Trimeresurus

Trimeresurus
	Bamboo pit viper, T. gramineus
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Order:	Squamata
Suborder:	Serpentes
Family:	Viperidae
Subfamily:	Crotalinae
Genus:	Trimeresurus; Lacépède, 1804
Synonyms
	Trimeresurus Lacépède, 1804; Craspedocephalus ; Kuhl & van Hasselt, 1822; Trimeresura – Fleming, 1822; Craspedocephalus – Gray, 1825; Megaera Wagler, 1830; Atropos Wagler, 1830; Trimesurus Gray, 1842[1];

Trimeresurus is a genus of venomous pit vipers found in Asia from the Indian Subcontinent throughout Southeast Asia, China and the Pacific Islands. Currently at least 50 species are recognized.[2] Common names include Asian palm pit vipers,[3] Asian lanceheads and Asian lance-headed vipers.[4]

Description

Most species in the genus Trimeresurus are relatively small, primarily arboreal species, with thin bodies and prehensile tails. However, Trimeresurus flavoviridis (the Okinawa habu) can reach a total length (including tail) of 242 cm (7 ft 9 in), and is one of the longest pit vipers in East Asia. Most Trimeresurus species are typically green in color, but some species also have yellow, black, orange, red, or gold markings.

Feeding

The diet of Trimeresurus species includes a variety of animals, including lizards, amphibians, birds, rodents, and other small mammals.

Reproduction

Like most viper species, many of the species in the genus Trimeresurus are ovoviviparous, bearing live young. However, some species such as T. flavoviridis, T. kaulbacki, and T. macrolepis are oviparous, laying eggs. Also, the reproductive biology of some Trimeresurus species is as yet unknown.

Venom

Trimeresurus venom varies in toxicity between species, but all are primarily hemotoxic and considered to be medically significant to humans.

Geographic range

Species in the genus Trimeresurus are found in Southeast Asia from India including regions of North Chotanagpur of Jharkhand to southern China and Japan, and the Malay Archipelago to Timor.[1]

Species

Species[2]	Taxon author[2]	Subsp.*[2]	Common name[5]	Geographic range[1]
T. albolabris	Gray, 1842	0	White-lipped pit viper	India (Assam), Nicobar Islands, Myanmar, Thailand, Cambodia, Laos, Vietnam, southern China (Fukien, Hainan, Kwangsi, Kwangtung), Hong Kong, West Malaysia, Indonesia (Sumatra, Borneo, Sulawesi, Java, Madoera, Lombok, Sumbawa, Komodo, Flores, Sumba, Roti, Timor, Kisar, Wetar).
T. andalasensis	David, Vogel, Vijaykumar & Vidal, 2006	0	Sumatran palm pit viper	Indonesia: Sumatra.
T. andersonii	Theobald, 1868	0	Anderson’s pit viper, Andaman pit viper	Andaman Islands, Nicobar Islands.
T. arunachalensis	Captain, Deepak, Pandit, Bhatt, and Athreya, 2019	0	Arunachal pit viper	India: Arunachal Pradesh
T. borneensis	W. Peters, 1872	0	Bornean pit viper	Indonesia: Borneo.
T. brongersmai	Hoge, 1969	0	Brongersma's pit viper	Indonesia: Simalur Island.
T. cantori	Blyth, 1846	0	Cantor's pit viper	India: Nicobar Islands, and possibly the Andaman Islands.
T. cardamomensis	Malhotra, Thorpe, Mrinalini, & Stuart, 2011	0	Cardamom Mountains green pit viper	Eastern Thailand, Koh Kong Province in Cambodia.
T. caudornatus	Chen, Ding, Vogel, & Shi, 2020	0	Ornamental-tailed pit viper	China (Yunnan).
T. davidi	Chandramouli, Campbell, & Vogel, 2020.[6]	0		Car Nicobar, India.
T. erythrurus	Cantor, 1839	0	Red-tailed bamboo pit viper	India (Assam and Sikkim), Bangladesh and Myanmar.
T. fasciatus	Boulenger, 1896	0	Banded pit viper	Indonesia: Djampea Island.
T. flavomaculatus	Gray, 1842	2	Philippine pit viper	Philippine Islands: Agutayan, Batan, Camiguin, Catanduanes, Dinagat, Jolo, Leyte, Luzon, Mindanao, Mindoro, Negros and Polillo.
T. gracilis	Ōshima, 1920	0	Kikushi habu	Central Taiwan.
T. gramineus^T	Shaw, 1802	0	Bamboo pit viper	Southern India.
T. gumprechti	David, Vogel, Pauwels & Vidal, 2002	0	Gumprecht's green pit viper	China (Yunnan), Laos, Thailand, Vietnam.
T. gunaleni	Vogel, David, & Sidik, 2014	0	Gunalen's pit viper	Sumatra.
T. hageni	Lidth de Jeude, 1886	0	Hagen's pit viper	Peninsular Thailand, West Malaysia, Singapore and Indonesia (Sumatra and the nearby islands of Bangka, Simalur, Nias, Batu and the Mentawai Islands.
T. honsonensis	Grismer, Ngo, & Grismer, 2008	0	Hon Son pit viper	Southern Vietnam.
T. insularis	Kramer, 1977	0	Sunda Island pit viper, White-lipped island pit viper	Indonesia, Timor-Leste
T. kanburiensis	M.A. Smith, 1943	0	Kanburi pit viper	Thailand.
T. labialis	Steindachner, 1867	0	Nicobar bamboo pit viper	India: Nicobar Islands.
T. macrolepis	Beddome, 1862	0	Large-scaled pit viper	The mountains of southern India.
T. macrops	Kramer, 1977	0	Large-eyed pit viper	Thailand, Cambodia and southern Vietnam.
T. malabaricus	Jerdon, 1854	0	Malabar rock pit viper	Southern and western India at 600-2,000 m elevation.
T. malcolmi	Loveridge, 1938	0	Malcolm's pit viper	Borneo.
T. mcgregori	Taylor, 1919	0	McGregor's pit viper, McGregor's tree viper, Philippine pit viper	Batan Island, Philippines.
T. medoensis	Zhao, 1977	0	Motuo bamboo pit viper	Northern India, northern Myanmar and China (southeastern Xizang).
T. mutabilis	Stoliczka, 1870	0	Central Nicobar pit viper, Central Nicobar bamboo pit viper	Central Nicobar Island.
T. nebularis	Vogel, David, & Pauwels, 2004	0	Cameron Highlands pit viper, Clouded pit viper	West Malaysia (Cameron Highlands), Thailand.
T. phuketensis	Sumontha, Kunya, S.G. Pauwels, Nitikul & Punnadee, 2011 [7]	0	Phuket pit viper	Thailand: Phuket Island.
T. popeiorum	M.A. Smith, 1937	2	Popes' pit viper	Northern India, Myanmar, Thailand, West Malaysia, Vietnam and Indonesia (Sumatra, the Mentawai Islands of Siberut, Sipora and North Pagai, and on the island of Borneo).
T. puniceus	Kuhl, 1824	0	Flat-nosed pit viper	Southern Thailand, West and East Malaysia (Sabah and Sarawak) and Indonesia (Borneo, Sumatra, the Mentawai Islands of Siberut and North Pagai, Simalur and Java.
T. purpureomaculatus	Gray, 1832	0	Mangrove pit viper	Bangladesh, Myanmar, Thailand, West Malaysia, Singapore and Indonesia (Sumatra).
T. rubeus	Malhotra, 2011	0	Ruby-eyed green pit viper	Cambodia (Mondulkiri), southern Vietnam.
T. sabahi	Regenass & Kramer, 1981	0	Sabah bamboo pit viper, Sabah pit viper	Sabah, Sarawak
T. salazar	Mirza, Bhosale, Phansalkar, Sawant, Gowande, & Patel (2020)	0	Salazar’s pit viper[8]	India: western lowlands of Arunachal Pradesh
T. schultzei	Griffin, 1909	0	Schultze's pit viper	Philippines: Palawan and Balabac.
T. septentrionalis	Kramer, 1977	0	Nepal pit viper, Himalayan white-lipped pit viper	Bangladesh, India, Nepal
T. sichuanensis	Guo & Wang, 2011	0	Sichuan pit viper	Sichuan, China
T. stejnegeri	Schmidt, 1925	2	Stejneger's pit viper	India (Assam), and Nepal through Myanmar and Thailand to China (Kwangsi, Kwangtung, Hainan, Fukien, Chekiang, Yunnan) and Taiwan.
T. strigatus	Gray, 1842	0	Horseshoe pit viper	The hills of southern India.
T. sumatranus	Raffles, 1822	0	Sumatran pit viper	Southern Thailand, West and East Malaysia (Sabah and Sarawak on Borneo) and Indonesia (Bangka, Billiton, Borneo, Sumatra and the nearby islands of Simalur, Nias, and possibly the Mentawai Islands [Sipora]).
T. tibetanus	Huang, 1982	0	Tibetan bamboo pit viper	China: Xiang (Tibet) Autonomous Region.
T. trigonocephalus	Donndorff, 1798	0	Sri Lankan pit viper	Throughout Sri Lanka from low elevations to about 1,800 m.
T. truongsonensis	Orlov, Ryabov, Thanh, & Cuc, 2004	0	Quang Binh pit viper	Central Vietnam
T. venustus	Vogel, 1991	0	Beautiful pit viper, Brown-spotted pit viper	Southern Thailand
T. vogeli	David, Vidal & Pauwels, 2001	0	Vogel's pit viper	Cambodia, Laos, Thailand, Vietnam.
T. wiroti	Trutnau, 1981	0	Wirot's pit viper	Thailand, West Malaysia.
T. yingjiangensis	Chen, Zhang, Shi, et al., 2019	0		Southwest China
T. yunnanensis	Schmidt, 1925	0	Yunnan bamboo pit viper	Southern China

*) Not including the nominate subspecies.
^T) Type species.[1]

Taxonomy

Additional species that may be recognized by other sources include:[9]

T. barati Regenass & Kramer, 1981. Commonly called Barat's bamboo viper, found in Indonesia.
T. fucatus Vogel, David & Pauwels, 2004. Commonly called the Siamese peninsula pit viper and found in southern Thailand, Myanmar, Malaysia.

The genus Trimeresurus (sensu lato) has been the subject of considerable taxonomic work since 2000, resulting in the recognition of additional genera within this complex. Most authors now recognise the genus Protobothrops for the species cornutus, flavoviridis, jerdonii, kaulbacki, mucrosquamatus, tokarensis, xiangchengensis,[10][11][12] since these have been shown not to be closely related to other Trimeresurus in recent phylogenetic analyses.

In addition, Malhotra and Thorpe (2004)[11] proposed a radical shake up of the entire genus, splitting Trimeresurus into seven genera. Their proposed arrangement (including species described since 2004) is shown in the table below:

Genus	Species included
Trimeresurus	andalasensis, borneensis, brongersmai, gramineus, malabaricus, puniceus, salazar, strigatus, trigonocephalus, wiroti
Cryptelytrops	albolabris, andersonii, cantori, erythrurus, fasciatus, honsonensis (Hon Son Pit Viper), insularis, kanburiensis, labialis, macrops, purpureomaculatus, rubeus, septentrionalis, venustus
Himalayophis	tibetanus
Parias	flavomaculatus, hageni, malcolmi, mcgregori, schultzei, sumatranus
Peltopelor	macrolepis
Popeia	barati, buniana, fucata, nebularis, popeiorum, sabahi
Viridovipera	gumprechti, medoensis, stejnegeri, truongsonensis, vogeli, yunnanensis

This new arrangement has been followed by many,[12][13] but not all[9][14] subsequent authors.

David et al. (2011) considered some of the genera of Malhotra & Thorpe to be subgenera of the genus Trimeresurus, creating new combinations such as "Trimeresurus (Parias) flavomaculatus", "Trimeresurus (Popeia) popeiorum", "Trimeresurus (Viridovipera) stejnegeri", etc.[15]

Gallery

A juvenile T. vogeli in Khao Yai National Park, Thailand
Trimeresurus fucatus from Khao Sok National Park, Thailand
Trimeresurus albolabris from Kaeng Krachan National Park, Thailand
Trimeresurus gumprechti, Gumprecht's green pit viper, from Phu Hin Rong Kla National Park, Thailand
Trimeresurus macrops from Khao Yai National Park, Thailand
Trimeresurus popeiorum from Kaeng Krachan National Park, Thailand

References

McDiarmid RW, Campbell JA, Touré T (1999). Snake Species of the World: A Taxonomic and Geographic Reference, Volume 1. Washington, District of Columbia: Herpetologists' League. 511 pp. ISBN 1-893777-00-6 (series). ISBN 1-893777-01-4 (volume).
"Trimeresurus ". Integrated Taxonomic Information System. Retrieved 27 September 2006.
Mehrtens JM (1987). Living Snakes of the World in Color. New York: Sterling Publishers. 480 pp. ISBN 0-8069-6460-X.
United States Navy (1991). Poisonous Snakes of the World. New York: U.S. Government / Dover Publications Inc. 203 pp. ISBN 0-486-26629-X.
Gumprecht A, Tillack F, Orlov NL, Captain A, Ryabov S (2004). Asian Pitvipers. First Edition. Berlin: GeitjeBooks. 368 pp. ISBN 3-937975-00-4.
Chandramouli, S. R.; Campbell, Patrick D.; Vogel, Gernot (1 November 2020). "A new species of green pit viper of the genus TrimeresurusLacépède, 1804 (Reptilia: Serpentes: Viperidae) from the Nicobar Archipelago, Indian Ocean" (PDF). Amphibian & Reptile Conservation. 14 (3): 169–176. Retrieved 9 November 2020.
Sumontha M, Kunya K, Pauwels OSG, Nitikul A, Punnadee S (2011). "Trimeresurus (Popeia) phuketensis, a New Pitviper (Squamata: Viperidae) from Phuket Island, Southwestern Thailand". Russian Journal of Herpetology 18 (3): 11-17.
Mirza, Zeeshan A; Bhosale, Harshal S; Phansalkar, Pushkar U.; Sawant, Mandar; Gowande, Gaurang G; Patel, Harshil (15 April 2020). "A New Species of Green Pit Vipers of the Genus Trimeresurus Lacépède, 1804 (Reptilia, Serpentes, Viperidae) from western Arunachal Pradesh, India". Zoosystematics and Evolution. 96 (1): 123–138. doi:10.3897/zse.96.48431. Retrieved 24 April 2020.
Trimeresurus at the Reptarium.cz Reptile Database. Accessed 2 February 2017.
Kraus F, Mink DG, Brown WM (1996). "Crotaline intergeneric relationships based on mitochondrial DNA sequence data". Copeia 1996: 763-773.
Malhotra A, Thorpe RS (2004). "A phylogeny of four mitochondrial gene regions suggests a revised taxonomy for Asian pitvipers (Trimeresurus and Ovophis)". Molecular Phylogenetics and Evolution 32: 83-100.
Castoe TA, Parkinson CL (2006). "Bayesian mixed models and the phylogeny of pitvipers (Viperidae: Serpentes)". Mol. Phylogenet. Evol. 39: 91-110.
Grismer LL, Grismer JL, McGuire JA (2006). "A new species of pitviper of the genus Popeia (Squamata: Viperidae) from Pulau Tioman, Pahang, West Malaysia". Zootaxa 1305: 1-19.
Vogel G (2006). Venomous Snakes of Asia / Giftschlangen Asiens. Frankfurt am Main: Terralog, Edition Chimaira.
David, Patrick; Vogel, Gernot; Dubois, Alain (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.