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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
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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).
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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.
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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.
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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)
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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
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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
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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.
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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.
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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.
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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
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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-
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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.
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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.
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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
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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.
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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.
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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).
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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
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● 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.
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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
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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.
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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
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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
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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.
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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
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(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.
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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
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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
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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).
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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
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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.
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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
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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).
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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
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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
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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.
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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.
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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.
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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
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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.
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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
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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,
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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.,
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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.
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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,
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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
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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.
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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.
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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
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& 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.
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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.*,
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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.
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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.
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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
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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
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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).
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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).
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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).
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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).
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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).
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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).
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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.
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and future directions in the taxonomy of Cyrtandra (Gesneriaceae),
with a new estimate of species number. Selbyana 31: 157–165.
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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,
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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;
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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;
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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,
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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.
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