A 'little dragon' from Kunming City: a new green pit viper from Yunnan Province, China (Squamata, Viperidae, Trimeresurus)
Authors/Creators
- 1. Qinghai University, Xining, China
- 2. Anhui Normal University, Wuhu, China
- 3. Duy Tan University, Da Nang, Vietnam
- 4. Institute of Plateau Biology of Xizang Autonomous Region, Lhasa, China
- 5. Lomonosov Moscow State University, Moscow, Russia
- 6. Society for Southeast Asian Herpetology, Heidelberg, Germany
- 7. Ailaoshan Subtropical Forest Ecosystem Observation and Research Station of Yunnan Province, Puer, China|Xi Shuang Ban Na Tropical Botanical Garden, Chinese Academy of Sciences, Yunnan, China
Description
A new species of the genus Trimeresurus Lacépède is described from Kunming City, Yunnan Province, China, based on an integrative analysis of morphological and molecular data. The new species, Trimeresurus loong sp. nov., is assigned to the subgenus Viridovipera Malhotra & Thorpe and can be distinguished from its congeners by the following combination of characters: the first supralabial completely separated from the nasal scale; a short and spinose hemipenis; moderate adult body size; dorsal scales in 19 (21 or 23)-19-15 rows; 150–151 ventral scales in males, and 157–158 in females; 67–68 subcaudals in males, and 57–64 in females; and distinct body color patterns. Molecular analyses based on the mitochondrial 16S, cyt b, and ND4 gene fragments indicate that Trimeresurus loong sp. nov. is genetically divergent from all currently recognized congeners, showing uncorrected p-distances ranging from 6.7–11.2% in cyt b and 6.1–8.3% in ND4. The discovery of this species brings the total number of recognized Trimeresurus species to 57, including nine recorded from Yunnan Province, further emphasizing the complex topography and exceptional level of endemism.
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References
- Akaike H (1973) Information theory and an extension of the maximum likelihood principle. In: Petrov BN, Csáki F (Eds) Second International Symposium on Information Theory, Tsahkadsor, Armenia, USSR. Akadémiai Kiadó, Budapest, 267–281.
- Akaike H (1974) A new look at the statistical model identification. IEEE Transactions on Automatic Control 19(6): 716–723. https://doi.org/10.1109/TAC.1974.1100705
- Ao JM, David P, Bordoloi S, Ohler A (2004) Notes on a collection of snakes from Nagaland, northeast India, with 19 new records for this state. Russian Journal of Herpetology 11(2): 155–162.
- Bu CF, Zheng XC, Zhao XT, Xu TY, Bai X, Jia YK, Chen ML, Hao LL, Xiao JF, Zhang Z, Zhao WM, Tang BX, Bao YM (2024) GenBase: A Nucleotide Sequence Database. Genomics, Proteomics & Bioinformatics 22(3): qzae047. https://doi.org/10.1093/gpbjnl/qzae047
- Burbrink FT, Lawson R, Slowinski JB (2000) Mitochondrial DNA phylogeography of the polytypic North American rat snake (Elaphe obsoleta): A critique of the subspecies concept. Evolution; International Journal of Organic Evolution 54(6): 2107–2118. https://doi.org/10.1554/0014-3820(2000)054[2107:MDPOTP]2.0.CO;2
- Burland TG (2000) DNASTAR's Lasergene sequence analysis software. Methods in Molecular Biology (Clifton, N.J. ) 132: 71–91. https://doi.org/10.1385/1-59259-192-2:71
- Captain A, Deepak V, Pandit R, Bhatt B, Athreya R (2019) A new species of pitviper (Serpentes: Viperidae: Trimeresurus Lacépède, 1804) from West Kameng District, Arunachal Pradesh, India. Russian Journal of Herpetology 26: 111–122. https://doi.org/10.30906/1026-2296-2019-26-2-111-122
- Che J, Jiang K, Yan F, Zhang YP (2020) Amphibians and Reptiles in Tibet: Diversity and Evolution. Science Press, Beijing, China, 800 pp. [in Chinese]
- Chen ZN, Zhang L, Shi JS, Tang YZ, Guo YH, Song ZB, Ding L (2019) A new species of the genus Trimeresurus from Southwest China (Squamata: Viperidae). Asian Herpetological Research 10: 13–23. https://doi.org/10.16373/j.cnki.ahr.180062
- Chen ZN, Yu JP, Vogel G, Shi SC, Song ZB, Tang YZ, Yang J, Ding L, Chen CS (2020) A new pit viper of the genus Trimeresurus (Lacépède, 1804) (Squamata: Viperidae) from Southwest China. Zootaxa 4768: 112–128. https://doi.org/10.11646/zootaxa.4768.1.7
- CNCB-NGDC Members and Partners (2025) Database Resources of the National Genomics Data Center, China National Center for Bioinformation in 2025. Nucleic Acids Research 53(D1): D30–D44. https://doi.org/10.1093/nar/gkae978
- Creer S, Malhotra A, Thorpe RS (2003) Assessing the phylogenetic utility of four mitochondrial genes and a nuclear intron in the Asian pit viper genus Trimeresurus: Separate, simultaneous, and conditional data combination analyses. Molecular Biology and Evolution 20: 1240–1251. https://doi.org/10.1093/molbev/msg136
- Darko YA, Voss O, Uetz P (2022) A dictionary of abbreviations used in reptile descriptions. Zootaxa 5219(5): 421–432. https://doi.org/10.11646/zootaxa.5219.5.2
- David P, Mathew R (2005) Notes on some noteworthy snake specimens deposited in the collections of Eastern Regional Station of the Zoological Survey of India. Records of the Zoological Survey of India, Occasional Paper 104: 83–90. https://doi.org/10.26515/rzsi/v104/i3-4/2005/159303
- David P, Captain A, Bhatt BB (2001a) On the occurrence of Trimeresurus medoensis Djao in Djao & Jiang, 1977 (Serpentes: Viperidae: Crotalinae) in India, with a redescription of this species and notes on its biology. Hamadryad 26(2): 222–238.
- David P, Vidal N, Pauwels OSG (2001b) A morphological study of Stejneger's pitviper Trimeresurus stejnegeri (Serpentes: Viperidae: Crotalinae) with the description of a new species from Thailand. Russian Journal of Herpetology 8(3): 205–222.
- David P, Vogel G, Pauwels OSG, Vidal N (2002) Description of a new species of the genus Trimeresurus from Thailand, related to Trimeresurus stejnegeri Schmidt, 1925 (Serpentes: Crotalidae). The Natural History Journal of Chulalongkorn University 2: 5–19. https://doi.org/10.58837/tnh.2.1.102858
- David P, Vogel G, Dubois A (2011) On the need to follow rigorously the Rules of the Code for the subsequent designation of a nucleospecies (type species) for a nominal genus which lacked one: the case of the nominal genus Trimeresurus Lacépède, 1804 (Reptilia: Squamata: Viperidae). Zootaxa 2992: 1–51. https://doi.org/10.11646/zootaxa.2992.1.1
- David P, Teynié A, Vogel G (2023) The Snakes of Laos. Edition Chimaira, Frankfurt am Main, Germany, 960 pp.
- Dawson K, Malhotra A, Thorpe RS, Guo P, Mrinalini , Ziegler T (2008) Mitochondrial DNA analysis reveals a new member of the Asian pitviper genus Viridovipera (Serpentes: Viperidae: Crotalinae). Molecular Phylogenetics and Evolution 49: 356–361. https://doi.org/10.1016/j.ympev.2008.05.044
- Dowling HG (1951) A proposed standard system of counting ventrals in snakes. British Journal of Herpetology 1: 97–99.
- Elangbam PS, Biakzuala L, Shinde P, Decemson HT, Vabeiryureilai M, Lalremsanga HT (2023) Addition of four new records of pit vipers (Squamata: Crotalinae) to Manipur, India. Journal of Threatened Taxa 15(6): 23315–23326. https://doi.org/10.11609/jott.8354.15.6.23315-23326
- Gumprecht A, Tillack F, Orlov NL, Captain A, Ryabov S (2004) Asian Pit Vipers. Geitje Books, Berlin, 368 pp.
- Guo P, Wang YZ (2011) A new genus and species of cryptic Asian green pitviper (Serpentes: Viperidae: Crotalinae) from southwest China. Zootaxa 2918(1): 1–14. https://doi.org/10.11646/zootaxa.2918.1.1
- Guo P, Malhotra A, Thorpe RS, Creer S, Pook CE (2009) Comments on the systematic status of specimens belonging to the genus Viridovipera (Serpentes: Viperidae: Crotalinae) from Sichuan and Yunnan provinces of southwestern China, with a redescription of V. yunnanensis. The Herpetological Journal 19: 151–162.
- Guo P, Liu Q, Zhong G, Zhu F, Yan F, Tang T, Xiao R, Fang M, Wang P, Fu X (2015) Cryptic diversity of green pitvipers in Yunnan, South-west China (Squamata: Viperidae). Amphibia-Reptilia 36: 265–276. https://doi.org/10.1163/15685381-00003004
- Guo P, Liu Q, Zhu F, Zhong GH, Chen X, Myers EA, Che J, Zhang L, Ziegler T, Nguyen TQ, Burbrink FT (2016) Complex longitudinal diversification across South China and Vietnam in Stejneger's pit viper, Viridovipera stejnegeri (Schmidt, 1925) (Reptilia: Serpentes: Viperidae). Molecular Ecology 25: 2920–2936. https://doi.org/10.1111/mec.13658
- Hoang DT, Chernomor O, Haeseler AV, Minh BQ, Vinh LS (2018) UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35(2): 518–522. https://doi.org/10.1093/molbev/msx281
- Huelsenbeck JP, Ronquist F, Nielsen R, Bollback JP (2001) Bayesian inference of phylogeny and its impact on evolutionary biology. Science 294: 2310–2314. https://doi.org/10.1126/science.1065889
- Idiiatullina SS, Pawangkhanant P, Tawan T, Worranuch T, Dechochai B, Suwannapoom C, Nguyen TV, Chanhome L, Poyarkov NA (2023) Limestone jewel: A new colourful karst-dwelling pitviper (Serpentes: Viperidae: Trimeresurus) from the poorly explored borderlands of southern Peninsular Thailand. Vertebrate Zoology 73: 697–716. https://doi.org/10.3897/vz.73.e109854
- Idiiatullina SS, Nguyen TV, Bragin AM, Pawangkhanant P, Le DX Vogel G, David P, Poyarkov NA (2024a) A new species of green pitviper of the Trimeresurus macrops complex (Reptilia: Serpentes: Viperidae) from South Central Coast Region of Vietnam. Zootaxa 5474(4): 375–411. https://doi.org/10.11646/zootaxa.5474.4.2
- Idiiatullina SS, Nguyen TV, Pawangkhanant P, Suwannapoom C, Chanhome L, Mirza ZA, David P, Vogel G, Poyarkov NA (2024b) An integrative taxonomic revision of the Trimeresurus popeiorum species complex (Reptilia: Serpentes: Viperidae), with descriptions of two new species from the Indo-Burma biodiversity hotspot. Vertebrate Zoology 74: 303–342. https://doi.org/10.3897/vz.74.e113347
- Idiiatullina SS, Pawangkhanant P, Suwannapoom C, Tawan T, Chanhome L, Nguyen TV, David P, Vogel G, Poyarkov NA (2024c) Another new species of karst-associated pitviper (Serpentes: Viperidae: Trimeresurus) from the Isthmus of Kra, Peninsular Thailand. European Journal of Taxonomy 930: 20–52. https://doi.org/10.5852/ejt.2024.930.2489
- Jiang K (2010) A method for evaginating the hemipenis of preserved snakes. Sichuan Journal of Zoology 29: 122–123. [in Chinese]
- Kalyaanamoorthy S, Minh BQ, Wong TKF, Haeseler AV, Jermiin LS (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14(6): 587–589. https://doi.org/10.1038/nmeth.4285
- Kumar S, Stecher G, Li M, Knyaz C, Tamura K (2018) MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms. Molecular Biology and Evolution 35: 1547–1549. https://doi.org/10.1093/molbev/msy096
- Li JN, Liang D, Wang YY, Guo P, Huang S, Zhang P (2020) A large-scale systematic framework of Chinese snakes based on a unified multilocus marker system. Molecular Phylogenetics and Evolution 148: 106807. https://doi.org/10.1016/j.ympev.2020.106807
- Liang YT, Ding L, Vogel G, Chen ZN, Wu ZJ (2025) A new species of the genus Trimeresurus Lacépède, 1804 (Reptilia: Squamata: Viperidae) from southwestern China. Zoosystematics and Evolution 101(4): 1653–1668. https://doi.org/10.3897/zse.101.162424
- Maki M (1931) A monograph of the snakes in Japan. Dai Ichi Shobo, Tokyo, Japan, pp. 1–7 + 1–240, Errata; (1), (1), 1–3 + 1–57, Pl. 1–39. [In English/Japanese] [The separate Japanese text volume and Pl. 40-86 appeared in 1933]
- Malhotra A, Thorpe RS (2000) A phylogeny of the Trimeresurus group of pit vipers: New evidence from a mitochondrial gene tree. Molecular Phylogenetics and Evolution 16: 199–211. https://doi.org/10.1006/mpev.2000.0779
- Malhotra A, Thorpe RS (2004a) A phylogeny of four mitochondrial gene regions suggests a revised taxonomy for Asian pitvipers (Trimeresurus and Ovophis). Molecular Phylogenetics and Evolution 32: 83–100. https://doi.org/10.1016/j.ympev.2004.02.008
- Malhotra A, Thorpe RS (2004b) Maximizing information in systematic revisions: A combined molecular and morphological analysis of a cryptic green pitviper complex (Trimeresurus stejnegeri). Biological Journal of the Linnean Society. Linnean Society of London 82: 219–235. https://doi.org/10.1111/j.1095-8312.2004.00354.x
- Mirza ZA, Lalremsanga HT, Bhosale H, Gowande G, Patel H, Idiiatullina SS, Poyarkov NA (2023) Systematics of Trimeresurus popeiorum Smith, 1937 with a revised molecular phylogeny of Asian pitvipers of the genus Trimeresurus Lacépède, 1804 sensu lato. Evolutionary Systematics 7(1): 91–104. https://doi.org/10.3897/evolsyst.7.97026
- Myers CW, Cadle JE (2003) On the snake hemipenis, with notes on Psomophis and techniques of eversion: A response to Dowling. Herpetological Review 34: 295–302.
- Nguyen LT, Schmidt HA, Haeseler AV, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution 32(1): 268–274. https://doi.org/10.1093/molbev/msu300
- Nguyen TQ, Nguyen TV, Pham CT, Ong AV, Ziegler T (2018) New records of snakes (Squamata: Serpentes) from Hoa Binh Province, northwestern Vietnam. Bonner Zoologische Beiträge 67(1): 15–24.
- Nguyen HN, Tran BV, Nguyen LH, Neang T, Yushchenko PV, Poyarkov NA (2020) A new species of Oligodon Fitzinger, 1826 from the Langbian Plateau, southern Vietnam, with additional information on Oligodon annamensis Leviton, 1953 (Squamata: Colubridae). PeerJ 8: e8332. https://doi.org/10.7717/peerj.8332
- Nguyen TV, Idiiatullina SS, Oo WP, Lee JL, Poyarkov NA, David P, Vogel G (2024) Range extension and expanded description of Trimeresurus caudornatus Chen, Ding, Vogel & Shi, 2020 (Serpentes: Viperidae), with the first country record from Myanmar. Zootaxa 5514(3): 201–231. https://doi.org/10.11646/zootaxa.5514.3.1
- Nguyen TV, David P, Ananjeva NB, Poyarkov NA, Vogel G (2025) A range extension of Trimeresurus mayaae Rathee et al., 2022 (Serpentes: Viperidae), with remarks on its pholidosis, natural history, and conservation status. Russian Journal of Herpetology (in press).
- Orlov NL, Ryabov SA, Bui TN, Ho CT (2004) A new species of Trimeresurus (Ophidia: Viperidae: Crotalinae) from the karst region in central Vietnam. Russian Journal of Herpetology 11(2): 139–149.
- Pawangkhanant P, Idiiatullina SS, Nguyen TV, Ruangsuwan T, Matsukoji T, David P, Suwannapoom C, Poyarkov NA (2025) A snake can change its finery: A new cryptic species of the Trimeresurus kanburiensis complex (Reptilia: Serpentes: Viperidae) from Central Thailand with an unusual ontogenetic color change. Zootaxa 5621(5): 514–546. https://doi.org/10.11646/zootaxa.5621.5.2
- Pope CH (1935) The reptiles of China. Turtles, crocodilians, snakes, lizards. Natural History of Central Asia, Vol. 10. American Museum of Natural History, New York, USA, 604 pp. https://doi.org/10.5962/bhl.title.12104
- Poyarkov NA, Nguyen TV, Popov ES, Geissler P, Pawangkhanant P, Neang T, Suwannapoom C, Ananjeva NB, Orlov NL (2023) Recent progress in taxonomic studies, biogeographic analysis, and revised checklist of reptiles in Indochina. Russian Journal of Herpetology 30(5): 255–476. https://doi.org/10.30906/1026-2296-2023-30-5-255-476
- Rambaut A (2018) FigTree v1.4.4. http://tree.bio.ed.ac.uk/software/figtree/ [accessed 10 April 2025]
- Rathee YS, Purkayastha J, Lalremsanga HT, Dalal S, Biakzuala L, Muansanga L, Mirza ZA (2022) A new cryptic species of green pitviper of the genus Trimeresurus Lacépède, 1804 (Serpentes: Viperidae) from northeast India. PLoS One 17: e0268402. https://doi.org/10.1371/journal.pone.0268402
- Ren JL, Jiang K, Huang JJ, David P, Li JT (2022) Taxonomic review of the genus Herpetoreas (Serpentes: Natricidae), with the description of a new species from Tibet, China. Diversity 14(2): 79. https://doi.org/10.3390/d14020079
- Ren JL, Huang JJ, Wu W, Jiang K, Li JT (2025) One mountain, two tigers: A new species of Gloydius (Serpentes: Viperidae) from the Upper Lancang Valley in Xizang, China, with comments on the diagnostic characters and evolution of the G. strauchi (Bedriaga, 1912) species complex. Asian Herpetological Research 16(1): 1–19. https://doi.org/10.3724/ahr.2095-0357.2024.0059
- Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542. https://doi.org/10.1093/sysbio/sys029
- Salvi D, Harris DJ, Kaliontzopoulou A, Carretero MA, Pinho C (2013) Persistence across Pleistocene ice ages in Mediterranean and extra Mediterranean refugia: Phylogeographic insights from the common wall lizard. BMC Evolutionary Biology 13: 147. https://doi.org/10.1186/1471-2148-13-147
- Sanders KL, Malhotra A, Thorpe RS (2006) Combining molecular, morphological and ecological data to infer species boundaries in a cryptic tropical pitviper. Biological Journal of the Linnean Society. Linnean Society of London 87: 343–364. https://doi.org/10.1111/j.1095-8312.2006.00568.x
- Schmidt KP (1925) New reptiles and a new salamander from China. American Museum Novitates (157): 1–5.
- Schmidt KP (1927) Notes on Chinese reptiles. Bulletin of the American Museum of Natural History 54(6): 467–551.
- Smith MA (1943) The Fauna of British India, Ceylon and Burma, Including the Whole of the Indo-Chinese Sub-Region. Reptilia and Amphibia. Volume 3: Serpentes. Taylor and Francis, London, 583 pp.
- Stephane G, Jean-franc OD, Vincent L, Maria A, Wim H, And OG (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: Assessing the performance of PhyML 3.0. Systematic Biology 59(3): 307–321. https://doi.org/10.1093/sysbio/syq010
- Sumontha M, Suntrarachun S, Pauwels OSG, Pawangkhanant P, Chomngam N, Iamwiriyakul P, Chanhome L (2021) A new karst-dwelling, colorful pitviper (Viperidae: Trimeresurus) from northern Peninsular Thailand. Zootaxa 4974(2): 307–332. https://doi.org/10.11646/zootaxa.4974.2.4
- Teynié A, David P (2014) Amphibiens et reptiles des formations karstiques du Laos. Bulletin de la Société Herpétologique de France 148: 451–489. [Amphibians and reptiles of the karst formations of Laos] [In French]
- Uetz P, Freed P, Aguilar R, Reyes F, Hošek J (2025) The Reptile Database. http://www.reptile-database.org [accessed 05 October 2025]
- Vogel G, Nguyen TV, David P (2023) A new green pitviper of the Trimeresurus albolabris complex (Reptilia: Serpentes: Viperidae) from central and southern Myanmar. Zootaxa 5357(4): 515–554. https://doi.org/10.11646/zootaxa.5357.4.3
- Wu YH, Hou SB, Yuan ZY, Jiang K, Huang RY, Wang K, Liu Q, Yu ZB, Zhao HP, Zhang BL, Chen JM, Wang LJ, Stuart BL, Chambers EA, Wang YF, Gao W, Zou DH, Yan F, Zhao GG, Fu ZX, Wang SN, Jiang M, Zhang L, Ren JL, Wu YY, Zhang LY, Yang DC, Jin JQ, Yin TT, Li JT, Zhao WG, Murphy RW, Huang S, Guo P, Zhang YP, Che J (2023) DNA barcoding of Chinese snakes reveals hidden diversity and conservation needs. Molecular Ecology Resources 23: 1124–1141. https://doi.org/10.1111/1755-0998.13784
- Xu YH, Nguyen TV, Wang ZQ, Zhang TR, Poyarkov NA, Wei C, Vogel G, Li JC, Deng JD, Sun FY, Peng LF, Weng SY (2025) A new green pitviper of the genus Trimeresurus Lacépède, 1804 (Squamata: Viperidae) from Xizang, China. Animals (Basel) 15: 2675. https://doi.org/10.3390/ani15182675
- Zhang FJ, Hu SQ, Zhao EM (1984) Comparative studies and phylogenetic discussions on hemipenial morphology of the Chinese Colubrinae (Colubridae). Acta Herpetologica Sinica 3: 23–44. [in Chinese]
- Zhang D, Gao FL, Jakovlić I, Zou H, Zhang J, Li WX, Wang GT (2020) PhyloSuite: An integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Molecular Ecology Resources 20(1): 348–355. https://doi.org/10.1111/1755-0998.13096
- Zhao EM (2006) Snakes of China. Volume I. Anhui Science and Technology Publishing House, Hefei, China, 372 pp. [in Chinese]
- Zhao EM, Huang MH, Zong Y (1998) Fauna Sinica: Reptilia. Volume 3. Squamata, Serpentes. Science Press, Beijing, China, 522 pp. [in Chinese]