensis, D. Girmansyah & M. Hughes 1499 (E), MH207212, MH207621, MH208035; B. kingiana, D.C. Thomas 08-102 (E), JF756374, JF756458, JF756542; B. kingiana, C.-I Peng P21226 (HAST), KR186479, KR186566, KR186739; B. kisuluana, Forrest 215 (E), MH207213, MH207622, MH208036; B. kisuluana, Lyon B.G. 120318 (LBG), –, MH207623, MH208037; B. klemmei, R.R. Rubite R182 (PNH), KR186480, KR186567, KR186740; B. koksunii, J. Durisseau 140138 (LBG), MH207214, MH207624, MH208038; B. komoensis, Tchimbélé 01/2012 (LBG), MH207215, MH207625, MH208039; B. komoensis, Tchimbélé 01/2012 (LBG), –, MH207626, MH208040; B. koordersii, D.C. Thomas & W.H. Ardi 08-62 (E), JF756407, JF756491, JF756575; B. krystofii, M. Tebbitt 701 (USZ), MH207216, MH207627, MH208041; B. kudoensis, C.P. Puglisi 67 (E), MH207217, MH207628, MH208042; B. kuhlmannii, Glasgow B.G. 00402907 (E), KP712942, KP713157, KP713272; B. labiensis, NHZ24 (SBG), MH207219, MH207630, –; B. lanceolata, Tebbitt MBG02 (E), KP713068, KP713101, KP713253; B. lanceolata, B. Sénéchal 20374 (LBG), KX756289, KX756332, MH208045; B. laruei, D.C. Thomas 08-138 (E), JF756403, JF756487, JF756571; B. laruei, Puglisi et al. CP225 (E), –, MH235412, MH235417; B. lasioura, D.C. Thomas & W.H. Ardi 09-110 (E), JN133315, JN133390, JN133419; B. leprosa, Permingeat, D. 274 (LBG), –, –, MH235433; B. letouzeyi, J. Duruisseau 110677 (LBG), MH207222, –, MH208046; B. limprichtii, J. Durisseau 60823 (LBG), MH207223, MH207633, MH208047; B. lindleyana, 20051412 (E), MH207224, MH207634, MH208048; B. listada, C.-I Peng 21053 (HAST), KP712980, KP713155, KP713296; B. lithophila, Y.M. Shui & al. B2012064 (KUN), MH207225, MH207635, MH208049; B. longifolia, Neale 11C (E), JF756368, JF756452, JF756536; B. longipetiolata, C.-I Peng 22519 (HAST), KP712970, KP713181, –; B. longiscapa, R.R. Rubite R298 (PNH), KR186482, KR186569, KR186742; B. longiscapa, R.R. Rubite R309 (PNH), KR186483, KR186570, KR186743; B. lophoptera, M. Tebbitt & A. Daza 807 (E), MH207226, MH207636, –; B. lophoptera, P.W. Moonlight & A. Daza 231 (E), MH207227, MH207637, MH208050; B. loranthoides subsp. loranthoides, E00205240 (E), KP713029, KP713213, –; B. lubbersii, L.L. Forrest 194 (E), KP712981, KP713100, KP713237; B. ludicra, P22052/RM08 (HAST), KP713009, KP713092, –; B. ludicra, A. Twyford 680 (E), MH207229, MH207639, MH208051; B. ludicra, A. Twyford 687 (E), MH207230, MH207640, MH208052; B. ludwigii, P.W. Moonlight & A. Daza 96 (E), MH207231, MH207641, –; B. ludwigii, M. Tebbitt 780 (QCA), MH207232, MH207642, –; B. ludwigii, M. Tebbitt 791 (QCA), –, MH207643, –; B. lugonis, Lyon B.G. 150110 (LBG), MH207233, MH207644, MH208053; B. lunaris, Lyon B.G. 140256 (LBG), MH207234, –, MH208054; B. lutea, JaraMuñoz AMB 332 (COL), KX756287, KX756320, –; B. luxurians, M. Tebbitt 158 (BKL), KP712940, KP713223, KP713288; B. luzonensis, C.-I Peng P22344 (HAST), KR186446, KR186533, KR186707; B. luzonensis, R.R. Rubite R316 (PNH), KR186484, KR186571, KR186744; B. luzonensis, R.R. Rubite R420 (PNH), KR186485, KR186572, KR186745; B. lyallii f. lyallii, J. Duruisseau 130264 (LBG), MH207236, –, MH208055; B. lyallii f. masoalensis, 20132223 (E), MH207237, –, MH208056; B. lyman-smithii, M. Tebbitt 116 (BKL), –, MH207646, MH208057; B. lyniceorum, RM04-01 (HAST), KP713041, KP713197, –; B. macduffieana, Glasgow B.G. 002-004-86, KP712928, KP713210, KP713276; B. macintyreana, D.C. Thomas 07-28 (E), JN133316, JN133391, JN133420; B. macrocarpa, Lyon B.G. 100726 (LBG), –, MH207647, MH208058; B. maculata, Glasgow B.G. 008-151-95, KP712930, KP713206, KP713274; B. madecassa, 20132231 (E), MH207238, MH207648, MH208059; B. madecassa, C.-I Peng 22318 (HAST), KP712973, KP713179, –; B. majungaensis, 20132228 (E), MH207239, –, MH208060; B. malabarica, L.L. Forrest 288 (E), JF756342, JF756426, JF756510; B. mamutensis, F. Lens 029 (SBG), MH207240, MH207649, MH208061; B. manicata, A. Twyford 590 (E), MH207241, MH207650, MH208062; B. manicata, Z. Badcock 8 (E), KP712989, KP713131, KP713259; B. manillensis, R.R. Rubite R256 (PNH), KR186486, KR186573, KR186746; B. mariannensis, J. Duruisseau 80396 (LBG), KX756299, KX756329, –; B. mariti, RM-08-352 (HAST), KP713035, KP713113, –; B. martabanica, Peng 24184 (HAST), –, MH244437, MH244440; B. masarangensis, D.C. Thomas 09-131 (E), JF756409, JF756493, JF756577; B. masoniana, L.L. Forrest s.n. (E), JF756372, JF756456, JF756540; B. masoniana, C.-I Peng P21411 (HAST), KR186487, KR186574, KR186747; B. maynensis, P.W. Moonlight & A. Daza 134 (E), MH207242, MH207651, –; B. maynensis, P.W. Moonlight & A. Daza 185 (E), MH207243, MH207654, MH208064; B. maynensis, Lyon B.G. 1050058 (LBG), MH207244, MH207655, MH208065; B. maynensis, P.W. Moonlight & A. Daza 162 (E), –, MH207652, MH208063; B. maynensis, P.W. Moonlight & A. Daza 176 (E), –, MH207653, –; B. maynensis, C.-I Peng s.n. (HAST), KP713063, KP713141, –; B. mazae, Hollands 008 E00668725 (E), KP713002, KP713116, KP713238; B. aff. mekongensis, D.C. Thomas & W.H. Ardi 09-108 (E), JN133308, JN133383, JN133412; B. mendumae, D.C. Thomas 07-27 (E), JN133317, JN133392, JN133421; B. meridensis, A. JaraMuñoz 2730 (COL), –, MH207656, MH208066; B. meridensis, L.L. Forrest 181 E00205112 (E), KP713057, KP713132, KP713308; B. meridensis, Glasgow Botanic Garden 014-124-82, KP713057, KP713132, KP713308; B. cf. meridensis, A. JaraMuñoz 2342 (COL), MH207095, MH207504, MH207921; 320 Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Appendix 1. Continued. B. merrilliana, C.-I Peng P23765 (HAST), KR186488, KR186575, KR186748; B. mexicana, J. Duruisseau 140145 (LBG), MH207245, –, MH208067; B. meyerijohannis, S. Barber & al. 38 (E), MH207246, MH207657, MH208068; B. micranthera, M. Tebbitt 702 (USZ), MH207247, MH207658, MH208069; B. micranthera, T. Sarkinen 2029 (E), KP713032, KP713204, –; B. micranthera, T. Sarkinen 2043 (E), KP713066, KP713198, –; B. micranthera, T. Sarkinen 2144 (E), KP713071, KP713199, –; B. microsperma, C.-I Peng s.n. (HAST), KP713010, KP713194, KP713325; B. mindorensis, R.R. Rubite R326 (PNH), KR186489, KR186576, KR186749; B. mindorensis, C.-I Peng P23456 (HAST), KR186490, KR186577, KR186750; B. minor, M. Tebbitt 74 (BKL), KP713053, KP713167, KP713265; B. minutifolia, J. Duruisseau 66060 (LBG), –, MH207659, –; B. molleri, D. Permingeat 327 (LBG), –, –, MH208070; B. mollicaulis, Glasgow B.G. 014-254-99 (E), KP713044, KP713212, KP713338; B. monadelpha, T. Sarkinen 2205 (E), KP713005, KP713117, MH208071; B. monadelpha, P.W. Moonlight & A. Daza 69 (E), MH207249, MH207661, MH208072; B. monadelpha, M. Tebbitt & A. Daza 833 (E), MH207250, MH207662, –; B. monadelpha, P.W. Moonlight & A. Daza 148 (E), MH207251, MH207663, MH208073; B. morsei, Unknown s.n. (E), JF756373, JF756457, JF756541; B. motozintlensis, A. Twyford 337 (E), MH207252, MH207664, MH208074; B. moyesii, Lyon B.G. 110675 (LBG), MH207253, MH207665, MH208075; B. multangula, D.C. Thomas & W.H. Ardi 08-90 (E), JF756364, JF756448, JF756532; B. aff. multangula, D.C. Thomas 09-85 (E), KP712963, KP713125, KP713239; B. multijugata, P. Wilkie 768 (E), JF756404, JF756488, JF756572; B. multinervia, M. Tebbitt 131 (BKL), KP713023, KP713163, KP713258; B. multinervia, M. Tebbitt 131 (BKL), KP713023, KP713163, KP713258; B. multistaminea, RBGE 20071056, MH207254, MH207666, MH208076; B. muricata, W.H. Ardi & D.C. Thomas WI27 (E), JF756378, JF756462, JF756546; B. murina, C.I Peng 24137 (HAST), MH207255, MH207667, MH208077; B. nana, Lyon B.G. 140495 (LBG), MH207256, MH207668, –; B. negrosensis, P. Wilkie 76 (E), JF756411, JF756495, JF756579; B. nelumbifolia, R. Hollands 009 (E), KP713040, KP713077, KP713230; B. nelumbifolia, TNY19 (HAST), MH235378, MH235398, MH235421; B. nelumbiifolia, C.-I Peng P20879 (HAST), KR186491, KR186578, KR186751; B. nepalensis, B. Sénéchal 41155 (LBG), –, –, MH208078; B. nepalensis, C.I Peng 20854 (HAST), MH207257, MH207669, MH208079; B. nigritarum, R.R. Rubite R419 (PNH), KR186492, KR186579, KR186752; B. nigritarum, R.R. Rubite R406 (PNH), KR186493, KR186580, KR186753; B. nigritarum, C.-I Peng P23855 (HAST), KR186494, KR186581, KR186754; B. nigritarum, C.-I Peng P23373 (HAST), KR186495, KR186582, KR186755; B. nigritarum, C.-I Peng P23451 (HAST), KR186496, KR186583, KR186756; B. nigritarum, C.-I Peng P23372 (HAST), KR186497, KR186584, KR186757; B. nigritarum, C.-I Peng P23586 (HAST), KR186498, KR186585, KR186758; B. nigritarum, C.-I Peng P23858 (HAST), KR186499, KR186586, KR186759; B. ningmingensis, C.-I Peng P20322 (HAST), KR186500, KR186587, KR186760; B. nitida, Lyon B.G. 920013 (LBG), MH207258, MH207670, –; B. nobmanniae, D.C. Thomas 09-123 (E), JN133318, JN133393, JN133422; B. nossibea, 110200080 (HAST), KP712974, KP713183, –; B. nosymangabensis, J. Duruisseau & B. Senechal s.n. (LBG), MH235384, MH235413, MH235424; B. nothobaramensis, S08 (SBG), MH207259, MH207671, MH208080; B. nurii, C. Salamone 50627 (LBG), MH207260, MH207672, MH208081; B. nurii, FRI 824 (FRIM), MH207261, MH207673, MH208082; B. oaxacana, no voucher, KX756280, KX756325, –; B. obliqua, Lyon B.G. 80132 (LBG), MH207262, MH207674, MH208083; B. oblongifolia, F. Lens 028 (SBG), MH207263, MH207675, MH208084; B. oblongifolia, F. Lens 014 (SBG), MH207264, MH207676, MH208085; B. obovoidea, T. Phutthai 244 (E), JF756386, JF756470, JF756554; B. obscura, E20030603 (E), KP712945, KP713219, KP713282; B. obtecticaulis, P.W. Moonlight & A. Daza 212 (E), MH207265, MH207677, MH208086; B. obtusifolia, C.-I Peng P23828 (HAST), KR186501, KR186588, KR186761; B. octopetala, P.W. Moonlight & A. Daza 67 (E), MH207266, MH207678, MH208087; B. octopetala, P.W. Moonlight & A. Daza 70 (E), MH207267, –, –; B. octopetala, M. Tebbitt 790 (QCA), MH207268, MH207679, MH208088; B. octopetala, M. Tebbitt & A. Daza 825 (E), MH207269, MH207680, –; B. octopetala, M. Tebbitt & A. Daza 842 (E), MH207270, MH207681, –; B. octopetala, M. Tebbitt & A. Daza 844 (E), MH207271, MH207682, MH208089; B. octopetala, P.W. Moonlight & A. Daza 208 (E), MH207272, –, MH208090; B. odetiantha, Glasgow B.G. 009-007-97 (E), KP712975, KP713190, KP713270; B. odorata, L.L. Forrest 158 E00205104 (E), KP713059, KP713168, KP713266; B. olbia, L.L. Forrest 185 E00205103 (E), KP712933, KP713186, KP713306; B. olivacea, C.P. Puglisi 239 (E), MH207273, MH207683, MH208091; B. olsoniae, J. Durisseau 60665 (LBG), MH207274, MH207684, MH208092; B. olsoniae, M. Tebbitt, Swensen & Yeadon 18 (BKL), KP713001, KP713156, KP713297; B. opuliflora, Lyon B.G. 21735 (LBG), MH207275, MH207685, MH208093; B. oxyanthera, Glasgow B.G. 00411378 (E), MH207276, MH207686, MH208094; B. oxyanthera, Lyon B.G. 11285 (LBG), –, MH207687, –; B. oxyloba, L.L. Forrest 279 E00205102 (E), JF756335, JF756419, JF756503; B. oxysperma, R.R. Rubite R213 (PNH), KR186502, KR186589, KR186762; B. oxysperma, C.-I Peng P23015 (HAST), KR186503, KR186590, KR186763; B. ozotothrix, D.C. Thomas & W.H. Ardi 08-58 (E), JN133319, JN133394, JN133423; B. pachypoda, Glasgow B.G. 015-107-89, KP712926, –, KP713268; B. palawanensis, C.-I Peng P23453 (HAST), KR186504, KR186591, KR186764; B. paleata, M. Peixoto 90309 (LBG), MH207277, MH207688, MH208095; B. palmata, M. Moller 01-127 (E), JF756360, JF756444, JF756528; B. papyraptera, NHZ08 (SBG), MH207278, MH207689, –; B. paracauliflora, C.I Peng 22309 (HAST), MH207279, MH207690, MH208096; B. paranaensis, M. Tebbitt, Swensen & Yeadon 25 (BKL), KP712947, KP713147, KP713298; B. parcifolia, M. Tebbitt 769 (QCA), MH207280, MH207691, MH208097; B. parcifolia, M. Tebbitt 779 (QCA), –, MH207692, MH208098; B. parcifolia, M. Tebbitt & A. Daza 845 (E), MH207281, MH207693, –; B. parcifolia, M. Tebbitt 783 (QCA), KX756277, KX756317, –; B. parilis, A. Mauriere 880035 (LBG), MH207282, MH207694, MH208099; B. parviflora, P.W. Moonlight & A. Daza 37 (E), MH207283, MH207695, MH208100; B. parviflora, P.W. Moonlight & A. Daza 155 (E), MH207284, MH207696, MH208101; B. pasamanensis, D. Girmansyah & M. Hughes 1506 (E), MH207285, MH207697, MH208102; B. pavonina, S. Neale 9C (E), JF756356, JF756440, JF756524; B. pearcei, Lyon B.G. 10128 (LBG), MH207286, MH207698, MH208103; B. pectennervia, A. JaraMuñoz 2621 (COL), MH207287, MH207699, MH208104; B. pedatifida, Lyon B.G. 21121 (LBG), MH207288, MH207700, MH208105; B. peltata, A. Mauriere 870118 (LBG), –, MH207701, –; B. peltata, 233332 (HAST), KP712988, KP713104, –; B. peltata, M. Tebbitt 153 (BKL), KP712992, KP713105, KP713235; B. peltatifolia, Y.M. Shui & al. Q031 (KUN), MH207289, MH207702, MH208106; B. pendula, DCT 09-03 (culta. SING), JF756395, JF756479, JF756563; B. perakensis, B. Sénéchal 110694 (LBG), MH207291, MH207704, MH208108; B. perdusenii, B. Sénéchal 130513 (LBG), MH207290, MH207703, MH208107; B. peruviana, P.W. Moonlight & A. Daza 173 (E), –, MH207705, –; B. peruviana, Moonlight 39 (E), –, –, MH208109; B. peruviana, M. Tebbitt & A. Daza 832 (E), –, MH207789, –; B. petastifolia, M. Tebbitt 83 (BKL), KP712935, KP713152, KP713269; B. phantasma, M. Tebbitt 757 (USZ), MH207292, MH207706, MH208110; B. phantasma, M. Tebbitt 721 (USZ), MH207293, MH207707, MH208111; B. picta, Chien-I Huang 5993 (E), MH207294, MH207708, MH208112; B. pilgeriana, M. Peixoto 120314 (LBG), MH207295, MH207709, MH208113; B. pinetorum, Hollands 006 E00668796 (E), KP713067, KP713091, KP713312; B. piurensis, Moonlight & Daza (E), MH207296, MH207710, MH208114; B. piurensis, P.W Moonlight & A. Daza 111 (E), KX756276, KX756318, –; B. platanifolia, Glasgow B.G.00604504 (E), MH207297, MH207711, MH208115; B. platanifolia, Lyon B.G. 42262 (LBG), MH207298, MH207712, MH208116; B. plebeja, A. Twyford 606 (E), MH207299, MH207713, MH208117; B. pleiopetala, M. Tebbitt & A. Daza 813 (E), MH207300, MH207714, MH208118; B. pleiopetala, P.W. Moonlight & A. Daza 295 (E), MH207301, MH207715, –; B. pleiopetala, P.W. Moonlight & A. Daza 297 (E), MH207302, MH207716, MH208119; B. plumieri, Lyon B.G. 100703 (LBG), MH207303, MH207717, MH208120; B. pluvialis, Lyon B.G. 940319 (LBG), MH207304, MH207718, MH208121; B. poculifera, L.L. Forrest 234 (E), JF756348, JF756432, JF756516; B. poilanei, Tran Huu Dang s.n. (E), MH207305, MH207719, MH208122; B. poliloensis, L.L. Forrest 139 (E), JF756412, JF756496, JF756580; B. polyandra, Lyon B.G. 140319 (LBG), MH207306, MH207720, MH208123; B. polygonata, M. Tebbitt 139 (BKL), KP712987, KP713106, KP713234; B. polygonata, Glasgow B.G. 01712370 (E), MH207308, MH207722, MH208125; B. polygonata, Glasgow B.G. 01712370 (E), MH207309, MH207723, MH208126; B. polygonoides, van der Burg 244 (WAG), JF756336, JF756420, JF756504; B. polypetala, M. Tebbitt & A.Daza MT839b (E), MH207310, MH207724, –; B. polypetala, P.W. Moonlight & A. Daza 113 (E), MH207311, MH207725, MH208127; B. polypetala, P.W. Moonlight & A. Daza 119 (E), MH207312, MH207726, MH208128; B. polypetala, Tebbitt 787 (USZ), MH235388, MH235403, MH235429; B. polypetala, Tebbitt 828 (USZ), MH235389, MH235409, –; B. popenoei, E20110200, KP713020, KP713095, KP713302; B. princeps, D. Permingeat 21701 (LBG), MH207313, MH207727, MH208129; B. pringlei, Glasgow B.G. 01800283 (E), MH207314, MH207728, –; B. pringlei, M. Tebbitt 130 (BKL), KP713074, KP713119, KP713319; B. prionota, D.C. Thomas & W.H. Ardi 09-97 (E), JN133320, JN133395, JN133424; B. pruniata, Glasgow B.G. 04010383 (E), MH207315, MH207729, MH208130; B. pseudodaedalea, A. Twyford 1066 (E), MH207316, MH207730, MH208131; B. pseudodryadis, M. Moeller 121890 (E), MH207317, MH207731, –; B. pseudolateralis, D.C. Thomas & W.H. Ardi 08-23 (E), JF756408, JF756492, JF756576; B. pseudolubbersii, Glasgow B.G. 045-023-92 (E), KP713072, KP713203, KP713329; Version of Record 321 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) TAXON 67 (2) • April 2018: 267–323 Appendix 1. Continued. B. pseudopleiopetala, M. Tebbitt & A. Daza 827 (E), MH207318, –, –; B. pteridiformis, Lyon B.G. s.n. (LBG), MH207319, MH207732, MH208132; B. pulchella, M. Peixoto 41666 (LBG), MH207320, MH207733, MH208133; B. pululahuana, A. JaraMuñoz 2659 (QCA), MH207321, MH207734, –; B. pulvinifera, J. Duruisseau 140148 (LBG), MH207322, MH207735, –; B. purpusii, M. Tebbitt 70 (BKL), KP713028, KP713087, KP713251; B. puspitae, C. Puglesi & al. 134 (E), MH207323, MH207736, MH208134; B. pustulata, MBG 1999-01-01, KP712998, KP713093, KP713227; B. puttii, S. Suddee 3375 (E), KP712959, KP713201, KP713326; B. quadrialata, Lyon B.G. 80424 (LBG), –, MH207737, MH208135; B. rabilii, S. Suddee 3371 (E), KP712968, KP713164, KP713328; B. racemosa, D. Girmansyah & M. Hughes 1509 (E), MH207324, MH207738, MH208136; B. racemosa, D. Girmansyah & M. Hughes 1509 (E), MH207325, –, –; B. rachmatii, Kreb s.n. (SBG), MH207326, MH207739, MH208137; B. radicans, Glasgow B.G. 009-089-95 (E), JF756345, JF756429, JF756513; B. radicans, Glasgow B.G. 009-089-95 (E), JF756345, JF756429, JF756513; B. rajah, Girmansyah & al. DEDEN1497 (E), –, MH207740, MH208138; B. rajah, Lyon B.G. 880168 (LBG), MH207327, MH207741, MH208139; B. rajah, FRIM 47082 (FRIM), MH207328, MH207742, MH208140; B. rantemarioensis, Kreb s.n. (SBG), MH207329, MH207743, MH208141; B. rantemarioensis, D.C. Thomas & W.H. Ardi 09-78 (E), JN133321, JN133396, JN133425; B. reginula, FRIM 82471 (FRIM), MH207330, MH207744, MH208142; B. reniformis, Suksuwan 26 E00198096 (E), KP712952, KP713149, KP713292; B. rhoephila, J. Durisseau 130617 (LBG), MH207331, MH207745, MH208143; B. rigida, Lyon B.G. 22147 (LBG), MH207332, MH207746, MH208144; B. robusta, D.C. Thomas & W.H. Ardi 08-133 (E), JF756363, JF756447, JF756531; B. rossmanniae, P. W. Moonlight & A. Daza 315 (E), KX756302, MH207747, MH208145; B. rossmanniae, D. Scherberich 1095 (LBG), KX756300, KX756330, MH208146; B. rotundifolia, Lyon B.G. 10553 (LBG), MH207335, –, MH208147; B. roxburghii, D.C. Thomas 08-103 (E), JF756371, JF756455, JF756539; B. rubella, Chien-I Huang 6000 (E), MH207336, MH207749, MH208148; B. rubida, S. Follin 110183 (LBG), MH207337, MH207750, MH208149; B. rubiteae, R.R. Rubite R356 (PNH), KR186505, KR186592, KR186765; B. rubriflora, M. Peixoto 41645 (LBG), MH207338, –, MH208150; B. rufa, Lyon B.G. 140648 (LBG), MH207339, MH207751, MH208151; B. rufipila, R.R. Rubite R265 (PNH), KR186506, KR186593, KR186766; B. rufoserica, M. Peixoto s.n. (LBG), MH207340, MH207752, MH208152; B. sageaensis, Kreb s.n. (SBG), MH207341, MH207753, MH208153; B. salaziensis, Lyon B.G. 100629 (E), MH207342, MH207754, MH208154; B. salesopolensis, Lyon B.G. 140322 (LBG), –, MH207755, –; B. salicifolia, Lyon B.G. 910693 (LBG), MH207343, MH207756, –; B. samhaensis, D.C. Thomas 09-01 (E), JF756339, JF756423, JF756507; B. sanguinea, C.-I Peng 21284 (HAST), KP712978, KP713188, KP713317; B. sanguineopilosa, D.C. Thomas & W.H. Ardi 09-125 (E), JN133322, JN133397, JN133426; B. santos-limae, C.-I Peng 21320 (HAST), KP713016, KP713126, –; B. sarcophylla, M. Tebbitt 153 (BKL), KP712986, KP713079, KP713236; B. sartorii, M. Tebbitt 153 (BKL), MH207344, MH207757, –; B. sartorii, A. Twyford 448 (E), MH207345, MH207758, MH208156; B. scabrida, Brown, K. 21551 (LBG), –, –, –; B. scharffiana, De Lacheisserie 890443 (LBG), MH207346, MH207759, MH208157; B. scharffii, M. Tebbitt 124 (BKL), KP712961, KP713224, KP713286; B. schmidtiana, E20080890 (E), KP713058, KP713120, KP713264; B. sciaphila, J. Duruisseau 120357 (LBG), –, MH207760, MH208158; B. scottii, M. Hughes 1569 (E), MH207347, MH207761, MH208159; B. scottii, C. Puglesi & al. 217 (E), MH207348, MH207762, MH208160; B. scutifolia, J. Duruisseau 60659 (LBG), –, MH207763, MH208161; B. scutifolia, C.I Peng 23324 (HAST), MH207349, MH207764, MH208162; B. sect. Gobenia, No voucher, MH235377, –, –; B. seemanniana, no voucher, MH207350, MH207765, MH208163; B. segregata, Duruisseau 140144 (LBG), KX756281, KX756314, –; B. semidigitata, Lyon B.G. 140324 (LBG), MH207351, MH207766, MH208164; B. semiovata, P.W. Moonlight & A. Daza 172 (E), MH207352, MH207767, –; B. semiovata, P.W. Moonlight & A. Daza 127 (E), MH207353, MH207768, MH208165; B. sericoneura, RBGE 20071052, MH207354, MH207769, MH208166; B. sericoneura, A. Twyford 854 (E), MH207355, MH207770, MH208167; B. sericoneura, A. Twyford 1110 (E), MH207356, MH207771, MH208168; B. sericoneura, M. Tebbitt 133 (BKL), KP713012, KP713089, KP713303; B. serotina, M. Tebbitt 773 (QCA), MH207357, MH207772, MH208169; B. serotina, M. Tebbitt 776 (QCA), KX756284, KX756319, MH208170; B. serratipetala, Woods, P.J.B. 1916 (E), JF756413, JF756497, JF756581; B. sibthorpoides, Tan PC 82520 (FRIM), MH207359, MH207774, MH208171; B. sibthorpoides, Mo. Hairul FRI 60095 (FRIM), MH207360, MH207775, MH208172; B. sibutensis, S11 (SBG), MH207361, –, MH208173; B. siccacaudata, D.C. Thomas 09-60 (E), JF756418, JF756502, JF756586; B. sikkimensis, D.C. Thomas 08-144 (E), JF756359, JF756443, JF756527; B. silletensis, D.C. Thomas 08-104 (E), JF756370, JF756454, JF756538; B. silverstonii, A. JaraMuñoz 2383 (COL), MH207362, MH207776, MH208174; B. simolapensis, C. Puglesi & al. 228 (E), –, MH207777, MH208175; B. sinofloribunda, Y.M. Shui & al. B2003009 (KUN), MH207363, MH207778, MH208176; B. sizemoreae, D.C. Thomas 08-111 (E), JF756361, JF756445, JF756529; B. smilacina, Cazaux, A. 0140454R (LBG), MH207364, MH207779, –; B. smithiae, Chamchamroon 3662 (E), JF756389, JF756473, JF756557; B. socotrana, T. Miller 19210/10 (E), JF756340, JF756424, JF756508; B. solananthera, Glasgow B.G. 021-070-04 (E), KP712999, KP713098, KP713243; B. solaniflora, A. JaraMuñoz 2564 (COL), MH207365, MH207780, MH208177; B. solimutata, E20110208 (E), KP713042, KP713207, KP713337; B. sousae, A. Twyford 20 (E), MH207366, MH207781, MH208178; B. sp. nov. sect. Baryandra, C.-I Peng P23859 (HAST), KR186444, KR186531, KR186705; B. sp. nov. sect. Baryandra, C.-I Peng P23508 (HAST), KR186463, KR186550, KR186723; B. sp. nov. sect. Casparya, A. JaraMuñoz 2732 (COL), MH207369, MH207783, –; B. sp. nov. sect. Casparya, A. JaraMuñoz 2740 (COL), MH207370, MH207784, MH208181; B. sp. nov. sect. Casparya, A. JaraMuñoz 2736 (COL), MH207371, MH207785, MH208182; B. sp. nov. sect. Casparya, A. JaraMuñoz 2754 (COL), MH207368, MH207782, MH208180; B. sp. nov. sect. Casparya, A. JaraMuñoz 2728 (COL), MH207367, –, MH208179; B. sp. nov. sect. Cyathocnemis, P.W. Moonlight & A. Daza 146 (E), MH207372, MH207786, MH208183; B. sp. nov. sect. Cyathocnemis, P.W. Moonlight & A. Daza 150 (E), MH207373, MH207787, –; B. sp. nov. sect. Cyathocnemis, P.W. Moonlight & A. Daza 144 (E), –, MH207790, –; B. sp. nov. sect. Gobenia, A. Jara-Muñoz 2622 (COL), –, MH207788, –; B. sp. nov. sect. Knesebeckia group II, P.W Moonlight & A. Daza 159 (E), KX756301, KX756321, MH208193; B. sp. nov. sect. Knesebeckia group II, P.W Moonlight & A. Daza 156 (E), KX756295, KX756322, MH208194; B. sp. sect. Baryandra, Kokubagata GK71 (HAST), KR186509, KR186596, KR186769; B. sp. sect. Baryandra, R.R. Rubite R136 (PNH), KR186510, KR186597, KR186770; B. sp. sect. Baryandra, R.R. Rubite R290 (PNH), KR186511, KR186598, KR186771; B. sp. sect. Baryandra, C.-I Peng P23452 (HAST), KR186512, KR186599, KR186772; B. sp. sect. Baryandra, C.-I Peng P23418 (HAST), KR186513, KR186600, KR186773; B. sp. sect. Baryandra, No voucher (AS balcony), MH235386, MH235402, MH235427; B. sp. sect. Bracteibegonia, D.C. Thomas & W.H. Ardi 08-132 (E), JN133328, JN133403, JN133432; B. sp. sect. Cyathocnemis, P.W. Moonlight & A. Daza 36 (E), MH207374, MH207791, MH208185; B. sp. sect. Cyathocnemis, P.W. Moonlight & A. Daza 95 (E), MH207375, –, MH208186; B. sp. sect. Cyathocnemis, P.W. Moonlight & A. Daza 86 (E), –, –, MH208184; B. sp. sect. Cyathocnemis, P.W. Moonlight & A. Daza 25 (E), MH235390, –, MH235430; B. sp. sect. Diploclinium, D.C. Thomas 08-145 (E), JF756352, JF756436, JF756520; B. sp. sect. Diploclinium, No voucher, MH235382, MH235416, MH235419; B. sp. sect. Erminea, Duruisseau, J. Senechal, B. LBG 100651, MH207376, MH207792, MH208187; B. sp. sect. Gireoudia, B. Sénéchal CRBS/EB201134 (LBG), MH207377, MH207793, MH208188; B. sp. sect. Gireoudia, Lyon B.G. 880193 (LBG), MH207378, MH207794, MH208189; B. sp. sect. Gireoudia, Glasgow B.G. 027-044-89 U195 (E), KP712993, KP713211, –; B. sp. sect. Gobenia, A. Jara-Muñoz 2620 (COL), –, MH207795, –; B. sp. sect. Gobenia, AJM 2623, –, MH235405, –; B. sp. sect. Jackia, D.C. Thomas 08-85 (E), JF756375, JF756459, JF756543; B. sp. sect. Jackia, MH1542 (E), MH235393, MH235399, MH235422; B. sp. sect. Loasibegonia, P20248 (HAST), –, MH235410, –; B. sp. sect. Parvibegonia, T. Phutthai 195 (E), JF756380, JF756464, JF756548; B. sp. sect. Petermannia, D.C. Thomas 08-135 (E), KP712934, KP713130, KP713336; B. sp. sect. Petermannia, DCT 07-1 (E), JN133323, JN133398, JN133427; B. sp. sect. Petermannia, DCT 08-146 (E), JN133327, JN133402, JN133431; B. sp. sect. Petermannia, DCT 09-136(E), JN133325, JN133400, JN133429; B. sp. sect. Petermannia, DCT 09-138 (E), JN133331, JN133406, JN133435; B. sp. sect. Platycentrum, L.L. Forrest 31 (E), JF756362, JF756446, JF756530; B. sp. sect. Pritzelia, Lyon B.G. 90450 (LBG), MH207379, MH207797, MH208190; B. speculum, P.W. Moonlight & A. Daza 158 (E), MH207380, MH207798, MH208191; B. speluncae, S. Follin 90457 (LBG), MH207381, MH207799, MH208192; B. squamulosa, C.-I Peng 21280 (HAST), KP712971, KP713182, –; B. squarrosa, RBGE 20071058, MH207384, MH207802, MH208195; B. squarrosa, J. Duruisseau 130620 (LBG), –, MH207803, –; B. staudtii, J. Duruisseau 120358 (LBG), –, MH207804, MH208196; B. stenogyna, S10 (SBG), MH207385, MH207805, MH208197; B. stevei, Scott, S. M. 04-315 (E), MH235381, MH235411, MH235420; B. stictopoda, M. Hughes 1409 (E), MH207387, MH207807, –; B. stictopoda, C. Puglesi & al. 239 (E), MH207386, MH207806, MH208198; B. stigmosa, M. Tebbitt 134 (BKL), KP712997, KP713096, KP713331; B. stigmosa, Adolfo Jara AJM 2729, MH235387, MH235404, –; B. aff. strigillosa, A. Twyford 346 E00619702 (E), KP713027, KP713161, KP713333; B. strigosa, Glasgow B.G. 004-137-91 (E), JF756416, JF756500, JF756584; B. suaveolens, 322 Version of Record TAXON 67 (2) • April 2018: 267–323 Moonlight & al. • Phylogenetic sectional classification of Begonia (Begoniaceae) Appendix 1. Continued. Lyon B.G. 80425 (LBG), MH207388, MH207808, MH208199; B. subacida, B. Sénéchal 120320 (LBG), MH207389, MH207809, MH208200; B. subciliata, P.W. Moonlight & A. Daza 189 (E), MH207390, MH207810, MH208201; B. subciliata, P.W. Moonlight & A. Daza 241 (E), MH207391, MH207811, MH208202; B. sublobata, Girmansyah & al. DEDEN1486 (E), KR186515, KR186602, KR186775; B. subnummarifolia, no voucher, KR186516, KR186603, KR186776; B. suborbiculata, R.R. Rubite R353 (PNH), KR186517, KR186604, KR186777; B. subscutata, Glasgow B.G. 05307997 (E), MH207392, MH207812, MH208203; B. subscutata, C.-I Peng s.n. (HAST), KP712958, KP713208, –; B. subspinulosa, M. Tebbitt & A. Daza 808 (E), MH207393, MH207813, MH208204; B. subvillosa, C.-I Peng 21291 (HAST), KP713046, KP713122, KP713262; B. subvillosa, Surin 25 E00198120 (E), KP713045, KP713121, KP713263; B. sudjanae, D.C. Thomas 08-109 (E), JF756377, JF756461, JF756545; B. susaniae, J. Duruisseau 060661 (LBG), –, MH207814, MH208205; B. sutherlandii, D.C. Thomas 08-140 (E), JF756337, JF756421, JF756505; B. sutherlandii, Jasper 1200-5 (HAST), KR186518, KR186605, KR186778; B. sykakiengii, C.-I Peng P23856 (HAST), KR186519, KR186606, KR186779; B. sykakiengii, C.-I Peng P23890 (HAST), KR186520, KR186607, KR186780; B. sylvestris, Senechal LBG 130370, MH235392, MH235400, MH235431; B. symsanguinea, L.L. Forrest 199 (E), JF756415, JF756499, JF756583; B. tagbanua, P. Blanc s.n. (E), KR186521, KR186608, KR186781; B. tagbanua, C.-I Peng P23472 (HAST), KR186522, KR186609, KR186782; B. taraw, P. Blanc s.n. taraw 2 (E), KR186523, KR186610, KR186783; B. taraw, P. Blanc s.n. taraw 1 (E), KR186524, KR186611, KR186784; B. tayabensis, R.R. Rubite R360 (PNH), KR186525, KR186612, KR186785; B. tenera, L. Kumarage 68 (E), MH207394, MH207815, MH208206; B. tenuifolia, D.C. Thomas 08-86 (E), JF756349, JF756433, JF756517; B. tetrandra, A. JaraMuñoz 2632 (COL), MH207395, MH207816, MH208207; B. teuscheri, Lyon B.G. 20483 (LBG), MH207396, MH207817, MH208208; B. thelmae, 20131424 (E), MH207397, –, MH208209; B. thiemei, A. Twyford 205 E00668802 (E), KP713003, KP713094, KP713229; B. thomsonii, Lyon B.G. 140496 (LBG), MH207398, MH207818, MH208210; B. thwaitesii, L. Kumarage 23 (E), MH207399, MH207819, MH208211; B. thyrsoidea, M. Tebbitt & A. Daza 809 (E), –, MH207820, MH208212; B. tigrina, C.-I Peng P22720 (HAST), KR186526, KR186613, KR186786; B. tiliifolia, A. Jara-Muñoz 2650A (COL), –, MH207821, –; B. toledana, A. JaraMuñoz 2750 (COL), MH207400, MH207822, MH208213; B. tominana, M. Tebbitt 719 (USZ), MH207401, MH207823, MH208214; B. tonduzii, Lyon B.G. 140248 (LBG), MH207402, MH207824, MH208215; B. torajana, D.C. Thomas & W.H. Ardi 09-104 (E), JN133332, JN133407, JN133436; B. trianae, A. JaraMuñoz 2668 (COL), MH207403, MH207825, MH208216; B. tribenensis, Chien-I Huang 6043 (E), MH207404, MH207826, MH208217; B. trichochila, C.-I Peng P20764 (HAST), KR186527, KR186614, KR186787; B. trichopoda, SUBOE 97 (E), MH207405, MH207827, MH208218; B. tumbezensis, M. Tebbitt 770 (QCA), –, MH207828, –; B. ulmifolia, E00198123, KP713048, KP713136, KP713341; B. ulmifolia, C.I Peng 22518 (HAST), MH207407, MH207830, MH208220; B. ulmifolia, L.L. Forrest 169 E00183958 (E), KP713043, KP713135, KP713273; B. ulmifolia, Brown, K. 21551 (LBG), MH235395, MH235406, MH235425; B. umbellata, A. JaraMuñoz 2762 (COL), MH207408, MH207831, MH208221; B. undulata, C.-I Peng 21275 (HAST), KP712938, KP713139, –; B. urticae, P.W. Moonlight & A. Daza 73 (E), MH207409, MH207832, –; B. urubambensis, M. Tebbitt & A. Daza 800 (E), MH207410, MH207833, MH208222; B. urubambensis, P.W Moonlight & A. Daza 244 (E), KX756298, KX756310, –; B. urubambensis, P.W. Moonlight & A. Daza 253 (E), MH207412, MH207835, –; B. vaccinioides, Sabah NP 25535 (SNP), MH207413, MH207836, MH208223; B. valerioi, B. Sénéchal 60678 (LBG), MH207414, MH207837, –; B. valida, Glasgow B.G. 007-066-75 (E), KP712964, KP713144, KP713289; B. vankerckhovenii, J.P. Biteau 070718 (LBG), MH207415, –, MH208224; B. variegata, J. Durisseau 100652 (LBG), –, –, MH208225; B. varipeltata, D.C. Thomas & W.H. Ardi 08-51 (E), JF756410, JF756494, JF756578; B. varipeltata, Kreb s.n. (SBG), MH207416, MH207838, MH208226; B. velata, M. Tebbitt & A. Daza 838 (E), MH207417, MH207839, –; B. velata, P.W. Moonlight & A. Daza 107 (E), MH207418, MH207840, MH208227; B. venosa, 20030608 (E), MH207419, –, MH208228; B. venosa, Suksuwan 34 E00198104 (E), KP712966, KP713150, KP713284; B. venusta, S. Neale 7 (E), JF756357, JF756441, JF756525; B. verecunda, D.C. Thomas 08-137 (E), JF756399, JF756483, JF756567; B. vermeulenii, Vermeulen 2301 (L), JN133333, JN133408, JN133437; B. versicolor, L.L. Forrest 2 (E), JF756358, JF756442, JF756526; B. violifolia, Lyon B.G. 880079 (LBG), MH207420, MH207841, –; B. viridiflora var. parviflora, P.W. Moonlight & A. Daza 216 (E), MH207421, MH207842, MH208229; B. viridiflora var. viridiflora, P.W. Moonlight & A. Daza 206 (E), MH207422, MH207843, MH208230; B. wadei, R.R. Rubite R699 (PNH), KR186528, KR186615, KR186788; B. watuwilensis, D.C. Thomas 09-55 (E), JF756406, JF756490, JF756574; B. weberbaueri, T. Sarkinen 2216 (E), KP713024, KP713102, KP713340; B. weberbaueri, Tebbit 829 (USZ), MH235396, MH235407, MH235428; B. weigallii, Pitisopa, Gardner, Herrington 10 (E), JN133334, JN133409, JN133438; B. wilsonii, Y.M. Shui & al. B2013151 (KUN), MH207423, MH207844, MH208231; B. wollnyi, M. Tebbitt s.n. (USZ), MH207424, MH207845, MH208232; B. woodii, C.-I Peng P23479 (HAST), KR186529, KR186616, KR186789; B. woodii, C.-I Peng P23496 (HAST), KR186530, KR186617, KR186790; B. yapenensis, ABEG 211 (E), MH207425, MH207846, MH208233; B. yunnanensis, C.I Peng 20941 (HAST), MH207426, MH207847, MH208234; H. sandwicensis, Edutg Monn 1245-2005, MH244433, MH244435, MH244438. Version of Record 323