Research Article |
Corresponding author: Dai-Ke Tian ( dktian@cemps.ac.cn ) Academic editor: Jan Wieringa
© 2021 Dai-Ke Tian, Wen-Guang Wang, Li-Na Dong, Yan Xiao, Min-Min Zheng, Bin-Jie Ge.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Tian D-K, Wang W-G, Dong L-N, Xiao Y, Zheng M-M, Ge B-J (2021) A new species (Begonia giganticaulis) of Begoniaceae from southern Xizang (Tibet) of China. PhytoKeys 187: 189-205. https://doi.org/10.3897/phytokeys.187.75854
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Begonia giganticaulis, a huge new species in Begonia sect. Platycentrum of Begoniaceae from southern Xizang (Tibet) of China, is described. Morphologically, it is mostly similar to B. longifolia and B. acetosella, but clearly differs from the former mainly by its dioecious and taller plants, sparse hairs on abaxial veins, longer inflorescence, unique shape of fruits, and differs from the latter mainly by its late and longer flowering time, 6-tepals of female flower and 3-loculed ovary. The phylogenetic analyses also support the separation of the new species from other taxa. Based on the current data, its conservation status is assigned to Endangered (B2a) according to the IUCN Red List Categories and Criteria.
Conservation status, molecular evidence, morphology, southern Tibet, taxonomy
Zangnan (southern Tibet) of China is located to the south of the Himalayas, including most parts of Cona, Lhünzê, Mêdog and Zayü counties, and some smaller parts of Nang and Mainling counties (
After a series of plant surveys recently, the authors have a better understanding of the diversity of Begonia in Tibet, particularly in its southern part (namely Zangnan) including Mêdog county. Up until now, 39 species and 4 varieties had been found in Tibet (
Species | Reference | County |
---|---|---|
Begonia aborensis Dunn |
|
Mêdog |
Begonia acetosella Craib |
|
Mêdog |
Begonia annulata K.Koch |
|
Mêdog |
Begonia asperifolia Irmsch. |
|
Bomê, Zayü, Lhünzê, Mêdog |
Begonia burkillii Dunn |
|
Mêdog |
Begonia cathcartii Hook.f. & Thomson |
|
Zayü |
Begonia dioica Buch.-Ham. ex D.Don |
|
Dinggyê |
Begonia difformis (Irmsch.) W.C.Leong, C.I Peng & K.F.Chung |
|
Mêdog |
Begonia flagellaris Hara |
|
Gyirong, Nyalam |
Begonia flaviflora var. flaviflora Hara |
|
Mêdog |
Begonia flaviflora var. gamblei (Irmscher) Golding & Karegeannes |
|
Mêdog |
Begonia giganticaulis D.K.Tian & W.G. Wang sp. nov. | In this study | Mêdog |
Begonia grandis Dryand. |
|
Zayü |
Begonia griffithiana (A.DC.) Warb. |
|
Mêdog |
Begonia handelii Irmsch. |
|
Mêdog |
Begonia hatacoa Buch.-Ham. ex D.Don |
|
Mêdog, Cona |
Begonia iridescens Dunn |
|
Mêdog, Zayü |
Begonia josephii A.DC. |
|
Cona, Dinggyê, Lhünzê, Mêdog, Yadong |
Begonia kekarmonyingensis Taram, D.Borah & M.Hughes |
|
Mêdog |
Begonia labordei H.Lév. |
|
Zayü |
Begonia limprichtii Irmsch.*(wrong identification, based on the distribution and morphological characteristics) |
|
Mêdog |
Begonia longifolia Blume |
|
Mêdog |
Begonia medogensis J.W.Li, Y.H.Tan & X.H.Jin |
|
Mêdog |
Begonia megaptera A.DC. |
|
Zayü |
Begonia nepalensis (A.DC.) Warb. |
|
Cona |
Begonia ovatifolia A.DC. |
|
Mêdog |
Begonia oyuniae M.Taram & N.Krishna |
|
Mêdog |
Begonia plamata var. plamata D.Don |
|
Mêdog |
Begonia palmata var. bowringiana (Champion ex Bentham) Golding & Karegeannes |
|
Mêdog |
Begonia palmata var. khasiana (Irmsch.) Golding & Kareg |
|
Mêdog |
Begonia pasighatensis D.Borah, Taram & Wahlsteen |
|
Mêdog |
Begonia picta Sm. |
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Gyirong, Mêdog, Nyalam |
Begonia pseudoheydei Y.M.Shui & W.H.Chen |
|
Mêdog |
Begonia rex Putz. |
|
Mêdog |
Begonia roxburghii (Miq.) A.DC. |
|
Mêdog |
Begonia scintillans Dunn |
|
Mêdog |
Begonia shilendrae Rekha Morris & P.D.McMillan |
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Cona |
Begonia sikkimensis var. sikkimensisA.DC. |
|
Mêdog |
Begonia sikkimensis var. kamengensis Rekha Morris, P.D.McMillan & Golding ex Golding |
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Cona |
Begonia silletensis Clarke |
|
Mêdog |
Begonia tessaricarpa C.B.Clarke |
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Mêdog |
Begonia thomsonii A.DC. |
|
Mêdog |
Begonia xanthina Hook.f. |
|
Mêdog |
Begonia zhongyangiana W.G.Wang et S.Z.Zhang |
|
Mêdog |
The field surveys were conducted on habitat, distribution, population size, morphology and specimen collection of the new species. Diagnosis of the morphological difference between the new species and its similar species was based on literature review, examination of herbarium specimens, and observation of both wild and cultivated plants.
The treatment on sections of Begonia follows
Sampled taxa and GenBank accession numbers of Begonia giganticaulis and the related taxa used for phylogenetic analysis.
Taxa | Collector, Voucher (Herbarium) | Origin | ITS | rpL16 | ndhA | References |
---|---|---|---|---|---|---|
Begonia acetosella Craib | Wang, W.G., WWG004 (HITBC) | Mengla, Yunnan, China | MW690105 | MW658199 | MW658212 | In this study |
B. acetosella Craib | Wang, W.G., WWG005 (HITBC) | Mengla, Yunnan, China | MW690106 | MW658200 | MW658213 | In this study |
B. acetosella var. hirtifolia Irmsch. | Wang, W.G., WWG0261 (HITBC) | Ruili, Yunnan, China | MW690107 | MW658201 | — | In this study |
B. acetosella var. hirtifolia Irmsch. | Wang, W.G., WWG0262 (HITBC) | Ruili, Yunnan, China | MW690108 | MW658202 | MW658214 | In this study |
B. acetosella var. hirtifolia Irmsch. | Wang, W.G., WWG0300 (HITBC) | Ruili, Yunnan, China | — | MW658203 | MW658215 | In this study |
B. aptera Blume | — | — | AJ491196 | — | JF756369 | Chiang (unpublised); |
B. balansana Gagnep. | — | — | AF485091 | KF707939 | MH207051 |
|
B. cathayana Hemsl. | — | — | AF280106 | KF707948 | KT599095 | Yang et al. (unpublished); |
B. cavaleriei H.Lév. | — | — | KF636430 | KF707949 | MK548079 |
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B. decora Stapf | — | — | KF636435 | KF707956 | JF756355 |
|
B. giganticaulis D.K.Tian & W.G.Wang | Wang, W.G., Li, Y.Y., Ma, X.D. & Shen, J.Y., WWG2014–1 (HITBC) | Mêdog, Tibet, China | MW690097 | MW658191 | MW658204 | In this study |
B. giganticaulis D.K.Tian & W.G.Wang | Wang, W.G., Li, Y.Y., Ma, X.D. & Shen, J.Y., WWG2015–1 (HITBC) | Mêdog, Tibet, China | MW690098 | MW658192 | MW658205 | In this study |
B. giganticaulis D.K.Tian & W.G.Wang | Wang, W.G., Li, Y.Y., Ma, X.D. & Shen, J.Y., WWG2015–2 (HITBC) | Mêdog, Tibet, China | MW690099 | MW658193 | MW658206 | In this study |
B. giganticaulis D.K.Tian & W.G.Wang | Wang, W.G., Li, Y.Y., Ma, X.D. & Shen, J.Y., WWG2014–3 (HITBC) | Mêdog, Tibet, China | MW690100 | MW658194 | MW658207 | In this study |
B. giganticaulis D.K.Tian & W.G.Wang | Wang, W.G., Li, Y.Y., Ma, X.D. & Shen, J.Y., WWG2014–2 (HITBC) | Mêdog, Tibet, China | MW690101 | MW658195 | MW658208 | In this study |
B. handelii Irmsch. | — | — | AF485093 | KF707969 | MH207176 | Forrest and Hollingsworth (unpublished); |
B. hatacoa Buch.-Ham. ex D.Don | — | — | AF485111 | KF707970 | JF756354 | Forrest and Hollingsworth (unpublished); |
B. longifolia Blume | Wang, W.G., WWG001 (HITBC) | Mengla, Yunnan, China | MW690102 | MW658196 | MW658209 | In this study |
B. longifolia Blume | Wang, W.G., WWG002 (HITBC) | Mengla, Yunnan, China | MW690103 | MW658197 | MW658210 | In this study |
B. longifolia Blume | Wang, W.G., WWG003 (HITBC) | Mengla, Yunnan, China | MW690104 | MW658198 | MW658211 | In this study |
B. nepalensis (A.DC.) Warb. | — | — | AY753726 | — | MH207257 |
|
B. obovoidea Craib | — | — | — | — | JF756386 |
|
B. pavonina Ridl. | — | — | KF636472 | KF708002 | JF756356 |
|
B. pedatifida H.Lév. | — | — | MK541092 | MK548068 | MK548115 |
|
B. roxburghii A.DC. | — | — | AF485092 | — | JF756371 |
|
B. versicolor Irmsch. | — | — | AF485090 | KF708023 | JF756358 | Forrest and Hollingsworth (unpublished); |
Sequences of each DNA region were aligned by MUSCLE online (https://www.ebi.ac.uk/Tools/msa/muscle/,
The parsimony analysis was conducted using PAUP v.4.0 b10 (
China. Xizang (Tibet) Autonomous Region: Mêdog county (墨脱县), Beibeng town (背崩乡), Baimu Xiri river (白母西日河), forest slope of river valley or water’s edge along stream, 29°21'9"N, 95°11'21"E, elev. 1320 m, 10 September 2020, Dai-Ke Tian, Fang Wen, Qing-Gong Mao, & Zhu Lu, TDK4773-A (holotype CSH! Barcode number: 0180561, ♀)
Habitat and large-sized plant of Begonia giganticaulis D.K.Tian & W.G.Wang, sp. nov. A habitat showing plants (arrows indicate) growing along stream bank B flowering plant growing along slope of valley C one of the tallest individuals with Dr. Dai-Ke Tian. (Photos A by Dai-Ke Tian B by Shi-Wei Guo C by Qing-Gong Mao).
Morphology of Begonia giganticaulis D.K.Tian & W.G.Wang, sp. nov. A one of the single tallest plants cut into four sections B main stem base C stems showing colour of nodal cross-sections D main stem with much expanded node and whitish-green lines or spots E expanded node on terminal branch F, G male plant branches showing inflorescences and different colours H female branches I adaxially (left) nearly glabrous and abaxially sparse hairs on veins (right, arrows indicate) on blade surfaces J stipules showing shape and colour. (Photo F by Wen-Guang Wang; others by Dai-Ke Tian).
Flower and fruit morphology of B. giganticaulis compared with its close species B. longifolia and B. acetosella A–H Begonia giganticaulis A staminate flowers with pinkish outer tepals B staminate flowers with white tepals C, D pistillate flower E ovary sections showing different colour F fruits on branch G, H dorsal and front views of fruits I–K B. longifolia I flowering and fruiting branch J fruits showing short horns K ovary dissection L–O B. acetosella L staminate flower M Pistillate flower N, O fruits with short horns or wings. (Photos C by Shi-Wei Guo E (left) L, M & O by Wen-Guang Wang; others by Dai-Ke Tian).
The new species is mostly similar to B. longifolia and B. acetosella, but clearly differs from the former mainly by its dioecious (vs. monoecious), taller (to 4 m vs. less than 2 m) plants, longer (vs. shorter) inflorescence, and unique shape of fruits, and differs from the latter mainly by its taller (to 4 m vs. less than 2 m) plants, late and longer (Jun. to Oct. vs. Mar. to Apr.) flowering time, longer (6–20 mm vs. 5–12 mm) pedicel, 6 (vs. 4) tepals of pistillate flower and 3 (vs. 4)-loculed ovary (Table
Morphological comparison of Begonia giganticaulis, B. longifolia and B. acetosella.
Character | B. giganticaulis | B. longifolia | B. acetosella |
---|---|---|---|
Plant | |||
sexuality | dioecious | monoecious | dioecious |
height (m) | up to 4 | less than 2 | less than 2 |
Petiole length (cm) | 0.7–7 | 1–12 | 1–10 |
Leaf blade surface | muriculate | glabrous to less muriculate | muriculate to hirsutulous |
Inflorescence | |||
peduncle length (mm) | 7–15 | 4–10 | 2–10 |
flower number | 1–11 | 1–11(15) | 1–3(5) |
Tepal number of pistillate flower | 6 | 6 | 4 |
Tepal colour | pinkish to white | white | pinkish to white |
Ovary | 3-loculed | 3-loculed | 4-loculed |
Pedicel length (mm) | |||
male flower | 10–20 | 5–12 | 8–12 |
female flower | 6–12 | 5–12 | 5–10 |
Fruit horn or wing | none to rarely short crest | none to short crest or horns | short to long horns or wings |
Flowering time | June-October | June-December | March-April |
Herb perennial, evergreen, to 4 m tall, dioecious. Rhizome short, stout, nearly unbranched, reddish brown, to 12 cm thick. Stem erect, reddish brown or green, glabrous, internodes to 5 cm thick, with many longitudinally fusiform whitish spots, cross section of stem often reddish brown, nodes notably enlarged, to 7 cm thick, with unequally oval to round whitish spots, many shrubby branches on the upper part of main stem. Stipule long-triangular, light green or pinkish green, 9–25 × 2–8 mm, glabrous, margin entire, dorsal ridge pinkish, apex acuminate with arista 4–6 mm long. Petioles green, pink to red, glabrous, 7–22 cm long, 1–3 mm thick. Leaf blade ovate-lanceolate to lanceolate, 4–19 × 0.8–8 cm, adaxial green, muriculate to nearly glabrous, adaxial veins slightly concave; abaxial greyish green, veins usually red, convex, main veins sparsely and obliquely strigose; base obliquely cordate, margin shallowly and remotely denticulate, apex long caudate; Inflorescence dichasial cyme, axillary, short, 3–5 cm long, unbranched to branched once, rachis glabrous, green, pinkish green to red, base usually red-brown, 7–15 mm long, 1–1.5 mm thick, 3–11 male flowers or 1–5 female per inflorescence. Bract often caducous, pinkish green, long triangular, glabrous, ca. 6 × 3 mm, apex acuminate; bracteoles smaller. Staminate flower: pedicel glabrous, white, whitish or pinkish green, 10–14 mm long, ca.1 mm thick; corolla 18–24 mm in diameter; tepals 4, subequal, glabrous, outer 2, obovate, 9–14 × 6–9 mm, apex obtuse, adaxially white and middle-upper part abaxially pink, or pure white for some individuals, longitudinal veins unapparent; inner 2, pure white, obovate to obovate-lanceolate, 8–13 × 5–7 mm, apex obtuse; androecium nearly actinomorphic, ca. 5 mm long, 6–7 mm in diam; stamens 48–60, filaments free, 1–2 mm long; anther yellow, 2–3 mm long, apex obtuse or nearly so. Pistillate flower: pedicel white or green-white, 6–12 mm long, 0.8–1 mm thick; corolla 20–25 mm, tepals 6, rarely 4, glabrous, outer 3 (rarely 2), obovate or long obovate, thick and rigid, 12–18 × 7–10 mm, adaxial surface nearly white, distinctly concave, abaxially pink on middle-upper part, inner 3 (rarely 2), obovate-lanceolate to oblanceolate or long elliptical, slightly narrower than outer tepals, 10–19 × 6–8 mm, white, glabrous, apex obtuse; styles + stigmas 5 mm long, 7–8 mm wide; styles 3, free; stigmas yellow, nearly U-shaped, each side spirally twisted 1.5 circles; ovary pink or green, with white convex spots; placentation axile, 3-loculed, each placenta 2-branched. Peduncle green to pinkish green, glabrous, 8–12 mm long, ca. 1 mm thick. Fruit red, pink or green, glabrous, triangular-gyroscopic, 8–11 × 1–12 mm wide, concave between two placentas, wingless to occasionally short ridged, apex with beak 3–4 mm long. Flowering Jun.–Oct., fruiting Jul.–Dec.
Illustration of Begonia giganticaulis D.K.Tian & W.G.Wang, sp. nov. (Drawn by Mr. Zhi-Min Li) A male flowering branches B female flowering branches C main stem line spots, much expanded node and internode base D expanded node and internode base on small upper branches E leaf blade F leaf (abaxial), showing sparse hairs on veins G stipule H staminate flowers I, J pistillate flower K side view of androecium L stamens M ovary and stigmas N fruit O dissection of ovary showing placentae.
China. Xizang: Mêdog County (墨脱县), Beibeng Town (背崩乡), Baimu Xiri River (白母西日河), forest slope of river valley or water’s edge along stream, 29°21'9"N, 95°11'21"E, elev. 1320 m, 10 September 2020, Dai-Ke Tian, Fang Wen, Qing-Gong Mao, & Zhu Lu TDK4773-B (paratype CSH!, ♂); 29°20'0"N, 95°10'49"E, elev. 1110 m, 10 September 2020, Dai-Ke Tian, Fang Wen, Qing-Gong Mao, & Zhu Lu TDK4765-A, TDK4765-B, (paratype CSH!); 29°18'32"N, 95°10'38"E, elev. 980 m, 10 September 2020, Dai-Ke Tian, Fang Wen, Qing-Gong Mao, & Zhu Lu TDK4777 (paratype CSH!); near Ani Bridge (阿尼桥), 29°17'8.41"N, 95°10'3.23"E, elev. 810 m, 3 July 2020, Wen-Guang Wang, You-Yun Li, Xing-Da Ma, & Jian-Yong Shen, WWG 2014 (paratype, HITBC!), WWG 2015 (paratype HITBC!); elev. 1100 m, 16 September 1974, anonymous 2608 (paratype PE!); elev. 800–1400 m, 30 June 1980, Wei-Lie Chen 10809 (PE!); near No. 2 Bridge, 29°16'42"N, 95°10'49"E, elev. 810 m, 1 October 2017, Dai-Ke Tian, Yan Xiao, Xin Zhong, Li-Zhi Tian & Zhu Lu TDK3429 (paratype CSH!); Beibeng to Hanmi (汗密), elev. 840 m, 7 August 2010, South Tibet Expedition Team (藏南队), Xiao-Hua Jin, Shu-Dong Zhang, Zhong-Yang Li, Bao-Cheng Wu, Xian-Yun Mu, Jing Li & Wei-Tao Jin, STET2304 (paratype PE!); Hanmi to Maniweng (马尼翁), elev. 800–1000 m, 6 August 1974, Qingzang Team 74-4114 (paratype PE!); elev. 1200 m, 24 June 1983, Bo-Sheng Li & Shu-Zhi Chen 05229 (paratype PE!); Maniweng to Ani Bridge, elev. 700–1000 m, 3 August 1972, Tibet Expedition Team, Institute of Biology 1631 (paratype HNWP!).
Begonia giganticaulis is currently found in at least two localities in Mêdog of Tibet. Additional populations might be discovered when more surveys are conducted in China-India border region. However, based on current data, it should be categorised as Endangered: B2a (
The specific epithet refers to the huge (very tall and thick stem) plant size of the new species, which is the tallest begonia in Asia.
We obtained 12 nrITS, 13 rpL16 intron, and 13 ndhA intron of the new species and related Begonia taxa. In order to reconstruct the phylogenetic relationship of the new species, 13 taxa within sect. Platycentrum were included and B. cavaleriei from sect. Coelocentrum was selected as outgroup. In total, the matrix was composed of 26 accessions and contained the 962 bp rpL16 intron, the 1109 bp ndhA intron and the 672 bp nrITS sequence. Of the total 2743 characters, 132 were parsimony informative.
Based on MP analysis, the new species was clustered with B. acetosella and B. acetosella var. hirtifolia (Fig.
Phylogenetic tree inferred by MP A and BI B analyses based on the combined matrix of two plastid loci (rpL16 intron and ndhA intron) and nuclear ITS. Maximum parsimony bootstrap A and Bayesian inference posterior probability values B are labelled on the branches; when the number is below 80 and 0.80 in maximum parsimony bootstrap and Bayesian inference posterior probability, respectively, the branches are labelled—.
Notes. – The earliest specimen of Begonia giganticaulis was collected in 1972 between Maliweng and Ani Bridge, Mêdog, Tibet, China. This species is similar to B. acetosella in appearance when its flowers are unavailable for observation, therefore, it was misidentified (24 June 1983, Bo-Sheng Li & Shu-Zhi Chen 05229, PE! was wrongly identified as B. acetosella by C.Z. Gu in March 2004). Also, due to its high similarity to B. longifolia particularly in morphology of flowers and fruits, B. giganticaulis was wrongly labelled as B. longifolia by
The study was supported by the funds from National Natural Science Foundation of China (31570199, 31860048), the Second Tibetan Plateau Scientific Expedition and Research (STEP) Program (2019QZKK0502), and Shanghai Administration Bureau of Landscape and City Appearance (F122416, G202401). The authors thank Dr. Qing-Gong Mao, Dr. Fang Wen, Mr. Li-Zhi Tian, Mr. Zhu Lu, Mr. Jian-Yong Shen, Mr. Xing-Da Ma and Mr. You-Yun Li, for supporting field survey, and Shi-Wei Guo for providing partial photos for use. The specimens from the Institute of Botany, Chinese Academy of Sciences were reviewed through Chinese Virtual Herbarium.