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Hebius modestus (GÜNTHER, 1875)

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Higher TaxaColubridae (Natricinae), Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Subspecies 
Common NamesE: Modest Keelback
Chinese: 腹斑腹链蛇 
SynonymTropidonotus modestus GÜNTHER 1875: 232
Tropidonotus modestus — BOULENGER 1893: 229
Tropidonotus johannis SMITH 1921 (non BOULENGER; fide SMITH 1943)
Nerodia modesta — WALL 1923
Natrix modesta — MELL 1929: 5
Natrix modesta — SMITH 1943: 290
Amphiesma modesta — MALNATE 1960
Paranatrix modesta — MAHENDRA 1984
Amphiesma modesta — COX et al. 1998: 45
Amphiesma modestum — ZHAO et al. 2000: 205
Amphiesma modesta — SHARMA 2004
Amphiesma modestum — ZHAO 2006
Hebius modestum — GUO et al. 2014
Amphiesma modestus — GUO et al. 2014: 432
Hebius modesta — GUO et al. 2014: 428
Hebius modestum — GUO et al. 2014: 437
Amphiesma modestum — WALLACH et al. 2014: 31
Hebius modestus — ZHOU et al. 2019
Hebius modestus — DAVID et al. 2021 
DistributionIndia (Meghalaya; Arunachal Pradesh), Myanmar (= Burma: Kachin; Shan), China (Yunnan)

Type locality: Khasi Hills, India  
Reproductionoviparous 
TypesHolotype: BMNH 1946.1.13.41 
DiagnosisDiagnosis. A moderately sized species of the genus Hebius characterized by the combination of (1) 19-19-17 dorsal scale rows, barely or weakly keeled at midbody and on the posterior part of the body, smooth on 1st DSR; (2) dorsal scales around the base of the tail weakly keeled; (3) head moderately distinct from the neck; (4) eye large; (5) maxillary teeth 27–30, the last two moderately enlarged; (6) tail long, with a ratio TaL/TL at least equal to 0.28 in females, up to 0.32 in males; (7) VEN 143–163; (8) SC 104–122; (9) prefrontal scales 2; (10) 1 or rarely 2 anterior temporals, rectangular and elongate; (11) venter pale ochre-brown or pale brown on a wide central area, broadly edged with dark brown or blackish on the outer quarter of ventrals, rarely entirely dark except on its most anterior part; (12) dorsal colour dark greyish-brown, brown or dark brown; (13) dorsum usually scattered with blackish-brown or black spots or blotches; (14) on each side, a more or less conspicuous, ochre-yellow, ochre-red, orange-brown or reddish-brown stripe, often reduced to a succession of elongate blotches on the anterior part of the body, extending from the nape to the base of the tail on 4th–7th or 5th–7th DSR, often but not always irregularly edged below and above by a series of large blackish-brown blotches; (15) no postocular streak; and (16) a short but broad, horizontal, pale yellowish-brown streak on the sides of the neck before the dorsolateral stripe, plus a streak on the nape behind the parietals (David et al. 2021).

Comparisons. Hebius modestus differs from the group of H. venningi, i.e., H. venningi, H. taronensis and H. nigriventer, with which it might be sympatric in northern India and Myanmar (Kachin State) by (1) the number of 19 DSR at midbody; (2) the general dorsal pattern, usually striped and with dark blotches above and below the dorsolateral stripe vs. not striped but chequered or, in H. nigriventer, with conspicuous, large dorsolateral blotches; (3) venter usually widely pale along its whole length, edged with blackish-brown, rarely entirely dark except on its most anterior part vs. (a) venter pale (coral red in life) mesially, on the anterior part of the body and dark, clouded posteriorly in H. venningi; (b) venter both pale and dark, i.e. with a pale background on its anterior part with irregular dark crossbands and nearly uniform on its posterior part in H. taronensis and (c) venter nearly entirely very dark in H. nigriventer; and (4) a conspicuous yellow streak on the sides of the neck (David et al. 2021).

Sexual dimorphism: (1) difference in the ratio TaL/TL: males: 0.320–0.326 (mean = 0.323, s = 0.003); females: 0.288–0.307 (mean = 0.299, s = 0.007). (2) Strongly by the difference in the number of subcaudals: males: 113–122 (mean = 117.0, s = 3.2); females: 104–109 (mean = 107.0, s = 2.2)(David et al. 2021). 
CommentType species: Mahendra (1984) designated Tropidonotus modestus Günther 1875 as the type species of his genus Paranatrix.

Distribution: possibly in Bhutan (Lenz 2012). Records from Vietnam are questionable according to DAVID et al. (2005). Not in Assam, Mizoram (India), not in Laos, Vietnam, Cambodia, not in Guizhou, Guangdong (China), not in Mizoram (India), the specimen depicted by Ahmed et al. (2009: 154) as Amphiesma cf. modestum is a specimen of Hebius venningi, fide P. David, pers. comm., 20 Apr 2021). 
References
  • Boulenger, G.A. 1893. Catalogue of the snakes in the British Museum (Nat. Hist.) I. London (Taylor & Francis), 448 pp. - get paper here
  • Boulenger, George A. 1890. The Fauna of British India, Including Ceylon and Burma. Reptilia and Batrachia. Taylor & Francis, London, xviii, 541 pp. - get paper here
  • Cox, Merel J.; Van Dijk, Peter Paul; Jarujin Nabhitabhata & Thirakhupt,Kumthorn 1998. A Photographic Guide to Snakes and Other Reptiles of Peninsular Malaysia, Singapore and Thailand. Ralph Curtis Publishing, 144 pp.
  • David, P., Bain, R. H., Nguyen Quang Truong, Orlov, N. L., Vogel, G., Vu Ngoc Thanh & T. Ziegler 2007. A new species of the natricine snake genus Amphiesma from the Indochinese Region (Squamata: Colubridae: Natricinae). Zootaxa 1462: 41-60 - get paper here
  • DAVID, PATRICK; GERNOT VOGEL, TRUONG QUANG NGUYEN, NIKOLAI L. ORLOV, OLIVIER S. G. PAUWELS, ALEXANDRE TEYNIÉ, THOMAS ZIEGLER 2021. A revision of the dark-bellied, stream-dwelling snakes of the genus Hebius (Reptilia: Squamata: Natricidae) with the description of a new species from China, Vietnam and Thailand. Zootaxa 4911: 1–61 - get paper here
  • Deuve, J. 1961. Liste annotee des Serpents du Laos. Bull. Soc. Sci. Nat. Laos 1:5-32.
  • Dowling, H.G., & Jenner, J.V. 1988. Snakes of Burma: checklist of reported species and bibliography. Smithsonian Herp. Inf. Serv. (76): 19 pp. - get paper here
  • Günther,A. 1875. Second report on collections of Indian Reptiles obtained bv the British Museum. Proc. Zool. Soc. London 1875: 224-234 - get paper here
  • GUO, PENG; FEI ZHU, QIN LIU, LIANG ZHANG, JIAN X. LI, YU Y. HUANG & R. ALEXANDER PYRON 2014. A taxonomic revision of the Asian keelback snakes, genus Amphiesma (Serpentes: Colubridae: Natricinae), with description of a new species. Zootaxa 3873 (4): 425–440 - get paper here
  • Lalremsanga, H.T.; Saipari Sailo and Chinliansiama 2011. Diversity of Snakes (Reptilia: Squamata) and Role of Environmental Factors in Their Distribution in Mizoram, Northeast India. Advances in Environmental Chemistry - get paper here
  • Lenz, Norbert 2012. Von Schmetterlingen und Donnerdrachen - Natur und Kultur in Bhutan. Karlsruher Naturhefte 4, Naturkundemuseum Karlsruhe, 124 pp.
  • Mahendra B C 1984. Handbook of the snakes of India, Ceylon, Burma, Bangladesh and Pakistan. ANNALS OF ZOOLOGY (AGRA) 22 (B): i-xvi, 1-412
  • Nguyen, S.V., Ho, C.T. and Nguyen, T.Q. 2009. Herpetofauna of Vietnam. Chimaira, Frankfurt, 768 pp.
  • Sharma, R. C. 2004. Handbook Indian Snakes. AKHIL BOOKS, New Delhi, 292 pp.
  • Smith, M.A. 1943. The Fauna of British India, Ceylon and Burma, Including the Whole of the Indo-Chinese Sub-Region. Reptilia and Amphibia. 3 (Serpentes). Taylor and Francis, London. 583 pp.
  • Wallach, Van; Kenneth L. Williams , Jeff Boundy 2014. Snakes of the World: A Catalogue of Living and Extinct Species. [type catalogue] Taylor and Francis, CRC Press, 1237 pp.
  • Wang, Kai; Jinlong Ren, Hongman Chen, Zhitong Lyu, Xianguang Guo Ke Jiang, Jinmin Chen, Jiatang Li, Peng Guo, Yingyong Wang, Jing Che 2020. The updated checklists of amphibians and reptiles of China. Biodiversity Science 28 (2): 189-218 - get paper here
  • Yang, J.-H., X.-Y. Huang, J.-F. Ye, S.-P. Yang, X.-.C. Zhang & B.P.-L. Chan 2019. A report on the herpetofauna of Tengchong Section of Gaoligongshan National Nature Reserve, China. Journal of Threatened Taxa 11(11): 14434–14451 - get paper here
  • Zhao, E.M. 2006. The snakes of China [in Chinese]. Hefei, China, Anhui Sience & Technology Publ. House, Vol. I, 372 pp., Vol. II (color plates), 280 pp.
  • ZHOU, ZHENGYAN; ZHIYONG SUN, SHUO QI, YUYAN LU, ZHITONG LYU, YINGYONG WANG, PIPENG LI, JIANZHANG MA 2019. A new species of the genus Hebius (Squamata: Colubridae: Natricinae) from Hunan Province, China. Zootaxa 4674 (1): 068–082 - get paper here
  • Ziegler, T., and Quyet, L.K. 2006. A new natricine snake of the genus Amphiesma (Squamata: Colubridae: Natricinae) from the central Truong Son, Vietnam. Zootaxa 1225:39-56 - get paper here
 
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