Trimeresurus
Trimeresurus is a genus of venomous pit vipers commonly known as Asian palm pit vipers or green pit vipers, belonging to the subfamily Crotalinae within the family Viperidae, characterized by their heat-sensing loreal pits and primarily arboreal lifestyle.[1] Comprising 57 species as of November 2025, these snakes are predominantly green or brownish in coloration, aiding camouflage in forested environments, and are equipped with hemotoxic venom that can cause significant medical issues in human populations.[2] The genus was established by Lacépède in 1804, with its type species being Trimeresurus gramineus.[3] These pit vipers are distributed across eastern and southeastern Asia, ranging from the Indian subcontinent (including the Western Ghats and northeastern regions) through southern China, Myanmar, Thailand, Laos, Vietnam, Cambodia, Malaysia, Indonesia, and extending to some Pacific islands.[4] Habitats typically include tropical rainforests, subtropical forests, karst landscapes, and mountainous areas up to elevations of 2,000 meters, where they often perch on vegetation or low branches during the day and become active at night.[5] Many species exhibit sexual dimorphism, with males possessing longer tails and brighter coloration, and their diet consists mainly of small mammals, birds, frogs, and lizards.[1] Trimeresurus species are medically significant, accounting for a substantial proportion of snakebites in Southeast Asia due to their proximity to human settlements and nocturnal habits.[6] Ongoing taxonomic research, driven by molecular phylogenetics, continues to reveal cryptic diversity and refine species boundaries, with several new species described in recent years from biodiversity hotspots like the Indo-Burma region.[7] Conservation concerns vary by species, with some facing threats from habitat loss and collection for the pet trade or traditional medicine.[4]Classification
Taxonomy
The genus Trimeresurus was established by Bernard-Germain-Étienne de La Ville-sur-Îllon, comte de Lacépède, in 1804, with the name derived from the Greek words trimerēs (meaning "three parts," referring to the three enlarged scales on the snout) and ourá (meaning "tail").[8] The type species is Trimeresurus gramineus (formerly placed in Crotalus), reflecting early classifications where Asian pit vipers were initially grouped under the New World genus Crotalus by earlier authors like Wagler in 1830.[9] Historically, the genus underwent significant revisions in the late 19th and 20th centuries. Species were separated from Crotalus due to distinct morphological features, such as the presence of heat-sensing pit organs and arboreal adaptations, with key contributions from Albert Günther's 1889 catalogue of snakes in the British Museum, which described several Trimeresurus taxa and clarified diagnostic traits like scale patterns and hemipenial morphology.[10] Further refinements occurred through works like Maslin's 1942 key to the genus, emphasizing separation from Bothrops and establishing Trimeresurus as a distinct Asian lineage within Viperidae.[11] Currently, Trimeresurus is recognized as a genus in the subfamily Crotalinae of the family Viperidae, encompassing 57 valid species as of November 2025, primarily arboreal green pit vipers distributed across Asia.[12] Taxonomic debates persist regarding its monophyly, with molecular phylogenetics from 2016–2023 proposing splits into up to seven genera based on clades defined by mitochondrial and nuclear DNA; for instance, larger terrestrial forms are often placed in Protobothrops (e.g., P. jerdonii), while some arboreal species align with Cryptelytrops (e.g., C. albolabris complex), and the T. albolabris group forms a distinct subclade supported by cytochrome b and 16S rRNA analyses.[13][14] These revisions highlight the genus's morphological heterogeneity, including variations in supralabial counts and dorsal scale rows.[15] Phylogenetically, Trimeresurus occupies a basal position among Asian pit vipers, forming a sister clade to New World crotalines alongside genera like Gloydius, Ovophis, and Protobothrops, as evidenced by multilocus analyses of mtDNA (e.g., ND4, cyt b) and nuclear genes (e.g., CMOS, PRLR).[13] Molecular dating indicates diversification began in the Miocene (approximately 23–5 million years ago), driven by tectonic uplift in Southeast Asia and habitat fragmentation, with Trimeresurus radiating into humid forest niches. Recent updates include 2023 revisions to the T. popeiorum complex, where integrative taxonomy using morphology, hemipenial structure, and mtDNA split it into multiple species, such as restricting T. popeiorum to central Vietnam and elevating T. caudornatus (originally described in 2020) as valid based on genetic divergence exceeding 5% in COI barcodes.[16] Additionally, 2024 studies proposed two new cryptic species from the Indo-Burma hotspot within this group, using ecological niche modeling and phylogenomics to resolve boundaries, while T. uetzi was described in 2023 from Nepal, emphasizing ongoing refinements via total-evidence approaches.[1][17]Species
The genus Trimeresurus currently comprises 57 recognized species, reflecting ongoing taxonomic revisions based on morphological, molecular, and phylogenetic analyses.[18] This number has increased significantly in recent years due to discoveries in biodiversity hotspots like the Himalayas, Western Ghats, and Southeast Asian islands, with at least 13 new species described since 2010. The type species is Trimeresurus gramineus Wagler, 1827, originally described from India.[15] Many species exhibit cryptic diversity, leading to splits from complexes like the T. albolabris and T. popeiorum groups. Note that some species listed in older classifications have been transferred to other genera such as Protobothrops based on recent phylogenetic studies. Below is a complete list of recognized species, including authors, years of description, type localities, and brief notes on synonyms or conservation status where applicable (IUCN statuses are noted for select species; many remain unassessed). This compilation draws from recent taxonomic syntheses and primary descriptions, emphasizing post-2010 additions to highlight dynamism. Species confirmed to be in other genera (e.g., Protobothrops) have been excluded.| Species | Author(s) and Year | Type Locality | Notes |
|---|---|---|---|
| T. albolabris | Gray, 1842 | Java, Indonesia | White-lipped pit viper; formerly included L. monticola as synonym; Least Concern (IUCN).[15] |
| T. andersonii | Boulenger, 1892 | Andaman Islands, India | Synonym: T. andamanensis; Vulnerable (IUCN). |
| T. arunachalensis | Captain et al., 2019 | Arunachal Pradesh, India | Recently described from eastern Himalayas; molecularly distinct from T. albolabris complex. |
| T. erythrurus | Boulenger, 1896 | Myanmar | Red-tailed pit viper; stable taxonomy. |
| T. flavoviridis | Hoge & Romano-Hoge, 1981 | Ryukyu Islands, Japan | Habu; includes subspecies like T. f. flavoviridis; Least Concern. |
| T. gracilis | Gray, 1842 | Malaysia | Slender green pit viper. |
| T. gramineus | Wagler, 1827 | India | Bamboo pit viper; type species; type locality Tamil Nadu. |
| T. hageni | Lidth de Jeude, 1888 | Sumatra, Indonesia | Hagen's pit viper. |
| T. insularis | Kramer, 1977 | Singapore and islands | Least Concern (IUCN). |
| T. kanburiensis | Smith, 1949 | Kanchanaburi, Thailand | Part of T. kanburiensis complex; recent splits include T. kuiburi (2011). |
| T. karanshola | Ganesh et al., 2020 | Western Ghats, India | Described from Karnataka; cryptic species in T. gramineus group. |
| T. macrolepis | Günther, 1889 | Nicobar Islands, India | Large-scaled pit viper; Data Deficient (IUCN). |
| T. malcolmi | Loveridge, 1944 | Luzon, Philippines | Philippine pit viper. |
| T. mcgregori | Taylor, 1917 | Philippines | McGregor's pit viper. |
| T. medogensis | Jiang & Li, 2022 | Medog County, Tibet, China | Recently described; molecular evidence supports separation from T. stejnegeri. |
| T. melanocephalus | Gray, 1842 | Malaysia | Black-headed pit viper. |
| T. monticola | Günther, 1864 | Sri Lanka | Mountain pit viper; synonym of T. trigonocephalus in some older classifications, now distinct. |
| T. muenchingeri | David et al., 2023 | Vietnam | Named for collector; from T. popeiorum complex. |
| T. nebularis | Vogel et al., 2014 | Borneo | Cloud forest pit viper. |
| T. obor | Hoge & Romano-Hoge, 1981 | Sumatra, Indonesia | Blotched pit viper. |
| T. otophiophilus | Mumpuni et al., 2015 | Sulawesi, Indonesia | Eared pit viper. |
| T. papei | Archer, 2021 | Malaysia | From T. popeiorum revision. |
| T. popeiorum | Smith, 1937 | Malaysia | Revised in 2023 into multiple species/subspecies, e.g., T. caudornatus as synonym; Vulnerable. |
| T. puniceus | Boie, 1827 | Java, Indonesia | Reddish pit viper. |
| T. rubeus | Ngoc & David, 2021 | Vietnam | Red pit viper; from T. albolabris complex. |
| T. salazar | Orlov et al., 2010 | Vietnam | Salazar's pit viper; described from Tam Dao; Least Concern. |
| T. schlegelii | Duméril et al., 1854 | Indonesia | Schlegel's pit viper; widespread. |
| T. sebelenyeri | David et al., 2023 | Laos | From T. popeiorum group. |
| T. stejnegeri | Stejneger, 1907 | Taiwan | Stejneger's pit viper; includes former T. monticola populations. |
| T. tibetanus | Huang, 1982 | Tibet, China | Tibetan pit viper; Data Deficient. |
| T. trigonocephalus | Günther, 1864 | Sri Lanka | Green pit viper; synonym issues with T. monticola resolved via phylogeny. |
| T. venustus | Vogel, 1991 | Thailand | Beautiful pit viper; part of T. kanburiensis complex. |
| T. vogeli | David et al., 2009 | Central Vietnam | Vogel's pit viper. |
| T. yatesi | David et al., 2023 | India | From T. popeiorum revision. |
| T. zonatus | Günther, 1862 | Andaman Islands, India | Zoned pit viper. |
| T. cardamomensis | Stuart & Heatwole, 2004 | Cambodia | Cardamom pit viper. |
| T. cernio | Günther, 1864 | Borneo | Cerro pit viper. |
| T. cryptus | Hoge & Romano-Hoge, 1981 | Philippines | Cryptic pit viper. |
| T. cyanolabris | Idiiatullina et al., 2024 | China | Recently described; blue-lipped.[19] |
| T. erythrochloris | Pawangkhanant et al., 2025 | Eastern Thailand | New species from T. albolabris complex; ontogenetic color change noted.[19] |
| T. flavomaculatus | Ishii et al., 2011 | Ryukyu Islands, Japan | Yellow-spotted pit viper; split from T. flavoviridis. |
| T. guangxiensis | Liang & Liu, 2007 | Guangxi, China | Guangxi pit viper. |
| T. honsonensis | David et al., 2012 | Vietnam | Hon Son pit viper. |
| T. kuiburi | Sumontha et al., 2011 | Thailand | Kuiburi pit viper; from T. kanburiensis. |
| T. labiatus | Günther, 1889 | Nicobar Islands, India | White-lipped variant; synonym debates with T. albolabris. |
| T. macrops | Kramer, 1977 | Malaysia | Large-eyed pit viper. |
| T. naejai | Sumontha et al., 2012 | Thailand | Naeja's pit viper. |
| T. phongsalyensis | David et al., 2012 | Laos | Phongsaly pit viper. |
| T. pretiosus | Xu et al., 2025 | Xizang, China | New species; morphologically similar to T. stejnegeri.[20] |
| T. sichuanensis | Guo & Wang, 1981 | Sichuan, China | Sichuan pit viper. |
| T. uetzi | Vogel et al., 2023 | Northeast India/Vietnam | Named for Reptile Database editor; recent range extension to India.[17] |
| T. cryptographicus | Pawangkhanant et al., 2025 | Thailand | Cryptic green pit viper; unusual ontogenetic color change.[21] |
| T. nujiang | Li et al., 2025 | Nujiang, China | New from Yunnan; likely extends to Myanmar.[5] |
| T. loong | Li et al., 2025 | Kunming City, Yunnan, China | 'Little dragon' green pit viper; described November 2025, raising total to 57.[22] |