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Begonia Myanmarica (Begoniaceae), a New Species from Myanmar, and Molecular Phylogenetics of Begonia Sect

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Begonia Myanmarica (Begoniaceae), a New Species from Myanmar, and Molecular Phylogenetics of Begonia Sect Tseng et al. Bot Stud (2017) 58:21 DOI 10.1186/s40529-017-0175-9 ORIGINAL ARTICLE Open Access Begonia myanmarica (Begoniaceae), a new species from Myanmar, and molecular phylogenetics of Begonia sect. Monopteron Yu‑Hsin Tseng1, Young‑Dong Kim2*, Ching‑I Peng1, Khin Myo Htwe3, Seong‑Hyun Cho4, Yoshiko Kono1 and Kuo‑Fang Chung1* Abstract Background: A new species, Begonia myanmarica, was discovered from Myanmar and herein documented. Char‑ acterized by a single developed wing in the ovary/fruit, this species would be assigned to sect. Monopteron (sensu Doorenbos et al. in The sections of Begonia including descriptions, keys and species lists: studies in Begoniaceae VI. Wageningen Agricultural University, Wageningen, 1998) that is known by B. grifthiana and B. nepalensis from the Himalaya. To confrm its sectional assignment, we conducted morphological, phylogenetic and cytological studies. Results: Morphological observations indicated that B. myanmarica was distinguishable from the two known spe‑ cies of sect. Monopteron by the leaf shape and size, 1-locular ovary, parietal placentation and chromosome number. Molecular phylogenetic analysis using nrITS sequences showed that B. myanmarica was not allied with the clade of sect. Monopteron, though both were nested within sect. Platycentrum-sect. Sphenanthera clade. Conclusions: Studies of morphology, molecular phylogenetics and cytology support the recognition of the new species, Begonia myanmarica, which is fully described and illustrated. Our results also indicate that B. myanmarica is not closely related to species previously assigned to sect. Monopteron, suggesting that the fruit morphology of a single developed wing in the ovary/fruit characterizing sect. Monopteron is homoplasious. Keywords: Begonia grifthiana, Begonia nepalensis, Chromosome, Morphology Background (A. DC.) Warb., Parvibegonia A. DC., Petermannia Begonia L. (Begoniaceae), comprising more than 1800 (Klotzsch) A. DC., Platycentrum (Klotzsch) A. DC., species classifed into 68 sections (Doorenbos et al. 1998; Putzeysia (Klotzsch) A. DC., Reichenheimia (Klotzsch) Hughes et al. 2015; Christenhusz and Byng 2016), is one A. DC., Ridleyella Irmscher, and Sphenanthera (Hassk.) of the largest genera of vascular plants. With more than Warb.]. Tereafter, four additional Asian sections were 760 Begonia species in Asia, Doorenbos et al. (1998) proposed [Leprosae (T.C. Ku) Y.M. Shui, Monolobium recognized 18 sections [Alicida C.B. Clarke, Apterobe- T.C. Ku, Pleiothece T.C. Ku, and Symbegonia (Warb.) G. gonia Warb., Baryandra A. DC., Bracteibegonia A. DC., Forrest & Hollingsw.] (Ku 1999; Shui et al. 2002; Forrest Coleocentrum Irmscher, Diploclinium (Lindl.) A. DC., and Hollingsworth 2003; Ku et al. 2007). Tese 22 Asian Haagea (Klotzsch) A. DC., Heeringia Irmscher, Lauchea sections are highly unequal in species numbers: eight of (Klotzsch) A. DC., Monophyllon A. DC., Monopteron the large sections (Petermannia, Platycentrum, Diplo- clinium, Reichenheimia, Coleocentrum, Parvibegonia, *Correspondence: [email protected]; [email protected] Sphenanthera, and Symbegonia) comprise 95% of Asian 1 Research Museum and Herbarium (HAST), Biodiversity Research Center, Begonia species and the rest 14 sections each with less Academia Sinica, Taipei 11542, Taiwan than fve species (Tomas 2010). Several molecular phy- 2 Department of Life Science, Hallym University, Chuncheon, Gangwon 24252, Republic of Korea logenetic studies have demonstrated the paraphyly or Full list of author information is available at the end of the article polyphyly of these large sections, suggesting homoplasy © The Author(s) 2017. This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. Tseng et al. Bot Stud (2017) 58:21 Page 2 of 9 of morphological characters used for current sectional Monopteron, B. grifthiana (Peng 20851) and B. nepalen- delimitations (Tebbitt et al. 2006; Tomas et al. 2011; sis (Peng 20854), cultivated in the greenhouse were also Chung et al. 2014). However, few studies have tested studied as a comparison. the monophyly of small Asian section of Begonia thus far [but see Rajbhandary (2010); Rubite (2010); Tomas Chromosome preparations (2010)]. Root tips were obtained from cultivated materials from Myanmar is botanically a most interesting country, but greenhouse of Academic Sinica. Somatic chromosome of there have been no critical foristic surveys for nearly the new species, B. myanmarica (Peng 23566), and two half a century. Tus far about 60 species of Begonia have species of sect. Monopteron: B. grifthiana (Peng 20851) been recorded from Myanmar (Hughes 2008; Tanaka and and B. nepalensis (Peng 20854), were examined using Hughes 2007; Tanaka and Hayami 2011; Peng et al. 2014b; root tips following the methods by Peng et al. (2014a). Tanaka and Peng 2016). During the feldwork in western Myanmar on 2 February 2012, the second author (YDK) Phylogenetic analyses collected an unknown Begonia with only one devel- DNA sequences of the nuclear ribosomal internal tran- oped wing in ovary/fruit, which is the key character of scribed spacer (nrITS) were used to evaluate the phyloge- Begonia sect. Monopteron sensu Doorenbos et al. (1998) netic relationship among new species and the two species frst delimited by de Candolle (1864) as Mezierea sect. of sect. Monopteron. DNA extraction, PCR amplifcation Monopteron. Presently, only two species, B. grifthiana and DNA sequencing followed Chung et al. (2014). To Warb. and B. nepalensis Warb., are recognized in sect. test the monophyly of sect. Monopteron and sectional Monopteron (de Candolle 1864; Doorenbos et al. 1998). assignment of new species, nrITS of 96 species used in Begonia nepalensis, the type species of sect. Monopteron, Chung et al. (2014) were adopted for phylogenetic analy- is native to Bhutan, Nepal and India (Fig. 5; Doorenbos sis (see Appendix for details). Alignment was conducted et al. 1998; Hughes et al. 2015). Its chromosome num- using MUSCLE implemented in MEGA5.2 (Tamura ber was reported to be 2n = 16 (Legro and Doorenbos et al. 2011) and verifed in Mesquite v3.03 (Maddison 1971), with an uncertain chromosome count 2n = 28–42 and Maddison 2015). Phylogenetic relationships were by Sharma and Bhattacharyya (1961). Begonia grifthi- constructed by Bayesian Inference (BI) method. Te ana, occurring in Bhutan and India (Fig. 5), is character- best nucleotide substitution models were determined by ized by lanceolate to oblong leaves with subcordate base. Modeltest v2.7 (Posada and Crandall 1998). For BI analy- Chromosome number of B. grifthiana was documented sis, the consensus topology was based on Markov chains as 2n = 22 (Doorenbos et al. 1998). Based on recent sys- algorithm implemented in MRBAYES 3.0b4 (Huelsen- tematics and phylogenetics of Begonia, sect. Monopteron beck and Ronquist 2001). Four chains of Markov chain is nested within the Platycentrum-Sphenanthera clade Monte Carlo (MCMC) simulation were carried out for (Rubite 2010; Tomas 2010; Rajbhandary et al. 2011; 1,500,000 generations each with trees sampled per 500 Leong 2017). generations. Te frst 500 trees of sampled trees were dis- Although morphology of the 1-winged ovary/capsule carded before the node probability was calculated (poste- of the undescribed Begonia should be assigned to sect. rior probability: PP). Monopteron, it difers from B. grifthiana and B. nepa- lensis signifcantly the leaf shape, leaf size and distribu- Taxonomic treatment tion. In this study, we described it as a new species. We Begonia myanmarica C.-I Peng & Y. D. Kim, sp. nov. also provide detailed morphological data and molecular (Figs. 1, 2) phylogenetic analysis to elucidate the sectional assign- ment for this species. Type MYANMAR. Sagaing Region, Alangdaw Kathapa Methods National Park, 22°18′ 47.7″ N, 94°28′32.7″ E, alt. 438 m, Morphological observations mixed deciduous forest, along the stream. Living collec- Rhizomes of Begonia myanmarica collected by YDK tion made by Seong-hyun Cho, Young-dong Kim, Yong-in from Myanmar were cultivated in the experimental Kim & Jeong-hun Lee MM-0611, 2 Feb 2012; type speci- greenhouse of the Biodiversity Research Center, Aca- mens (with fowers and fruits) pressed from plants culti- demia Sinica, Taipei, Taiwan. Fully grown plants with vated in the experimental greenhouse, Academia Sinica, fowers and fruits (Peng 23565, 23566) were used for Taipei, Taiwan, 20 Mar 2016, Ching-I Peng 23566 (holo- morphological observation. Te two species of sect. type: RAF; isotypes: HAST, KB). Tseng et al. Bot Stud (2017) 58:21 Page 3 of 9 Fig. 1 Begonia myanmarica C.‑I Peng & Y. D. Kim. A Habit. B Leaf adaxial surface. C Stipule. D Male fower, face view, D′ Male fower, side view. E, E′, E″, E‴ Stamen. F Female fower, face view, F′ Female fower, side view. G Style and stigma. H, H′, H″, H‴ Cross section of ovary. I Capsule Diagnosis Herbs monoecious, perennial. Rhizomes stout, 2–4 cm Begonia myanmarica is a unique species with an erect across, to 9 cm long; erect stem 50–90 cm tall, 1–2 cm habit; large, ovate to broadly ovate leaves (ca. 20–40 cm thick, internodes 5–15 cm, glabrous. Stipules caducous, long, 22–30 mm across); sole, much protruded wing in glabrous, triangular, apex aristate or
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