Begonia Pulchrifolia (Sect. Platycentrum), a New Species of Begoniaceae from Sichuan of China

http://dx.doi.org/10.11646/phytotaxa.207.3.2

Begonia pulchrifolia (sect. Platycentrum), a new species of Begoniaceae from Sichuan of China

DAI-KE TIAN1,2*, CHUN LI1,2**, CE-HONG LI3 & XIAO-JIE LI3 1Shanghai Chenshan Plant Science Research Center, Chinese Academy of Sciences, 3888 Chenhua Road, Songjiang, Shanghai 201602, China; 2Shanghai Key Laboratory for Plant Functional Genomics and Resources, Shanghai Chenshan Botanical Garden, 3888 Chenhua Road, Songjiang, Shanghai 201602, China; 3Emeishan Experimental Station of Biological Resources, Sichuan Provincial Academy of Natural Resources, Emeishan, Sichuan 614200, China. *Corresponding author; e-mail: [email protected], **Co-first author.

Abstract

Begonia pulchrifolia D.K.Tian & C.H.Li, a new species in Begonia sect. Platycentrum A.DC. (Begoniaceae) from Sichuan of China, is described and illustrated. Morphologically, it is only similar to the young or smaller individuals of B. dielsiana and B. emeiensis in plant shape and some leaf characters but differs easily by its foliage color and deeper divided leaf lobes, small-sized flowers, hairy outer petals of the male flowers, and long abaxially winged fruits. Molecular phylogenetic analysis based on nrDNA also supported the distinction of B. pulchrifolia from B. dielsiana and B. emeiensis.

Key words: China, Sichuan, Begonia, new species

Introduction

Begonia is considered the world sixth largest genus in vascular plants (Hoover et al. 2004) and about 1600 species have been described so far (Thomas, 2010; Aitawade et al. 2012). The species occur in subtropical and tropical regions, with the greatest diversity in America and Asia (> 600 species each), whilst being relatively poor in Africa (160 species) and absent in Australia (Goodall-Copestake et al., 2010). In Asia, China has the largest number of Begonia species. Based on Flora of China (Gu et al. 2007), 173 species are recorded in China. Since publication of the Flora of China, nearly 20 new species from China have been published (Ding et al. 2014, Ku et al. 2008, Li et al. 2008, Liu et al. 2007, Ma et al. 2006, Peng et al. 2008a,b, 2009, 2010, 2012, 2013, 2014a,b, Shui 2007, Tian et al. 2014, Wei et al. 2007). According to our investigation in the past years, there are still some potential new taxa needing to be studied and published in the future. Here, we described and illustrated Begonia pulchrifolia, a new species from Sichuan of China. In November 2013, Daike Tian, traveled to Beijing and found one specimen of a potential new taxon of Begonia at the Herbarium of the Health Science Center, Peking University (PEM). This specimen was collected from Mount Emei in Sichuan Province but did not include collector’s name and collection date (possibly 30 years ago). To investigate this doubtful species, Daike Tian traveled to Mount Emei in September 2014 and met with Cehong Li, a staff member of Mount Emei Experimental Station of Biological Resources, Sichuan Provincial Academy of Natural Resources. Cehong Li turned out to be the one who first found the taxon in Mount Emei in July 1990, and he had introduced wild plants to glasshouse for cultivation in October 2003. Through a further survey, we determined several populations of this potential new species in both Meinvfeng of Leshan and Mount Emei of Emeishan, Sichuan. Morphologically, Begonia pulchrifolia is only similar to the immature or small-sized individuals of B. dielsiana E. Pritzel ex Diels (1900: 479) and B. emeiensis C.M. Hu ex C.Y. Wu & T.C. Ku (1995: 273) in plant shape and some leaf characters, but differs clearly by its foliage color and usually deeper lobed leaves, small-sized flowers, hairy outer petals of the male flowers, and long abaxially winged fruits. In order to understand more about its taxonomic status and similarity with other species, a molecular phylogenetic analysis based on the nuclear ribosomal DNA (nrDNA) internal transcribed spacer (ITS) was conducted using 33 accessions representing a total of 30 species from the five main sections of Begonia delimited in China and one section from Africa based on the treatments by Doorenbos et al. (1998) and Shui et al. (2002).

242 Accepted by Hugo de Boer: 17 Apr. 2015; published: 11 May 2015 20484 (KUN) 40778 (KUN) 40782 (KUN) TDK2243 (CSH) et al. et al. et al. et al. Tian, D.K., Li, C. TDK1272 (CSH) D.K., Li, C. Tian, X.J.Y.01004 (KUN) X.J.Y.01004 Li, J.X. TDK2279 (CSH) Li, J.X. X.J.Y.01012 (KUN) X.J.Y.01012 (KUN) X.J.Y.01010 Tian, D.K., Li, C. TDK785 (CSH) D.K., Li, C. Tian, TDK866 (CSH) D.K., Li, X.P. Tian, Shui, Y.M. Shui, Y.M. X.J.Y.01001 (KUN) X.J.Y.01001 Tian, D.K., Tian, L.Z. TDK2111 (CSH) TDK2111 L.Z. Tian, D.K., Tian, Tian, D.K., Tian, L.Z. TDK2356 (CSH) L.Z. Tian, D.K., Tian, no voucher available Shui, Y.M. Shui, Y.M. Forrest, L.L.241 (E) Forrest, L.L. 24 (E) Tian, D.K., Li, C. TDK757 (CSH) D.K., Li, C. Tian, Tian, D.K., Mo, H.B. TDK158 (CSH) D.K., Mo, H.B. Tian, TDK774 (CSH) D.K., Li, C. Tian, Tian, D.K., Tian, L.Z. TDK2250 (CSH) L.Z. Tian, D.K., Tian, Peng 18779 Li, C. Yang, L.H. TDK2268 (CSH) L.H. Yang, Li, C. Li, X.J. LXJ022 (CSH) Li, X.P. TDK763 (CSH) D.K., Li, C. Tian, Tian, D.K., Tian, Tian, D.K., Li, C. TDK1300 (CSH) D.K., Li, C. Tian, LI, C.H., LCH001 (CSH) Tian, D.K., Tian, L.Z. TDK2249 (CSH) L.Z. Tian, D.K., Tian, LI, C.H., LCH002 (CSH) D-38 (KUN) Y.M. Shui, TDK1280 (CSH) D.K., Li, C. Tian, Shui, Y.M. Shui, Y.M. Tian, D.K., Li, C. TDK1338 (CSH) D.K., Li, C. Tian, Tian, D.K., Tian, L.Z. TDK2250 (CSH) L.Z. Tian, D.K., Tian, Diploclinium Sphenanthera Sphenanthera Diploclinium Reichenheimia Platycentrum Coelocentrum Section (Herbarium) Voucher Collector, Sphenanthera Reichenheimia Platycentrum Coelocentrum Sphenanthera Augustia Platycentrum Sphenanthera Reichenheimia Platycentrum Platycentrum Platycentrum Diploclinium Sphenanthera Diploclinium Platycentrum Platycentrum Platycentrum Reichenheimia Diploclinium Diploclinium Platycentrum Coelocentrum Platycentrum Platycentrum Reichenheimia AF048975 AF048988 AF048974 JF975965 Genbank accession no. AY048980 AF485130 JF976018 AF469126 AF485114 KF636473 JF976047 JF976001 Lincang, Yunnan, China Lincang, Yunnan, ChinaLvchun, Yunnan, KP710824 Lincang, Yunnan, China Lincang, Yunnan, Kunming Botanic Garden, China Napo, Guangxi, ChinaXinyi, Guangdong, China KP710820 KP710815 Malipo, Yunnan, China Malipo, Yunnan, Xishuangbanna, Yunnan, China Xishuangbanna, Yunnan, Nanchuan, Chongqing, China KP710819 Wulong, Chongqing, ChinaWulong, KP710805 Beijing Botanic Garden, China Hekou, Yunnan, China Hekou, Yunnan, South Africa Maguan, Yunnan, China Maguan, Yunnan, Bama, Guangxi, China KP710813 Yuxi, Yunnan, China Yunnan, Yuxi, KP710821 Tianlin, Guangxi, ChinaTianlin, KP710810 Emeishan, Sichuan, China KP710816 Guizhou, China Yichang, Hubei, ChinaYichang, Fengshan, Guangxi, China KP710828 KP710818 Fangchenggang, Guangxi, China KP710826 Meinvfeng, Leshan, Sichuan, China KP710811 Pingbian, Yunnan, ChinaPingbian, Yunnan, KP710817 Meinvfeng, Leshan, Sichuan, China KP710812 Leshan, Sichuan, China KP710822 Hekou, Yunnan, China Hekou, Yunnan, ChinaLvchun, Yunnan, KP710827 Mount Emei, Emeishan, Sichuan, China KP751374 Gejiu, Yunnan, China Gejiu, Yunnan, Pingbian, Yunnan, ChinaPingbian, Yunnan, KP710814 Changning, Sichuan, China KP715148 Yiliang, Yunnan, China Yunnan, Yiliang, KP710823 Craib C.M.Hu ex C.Y.Wu & T.C.Ku & C.M.Hu ex C.Y.Wu Hance Hance J.D.Hooker D.K.Tian & C.H.Li D.K.Tian D.K.Tian & C.H.Li D.K.Tian D.K.Tian & C.H.Li D.K.Tian Irmscher H.Léveillé Irmscher (A.DC.) C.B.Clarke Blume H.Léveillé

Y.M.Shui & W.H.Chen Y.M.Shui C.M.Hu ex C.Y.Wu & T.C.Ku & C.M.Hu ex C.Y.Wu E.Pritzel C.Y.Wu D.Don C.M.Hu ex C.Y.Wu & T.C.Ku & C.M.Hu ex C.Y.Wu Y.M.Shui Gagnepain Dryander Irmscher Irmscher H.Léveillé & Vaniot H.Léveillé & Y.M.Shui & W.H.Chen Y.M.Shui Species used for phylogenetic analysis. Irmscher T.C.Ku Irmscher Hemsley Otto & Dietr. H.Léveillé B. biflora Begonia acetosella B. arboreta Taxon Origin B. longifolia B. wenshanensis B. wilsonii B. chingii B. circumlobata B. dielsiana B. longistyla Note: Genbank accession number in bold represent sequences downloaded from NCBI. B. dregei B. morsei B. palmata B. edulis B. parvula B. emeiensis B. pedatifida B. pedatifida B. fimbristipula B. pulchrifolia B. grandis B. handelii B. hemsleyana B. pulchrifolia B. henryi B. pulchrifolia B. huangii B. ruboides B. setifolia B. silletensis B. laminariae B. limprichtii B.lithophila TABLE 1. TABLE

Taxonomic Sampling

The putative new species was assigned to Begonia sect. Platycentrum A.DC. (1859: 134) according to morphological traits of fruits and placenta. Several species in sect. Platycentrum and 4 to 6 representative species of five main sections of Begonia in China were chosen to assess the placement of this potential new species in the phylogeny (Table 1). Begonia dregei Otto & Dietrich (1836: 357) of sect. Augustia (Klotzsch) A. DC. (1864: 384) from South Africa was used as outgroup in the phylogenetic analysis. The morphological data were collected from the wild living plants during field surveys. Meanwhile, fresh leaves of Begonia species were sampled and immediately preserved in the sealed bags with silica gel for later lab work.

DNA Sequencing and Molecular Analyses

The genomic DNA was extracted from leaf samples using DNAsecure Plant Kit (Tiangen Biotech, Beijing, China), following the manufacturer’s protocols. The nuclear ribosomal DNA (nrDNA) internal transcribed spacer (ITS) region was amplified and sequenced using primers of Thomas (2010). Amplification reactions were performed in 20 μL reactions containing 11.0 μL of ddH2O, 4.0 μL of 5 × Fast HiFidelity PCR buffer, 1.0 μL of 20 × Fast PCR Enhancer, 0.8 μL of each forward and reverse primer (10 μM), 0.4 μL of Fast HiFidelity Polymerase (Tiangen Biotech), 2.0 μL of DNA template. PCR reactions were conducted using the following program: an initial denaturation at 94°C for 2 min, followed by 35 cycles of 15 s denaturation at 94°C and 30 s annealing and elongation at 68°C, and a final elongation step at 68°C for 5 min. Phylogenies for ITS region were reconstructed using Bayesian inference. The GTR+G model was chosen as the optimal model of nucleotide substitution using the Akaike information criterion (AIC; Burnham & Anderson 2002) as implemented in MrModeltest 2.3 (Nylander 2004). Bayesian Markov Chain Monte Carlo (MCMC) inference analysis was performed using MrBayes version 3.1.2 (Ronquist & Huelsenbeck 2003) at the CIPRES Science Gateway (Miller et al. 2010). Two independent MrBayes analyses were performed with four Markov chains and five million generations. Trees were sampled every 100 generations, resulting in 50,000 trees for each run, with the first 25% discarded as burn- in. The posterior probabilities for individual clades obtained from separate analysis were compared for congruence and then combined and summarized on a 50% majority-rule consensus tree.

Results

Taxonomy

Begonia pulchrifolia D.K.Tian & C.H.Li, sp. nov. (Fig. 1 & 2). Type:—China. Meinvfeng, Leshan, Sichuan, 29°20′57′′N, 103°31′47′′E, 1160 m, 14 September 2014, D.K. Tian et al., TDK-2241 (holotype CSH!; isotype CSH!). Topotype:—at the same place, 29°21′26′′N, 103°32′00′′E, 800 m, 14 September 2014, D.K. Tian et al., TDK-2243 (CSH!). Begonia pulchrifolia D.K. Tian & C.H. Li can be easily separated from other Begonia species. Morphologically, it is only similar to the young or smaller individuals of B. dielsiana and B. emeiensis but clearly differs from them by its leaf color and usually deeper divided leaf lobes, small-sized flowers, hairy outer petals of male flowers, 5 petals of female flowers and longer abaxial fruit wings (Table 2, Fig. 5). Herb rhizomatous, perennial, deciduous in winter, 10–20 cm tall. Rhizome creeping, short, rarely branched, 3–10 cm long, 4–15 mm thick. Leaves 2–9 (12) per plant, basal or occasionally on 1–3 nodal stem during inflorescence development. Petiole green or pinkish, cylindrical, glabrous without ditches and spots on surface, (3)7–30 cm long, (2)3–7 mm thick. Blade nearly oval, 6–25 cm long, 4–20 cm wide, both sides sparsely short gray or red rigid hairy, hair less than 0.5 mm long; adaxial surface dark green with light-green areas particularly at base; abaxial surface red to deep red with light-green areas, rarely entire deep-red; base oblique, cordate, near overlapped to slightly overlapped; margin usually triangular, shallow lobed to rarely deep divided in large individuals, with denticulate, rigid hairs; venation palmate, 9–11 veined, adaxial slightly impressed, abaxial distinctly prominent.

244 • Phytotaxa 207 (3) © 2015 Magnolia Press TIAN ET AL. FIGURE 1. A–K Begonia pulchrifolia (TDK-2241, 2243, CSH). A. Habit; B. Commonly seen short rigid hairs on leaf surface; C. Longer hairs rarely seen in few individuals; D–F. Male flower: front view and stamens; G–I. Female flower (G—front view, H—style and stigma, I—middle cross-section of ovary); J–K: Capsule with three unequal wings. (Drawn by Yanxin Piao)

A NEW SPECIES OF BEGONIACEAE FROM CHINA Phytotaxa 207 (3) © 2015 Magnolia Press • 245 Inflorescences 1–4, basal or occasionally terminal on short stem with 1–2 internodes, 6–30 cm long. Primary peduncle green or pinkish, glabrous, 5–21 cm long, 2–5 mm in diameter. Bracts ovate-lanceolate, deciduous, glabrous, margin entire. Flowers 5–15 per inflorescence; male flowers open earlier at the same node, about 1.5 folds of female flowers in number. Staminate flower: pedicel green or slightly pink, glabrous, 20–27 mm long and up to 1 mm in diameter; corolla 32–36 × 26–31 mm; tepals 4, outer 2 larger, color uneven, upper pink, lower near white, broadly ovate, 15–18 × 12–19 mm, stripes distinct on adaxial surface, sparse hairs on abaxial surface around axis; inner 2 smaller, white-pinkish, oblanceolate or ovate-lanceolate, 12–15 × 4.5–7 mm, stripes distinct. Androecium 3–4 mm long, 4–5 mm in diameter; stamens about 40–60, filaments about 3 mm long, upper 1/3 free, lower 2/3 fused; anthers yellow, obovate, nearly 1 mm long, apex slightly depressed, fewer obtuse.

FIGURE 2. A–M. B. pulchrifolia (TDK-2241, 2243, Photos by Daike Tian). A. Habit and foliage; B. Large blooming mature individual; C. Diversity in leaf color and shallow lobes to rarely deep-divided in large individuals; D. Commonly seen short adaxial hairs; E. Rarely seen longer adaxial hairs; F. Side-view of male flower with hairs on outer petals; G. Front view of male flower; H. Stamens; I–J. Side and front views of female flower; K. Cross-section of ovary; L. Fruits with unequal wings; M. Mature seeds.

246 • Phytotaxa 207 (3) © 2015 Magnolia Press TIAN ET AL. Pistillate flower: pedicel pink, glabrous, 13–20 mm long, 1 mm in diameter. Corolla 18–27 mm in diameter; tepals 5, nearly ovate, grabrous, upper pink, lower nearly white, 8–15 × 3–12 mm, inner smaller, stripes distinc on adaxial surface. Styles 2, free, 1.5–2 mm long; stigma 2, golden yellow, U-shaped, 1–2 mm long, 2 circles spiralled. Ovary 3-loculed, placentae axile, bifid. Capsule nodding, ovoid, glabrous, unequally 3-winged; abaxial wing nearly round- rectangular, 12–16 × 8–11 mm; lateral wings shorter, 1.5–5 × 9–12 mm. Seeds numerous, brown, ellipsoid, ovoid to ovoid-conical, 0.3–04 × 0.20–0.25 mm. Flowering July to Sept., fruiting Aug. to Nov. Etymology:—The epithet is named for its beautiful foliage color and elegant plants. Phenology:—The plants bloom between July and September, the fruits ripe from August to November. Distribution and habitat:—In Meinvfeng, Leshan City, at an elevation of 800–1160 m, and in Jiushijiudaoguai, Emeishan City at an elevation of 1235–1420 m, both in Sichuan Province, China (Fig. 3). Growing on hilly slopes under forest cover or on rocks in moist valleys. In both sites, no other Begonia species have been found to occur together with this new taxa.

FIGURE 3. Distribution of B. pulchrifolia in Sichuan Province, China.

Additional specimens examined:—China. Mount Emei, Emeishan City, Sichuan Province, 29°33′00.59′′N, 103°22′00.11′′E, 1230–1420 m, 20 Sept. 2014, C.H. Li 14001 (CSH; CDBI); Emei, Emeishan City, Sichuan Province, anonymous and unknown date (1980 ?), collection No. 80061, herbarium specimen routing number as PEM-15460 (PEM). Proposed IUCN Conservation assessment:—VN. Based on the area of occupancy estimated to be <500 km2 (EN: B2), species existing at two separated locations (EN: B2a), and quality of habitat continuing decline due to

A NEW SPECIES OF BEGONIACEAE FROM CHINA Phytotaxa 207 (3) © 2015 Magnolia Press • 247 increasing tourism and potential exploitation for ornamental and medicinal uses (EN: B2b(iii)), B. pulchrifolia should be considered Endangered (EN: B2ab(iii)) according to the IUCN red list criteria (IUCN 2014). Therefore, a sustainable conservation strategy should be made.

FIGURE 4. Bayesian inference of the phylogenetic position of the newly described Begonia pulchrifolia within the Sect. Platycentrum based on nuclear ITS sequences. Nodes with bpp < 0.50 have been collapsed. Sectional placement of taxa is indicated by the following abbreviations: Aug: Augustia, Coe: Coelocentrum, Dip: Diploclinium, Pla: Platycentrum, Rei: Reichenheimia, Sph: Sphenanthera. Numbers after the species names indicate different populations. The putative new species is indicated in bold.

Molecular Analysis

The ITS data set containing 33 accessions represented 30 species, five main sections of Begonia in China and one section from Africa (Table 1). The aligned matrix of ITS region was 705 bp, of which 345 (48.9%) were variable and 201 (28.5%) were informative. Result of Bayesian inference analysis was displayed in Figure 4. The phylogenetic analysis showed that all species in sect. Platycentrum appeared monophyletic with a high Bayesian posterior probability (bpp = 0.89), the putative new species was placed in a well-supported clade (bpp = 1) with B. pedatifida H. Léveillé (1909: 21) and B. limprichtii Irmscher (1922: 12). The morphologically most similar species B. dielsiana and B. emeiensis formed

248 • Phytotaxa 207 (3) © 2015 Magnolia Press TIAN ET AL. another clade elsewhere in sect. Platycentrum (bpp = 1), which supports that the new species is a different from both B. dielsiana and B. emeiensis. Three individuals of the new species (bpp = 1) and two individuals of B. pedatifida (bpp = 0.98) from different populations formed two well-supported clades respectively, which indicated this new species was a lineage independent from B. pedatifida and other species.

Discussion

B. pulchrifolia is confirmed as a new species based on careful investigation on literature and both herbarium and living specimen, as well as results of molecular phylogenetic analysis. It is easily distinguished from all other Chinese Begonia species which have been published before. However, it may be morphologically confused with the young or smaller individuals of B. dielsiana and B. emeiensis when these three species don’t show flowers and fruits (Fig. 5). Leaf color is a key character to separate B. pulchrifolia from B. dielsiana and B. emeiensis. Additionally, the latter two species have much larger size of plants, leaves and flowers, shallower lobes, 6 female-petals, highly developed stigma, and shorter abaxial fruit wings (Table 2, Fig. 5). Due to leaf color similarity, B. pulchrifolia is sometimes mistreated by local people as the variant plants of Dysosma delavayi (Franchet) Hu (1937: 37) in Berberidaceae, a traditional Chinese medicinal herb. It can be used for an ornamental because of its elegant plants and beautiful leaf color.

TABLE 2. Comparison of Begonia pulchrifolia with B. dielsiana and B. emeiensis B. pulchrifolia B. dielsiana B. emeiensis

Habit Deciduous in winter; isolated Evergreen; isolated plants to Evergreen; isolated plants to plants to small patch large patch large patch

Plant height (cm) 10–20 15–70 Up to 130

Rhizome 3–10 cm long, 4–15 mm thick, 10–20 cm long, 1–3.5 cm thick, Up to 1 m long, 4.5 cm thick, rarely branched usually branched highly branched

Stem Rarely seen, short Common seen, long Common seen, long

Leaf color Upper surface dark-green with Upper surface green, abaxially Upper surface green, abaxially green areas, abaxially red with light-green light-green light green areas Blade size (cm) 6–25 × 4–20 9–31 × 7–22 12–31 × 11–27

Inflorescence height Usually above leaf surface Under leaf surface Under leaf surface Flower corolla size (mm) Small, male flower 32–36 × 26–31; Larger, male flower 38–50 × Largest, male flower 40–60 × female 18–27 35–47; female 40–50 41–58; female 55–65 Petal thickness Thin Thick Thick Female petal number and Petal 5; stigma less developed Petal 6, rarely 5; stigma highly Petal 6 more than 5; stigma stigma developed usually with many highly developed upper branches Abaxial fruit wing size 12–16 × 8–11 8–14 × 10–16 Unmeasured, shorter than (mm) that of B. dielsiana based on observation Note: Data are mainly based on observation on living plants in the wild and partially from literature.

Acknowledgements

This work is supported by the funds from Shanghai Administration Bureau of Landscape and City Appearance (funded project code: F122416) and the specific project for strategic biological resources and technology supporting system from Chinese Academy of Sciences (CAS) (funded project code: CZBZX-1). The authors thank Ms. Yanxin Piao from Shanghai Institute of Visual Art for illustrating this new species.

A NEW SPECIES OF BEGONIACEAE FROM CHINA Phytotaxa 207 (3) © 2015 Magnolia Press • 249 FIGURE 5 A–C. B. dielsiana (TDK-2119): A. Habit and foliage; B. Medium-sized individual with fruits; C. Flowers and ovary dissection (upper: 4-petalled male flowers; lower two: 6-petalled female flowers at different stages). D–F. B. emeiensis (TDK-2249): D. Habit and foliage; E. Flower front view (upper three: 6-petalled female flowers; lower one: 4-petalled male flower); F. Female flowers with 6, 5 petals. (Photos by Daike Tian)

Reference

Aitawade, M.M. & Yadav, S.R. (2012) Taxonomic status of Begonia aliciae (Begoniaceae). Rheedea 22: 111–115. Burnham, K.P. & Anderson, D.R. (2002) Model Selection and Multimodel Inference: a practical information-theoretic approach. Springer, New York, 488 pp. De Candolle, A. (1864) Ordo CLXXII. Begoniaceae. Prodromus systematis naturalis regni vegetabilis 15. Victoris Masson et filii, Parisiis,

250 • Phytotaxa 207 (3) © 2015 Magnolia Press TIAN ET AL. pp. 266–408. De Candolle, A. (1859) Memorire sur la famille des Begoniaceae. Annales des Science Naturelles- Botanique IV 11: 92–149. Diels, L. (1900) Die Flora von Central-China: Begoniaceae. Botanische Jahrbücher fur Systematik 29: 479. Ding, B., Nakamura, K., Kono, Y., Ho, M.J. & Peng, C.I. (2014) Begonia jinyunensis (Begoniaceae, section Platycentrum), a new palmately compound leaved species from Chongqing, China. Botanical Studies 55: 62–69. http://dx.doi.org/10.1186/s40529-014-0062-6. Doorenbos, J., Sosef, M.S.M., & Wilde, J.J.F.E. (1998) The sections of Begonia. Wageningen Agricultural University, Wageningen, 66 pp. Goodall-Copestake, W., Pérez-Espona, S., Harris, D.J. & Hollingsworth, P.M. (2010) The early evolution of the mega-diverse genus Begonia (Begoniaceae) inferred from organelle DNA phylogenies. Biological Journal of the Linnean Society 101 (2): 243–250. http://dx.doi.org/10.1111/j.1095-8312.2010.01489.x. Gu, C.Z., Peng, C.I. & Turland, N.J. (2007) Begoniaceae. In Wu, C.Y. & Raven, P.H. (Eds.) Flora of China. Science Press, Beijing, pp. 153–207. Hoover, W.S., Karegeannes, C., Wiriadinata, H. & Hunter, J.M. (2004) Notes on the geography of South-East Asian Begonia and species diversity in montane forests. Telopea 10: 749–764. Hu, H.-H. (1937) Notulae sytanticae ad Florem Sinensium. Bulletin of the Fan Memorial Institute of Biology 8: 31–44. Irmscher, E. (1922) Centralblatt für Sammlung und Veroffentlichung von Einzeldiagnosen neuer Pflanzen. Repertorium Specierum Novarum Regni Vegetabilis, Beihefte 12: 440. IUCN (2014) Guidelines for Using the IUCN Red List Categories and Criteria. Version 11. Prepared by the Standards and Petitions Subcommittee. Available from: http://jr.iucnredlist.org/documents/RedListGuidelines.pdf (accessed 28 January 2015). Ku, S.M., Kono, Y. & Liu, Y. (2008) Begonia pengii (sect. Coelocentrum, Begoniaceae), a new species from limestone areas in Guangxi, China. Botanical Studies 49: 167–175. Léveillé, H. (1909) Decades plantarum novarum. XVI. Repertorium Specierum Novarum Regni Vegetabilis 7: 21. Li, H.Z., Ma, H., Zhou, Z.K. & Guan, K.Y. (2008) A new species of Begonia (Begoniaceae) from Guangxi, China. Botanical Journal of the Linnean Society 157: 83–90. http://dx.doi.org/10.1111/j.1095-8339.2008.00771.x Liu, Y., Ku, S.M. & Peng, C.I. (2007) Begonia bamaensis (sect. Coelocentrum, Begoniaceae), a new species from limestone areas in Guangxi, China. Botanical Studies 48: 465–473. Ma, H. & Li, H.Z. (2006) Begonia guaniana (Begoniaceae), a new species from China. Annales Botanici Fennici 43: 466–477. Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Gateway Computing Environments Workshop (GCE). IEEE, New Orleans, LA, pp. 1–8. http://dx.doi.org/10.1109/GCE.2010.5676129. Nylander, J.A.A. (2004) MrModeltest v2. Program distributed by the author. Evolutionary Biology Centre, Uppsala University. Available from: http://www.softpedia.com/get/Science-CAD/MrModeltest.shtml (accessed 28 January 2015). Otto, F. & Dietrich, A. (1836) Die Begonien. Berliner Allgemeine Gartenzeitung 4: 357. Peng, C.I., Jin, X.H., Ku, S.M., Kono, K., Huang, H.Y. & Yang, H.A. (2014a) Begonia wuzhishanensis (sect. Diploclinium, Begoniaceae), a new species from Hainan Island, China. Botanical Studies 55: 24–30. Peng, C.I., Ku, S.M., Kono, Y. & Liu, Y. (2012) Begonia chongzuoensis (sect. Coelocentrum, Begoniaceae), a new calciphilefrom Guangxi, China. Botanical Studies 53: 283–290. Peng, C.I., Ku, S.M., Yang, H.A., Leong, W.C., Liu, Y., Nguyen, T.H., Kono, Y. & Chung, K.F. (2014b) Two new species of Begonia sect. Coelocentrum, B. guixiensis and B. longa, from Sino-Vietnamese limestone karsts, Botanical Studies 55: 52–63. http://dx.doi.org/10.1186/s40529-014-0052-8. Peng, C.I., Yang, H.A., Kono, Y., Chung, K.F., Huang, Y.S., Wu, W.H. & Liu, Y. (2013) Novelties in Begonia sect. Coelocentrum: B. longgangensis and B. ferox from limestone areas in Guangxi, China. Botanical Studies 54: 1–9. http://dx.doi.org/10.1186/1999-3110-54-44 Peng, C.I., Ku, S.M., Kono, Y., Chung, K.F. & Liu, Y (2008a) Two new species of Begonia (sect. Coelocentrum, Begoniaceae) from limestone areas in Guangxi, China: B. arachnoidea and B. subcoriacea. Botanical Studies 49: 405–418. Peng, C.I., Liu, Y. & Kono, Y. (2010) Begonia × breviscapa (Begoniaceae), a new intersectional natural hybrid from limestone areas in Guangxi, China. Botanical Studies 51: 107–117. Peng, C.I., Liu, Y. & Ku, S.M. (2008b) Begonia aurantiflora (sect. Coelocentrum, Begoniaceae), a new species from limestone areas in Guangxi, China. Botanical Studies 49: 83–92. Peng, C.I. & Ku, S.M. (2009) Begonia × chungii (Begoniaceae), a new natural hybrid in Taiwan. Botanical Studies 50: 241–250. Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19 (12): 1572–1574.

A NEW SPECIES OF BEGONIACEAE FROM CHINA Phytotaxa 207 (3) © 2015 Magnolia Press • 251 Shui, Y.M. (2007) Begonia tetralobata (Begoniaceae), a new species from China. Annales Botanici Fennici 44: 76–79. Shui, Y.M., Peng, C.I. & Wu, C.Y. (2002) Synopsis of the Chinese species of Begonia (Begoniaceae), with a reappraisal of sectional delimitation. Botanical Bulletin of Academia Sinica 43: 313–327. Tian, D.K., Li, C., Yan, Y.H., Li, X.P. & Meng, J. (2014) Begonia intermedia, a new species of Begoniaceae from Hainan, China. Phytotaxa 166 (2): 114–122. http://dx.doi.org/10.11646/phytotaxa.166.2.2. Thomas, D.C. (2010) Phylogenetics and Historical Biogeography of Southeast Asian Begonia L. (Begoniaceae). PhD dissertation of The University of Glasgow, UK. Wei, Z.D., Shui, Y.M., Zhang, M.D. & Zhang, R.M. (2007) Begonia coelocentroides YM Shui & ZD Wei, a new species of Begoniaceae from Yunnan, China. Acta Phytotaxonomica Sinica 45: 86–89. Wu, C.Y. & Ku, T.C. (1995) New taxa of Begonia from China. Acta Phytotaxonomica Sinica 33: 251–280.