Mangshan Pit Viper (Protobothrops mangshanensis) Fact Sheet: Bibliography & Resources

Alencar LRV et al. 2016. Diversification in vipers: Phylogenetic relationships, time of divergence and shifts in speciation rates. Molecular Phylogenetics and Evolution. 2016;105:50–62. https://doi.org/10.1016/j.ympev.2016.07.029

Baldwin B. 2025. Management and reproduction of Mangshan pitvipers at the San Diego Zoo. Oral presentation, Biology of Pitvipers 5 (research conference). Rodeo, New Mexico, USA. 24 Jul 2025.

Bogert CM. 1947. Rectilinear locomotion in snakes. Copeia 1947(4):253-254. doi:10.2307/1438921.

Boyer D. 2008. Mangshan pitviper: husbandry and conservation initiative. Snake TAG mid-year group meeting. Oklahoma City, 24 March 2008. Accessed 2011 May 19. http://www.snaketag.org/Documents/SAGMeetingMinutes2009/SAGminutes2008.pdf (no longer available online).

Boyer D. 2010. Mangshan pitviper: husbandry and conservation initiative. PowerPoint presentation; 35 slides. San Diego (CA): San Diego Zoo Global.

Boyer DA. 2007. Step inside this old house. ZOONOOZ, Vol. LXXX (No. 7); p. 12–16.

Böhme M, Ilg A. 2003: fosFARbase [online database]. [last updated 2022; accessed 02 Jul 2025]. https://www.fosfarbase.org.

Campbell JA, Brodie ED. 1992. Introduction. In: Campbell JA, Brodie ED (eds.). Biology of the pitvipers. Tyler (TX): Selva.

Chen S, Yang D, Yang W, Chen Y. 2013. Artificial incubation and snakelet captive breeding of Mangshan pitviper (Protobothrops mangshanensis). Chinese Journal of Ecology. 2013. https://www.cje.net.cn/EN/abstract/abstract21210.shtml

Chen YH. 1990. A new species of snake in China — Trimeresurus mangshanensis [Translated in: David P, Tong H. 1997. Translations of recent descriptions of Chinese pitvipers of the Trimeresurus complex (Serpentes Viperidae), with a key to the complex in China and adjacent areas. Smithsonian Herpetol. Inf. Serv. 112, 14.]

Chiszar D et al. 1982. Strike-induced chemosensory searching in Old World vipers and New World pit vipers. Animal Learning & Behavior. 1982;10(2):121–125. https://doi.org/10.3758/bf03212258

Chiszar D, Lee RKK, Radcliffe CW, Smith HM. 1992. Searching behaviors by rattlesnakes following predatory strikes. In: Campbell JA, Brodie, Jr. ED, editors. Biology of the pitvipers. Selva. 1992. p 369–382.

Cundall D et al. 2012. Drinking in snakes: resolving a biomechanical puzzle. Journal of Experimental Zoology Part A: Ecological Genetics and Physiology. 2012;317(3):152–172. https://doi.org/10.1002/jez.1710

Daltry JC, Wuster W, Thorpe RS. 1997. The role of ecology in determining venom variation in Calloselasma rhodostoma. In: Thorpe RS, Wuster W, Malhoutra A (eds.). Venomous snakes: ecology, evolution and snakebite. Oxford: Zoological Society of London.

Deng Z, Ding X, et al. 2024. Behavior and activity patterns of the critically endangered Mangshan pit viper (Protobothrops mangshanensis) determined using remote monitoring. Animals. 2024;14(15):2247. https://doi.org/10.3390/ani14152247

Deng Z, Xia X, et al. 2024. Predicting the spatial distribution of the Mangshan pit viper (Protobothrops mangshanensis) under climate change scenarios using MaxEnt modeling. Forests. 2024;15(4):723. https://doi.org/10.3390/f15040723

Duan Yan. 2011. 'Dr Snake' bears fangs for conservation. China Daily News. [updated 17 Feb 2011; accessed 24 Sep 2024]. https://www.chinadaily.com.cn/china/2011-02/17/content_12029422.htm.

Ernest CH, Zug GR. 1996. Snakes in question. Washington D.C.: Smithsonian Press.

Gardner JD, Cifelli RL. 1999. A primitive snake from the Cretaceous of Utah. In: Unwin, D. (ed.). Cretaceous fossil vertebrates. Special Papers in Palaeontology 60. Paleontological Association.

Ghanbari A, Rostami A, Noorani SMRS, Fakhrabadi MMS. 2008. Kinematics and kinetics analysis of rectilinear locomotion gait. In: Marques L, de Almeida A, Tokhi MO (eds.). Advances in mobile robotics: proceedings of the eleventh international conference on climbing and walking robots. Coimbra, Portugal: World Scientific Books.

Gong S et al. 2013. Population status, distribution and conservation needs of the endangered Mangshan pit viper Protobothrops mangshanensis of China. Oryx. 2013;47(01):122–127. https://doi.org/10.1017/S0030605311001037

Goris RC. 2011. Infrared organs of snakes: an integral part of vision. Journal of Herpetology. 2011;45(1):2–14. https://doi.org/10.1670/10-238.1

Gray J. 1946. The mechanism of locomotion in snakes. Journal of Experimental Biology. 1946;23(2):101–120. https://doi.org/10.1242/jeb.23.2.101

Green HW. 1992. The ecological and behavioral context for pitviper evolution. In: Campbell JA, Brodie ED (eds.). Biology of the pitvipers. Tyler (TX): Selva. p. 107-118.

Gumprecht A, Boyer DM, Tepedelen K. 2002. Die Grubenottern der Gattung Trimeresurus (sensu lato) Lacépède. Teil I: Anmerkungen zur Biologie, Haltung und Nachzucht von Ermia mangshanensis (Zhao, 1990) [The Pitvipers of Genus Trimeresurus (sensu lato) Lacépède. Part I: Notes on the biology, husbandry and breeding of Ermia mangshanensis (Zhao, 1990)]. Sauria. 2002;24:3–11.

Gumprecht A, Tillack F. 2004. Proposal for a replacement name of the snake genus Ermia Zhang, 1993. Rus J Herpetol 11: 73–76.

Guo P et al. 2016. Multilocus phylogeny of the Asian lance-headed pitvipers (Squamata, Viperidae, Protobothrops). Zootaxa. 2016 [accessed 2025 Feb 26];4093(3). https://doi.org/10.11646/zootaxa.4093.3.5

Guo P, Li J, Chen Y, Wang Y. 2012. Designation of a neotype for Protobothrops mangshanensis (Zhao, 1990). Asian Herpetological Research. 2012;3(4):340–341. https://doi.org/10.3724/SP.J.1245.2012.00340

Guo P, Malhotra A, Li PP, Creer S, Pook CE. 2007. New evidence on the phylogenetic position of the poorly known Asian pitviper Protobothrops kaulbacki (Serpentes: Viperidae: Crotalinae) with a redescription of the species and a revision of the genus Protobothrops. Herpetol J, 17:237–246.

Hayes WK, Herbert SS, Rehling GC, Gennaro JF. 2002. Factors that influence venom expenditure in viperids and other snake species during predatory and defensive contexts. In: Schuett GW, Hoggren M, Douglas ME, Greene HW (eds.). Biology of the vipers. Eagle Mountain (UT): Eagle Mountain Publishing, LLC. pp. 207-234.

Head JJ, Mahlow K, Müller J. 2016. Fossil calibration dates for molecular phylogenetic analysis of snakes 2: Caenophidia, Colubroidea, Elapoidea, Colubridae. Palaeontologia Electronica 19.2.2FC: 1–21.

Heatwole H, Davison E. 1976. A review of caudal luring in snakes with notes on its occurrence in the Saharan sand viper, <i. Herpetologica, 32(3), 332–336. http://www.jstor.org/stable/3891463.

Hoge AR, Romano-Hoge SARWL. 1983. Notes on micro and ultrastructure of "Oberhäutschen" in Viperoidea. Memórias do Instituto Butantan São Paulo (1980-1981), 44-45: 81–118. https://www.biodiversitylibrary.org/page/55679378.

Home E. 1812. IX. Observations intended to show that the progressive motion of snakes is partly performed by means of the ribs. Philosophical Transactions of the Royal Society of London. 1812;102:163–168. https://doi.org/10.1098/rstl.1812.0011

Ivanov M. 1999. The first European pit viper from the Miocene of Ukraine. Acta Palaeontologica Polonica, 44(3), 327-334. https://bibliotekanauki.pl/articles/20013.pdf.

Ivanov M, Vasilyan D, Böhme M, Zazhigin VS. 2019. Miocene snakes from northeastern Kazakhstan: new data on the evolution of snake assemblages in Siberia. Historical Biology 31(10):1284–1303. https://doi.org/10.1080/08912963.2018.1446086

Ivanov M. 2022. Miocene snakes of Eurasia: a review of the evolution of snake communities. In: Gower DJ, Zaher H, editors. The Origin and Early Evolutionary History of Snakes. 1st ed. Cambridge University Press; 2022. p 85–110. https://doi.org/10.1017/9781108938891.007

Jiang ZG, Jiang JP, Wang YZ, Zhang E, Zhang YY, Li LL, Xie F, Ji B, Cao L, Zheng GM. 2016. Red list of China’s vertebrates. Biodiv Sci 24: 500–551 (in Chinese).

Kuch U, Müller J, Mödden C, Mebs D. 2006. Snake fangs from the Lower Miocene of Germany: evolutionary stability of perfect weapons. Naturwissenschaften. 2006;93(2):84–87. https://doi.org/10.1007/s00114-005-0065-y

Lajoye Q et al. 2024. Paraspecificity of Mexican antivipmyn TRI antivenom in envenomation by Chinese Protobothrops mangshanensis (Mangshan pit viper) in France: A case report and experimental neutralization of venom procoagulant effect. Toxicon. 2024;247:107826. https://doi.org/10.1016/j.toxicon.2024.107826

Lissmann HW 1950. Rectilinear locomotion in a snake (Boa occidentalis). Journal of Experimental Biology 26(4):368. http://jeb.biologists.org/content/26/4/368.full.pdf.

Lynch VJ 2009. Live-birth in vipers (Viperidae) is a key innovation and adaptation to global cooling during the Cenozoic. Evolution 63(9):2457-2465.

Lynch VJ. 2009. Live-birth in vipers (Viperidae) is a key innovation and adaptation to global cooling during the cenozoic. Evolution. 2009;63(9):2457–2465. https://doi.org/10.1111/j.1558-5646.2009.00733.x

Malhotra A, Creer S, Pook CE, Thorpe RS. 2010. Inclusion of nuclear intron sequence data helps to identify the Asian sister group of New World pitvipers. Molecular Phylogenetics and Evolution. 2010;54(1):172–178. https://doi.org/10.1016/j.ympev.2009.09.007

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.

Maritz B et al. 2016. Identifying global priorities for the conservation of vipers. Biological Conservation. 2016;204:94–102. https://doi.org/10.1016/j.biocon.2016.05.004

McDiarmid RW, Campbell JA, Touré T'SA. 1999. Snake species of the world. a taxonomic and geographic reference. Volume 1. Washington D.C.: The Herpetologists' League. pp. 229-352.

Mebs D et al. 2006. Biochemical and biological activities of the venom of the Chinese pitviper Zhaoermia mangshanensis, with the complete amino acid sequence and phylogenetic analysis of a novel Arg49 phospholipase A2 myotoxin. Toxicon. 2006;47(7):797–811. https://doi.org/10.1016/j.toxicon.2006.01.031

Messenger KR, Malhotra A. 2016. East Asia region of the VSG. Vipera: the IUCN Viper Specialist Group Bulletin. 1(Dec 2016):16-19. https://viperconservation.org/wp-content/uploads/2021/01/VSG-newsletter-Vipera-2016b.pdf

NFGA (National Forestry and Grasslands Administration). 2021. The 14th five-year plan for protection and development in the forest and grassland sector of the People's Republic of China (2021-2025). Beijing: National Forestry and Grassland Administration (Ministry of Natural Resources). [accessed 30 Jul 2025] http://www.forestry.gov.cn. https://montreal-process.org/documents/publications/general/2021/China5YrForestGrasslandPlan.pdf.

O'Connell BO, Greenlee R, Bacon J, Chiszar D. 1982. Strike-induced chemosensory searching in Old World vipers and New World pit vipers at San Diego Zoo. Zoo Biology 1:287-294.

O’Shea M. 2023. Protobothrops mangshanensis Mangshan pitviper (Zhao in Zhao and Chen, 1990). In: The book of snakes: a life-size guide to six hundred species from around the world. Second edition. The University of Chicago Press; 2023. p 595.

Olives TD et al. 2016. Complete clinical course of envenomation by Protobothrops mangshanensis: delayed coagulopathy and response to Trimeresurus albolabris antivenom. Clinical Toxicology. 2016;54(3):290–292. https://doi.org/10.3109/15563650.2016.1138227

Orlov NL, Ryabov SA, Nguyen TT. 2009. Two new species of genera Protobothrops Hoge et Romano-Hoge, 1983 and Viridovipera Malhotra et Thorpe, 2004 (Ophidia: Viperidae: Crotalinae) from karst region in northeastern Vietnam. Part I. Description of a new species of Protobothrops genus. Russian Journal of Herpetology, 16(1), 69-82.

Paclík V, Ivanov M. 2022. Squamates from the early Miocene (Aquitanian) of Weißenburg 6 (Bavaria, Germany). Acta Musei Moraviae, Scientiae geologicae 107(1):73-89.

Parham D. 2018 Oct 08. Super sensors. SDZWA Stories (blog). https://sandiegozoowildlifealliance.org/story-hub/zoonooz/super-sensors.

Parkinson DL, Campbell JA, Chippindale PT. 2002. Multigene phylogenetic analysis of pitvipers, with comments on their biogeography. In: Schuett GW, Hoeggren M, Douglas ME, Greene HW (Eds.). Biology of the Vipers. Utah: Eagle Mountain Publishing, LC. pp. 93-111.

Perkins W. 2013. For goodness snakes. ZOONOOZ, Vol. LXXXVI (No. 8; digital only); p. 38–46. https://issuu.com/sdzglobal/docs/zoonooz-august-2013.

Pontes‐Nogueira M, Alencar LRV, Sawaya RJ. 2025. From Asia to the Americas: a new cross‐continental perspective on viper biogeography and habitat dynamics. Journal of Biogeography. 2025 [accessed 2025 Jul 15];52(8). https://doi.org/10.1111/jbi.15153

Pough, FH and Groves JD. 1983. Specializations of the body form and food habits of snakes. American Zoologist 23(2):443-454.

Pough FH, Kwiecinski G, Bemis W. 1978. Melanin deposits associated with the venom glands of snakes. Journal of Morphology 155(1):63-71. doi:10.1002/jmor.1051550105.

Pyron R, Burbrink FT, Wiens JJ. 2013. A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology. 2013;13(1):93. https://doi.org/10.1186/1471-2148-13-93

Pyron RA et al. 2011. The phylogeny of advanced snakes (Colubroidea), with discovery of a new subfamily and comparison of support methods for likelihood trees. Molecular Phylogenetics and Evolution. 2011;58(2):329–342. https://doi.org/10.1016/j.ympev.2010.11.006

Quijada-Mascareñas A, Wüster W. 2010. Recent advances in venomous snake systematics. In: Mackessy SP (ed.). Handbook of venoms and toxins of reptiles. Boca Raton (FL): CRC Press. pp. 25-64.

Rage J-C. 2013. Mesozoic and Cenozoic squamates of Europe. Palaeobiodiversity and Palaeoenvironments. 2013;93(4):517–534. https://doi.org/10.1007/s12549-013-0124-x

Szyndlar Z, Rage JC. 1999. Oldest fossil vipers (Serpentes: Viperidae) from the Old World. Kaupia 8: 9–20.

Szyndlar Z, Rage J-C. 2002. Fossil record of the true vipers. In: Schuett GW, Hoeggren M, Douglas ME, Greene HW (eds.). Biology of the vipers. Utah: Eagle Mountain Publishing, LC. pp. 419-444.

Thomas RG, Pough FH. 1979. The effect of rattlesnake venom on digestion of prey. Toxicon. 1979;17(3):221–228. https://doi.org/10.1016/0041-0101(79)90211-3

Uetz P, Freed P, Aguilar R, Reyes F, Kudera J, Hošek J (editors). 2025. Protobothrops mangshanensis (Zhao, 1990). In: The Reptile Database [Internet database]. [last updated 25 May 2025; accessed 2025 Jul 15]. http://www.reptile-database.org/.

UNEP. 2025. Protobothrops mangshanensis. Species+ [online database]. Nairobi, Kenya: UNEP [compiled by UNEP-WCMC, Cambridge, UK]; [accessed 2025 May 28]. https://www.speciesplus.net/species#/taxon_concepts/101403/legal.

UNEP. 2019. Species+: Trimeresurus mangshanensis [online database]. Nairobi, Kenya: UNEP [compiled by UNEP-WCMC, Cambridge, UK]. [accessed 2019 Oct 09]. https://www.speciesplus.net/species#/taxon_concepts/7472/legal.

Valenta J, Stach Z, Michalek P. 2014. Exotic snake bites in the Czech Republic—epidemiological and clinical aspects during 15-year period (1999–2013). Clinical Toxicology. 2014;52(4):258–264. https://doi.org/10.3109/15563650.2014.902066

Valenta J, Stach Z, Otahal M. 2012. Protobothrops mangshanensis bite: first clinical report of envenoming and its treatment. Biomedical Papers. 2012;156(2):183–185. https://doi.org/10.5507/bp.2012.021

Vitt LJ, Caldwell JP. 2014. Herpetology: an introductory biology of amphibians and reptiles. Fourth edition. Elsevier, AP, Academic Press is an imprint of Elsevier; 2014.

Wang S, Xie Y (editors). 2004. China Species Red List. Vol. 1. Higher Education Press, Beijing, China.

Wang YZ. 2021. China’s red list of biodiversity: Vertebrates, Vol. III, Reptiles (I). Beijing: Science Press (in Chinese).

Weinstein SA, Smith TL, Kardong KV. 2010. Reptile venom glands: form, function, and future. In: Mackessy SP, editor. Handbook of venoms and toxins of reptiles. Taylor & Francis; 2010. p 65–91.

Wilson LM et al. 2024. Prolonged venom-induced consumptive coagulopathy following Mangshan pit viper (Protobothrops mangshanensis) envenomation despite Hemato Polyvalent antivenom administration. Toxicon. 2024;238:107563. https://doi.org/10.1016/j.toxicon.2023.107563

Wüster W, Peppin L, Pook CE, Walker DE. 2008. A nesting of vipers: phylogeny and historical biogeography of the Viperidae (Squamata: Serpentes). Molecular Phylogenetics and Evolution. 2008;49(2):445–459. https://doi.org/10.1016/j.ympev.2008.08.019

Wuster W, Salomao MG, Quijada-Mascarenas, JA, Thorpe RS. 2002. Origins and evolution of the South American pitviper fauna. In: Schuett GW, Hoeggren M, Douglas ME, Greene HW (eds.). Biology of the vipers. BBBSP (Bitamtam-British Bothrops Systematics Project). Utah: Eagle Mountain Publishing, LC. pp. 111-128.

Yang D, Chen S, Chen Y, Yan Y. 2013. Using head patch pattern as a reliable biometric character for noninvasive individual recognition of an endangered pit viper Protobothrops mangshanensis. Asian Herpetological Research. 2013;4(2):134–139. https://doi.org/10.3724/SP.J.1245.2013.00134

Zhang B et al. 2020. Habitat association in the critically endangered Mangshan pit viper (Protobothrops mangshanensis), a species endemic to China. PeerJ. 2020;8:e9439. https://doi.org/10.7717/peerj.9439

Zhang B et al. 2023. Habitat choice for narrowly distributed species: multiple spatial scales of habitat selection for the Mangshan pit viper. Global Ecology and Conservation. 2023;45:e02512. https://doi.org/10.1016/j.gecco.2023.e02512

Zhang FJ. 1993. Division of the genus Trimeresurus (Serpentes: Viperidae), based on the morphology of their skulls. In Zhao EM, Chen BH, Papenfuss T (eds.). Proceedings of the First Asian Herpetological Meeting. Beijing: China Forestry Press; p. 48–57.

Zhang FJ. 1993. Division of the genus Trimeresurus (sensu lato) (Serpentes: Viperidae), based on the morphology of their skulls. In: Zhao EM, Chen BH, Papenfuss TJ (eds.). Proceedings of the First Asian Herpetology Meeting, Huangshan. Beijing, China: Forestry Press. Herpetological Series 4:48–57.

Zheng Y, Wiens JJ. 2016. Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species. Molecular Phylogenetics and Evolution. 2016;94:537–547. https://doi.org/10.1016/j.ympev.2015.10.009

Zhao EM. 1998. China red data book of endangered animals: Amphibia and reptilia. Science Press, Beijing, China.

Zhao EM, Adler K. 1993. Herpetology of China. Society for the Study of Amphibians and Reptiles in cooperation with the Chinese Society for the Study of Amphibians and Reptiles.

Zhao EM and Chen YH. 1990. Description of a new species of the genus Trimeresurus. [Translated in: David P, Tong H. 1997. Translations of recent descriptions of Chinese pitvipers of the Trimeresurus complex (Serpentes Viperidae), with a key to the complex in China and adjacent areas. Smithsonian Herpetol. Inf. Serv. 112, 12–13, 21–22.]

Zhou Z. 2022. Protobothrops mangshanensis (amended version of 2012 assessment). The IUCN red list of threatened species. Version 2024-1; e.T192140A217768687. doi:10.2305/IUCN.UK.2016-3.RLTS.T22679836A92831716.en.

Zug GR. 1993. Herpetology: an introductory biology of amphibians and reptiles. San Diego (CA): Academic Press.

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