Research Article |
Corresponding author: Yun-Hong Tan ( tyh@xtbg.org.cn ) Academic editor: Xiao-Hua Jin
© 2020 Mya Bhone Maw, Hong-Bo Ding, Bin Yang, Pyae Pyae Win, Yun-Hong Tan.
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:
Maw MB, Ding H-B, Yang B, Win PP, Tan Y-H (2020) Taxonomic studies on Begonia (Begoniaceae) in Myanmar I: three new species and supplementary description of Begonia rheophytica from Northern Myanmar. In: Jin X-H, Xia N-H, Tan Y-H (Eds) Plant diversity of Southeast Asia-II. PhytoKeys 138: 203-217. https://doi.org/10.3897/phytokeys.138.38721
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Three new species of Begonia (B. chenii, B. putaoensis and B. crassitepala) belonging to Begonia section Platycentrum and a supplementary description of B. rheophytica with a detailed description of female flowers from Putao, Kachin State, Northern Myanmar, are described and illustrated. All the new species are endemic to Northern Myanmar and can be easily distinguished from other species among the section Platycentrum. A detailed description, photographs, habitat, distribution and a comparison with the most related allied species for all new species are provided.
Kachin State, Sect. Platycentrum, Putao District, Sect. Sphenanthera
Begonia
During floristic surveys of northern Myanmar from 2016 to 2018, some interesting Begonia specimens were collected. After conducting a detailed examination of the morphological characteristics of the collected material, reviewing the type specimens and taxonomic publications, the authors have confirmed that the specimen of Begonia collected from northern Myanmar belong to species new to science, which are described and illustrated below.
Historically, based on the characters of axial placentation, 3 or 4-locular ovary, berry-like and wingless fruit, Begonia chenii should belong to Begonia sect. Sphenanthera (
Measurements and morphological character assessments of the new species have been examined based on fresh materials and dried specimens. They have been compared with morphologically similar species by affinities inferred using descriptions (
Begonia chenii Y.H. Tan, M.B. Maw & H.B. Ding is mostly similar to B. mariachristinae
Myanmar. Kachin State: Putao District, on the way from Putao to Upper Shankhaung, in tropical rain forest, 27°25'36.87"N, 97°16'13.56"E, 512 m, 4 May 2017, Y.H. Tan, B. Yang, H.B. Ding, X.D. Zeng, M.B. Maw & T.S. Tin M1378 (holotype: HITBC!; isotypes: RAF!).
Perennial herb, dioecious or rarely monoecious, lacking rhizome or tuber. Stem erect, 40−60 cm tall, reddish brown, densely white pilose, internode 2−11 cm long, branching. Stipule persistent, ovate, 1–15 × 3–5 mm, papery, keeled, apex cuspidate (1–4 mm), margin entire, slightly pilose. Leaf alternate, petiole 1.5–3 cm long, reddish-brown, densely reddish pilose; blade asymmetric, lanceolate-ovate, 8–11 × 2.5–4 cm, apex attenuate, base oblique, margin serrate and with reddish hispid, venation palmate-pinnate, 5−6 pairs of veins; upper surface green or dark green with white patches and dots between the veins, bright green shot with metallic blue depending on the angle of the light, especially on young leaves, sparsely reddish hispid, especially along the midrib and lateral veins; lower surface deep red or deep red with light green areas both margin linings, scattered reddish hispid and densely along the midrib and lateral veins. Inflorescence axillary, flower solitary or in a simple cyme, pendulous; bract persistent, ovate to narrow lanceolate, 4–8 × 2–3 mm. Staminate flower: pedicel 0.8–1.1 cm, reddish, glabrous or sparsely pilose; tepals 4 (rarely 5 or 6), reddish with white margins, unequal, inner 2 (rarely 3), ovate, 7–1 × 6–1 mm, glabrous, outer 2 (rarely 3), ovate, 7–9 × 7–1 mm, reddish or whitish pilose on the outer surface, margin entire; androecium actinomorphic, stamens numerous, filament free, anther oblong, golden yellow. Pistillate flower: pedicel 0.6–0.8 cm, tepals 6 (rarely 4 or 7), unequal, inner 3, elliptic, 8–10 × 3–4 mm, pink to white, glabrous, outer 3, ovate or elliptic, 7–11 × 5–7 mm, reddish or whitish pilose on outer surface; ovary red, slightly or densely pilose on the surface, triangular or rhomboid winged, 3–5 × 2–5 mm, placentation axillary, locules 3 or 4, placentae 2 per locule; styles 3, fused at base, stigma bifid with twisted bands, highly convolute, yellow or golden yellow. Fruit berry-like, red, reddish or whitish pilose, triangular, rhomboid (8–15 × 6–9 mm) or suboblate (8–15 mm in diam.), 3 or 4 horned, rarely wingless.
Begonia chenii Y.H.Tan, M.B.Maw & H.B.Ding, sp. nov. (photographed by H.B. Ding and Y.H. Tan) A habitat B leaves (back view) C inflorescence D staminate flower showing 6 tepals E pistillate flower F infructescence showing monoecious G infructescence showing stigmas H staminate flower showing variation of 4, 5, 6 tepals I tepals of staminate flower J androecium K tepals of pistillate flower L ovary and stigma M ovary (3-winged) N ovary (4-winged) O–P serial cross section of ovary (locules 4) Q serial cross section of ovary (locules 4 and wingless) R serial cross section of ovary (locules 3).
Flowering from April to May; fruiting from May to June.
The species is only known from the type locality, Putao District, Kachin State, Northern Myanmar.
The species grows in the moist shaded environment of tropical rain forest, elevation about 512 m.
The species epithet “chenii” is named after Professor Chen Jin, the director of Southeast Asia Biodiversity Research Institute, Chinese Academy of Sciences, who gave us the opportunity to study the Myanmar flora, which led to the discovery of this new species.
Data Deficient (DD). Begonia chenii was collected along the path on the way from Putao to Upper Shankhaung where any signs of major anthropogenic disturbance were noticed in the type locality. However, further explorations are needed for a proper assessment of conservation due to insufficient information on its distribution and population status. Therefore, the species has been preliminarily assigned to Data Deficient (DD) category according to The Guidelines for Using The IUCN Red List Categories and Criteria (
(paratypes). Myanmar. Kachin State: Putao District, Upper Shankhaung, in tropical montane forest, 27°25'36.87"N, 97°16'13.56"E, 512 m, 4 May 2017, Y.H. Tan, B. Yang, H.B. Ding, X.D. Zeng, M.B. Maw & T.S. Tin M1379 (HITBC!); Kachin State: Putao District, Upper Shankhaung, 27°25'35"N, 97°16'14"E, 500 m, 29 April 2016, Y.H. Tan & S.S. Zhou M201627 (HITBC!); Kachin State: Putao District, Upper Shankhaung, 27°25'34"N, 97°16'13"E, 520 m, 7 May 2017, S.S. Zhou & X.D. Zeng M2030 (HITBC!; RAF!)
Affinities. Begonia chenii is morphologically similar to B. mariachristinae. But it can be easily distinguished in having 3 or 4 locules (vs. 2 locules). See Table
Comparison of key morphological characters of Begonia chenii and B. mariachristinae.
Attributes | B. chenii | B. mariachristinae |
Stem | 40–60 cm tall, reddish brown densely white pilose | 40–60 cm tall dark red to maroon, hairs |
Stipules | persistent, slightly pilose | persistent, glabrous |
Petiole | 1.5–3 cm long, densely reddish pilose | 1–5 cm long, sparsely puberulous |
Leaves | lanceolate-ovate, 8–11 × 2.5–4 cm | lanceolate-ovate 6–11.5 × 2.5–4.5 cm |
Upper surface | sparsely reddish hispid especially along the midrib and lateral veins | slightly reddish hispid |
Lower surface | scattered reddish hispid and densely along the midrib and lateral veins | slightly hairy along midrib |
Male flower | tepals 4 (rarely 5 or 6) reddish with white linings | tepals 4 pink or white |
Female flower | tepals 6 (rarely 4 or 7) pink to white | tepals 4 pink or white |
Ovary | locules 3 or 4 red, slightly or densely pilose | locules 2 green, hispid |
Style | 3 | 2 (or 3) |
Begonia putaoensis Y.H. Tan, M.B. Maw & H.B. Ding is morphologically similar to B. scintillans
Myanmar. Kachin State: Putao District, on the way from Camp 2 to Camp 3 along Putao to Madwel, on moist rocky slope in tropical rain forest, 27°39'35"N, 97°22'30"E, 505 m, 29 November 2017, Y.H. Tan, B. Yang, H.B. Ding, X.D. Zeng, M.B. Maw & P.K. Linn M2923 (holotype: HITBC!; isotypes: RAF!).
Perennial herb, monoecious, 10–25 cm tall. Rhizome creeping with adventitious roots, sometimes branched, 3–30 cm long, 5–12 mm thick, reddish brown, densely pubescent or rusty villous, internode short, 3–8 mm long. Stipule triangular, 6–12 × 5–6 mm, apex cuspidate (3–5 mm), margin entire, rusty colored, densely rusty tomentose on both surfaces, persistent; petiole scarlet red to crimson, cylindrical, 3–15 cm long, densely rusty tomentose. Blade ovate to widely ovate, 6.5–15 cm long, 6–11cm wide, asymmetric, adaxially dark green with gray or light-green areas, slightly pubescent, abaxially light-green, deep red along veins, rarely entirely dark red or red with light-green areas, slightly pubescent, rusty villous on the veins; base cordate, apex acuminate to attenuate, margin sinuate, with sparse hairs; venation palmate, 7–9 veined, adaxial slightly impressed, abaxial distinctly prominent. Inflorescence axillary, sub-corymb, erect, branching 2–3 times, 7–19 cm long. Primary peduncle 5.5–14.5 cm long, densely rusty tomentose, dark red, secondary 0.6–1.5 cm long; bract ovate to lanceolate or obovate, 8–13 × 3–7 mm, glabrous, apex acute, margin entire, sometimes with ciliate, 2–10 flowers per inflorescence, male flowers open earlier at the same node. Staminate flower: pedicel white or pink, glabrous, 0.9–3.7 cm long, tepals 4, rarely 6, outer 2 (or 3) larger, pink, ovate to suborbicular, 1.3–1.8 × 1.1–2.1 cm, glabrous or abaxially strigose; inner 2 (rarely 3), smaller, white-pinkish, ovate or obovate, 9–17 × 7–11 mm., glabrous; androecium 4–6 mm long, 5–7 mm in diameter; stamens numerous, filaments ca. 1.7 mm long, anthers yellow, obovate, nearly 1.2 mm long, apex obtuse. Pistillate flower: pedicel dark red, glabrous, 2.6–3.3 cm long, tepals 5, rarely 6, outer 3, larger, pink, ovate, 14–16 × 1–13 mm, inner 2 (rarely 3), smaller, white-pinkish, ovate to suborbicular, 12–16 × 8–11 mm; ovary glabrous, 2-loculed, placentae axillary, placentae 2 per locule, styles 2 or 3, fused at base, stigma bifid with twisted bands, highly convolute, yellow or golden yellow. Capsule nodding, ovoid, glabrous, unequally 3-winged; abaxial wing nearly round-rectangular, 13–17 mm broad, lateral wings shorter, 2.5–3.0 mm broad.
Begonia putaoensis Y.H.Tan, M.B.Maw & H.B.Ding, sp. nov. (wild, photographed by H.B. Ding) A–B habitat C rhizome D inflorescence E stipule on stem F–H single leaf (front and back view) I flowers (front view) J flowers (back view) K outer tepals of male flower (back view) L inner tepals of male flower (back view) M androecium with pedicel.
Begonia putaoensis Y.H.Tan, M.B.Maw & H.B.Ding, sp. nov. (cultivated plants, photographed by H.B. Ding) A habit B staminate flower showing 4 tepals C staminate flower showing 6 tepals D pistillate flower showing 6 tepals and 3 styles E pistillate flower showing 5 tepals and 2 styles F rhizome G stipule (back view) H stipule (front view) I bract J uppermost bract K stigma (3, front view) L stigma (2, side view) M androecium (front view) N fruit with unequal wings O serial cross section of ovary.
Flowering from November to December; fruiting from December to February.
The species is known from the single locality in Putao District, Kachin State, Northern Myanmar.
The species grows on moist rocky slopes of tropical montane forest, elevation 500–900 m.
The species epithet refers to the type locality of the species, Putao District, Kachin State, Northern Myanmar.
Data Deficient (DD). The species might not confront strong human pressures because of the remoteness of its type locality. But we cannot assess the species’ risk of extinction due to lack of data. Therefore, the species is temporarily assigned a status Data Deficient (DD) according to The Guidelines for Using The IUCN Red List Categories and Criteria (
Begonia putaoensis is mostly similar in morphological characters to B. scintillans and B. annulata K.
Comparison of key morphological characters of Begonia putaoensis, B. scintillans and B. annulata.
Attributes | B. putaoensis | B. scintillans | B. annulata |
Habit | rhizomatous, creeping 15–25 cm tall | rhizomatous, creeping 7–15 cm tall | rhizomatous, erect 15–30 cm tall |
Internode | short, 0.3–0.8 cm long | 3–5 cm long | 0.7–1.5 cm long |
Stipule | triangular, 6–12 × 5–6 mm densely rusty tomentose on both surfaces | lanceolate, 6–11 × 4–6 mm villose on outer surface | lanceolate, 4 –13 × 2 – 6 mm tomentose on outer surface |
Leaf | ovate to widely ovate 6.5–15 × 6–11cm | ovate-orbicular 4.5–10 × 3.5–7 cm | ovate 9–15 × 5–10 cm |
Upper surface | dark green with silver or light green areas sparsely pubescent | dark green with small silver spots, densely strigose | dark green with white/silver bands slightly tomentose or strigose |
Lower surface | light green, deep red along veins entire dark-red or red with light green area slightly pubescent, rusty villous on the veins | red, densely red tomentose | red and green, strigose |
Staminate flower | tepals 4 (rarely 6), pink or white-pinkish | tepals 4, coral pink | tepals 4, white to pink |
Pistillate flower | tepals 5 (rarely 6), glabrous pink or white-pinkish | tepals 4–5, coral pink, pilose on outer surface | tepals 4–5, white to pale pink, puberulous on outer surface to glabrous |
Style | 2 or 3 | 3 | 2 |
Capsule | ovoid, glabrous, 3-winged longest one round-rectangular 13–17 mm broad | obovoid, red villose, 3-winged longest one rounded oblong 4–6 mm broad | ellipsoid, tomentose, 3-winged longest one rounded oblong 5–9 mm broad |
(paratypes). Myanmar. Kachin State: Putao District, near around Camp 5, along Putao to Madwel, on moist rocky slopes in tropical rain forest, 27°43'59.51"N, 97°22'52.27"E, 873 m, 3 December 2017, Y.H. Tan, B. Yang, H.B. Ding, X.D. Zeng, M.B. Maw & P.K. Linn M3168 (HITBC!; RAF!); MYANMAR. Kachin State: Putao District. Voucher from the cultivated plant in the Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, 12 November 2018, H.B. Ding XTBG-0050 (HITBC!).
Begonia crassitepala Y.H. Tan & M.B. Maw is morphologically similar to B. dryadis
Myanmar, Kachin State, Putao District, on the way from Ratbaw to Alanga, 27°17'13.73"N, 97°44'24.28"E, 836 m, 15 June 2018, Y.H. Tan, B. Yang, H.B. Ding, X.D. Zeng, M.B. Maw & H.L. Naing M4495 (holotype: HITBC!; isotypes: RAF!).
Perennial herb, monoecious, rhizomatous. Stem erect, 40−60 cm tall, reddish, densely rusty wooly tomentose, and sparsely covered by whitish soft spine-like hairs, internodes 1−2 cm long. Stipule lanceolate, 1−1.3 × 0.3−0.5 cm, slightly or densely rusty tomentose, deciduous. Leaf petiole 4.5−15 cm, densely rusty tomentose, slightly whitish stiff hairs; blade ovate to broadly ovate, base cordate with overlapping lobes, 12−20 × 15−19.5 cm, asymmetric, adaxially green, densely pinkish or grey hirsute, abaxially green, rusty puberulous, venation palmate-pinnate, densely rusty wooly tomentose along midrib and veins, margin denticulate, reddish hirsute along the margin, apex acuminate. Inflorescence nearly terminal, cymose, peduncle 4.4–6.0 cm long, reddish, slightly rusty wooly tomentose and whitish soft spine-like hairs. Staminate flower: pedicel 1.8–2.5 cm long, whitish villose, tepals 4, unequal, outer 2, ovate, 2.2−2.7 × 2.6−2.8 cm, whitish or pinkish with pink lining, thick, ca. 2 mm, inner 2, smaller, 1.7 × 2.1 cm, whitish or pinkish (sometimes with pinkish lining), densely whitish villose on the outer surface; stamen numerous, ca. 200; filaments ca. 2 mm long, fused at base; anther oblong to elliptic, 1–2 mm long. Pistillate flower: pedicel 2.2–2.5 cm, densely whitish or rusty villose, bracteoles absent; tepals 5, equal, obovate, outer 2, 2.0–2.9 × 1.8–2.0 cm, pure white or sometimes with pink lining, densely pinkish or whitish strigose on outer surface, margin entire, inner 3, similar to outer ones but smaller, 1.8–2.1 × 1.6–1.8 cm, ovary 2-locular, placentation axillary, placentae 2 per locule, densely reddish or whitish succulent strigose and tuberculate, styles 2, fused at base, stigma bifid with twisted bands, highly convolute, yellow or golden yellow. Fruit berry-like, elliptic, pale green to pink, 3 wings, unequal, with whitish or reddish succulent strigose and tuberculate (especially on wings).
Begonia crassitepala Y.H.Tan & M.B.Maw, sp. nov. (photographed by H.B. Ding) A habit B inflorescence C stipule and stem (showing the whitish soft spine-like hairs) D rhizome E single leaf (front view) F single leaf (back view) G hirsute leaf margins H flowers (front view staminate flower) I staminate flower (back view) J dried fruits K pistillate flower (front view) M staminate flower showing 4 tepals N pistillate flower (side view) L capsule and cross section of ovary.
Flowering from June−July; fruiting from July−August.
Endemic to the type locality, Putao District, Kachin State, Northern Myanmar.
The species epithet refers to its thick tepals.
In the tropical montane forest up to about 577 m elevation, on the moist soil slope.
Data Deficient (DD). Begonia crassitepala have been collected along the roadside from Langsa to Gawlaw village where no signs of major anthropogenic disturbance were noticed. Further exploration is required to access the current range of the species (
(paratypes). Myanmar. Kachin State: Putao District, along Langsa to Gawlei, tropical montane forest, 27°32'28.94"N, 97°56'36.09"E, 577 m, 2 June 2018, Myanmar Exped. M3952 (HITBC!; RAF!); Kachin State, Putao District, Gathu to Tongwang Cave, 27°29'53.48"N, 97°58'30.84"E, 664 m, 4 June 2018, Myanmar Exped. M4008 (HITBC!; RAF!); Kachin State, Putao District, Gathu to Tangsa, 27°28'41.17"N, 97°56'46.40"E, 550 m, 31 May 2018, Myanmar Exped. M3797 (HITBC!; RAF!); Kachin State, Putao District, Putao Township, Maliraing area, buffer zone of Hkakaborazi National Park, between camp 1 and camp 2, 27°38'03.6"N, 97°22'11.2"E, 552 m, 6 November 2016, Kate et al. 2253 (NY02653741!); Kachin State, Putao District, Naungmung Township, buffer zone of Hkakaborazi National Park, Hill next to Naungmung village, 27°31'02.2"N, 97°50'46.2"E, 845 m, 12 June 2017, Kate et al. 2880 (NY02653917!); Between N Dung Ga and Ting Pru Ting Sar, 27 August 1953, Thar Hla & Chit Ko Ko 4447 (RAF!).
Begonia crassitepala is the most distinct species in the section Platycentrum thanks to its thicken tepals and succulent strigose and tuberculate ovary. The new species shares similar characteristics with B. dryadis in ovate to broadly ovate leaf and 4 tepals of staminate flower. However, it can be easily distinguished by its stem and petiole having rusty tomentose and whitish soft spine-like hairs (vs. puberulous), adaxially leaf having densely pinkish or grey hirsute (vs. subglabrous), adaxially outer 2 tepals of pistillate flower having densely pinkish or whitish strigose (vs. puberulous), ovary having densely reddish or whitish succulent strigose and tuberculate (vs. puberulous).
Myanmar. Hills east of the Mali Hka, 2000−3000 ft, xii 1930, Kingdon-Ward 9067 (holotype: BM000896328; isotypes: BM000896327, NY02652766).
Herb, rhizomatous, firmly rooted to rock. Rhizome 2.5−4.0 cm long and 0.5−1.0 cm in diam., internode 0.2−0.7 cm long. Stipule reddish brown, eventually deciduous, narrowly triangular, 0.7−1.0 × 0.3−0.5 cm, keeled, margin entire, glabrous. Leaf petiole deep red or deep red to green, turns to brown in mature leaves, sparsely or densely white pilose, 4−17 cm in length, deeply grooved above; blade symmetric, narrowly lanceolate, 13.4−18.2(−21) × 2.2−4.0 cm, base attenuate, sometimes unequal, margin red, toothed, teeth tipped by a short ciliate, sometimes undulate, apex elongate; adaxially dark green, glabrous; abaxially pale green, veins densely or sparsely white pubescent; venation pinnate, 5−7 pairs of veins, reddish green in young leaves, red in mature leaves. Inflorescence axillary, cymose, peduncle erect, 14−20 cm long, reddish green or pale reddish brown, sparsely hairy; bract caducous, broadly ovate−triangular (or ovate−lanceolate), 6−8 × 3−5 mm, purplish red to dark yellow-green, margin entire, hairless. Staminate flower: pedicel 1.9−2.1 cm, pale pink (or) pinkish white, sparsely hairy; bracteoles ca. 3 mm, narrowly ovate, dark yellow green, margin entire, hairless, soon falling, tepals 4, unequal, inner 2, elliptic, 1.4−1.7 × 0.8−1.0 cm, pure white to pinkish, margin entire, outer tepals 2, broadly ovate, 1.1−1.8 × 1.1−1.5 cm, pure white or pinkish, tip rounded, margin entire; androecium actinomorphic; stamens numerous, filaments fused at base; anther golden yellow, narrowly oblong, apex rounded. Pistillate flower: pedicels 2−3 cm, reddish or purplish red, hairless; bracteole narrowly ovate 6−8 × 3−5 mm, dark yellow green (sometimes crystal white), soon falling; tepals 5−6, unequal, outer 2, equal, broadly ovate, tip rounded, 0.7−1.2 × 0.8−1.2 cm, pure white or white to rosy pink, inner 3 or 4, unequal, 3 larger, 0.8−1.2 × 0.7−1.0 cm, elliptic, pure white or pinkish, tip rounded, 1 smaller, ca. 0.8 × 0.3 cm, pure white, crescent, styles 2, free, stigma bifid with twisted bands, greenish yellow, 4−5 mm long, ovary purplish red, ca. 9−15 mm long, 2−4 mm in diam., wings 3, unequal, placentation axile, locules 2, placentae 2 per locule. Capsule nodding, 3-winged, unequal, major wing 8−12 mm long, lateral wings 2−3 mm long.
Begonia rheophytica M. Hughes (photographed by H.B. Ding and Y.H. Tan) A habitat B staminate flower (front view) C pistillate flowers D inflorescences E staminate flowers (front and back view) F single leaf (back view) G single leaf (front view) H pistillate flower I ovary with gynoecium, pedicel and bracts J ovary with gynoecium K bracts L pedicel M androecium with pedicel N tepals of staminate flower O tepals of pistillate flower P capsule Q–R cross section of ovary.
Only found in Putao District, Kachin State, Northern Myanmar.
Myanmar, Kachin State, Putao District, Camp 1 to Namti (Camp 2), understory herbs in tropical rain forest, 27°24'36.80"N, 97°39'24.38"E, 801 m, 12 December 2018, Myanmar Exped. M3427 (HITBC!; RAF!); Kachin State, Putao District, near around Camp 1, understory herbs in tropical rain forest, 27°24'18.70"N, 97°36'24.18"E, 850 m, 11 December 2018, Myanmar Exped. M3334 (HITBC!; RAF!); Kachin State, Putao District, humid rocks near streams or near caves by waterfall of tropical montane forest, 27°24'46.31"N, 93°39'36.28"E, 808 m, 16 December 2017, Myanmar Exped. M3747 (HITBC!, RAF!).
This species was originally described by
The authors would like to express their gratitude to the Forest Research Institute of Myanmar for permission to conduct this study in the Northern Myanmar, and for their support and collaboration. We are grateful to Xiao-Dong Zeng, Kyaw Swar and Myint Zaw for their kind help in fieldwork. This work was financially supported by a grant from the National Natural Science Foundation of China (Grant no. 31970223) awarded to Yun-Hong Tan, a project of the Lancang-Mekong Cooperation (LMC) Special Fund (Biodiversity Monitoring and Network Construction along Lancang-Mekong River Basin project), CAS “the Belt and Road” Master Fellowship Program and CAS President’s International Fellowship Initiative (PIFI) and the Southeast Asia Biodiversity Research Institute, Chinese Academy of Sciences (Y4ZK111B01) and the CAS 135 program (No. 2017XTBG-F03).