Begonia jinyunensis (Begoniaceae, section Platycentrum), a new palmately compound leaved species from Chongqing, China
© Ding et al.; licensee Springer 2014
Received: 30 June 2014
Accepted: 21 July 2014
Published: 2 August 2014
Continental China is the center of Begonia species diversity in Asia and contains more than 60 species out of about 110 named species of section Platycentrum. Mt. Jinyun, located in Chongqing City at the upper reaches of the Yangtze River, harbors a subtropical broadleaved forest with high species diversity. During a botanical survey in Mt. Jinyun, an unknown Begonia species of sect. Platycentrum with palmately compound leaves was collected and studied based on detailed morphological observations and cytological and molecular phylogenetic analyses.
The unknown Begonia bears a superficial resemblance to B. hemsleyana in having palmately compound leaves, a feature unseen in other species of sect. Platycentrum in China. It is however sharply distinct from the latter in the acaulous habit with aerial stems seen only at anthesis and long rhizomes (vs. erect stems to 70 cm or taller with short rhizomes), 4–6 pinnatilobed leaflets with indistinct, decurrent petiolules (vs. 7–10 serrate leaflets with distinct petiolules), and white (vs. pink) tepals. Molecular phylogenetic analyses based on nuclear ribosomal DNA and chloroplast DNA sequences indicated that this species was allied to Platycentrum species occurring in Southwest and South-central China and Vietnam, including B. hemsleyana, and clearly separable from these species. Somatic chromosome number of 2n = 22 was reported for this unknown species. The diploid chromosome number is agreeable with those published for Begonia sect. Platycentrum.
The unknown Begonia is confirmed to be a new species of sect. Platycentrum and hereto described as Begonia jinyunensis C.-I Peng, B. Ding & Q. Wang.
The mega-diverse genus Begonia L. (Begoniaceae) comprises ca. 1,500–1,600 species widely distributed in the tropical and subtropical regions of the world except Australia (Kiew , Tebbitt , Chung et al. ). Section Platycentrum (Klotzsch) A. DC. comprises about 110 species of herbs and subshrubs that are widely distributed in Asia, ranging from India to the Himalayas, Indo-China, China, Taiwan, and Malesia (Doorenbos et al. , Shui et al. ). Continental China is the center of Begonia species diversity in Asia and contains more than 60 Platycentrum species predominantly in South of Yangtze River (Shui et al. , Gu , Wei et al. ). They are either terrestrial on thick humus, epiphytic on trees, or epipetric on rock faces or crevices at 100–2,200 m altitude (Shui et al. ). Many Begonia species are local- or site-endemics and even after the publication of Flora of China (Gu et al. ) many new Begonia species have been discovered (Liu et al. , Shui , Wei et al. , Ku et al. , Li et al. , Peng et al. [2008a], [2008b], , , , ), suggesting that the species-rich genus has not been fully explored in China.
Living collections of Begonia jinyunensis (Bo Ding 20130701, Ching-I Peng 24108, 24109, 24110) were used for morphological observations. In addition, herbarium specimens representative of B. jinyunensis were carefully studied in IBSC.
Molecular phylogenetic analyses
Molecular phylogenetic analyses of B. jinyunensis incorporated B. hemsleyana and 21 species (Appendix 1); these 21 species fell in the same clade with B. hemsleyana in a previous phylogenetic study on Asian Begonia based on concatenated sequences of the internal transcribed spacer (ITS) region (including ITS1 and ITS2 and the 5.8S rRNA gene) of 18S-26S nuclear ribosomal DNA (nrDNA) and an intron of rpl16 gene (rpl16) of chloroplast DNA (cpDNA) (Chung et al. ). These species were mostly sect. Platycentrum (14 species including B. hemsleyana) but also included five species of sect. Diploclinium (Lind.) A. DC., two of sect. Sphenanthera (Hassk.) Warb., and B. longicarpa K. Y. Guan & D. K. Tian, currently not assigned to any section (Chung et al., ). Note that Asian sections of Begonia are largely non-monophyletic (Thomas et al. , Chung et al. ). Outgroup taxa were B. parvula H. Lév. & Vaniot of sect. Reichenheimea (Klotzsch) A. DC. and B. ravenii C.-I Peng & Y. K. Chen of sect. Diploclinium, which were placed outside of the clade (Chung et al. ). For B. jinyunensis three plants (Ching-I Peng 24108, 24109, 24110) from three localities in Mt. Jinyun were examined; for B. hemsleyana four plants, three from Yunnan (Ching-I Peng 17590, 20933, 22778) and one from Vietnam (Ching-I Peng 20204), were studied. For the other species one sample each was used. Samples sequenced in this study are indicated with asterisks (Appendix 1) and sequences of the other samples were cited from Chung et al. (). Methods of DNA extraction, PCR, and DNA sequencing followed Nakamura et al. (). DNA sequences were aligned using ClustalX ver. 1.8 (Thompson et al. ) and then manually adjusted. Phylogenetic analyses were based on a Bayesian approach using MrBayes ver. 3.1.2 (Ronquist & Huelsenbeck ) and a maximum parsimony (MP) criterion using PAUP* ver. 4.0b10 (Swofford ).
In the Bayesian phylogenetic analysis, the best substitution models for the ITS and rpl16 data were selected as SYM + I and GTR + I, respectively, using KAKUSAN4 (Tanabe ) based on Bayesian information criterion (BIC). Two separate runs of Metropolis coupled Markov chain Monte Carlo analyses were performed, each with a random starting tree and four chains (one cold and three heated). The chain length was ten million generations, and the chain was sampled every one thousandth generation from the cold chain. The mixing and convergence of the chains of the two runs was assessed using Tracer ver. 1.5.0 (Drummond & Rambaut ). The first 10% of the total 10,000 sample trees were discarded as burn-in. After the burn-in, the effective sample sizes of all parameters were > 200, indicating that the analyses sampled the posterior distributions of each parameter satisfactorily, and the values of Average Standard Deviation of Split Frequency (ASDSF) were below 0.005. The 50% majority rule consensus tree of all the post-burn-in trees with Bayesian posterior probabilities (PP) was visualized with FigTree ver. 1.3.1 (Drummond & Rambaut ).
In the MP phylogenetic analysis, indels were treated as missing data. The characters were treated as unordered, and the character transformations were equally weighted. The branch collapse option was set to collapse at a minimum length of zero. A heuristic parsimony search was performed with 1,000 replicates of random additions of sequences with ACCTRAN character optimization, tree bisection– reconnection (TBR) branch swapping, and MULTREES and STEEPEST DESCENT options on. Statistical support for each clade was assessed by bootstrap analysis (Felsenstein ). Ten thousand replicates of heuristic searches, with the TBR branch swapping switched on and MULTREES options off, were performed to calculate bootstrap percentages (BP).
Somatic chromosome morphology was studied for one plant of B. jinyunensis (Ching-I Peng 24110, HAST) using root tips. The methods of pretreatment, fixation, and staining for chromosome observations followed our previous paper (Peng et al. ). Classification of the chromosome complements based on centromere position at mitotic metaphase followed Levan et al. ().
Results and discussion
Molecular phylogeny based on nrDNA and cpDNA
Type:—CHINA. Chongqing Municipality, Beibei, Mt. Jinyun, Jinyun Mountain National Nature Reserve, 29° 49' N, 106° 20' E, ca. 789 m alt., 10 July 2013, Bo Ding 20130701, with Qian Wang and Hai-Jun Wen (holotype, HAST; isotype, CDBI, E, IBSC, KUN, PE) 縉雲秋海棠.
Begonia jinyunensis bears a superficial resemblance to B. hemsleyana in having palmately compound leaves, a feature unseen in other species of sect. Platycentrum in China. It is however sharply distinct from the latter in the acaulous habit with short aerial stems seen only at anthesis and long rhizomes (vs. erect stems to 70 cm or taller with short rhizomes), 4–6 pinnatilobed leaflets with indistinct, decurrent petiolules (vs. 7–10 serrate leaflets with distinct petiolules), and white (vs. pink) tepals.
Herbs, monoecious; epipetric; perennial; rhizomatous. Rhizomes reddish-brown, 5–15 cm or longer, 6–16 mm across, subglabrous, internodes 2–12 mm. Stipules reddish-brown, widely ovate, ca. 3–6 × 4–6 mm, slightly keeled, apex shortly aristate, subglabrous, deciduous. Leaves many, arising directly from the rhizome, with 1–3 cauline leaves at anthesis; petioles 10–22 cm long, subglabrous or sparely hispidulous. Blade pale to dark green above, tinged reddish (pronounced on veins) beneath, palmately compound, leaflets 4–6, obliquely oblong-lanceolate or obovate-lanceolate, 4–10 × 1.5–3 cm, abaxially sparsely setulose, adaxially sparsely hirsute, margins pinnatilobed, apex caudate-acuminate or acuminate, base cuneate or attenuate into a short, winged petiolule 2–5 mm long. Inflorescence 10–35 cm long, subglabrous; pedicel 1–4 cm; bracts ovate-oblong, 4–5 × 2–3 mm, apex acute. Staminate flowers: pedicel 11–15 mm, tepals 4, white, outer 2 broadly ovate, nearly as long as wide, 12–15 × 11–15 mm, inner 2 oblong-obovate, 8–10 × 4–6 mm; stamens numerous. Pistillate flowers: pedicel 10–40 mm, tepals 5, white, obovate to broadly obovate, 7–15 × 5–10 mm; ovary 2-loculed; placentae axile, bilamellate; styles 2, ca. 5 mm long, fused to halfway. Capsule nodding, ellipsiod, fruit body ca. 10–20 mm long, 7–12 mm thick (wings excluded), unequally 3-winged; abaxial wing falcate-ligulate, 12–18 mm wide. Somatic chromosome number, 2n = 22 (Figure 3).
Additional specimens examined
CHINA. Chongqing Municipality. Mt. Jinyun, Pokong Tower, under shady rocks, 780 m alt., 5 August 1934, T.-H. Tu 5087 (IBSC); Mt. Jinyun ("Tsing-yun Shan"), s.d., received by IBSC in May 1940, L.-Y. Lin 1082 (IBSC); Mt. Jinyun, Jinyun Elementary School, collected and presented by Bo Ding, 29 July, 2013, Ching-I Peng 24108 (HAST); Mt. Jinyun, Qinglongzhai, collected and presented by Bo Ding, 29 July, 2013, Ching-I Peng 24109 (HAST); Mt. Jinyun, Shaolongguan, collected and presented by Bo Ding, 29 July, 2013, Ching-I Peng 24110 (HAST).
Distribution, habitat and ecology
Begonia jinyunensis is currently known only from Jinyun Mountain National Nature Reserve. The species usually grows on limestone faces of shaded moist environments such as valleys and broadleaved forests. Flowering in July; fruiting in August.
The specific epithet is derived from the type locality, Mt. Jinyun.
IUCN Red list category
Vulnerable (VU D2). Begonia jinyunensis is known only from a narrow area of Mt. Jinyun, Chongqing Municipality. Although the area is under protection as a national nature reserve, habitat disturbance brought about by human activities such as tourism and maintenance/building of roads/walking trails may have a negative impact on the species.
Detailed morphological observations and molecular phylogenetic analyses based on nuclear ribosomal DNA and chloroplast DNA sequences supported the recognition of the new species Begonia jinyunensis. Somatic chromosome number of 2n = 22 was reported for this new species. Begonia jinyunensis is currently known only from the type locality, Mt. Jinyun, Chongqing Municipality, Southwest China.
Begonia species included in the molecular phylogenetic analyses. Shown here are species name, section name (Diploclinium–DIP; Platycentrum–PLA; Reichenheimia–REI; Sphenanthera–SPH), voucher specimen (HAST), and GenBank accession numbers for rpl16 and ITS. Asterisks indicate samples sequenced in this study; sequences of the other samples were cited from Chung et al. ().
Begonia alveolata T. T. Yu, DIP, Ching-I Peng 20421, KF707933, AY048977. Begonia augustinei Hemsl., PLA, Ching-I Peng 20759, KF707936, KF636421. Begonia balansana Gagnep., SPH, Ching-I Peng 21928, KF707939, AF485091. Begonia cathayana Hemsl., PLA, Ching-I Peng 20288, KF707948, AF280106. Begonia decora Stapf, PLA, Ching-I Peng 20261, KF707956, KF636435. Begonia dryadis Irmsch., PLA, Ching-I Peng 18016, KF707959, KF636436. Begonia edulis H. Lév., PLA, Ching-I Peng 18747, KF707960, KF636437. Begonia handelii Irmsch., SPH, Ching-I Peng 17513, KF707969, AY048982. Begonia hatacoa Buch.-Ham. ex D. Don, PLA, Ching-I Peng 20861, KF707970, KF636444. Begonia hemsleyana Hook. f., PLA, Ching-I Peng 17590, KF707971, AF485099; Ching-I Peng 20204*, AB972933, AB972927; Ching-I Peng 20933*, AB972934, AB972928; Ching-I Peng 22778*, AB972935, AB972929. Begonia jinyunensis C.-I Peng, B. Ding & Q. Wang, PLA, Ching-I Peng 24108*, AB972936, AB972930; Ching-I Peng 24109*, AB972937, AB972931; Ching-I Peng 24110*, AB972938, AB972932. Begonia laminariae Irmsch., PLA, Ching-I Peng 17447, KF707982, KF636455. Begonia longicarpa K. Y. Guan & D. K. Tian, No section designation, Ching-I Peng 18651, KF707987, AY048985. Begonia longifolia Blume, SPH, Ching-I Peng 16795, KF707988, AF485105. Begonia oreodoxa Chun & F. Chun ex G. Y. Wu & T. C. Ku, PLA, Ching-I Peng 20454, KF707996, KF636467. Begonia palmata D. Don, PLA, Ching-I Peng 20993, KF707998, KF636468. Begonia parvula H. Lév. & Vaniot, REI, Ching-I Peng 20396, KF708001, KF636471 Begonia pavonina Ridl., PLA, Ching-I Peng 20239, KF708002, KF636472. Begonia pedatifida H. Lév., PLA, Ching-I Peng 18779, KF708003, KF636473. Begonia ravenii C.-I Peng & Y. K. Chen, DIP, Ching-I Peng 14855, KF708009, KF636479. Begonia ruboides C. M. Hu ex C. Y. Wu & T. C. Ku, DIP, Ching-I Peng 18705, KF708011, KF636481. Begonia sikkimensis A. DC., PLA, Ching-I Peng 20848, KF708015, KF636485. Begonia taiwaniana Hayata, DIP, Ching-I Peng 18111, KF708020, KF636488. Begonia versicolor Irmsch., PLA, Ching-I Peng 18688, KF708023, AF485090. Begonia wenshanensis C. M. Hu ex C. Y. Wu & T. C. Ku, DIP, Ching-I Peng 20516, KF708025, AY048974.
We thank Qian Wang and Hai-Jun Wen for the help in field collection; the curator and staff of the herbarium IBSC for allowing us to study their specimens; Che-Wei Lin for the fine drawing, and our lab members in Academia Sinica for assistance in cultivation of plant materials in the experimental greenhouse. This study was supported in part by a series of research grants from National Science Council and Academia Sinica, Taiwan to CIP.
- Chung K-F, Leong W-C, Rubite RR, Repin R, Kiew R, Liu Y, Peng C-I: Phylogenetic analyses of Begonia sect. Coelocentrum and allied limestone species of China shed light on the evolution of Sino-Vietnamese karst flora. Bot Stud 2014, 55: 1. doi:10.1186/1999–3110–55–1 doi:10.1186/1999-3110-55-1 10.1186/1999-3110-55-1View ArticleGoogle Scholar
- Doorenbos JM, Sosef MS, de Wilde JJFE: The sections of Begonia including descriptions, keys and species lists. Wageningen Agricultural University, Wageningen; 1998.Google Scholar
- Drummond AJ, Rambaut A: BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evol Biol 2007, 7: 214. doi:10.1186/1471–2148–7-214 doi:10.1186/1471-2148-7-214 10.1186/1471-2148-7-214View ArticlePubMedPubMed CentralGoogle Scholar
- Felsenstein J: Confidence limits on phylogenies: an approach using the bootstrap. Evolution 1985, 39: 783–791. 10.2307/2408678View ArticleGoogle Scholar
- Gu C-Z: Infrageneric classification of Begonia . In Flora of China. Edited by: Wu Z-Y, Raven P, Hong D-Y. Beijing & Missouri Botanical Garden Press, St. Louis; 2007:205–207.Google Scholar
- Gu C-Z, Peng C-I, Turland NJ: Begoniaceae. In Flora of China. Edited by: Wu Z-Y, Raven P, Hong D-Y. Beijing & Missouri Botanical Garden Press, St. Louis; 2007:153–207.Google Scholar
- Kiew R: Begonias of Peninsular Malaysia. National History Publications (Borneo), Kota Kinabalu, Malaysia, and Singapore Botanic Gardens National Parks Board, Singapore; 2005.Google Scholar
- Ku S-M, Kono Y, Liu Y: Begonia pengii (sect. Coelocentrum , Begoniaceae), a new species from limestone areas in Guangxi, China. Bot Stud 2008, 49: 167–175.Google Scholar
- Levan A, Fredga K, Sandberg AA: Nomenclature for centromeric position on chromosomes. Hereditas 1964, 52: 201–220. 10.1111/j.1601-5223.1964.tb01953.xView ArticleGoogle Scholar
- Li H-Z, Ma H, Zhou Z-K, Guan K-Y: A new species of Begonia (Begoniaceae) from Guangxi, China. Bot J Linn Soc 2008, 157: 83–90. 10.1111/j.1095-8339.2008.00771.xView ArticleGoogle Scholar
- Liu Y, Ku S-M, Peng C-I: Begonia bamaensis (sect. Coelocentrum , Begoniaceae), a new species from limestone areas in Guangxi, China. Bot Stud 2007, 48: 465–473.Google Scholar
- Nakamura K, Chung K-F, Huang C-J, Kono Y, Kokubugata G, Peng C-I: Extreme habitats that emerged in the Pleistocene triggered divergence of weedy Youngia (Asteraceae) in Taiwan. Mol Phylo Evol 2012, 63: 486–499. 10.1016/j.ympev.2012.01.023View ArticleGoogle Scholar
- Nakata M, Guan K-Y, Godo T, Lu Y-X, Li J-X: Cytological studies on Chinese Begonia (Begoniaceae) I. Chromosome numbers of 17 taxa of Begonia collected in 2001 field studies in Yunnan. Bull Bot Gard Toyama 2003, 8: 1–16.Google Scholar
- Peng C-I, Liu Y, Ku S-M: Begonia aurantiflora (sect. Coelocentrum , Begoniaceae), a new species from limestone areas in Guangxi, China. Bot Stud 2008, 49: 83–92.Google Scholar
- Peng C-I, Ku S-M, Kono Y, Chung K-F, Liu Y: Two new species of Begonia (sect. Coelocentrum , Begoniaceae) from limestone areas in Guangxi, China: B. arachnoidea and B. subcoriacea . Bot Stud 2008, 49: 405–418.Google Scholar
- Peng C-I, Liu Y, Kono Y: Begonia × breviscapa (Begoniaceae), a new intersectional natural hybrid from limestone areas in Guangxi, China. Bot Stud 2010, 51: 107–117.Google Scholar
- Peng C-I, Ku S-M, Kono Y, Liu Y: Begonia chongzuoensis (sect. Coelocentrum , Begoniaceae), a new calciphile from Guangxi, China. Bot Stud 2012, 53: 283–290.Google Scholar
- Peng C-I, Yang H-A, Kono Y, Chung K-F, Huang Y-S, Wu W-H, Liu Y: Novelties in Begonia sect. Coelocentrum: B. longgangensis and B. ferox from limestone areas in Guangxi, China. Bot Stud 2013, 54: 44. doi:10.1186/1999–3110–54–44 doi:10.1186/1999-3110-54-44 10.1186/1999-3110-54-44View ArticleGoogle Scholar
- Peng C-I, Wang H, Kono Y, Yang H-A: Begonia wui-senioris (sect. Platycentrum , Begoniaceae), a new species from Myanmar. Bot Stud 2014, 55: 13. doi:10.1186/1999–3110–55–13 doi:10.1186/1999-3110-55-13 10.1186/1999-3110-55-13View ArticleGoogle Scholar
- Qing L, Li Y-X, Zhong Z-C: Effects of moisture availability on clonal growth in bamboo Pleioblastus maculata . Plant Ecol 2004, 173: 107–113. 10.1023/B:VEGE.0000026334.40661.06View ArticleGoogle Scholar
- Ronquist F, Huelsenbeck JP: MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 2003, 19: 1572–1574. 10.1093/bioinformatics/btg180View ArticlePubMedGoogle Scholar
- Shui Y-M: Begonia tetralobata (Begoniaceae), a new species from China. Ann Bot Fenn 2007, 44: 76–79.Google Scholar
- Shui Y-M, Peng C-I, Wu C-Y: Synopsis of the Chinese species of Begonia (Begoniaceae), with a reappraisal of sectional delimitation. Bot Bull Acad Sin 2002, 43: 313–327.Google Scholar
- Swofford DL: PAUP: phylogenetic analysis using parsimony, version 4.0b10. Sinauer Associates, Sunderland; 2002.Google Scholar
- Tanabe AS: Kakusan4 and Aminosan: two programs for comparing nonpartitioned, proportional and separate models for combined molecular phylogenetic analyses of multilocus sequence data. Mol Ecol Resour 2011, 11: 914–921. 10.1111/j.1755-0998.2011.03021.xView ArticlePubMedGoogle Scholar
- Tebbitt MC: Begonia: cultivation, identification, and natural history. Timber Press, Portland; 2005.Google Scholar
- Thomas DC, Hughes M, Phutthai T, Rajbhandary S, Rubite R, Ardi WH, Richardson JE: A non-coding plastid DNA phylogeny of Asian Begonia (Begoniaceae): Evidence for morphological homoplasy and sectional polyphyly. Mol Phyl Evol 2011, 60: 428–444. 10.1016/j.ympev.2011.05.006View ArticleGoogle Scholar
- Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG: The ClustalX windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Res 1997, 24: 4876–4882. 10.1093/nar/25.24.4876View ArticleGoogle Scholar
- Tian D-K, Guan K-Y, Zhou Q-X, Gu Z-J: Chromosome numbers of eight taxa of Begonia from Yunnan. Acta Bot Yunn 2002, 24: 245–249.Google Scholar
- Wei Z-D, Shui Y-M, Zhang M-D, Zhang R-M: Begonia coelocentroides YM Shui & ZD Wei, a New Species of Begoniaceae from Yunnan, China. Acta Phytotax Sin 2007, 45: 86–89.Google Scholar
This article is published under license to BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly credited.