Skip to main content Skip to main navigation menu Skip to site footer
Type: Monograph
Published: 2004-11-15
Page range: 1–63
Abstract views: 125
PDF downloaded: 3

A review of morphological variation in Trimeresurus popeiorum (Serpentes: Viperidae: Crotalinae), with the description of two new species

Society for Southeast Asian Herpetology, Im Sand 3, D-69115 Heidelberg, Germany
Département Systématique et Evolution, USM 602 Taxonomie-collection Reptiles & Amphibiens, Case postale 30, Muséum National d’Histoire Naturelle, 25 rue Cuvier, F-75231 Paris Cedex 05, France
Department of Recent Vertebrates, Institut Royal des Sciences Naturelles de Belgique Rue Vautier 29, B-1000 Brussels, Belgium
Reptiles Thailand West Malaysia Sumatra Borneo Serpentes Viperidae Trimeresurus Trimeresurus popeiorum Trimeresurus fucatus spec. nov. Trimeresurus nebularis spec. nov. Trimeresurus sabahi Trimeresurus barati

Abstract

Variation in morphological characters were investigated among 136 specimens (128 specimens examined by us and eight specimens described in the literature) from 44 populations of the whole range of the pitviper currently known as Trimeresurus popeiorum Smith, 1937. Univariate and mul-tivariate analyses of these morphological characters allowed us to recognize six clusters of populations that are morphologically diagnosable, and that are here considered to represent independent lineages. Five of these clusters are considered to be distinct species following the Biological Species Concept and the Phylogenetic Species Concept. Two of them are described as new. Trimeresurus fucatus spec. nov. includes populations from southern Thailand and most of West Malaysia. Trimeresurus nebularis spec. nov. is described for populations from Cameron Highlands of West Malaysia. A population from Toba Massif, northern Sumatra, is referred to this complex, but cannot be assigned to a species at the present time. Trimeresurus popeiorum sabahi is raised to specific status, Trimeresurus sabahi new comb., to accommodate the populations from Borneo, whereas Trimeresurus barati new comb. includes the populations from western Sumatra and the Mentawai Archipelago. Separate keys to the two sexes of the recognised species of the T. popeiorum complex are provided.KEY WORDS: Thailand, West Malaysia, Sumatra, Borneo, Serpentes, Viperidae, Trimeresurus, Trimeresurus popeiorum, Trimeresurus fucatus spec. nov., Trimeresurus nebularis spec. nov., Trimeresurus sabahi, Trimeresurus baratiBefore the paper by Pope & Pope (1933), all green Trimeresurus species were gathered under the name Trimeresurus gramineus (Shaw, 1802). In a first step towards understanding the systematics of the genus, these authors split the nominal taxon gramineus into six species. The specific nomen gramineus was applied to a widespread species, ranging from northeastern India to western Indonesia. Indian populations were referred to a new species described as Trimeresurus occidentalis. Subsequently, Smith (1937) correctly showed that Pope & Pope (1933) misunderstood the type locality of gramineus, and showed that the type locality for T. gramineus was within the range of T. occidentalis. Therefore, Trimeresurus occidentalis Pope & Pope 1933 became a subjective junior synonym of T. gramineus (Shaw, 1802), leaving unnamed the distinct eastern taxon. Smith (1937) named it as Trimeresurus popeiorum. Unfortunately, he failed to designate a type specimen and a type locality for this new taxon. This interpretation was accepted by most subsequent authors except Hoge & Romano Hoge (1981) and Welch (1988). Another issue affecting the specific nomen is its spelling. Smith (1943) corrected the original spelling as popeorum on the basis that it was indeed a clerical error. This spelling was largely accepted, and was the subject of recent controversies. This problem will be addressed in another paper (David & Vogel, submitted). We consider that the correct spelling is indeed popeiorum. Eventually, Taylor & Elbel (1958), regarded as syntypes of Trimeresurus popeiorum Smith, 1937 all specimens referred by Pope & Pope (1933) to as T. gramineus, and designated the specimen BMNH 72.4.17.137 as the lectotype of the species. Consequently, the type locality was restricted to Khasi Hills, Assam , now in the State of Meghalaya, India.

References

  1. Ahmed, S. & Dasgupta, G. (1992) Fauna of West Bengal. Part 2. Reptilia, Zoological Survey of India, Calcutta, State Fauna Series, 3, 1–65.

    Anonymous. (2000) Kaeng Krachan. Amazing Forest of Phetchaburi River, The Tourism Authority of Thailand, Bangkok, 159 pp.

    Baumann, F. (1913) Reptilien und Batrachier des Berner Naturhistorischen Museums aus dem Battak-Gebirge von West-Sumatra. Zoologische Jahrbücher Jena, Abteilung für Systematik, Ökologie und Geographie der Tiere, 34 (3), 257–278.

    Böhme, W. (1988) Zur Genitalmorphologie der Sauria: funktionelle und stammesgeschichtliche Aspekte. Bonner zoologische Monographien, 27, 1–176.

    Boulenger, G.A. (1896) Catalogue of the Snakes in the British Museum (Natural History). Volume III. Containing the Colubridae (Opisthoglyphae and Proteroglyphae), Amblycephalidae and Viperidae, British Museum (Natural History), London, xiv + 727 pp., Pls. 1–25.

    Boulenger, G.A. (1912) A vertebrate fauna of the Malay Peninsula from the Isthmus of Kra to Singapore, including the adjacent islands. Reptilia and Batrachia, Taylor & Francis, London, xiii + 294 pp.

    Campden-Main, S.M. (1970) A field guide to the snakes of South Vietnam, Smithsonian Institution, Washington, v + 112 pp., 1 plate.

    Chan-ard, T., Grossmann, W., Gumprecht, A. & Schulz, K.-D. (1999) Amphibians and Reptiles of Peninsular Malaysia and Thailand. An illustrated checklist, Bushmaster Publ., Wuerselen, 240 pp.

    Cantor, T.E. (1839) Spicilegium Serpentium Indicorum. Proceedings of the Zoological Society of London, 1839, 31–34 & 49–55.

    Cox, M.J. (1991) The snakes of Thailand and their husbandry, Krieger Publishing Co., Malabar, Florida, xxxviii + 526 pp.

    Cox, M.J., van Dijk, P.P., Nabhitabhata, J. & Thirakhupt, K. (1998) A Photographic Guide to Snakes and other Reptiles of Peninsular Malaysia, Singapore and Thailand, New Holland Publishers, London–Cape Town–Sydney–Singapore, 144 pp.

    Cracraft, J. (1997) Species concept in systematics and conservation biology – an ornithological viewpoint. In: Claridge, M.F., Dawah, H.A. & Wilson, M.R. (eds.) Species. The units of biodiversity, Chapman & Hall, London, The Systematics Association Special Volume Series, 54, 325–339.

    David, P. & Ineich, I. (1999) Les serpents venimeux du monde: systématique et répartition. Dumerilia, Paris, 3, 3–499.

    David, P., Vidal, N. & Pauwels, O.S.G. (2001) 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. (1996) The snakes of Sumatra. An annotated checklist and key with natural history notes, Edition Chimaira, Frankfurt–am–Main, 260 pp.

    David, P. & Vogel, G. (In prep.) Trimeresurus popeiorum or Trimeresurus popeorum? On the correct spelling of the specific nomen of the Pope’s Pitviper (Reptilia, Serpentes, Crotalidae). Hamadryad.

    David, P., Vogel, G., Pauwels, O.S.G. & Vidal, N. (2002) Description of a new species of the genus Trimeresurus from Thailand, related to Trimeresurus stejnegeri Schmidt, 1925 (Serpentes, Viperidae, Crotalinae). The Natural History Journal of Chulalongkorn University, 2 (1), 25–30.

    Dowling, H.G. (1951) A proposed standard system of counting ventrals in snakes. British Journal of Herpetology, 1 (5), 97–99.

    Dring, J.C.M. (1979) Amphibians and reptiles from northern Trengganu, Malaysia, with descriptions of two new geckos: Cnemaspis and Cyrtodactylus. Bulletin of the British Museum of Natural History, (Zoology), 34 (5), 181–241.

    Dring, J.C.M., McCarthy, C.J. & Whitten, A.J. (1990) The terrestrial herpetofauna of the Mentawai Islands, Indonesia. Indo-Malayan Zoology, 6 [1989], 119–132.

    Dubois, A. (1977) Les problèmes de l’espèce chez les Amphibiens Anoures. Mémoires de la Société Zoologique de France, 39, 161-284.

    Flower, S.S. (1896) Notes on a collection of reptiles and batrachians made in the Malay Peninsula in 1895–96; with a list of the species recorded from that region. Proceedings of the Zoological Society of London, 1896, 856–914, Pls. 44–46.

    Flower, S.S. (1899) Notes on a second collection of reptiles made in the Malay Peninsula and Siam, from November 1896–September 1898, with a list of the species recorded from those countries. Proceedings of the Zoological Society of London, 1899, 600–696, Pls. 36–37.

    Frost, D.R. & Hillis, D.M. (1990) Species in concept and practice: herpetological implications. Herpetologica, 46 (1), 87–104.

    Giannasi, N., Malhotra, A. & Thorpe, R. S (2001) Nuclear and mtDNA phylogenies of the Trimeresurus complex: implications for the gene versus species tree debate. Molecular Phylogenetics and Evolution, 19 (1), 57–66.

    Golay, P., Smith, H.M., Broadley, D.G., Dixon, J.R., McCarthy, C.J., Rage, J.-C., Schätti, B. & Toriba, M. (1993) Endoglyphs and other major venomous snakes of the world. A checklist, Azemiops S.A. Herpetological Data Center, Aïre–Genève, xv + 478 pp.

    Grandison, A.G.C. (1978) Snakes of West Malaysia and Singapore. Annalen des Naturhistorischen Museums in Wien, 81, 283–302.

    Grossmann, W. & Tillack, F. (2001) Bemerkungen zur Herpetofauna des Khao Lak, Phang Nga, thailändische Halbinsel. Teil III: Ergebnisse der Jahre 1999 und 2000. Sauria, Berlin, 23 (3), 21–34.

    Gumprecht, A. (2001) Die Bambusottern der Gattung Trimeresurus Lacépède. Teil IV: Checkliste der Trimeresurus-Arten Thailands. Sauria, Berlin, 23 (2), 25–32.

    Hoge, A.R. & Romano Hoge, S.A.R.W.L. (1981) Poisonous snakes of the world. Part I. Check-list of the pit vipers Viperoidea, Crotalinae. Memorias do Instituto Butantan, 42/43 [1978/1979], 179–310.

    How, R.A., Schmitt L.H. & Suyanto, A. (1996) Geographical variation in the morphology of four snake species from the Lesser Sunda Islands, Eastern Indonesia. Biological Journal of the Linnean Society, 59, 439–456.

    ICZN (International Commission on Zoological Nomenclature) (1999) International Code of Zoological Nomenclature. Fourth Edition adopted by the International Union of Biological Sciences, International Trust of Zoological Nomenclature, London, xxix + 306 pp.

    Inger, R.F. & Colwell, R.K. (1977) Organization of contiguous communities of amphibians and reptiles in Thailand. Ecological Monographs, 47, 229–253.

    Inger, R.F. & Voris, H.K. (2001) The biogeographical relations of the frogs and snakes of Sundaland. Journal of Biogeography, 28, 863–891.

    Iskandar, D.T. & Colijn, E. (2001) A checklist of Southeast Asian and New Guinean reptiles. Part I: Serpentes. Biodiversity Conservation Project (Indonesian Institute of Sciences – Japan International Cooperation Agency – The Ministry of Forestry), The Gibbon Foundation and Institut of Technology, Bandung, 195 pp.

    Jintakune, P. & Chanhome, L. (1995) Ngoo phit nai Prathet Thai, Queen Saovabha Memorial Institute, Thai Red Cross Society, Bangkok, 175 (+ 3 unnumbered) pp. (In Thai). Note. – Translation of the title: The venomous snakes of Thailand.

    Leong, T.M. & Lim, K.K.P. (2003) Herpetofaunal records from Fraser’s Hill, Peninsular Malaysia, with larval descriptions of Limnonectes nitidus and Theloderma asperum (Amphibia: Ranidae and Rhacophoridae). Raffles Bulletin of Zoology, 51 (1), 123–136.

    Leviton, A.E., Wogan, G.O.U., Koo, M.S., Zug, G.R., Lucas, R.S. & Vindum, J.V. (2003) The dangerously venomous snakes of Myanmar. Illustrated checklist with keys. Proceedings of the California Academy of Sciences, 54 (2), 407–462.

    Lim, B.L. (1982) Poisonous snakes of Peninsular Malaysia. Second Edition, Art Printing Works Kuala Lumpur, viii + 73 pp.

    Lim, B. L. (1990) Venomous land snakes of Malaysia. In: Gopalakrishnakone, P. & Chou, L. M. (ed.) Snakes of medical importance (Asia–Pacific region), National University of Singapore, Singapore, 387–417.

    Lim, B. L. (1991) Poisonous snakes of Peninsular Malaysia. Third Edition, Malayan Nature Society, Kuala Lumpur, 73 pp.

    Lim, B.L., Ratnam, L. & Hussein, N. A. (1995) Snakes examined from the Sungai Singgor area of Temengor, Hulu Perak, Malaysia. Malayan Nature Journal, 48, 357–364.

    Lim, K.K.P. & Lim, L.J. (1999) The terrestrial herpetofauna of Pulau Tioman, Peninsular Malaysia. Raffles Bulletin of Zoology, Supplement, 6, 131–155.

    Malhotra, A. & Thorpe, R.S. (2004) A phylogeny of four mitochondrial gene regions suggests a revised taxonomy for Asian pitvipers (Trimeresurus and Ovophis), Molecular Phylogenetics and Evolution, 32 (1), 83–100.

    Malkmus, R. (1992) Herpetologische Beobachtungen am Mount Kinabalu, Nord-Borneo. III. Mitteilungen aus dem Zoologischen Museum in Berlin, 68 (1), 101–138.

    Malkmus, R. (1994) Herpetologische Beobachtungen am Mount Kinabalu, Nord-Borneo. IV. Mitteilungen aus dem Zoologischen Museum in Berlin, 70 (2), 217–249.

    Malkmus, R., Manthey, U., Vogel, G., Hoffmann, P. & Kosuch, J. (2002) Amphibians & reptiles of Mount Kinabalu (North Borneo), A.R.G. Gantner Verlag Kommanditgesellschaft, Ruggell, 424 pp.

    Manthey, U. & Grossmann, W. (1997) Amphibien & Reptilien Südostasiens, Natur und Tier-Verlag, Münster, 512 pp.

    Maslin, T.P. (1942) Evidence for the separation of the crotalid genera Trimeresurus and Bothrops, with a key to the genus Trimeresurus. Copeia, 1942 (1), 18–24.

    Mayden, R.L. (1997) A hierarchy of species concept: the denouement in the saga of the species problem. In: Claridge, M.F., Dawah, H.A. & Wilson, M.R. (ed.) Species. The units of biodiversity, Chapman & Hall, London, The Systematics Association Special Volume Series, 54, 381–424.

    Mayr, E. (2000) The Biological Species Concept. In: Wheeler, Q.D & Meier, R. (ed.) Species concepts and phylogenetic theory. A debate, Columbia University Press, New York, 17–29.

    Mayr, E. & Ashlock, P.D. (1991) Principles of systematic zoology, Second edition, McGraw-Hill Book Co., Singapore, xx + 475 pp.

    McDiarmid, R.W., Campbell, J.A. & Touré, T’S.A. (1999) Snake species of the world. A taxonomic and geographical reference. Volume 1, The Herpetologists’ League, Washington DC, xi + 511 pp.

    Mouhot, H. (1864) Travels in the Central parts of Indo-China (Siam), Cambodia, and Laos, during the years 1858, 1859 and 1860. Vol. II, John Murray, London, viii + 301 pp., map.

    Nootpand, W. (1971) Poisonous snakes of Thailand, Thai Zoological Centre, Bangkok, 68 pp. (In Thai & English).

    Nutphand, W. (2001) Snakes in Thailand, Amarin Printing and Publishing Public Co., Ltd, Bangkok, 319 pp. (In Thai).

    Orlov, N. (1997) Viperid snakes (Viperidae Bonaparte, 1840) of Tam-Dao mountain ridge (Vinh–Phu and Bac–Thai provinces, Vietnam). Russian Journal of Herpetology, 4 (1), 64–74, Figs. I–VI.

    Orlov N., Ananjeva, N., Barabanov, A., Ryabov, S. & Khalikov, R. (2002a) Diversity of vipers (Azemiopinae, Crotalinae) in East, Southeast, and South Asia: Annotated checklist and natural history data (Reptilia: Squamata: Serpentes: Viperidae). In: Fritz, U. (Ed.), Collectanea Herpetologica. Essays in honour of Fritz Jürgen Obst. Faunistische Abhandlungen des Staatlichen Museums für Tierkunde Dresden, 23, 177–218.

    Orlov, N., Ananjeva, N. & Khalikov, R. (2002b) Natural history of Pitvipers in eastern and southeastern Asia. In: Schuett, G.W., Höggren, M., Douglas, M.E. & Greene, H.W. (Ed.) Biology of the Vipers, Eagle Mountain Publishing, Eagle Mountain, Utah, 345–359.

    Pauwels, O.S.G., David, P., Chimsunchart, C. & Thirakhupt, K. (2003) Reptiles of Phetchaburi Province, Western Thailand: a list of species, with natural history notes, and a discussion on the biogeography at the Isthmus of Kra. The Natural History Journal of Chulalongkorn University, 3 (1), 23–53.

    Pope, C.H. & Pope, S.H. (1933) A study of the green pit-vipers of southeastern Asia and Malaysia, commonly identified as Trimeresurus gramineus (Shaw), with description of a new species from Peninsular India. American Museum Novitates, 620, 1–12.

    Raffles, T.S. (1822) Second part of the descriptive catalogue of a zoological collection made in the island of Sumatra and its vicinity. Transactions of the Linnean Society of London, 13 (2), 277–340.

    Regenass, U. & Kramer, E. (1981) Zur Systematik der grünen Grubenottern der Gattung Trimeresurus (Serpentes, Crotalidae). Revue suisse de Zoologie, 88 (1), 163–205.

    Siegel, S. (1956) Nonparametric statistics for the behavioral sciences, McGraw-Hill Kogakusha, New York, Toronto, London and Tokyo, xvii + 312 pp.

    Shaw, G. (1802) General Zoology, or Systematic Natural History. With plates from the first authorities and most select specimens Engraved principally by Mr. Heath. Volume III, Part II. Amphibia, G. Kearsley, London, i–vi + 313–615, Pls. 87–140.

    Smedley, N. (1932) Amphibians and reptiles from the Cameron Highlands, Malay Peninsula. Bulletin of the Raffles Museum, 6 [1931], 105–123, Pl. 2.

    Smith, M.A. (1922) On a collection of reptiles and batrachians from the mountains of Pahang, Malay Peninsula. Journal of the Federated Malay States Museums, 10, 263–282.

    Smith, M.A. (1930) The reptilia and amphibia of the Malay Peninsula. Bulletin of the Raffles Museum, 3, i–xviii + 1–149.

    Smith, M.A. (1937) The names of two Indian vipers. Journal of the Bombay Natural History Society, 39 (4), 730–731.

    Smith, M.A. (1943) The Fauna of British India, Ceylon and Burma, including the whole of the Indo–chinese subregion. Reptilia and Amphibia. Vol. III, Serpentes, Taylor & Francis, London, xii + 583 pp.

    Stuebing, R.B. (1991) A checklist of the snakes of Borneo. Raffles Bulletin of Zoology, 39 (2), 323–362.

    Stuebing, R.B. & Inger, R.F. (1999) A fieldguide to the snakes of Borneo, Natural History Publications, Kota Kinabalu, 254 pp.

    Taylor, E.H. (1965) The serpents of Thailand and adjacent waters. University of Kansas Science Bulletin, 45 (9), 609–1096.

    Taylor, E.H. & Elbel, R.E. (1958) Contribution to the herpetology of Thailand. University of Kansas Science Bulletin, 38 (2), 1033–1189.

    Tu, M.-C., Wang, H.-Y., Tsai, M.-P., Toda, M., Lee, W.-J., Zhang. F. J. & Ota, H. (2000) Phylogeny, taxonomy, and biogeography of the Oriental Pitvipers of the genus Trimeresurus (Reptilia: Viperidae: Crotalinae): A molecular perspective. Zoological Science, 17, 1147–1157.

    Thumwipat, B. & Nutphand, W. (1982) Treatment of patients bitten by venomous snakes and venomous snakes of Thailand, Thai Zoological Center, Bangkok, (5 + 7 unnumbered pages), 162 pp. (In Thai).

    Tweedie, M.W.F. (1941) The poisonous snakes of Malaya. Part III. The Vipers and Sea Snakes. Malayan Nature Journal, 1 (4), 128–134.

    Tweedie, M.W.F. (1954) The snakes of Malaya. Government Printing Office, Singapore, 139 pp., Pls. 1–12.

    Tweedie, M.W.F. (1957) The snakes of Malaya, Second edition, Government Printing Office, Singapore, 143 pp., Pls. 1–16.

    Tweedie, M.W.F. (1983) The snakes of Malaya, Third edition, Singapore National Printers, Singapore, 167 pp., Pls. 1–12.

    Vogel, G. (1990) Über eine verkannte Trimeresurus-Art aus Thailand. Sauria (Berlin), 12 (2), 11–13.

    Voris, H.K. (2000) Maps of Pleistocene sea levels in Southeast Asia: shorelines, river systems and time durations. Journal of Biogeography, 27, 1153–1167.

    Wall, F. (1909) Notes on snakes from the neighbourhood of Darjeeling. Journal of the Bombay Natural History Society, 19, 337–357.

    Welch, K.R.G. (1988) Snakes of the Orient: a checklist, Robert F. Krieger Publ. Co., Malabar, Florida, vii + 183 pp.

    Wheeler, Q.D. & Platnick, N.I. (2000) The Phylogenetic Species Concept (sensu Wheeler & Platnick). In: Wheeler, Q.D & Meier, R. (ed.) Species concepts and phylogenetic theory. A debate, Columbia University Press, New York, 55–69.

    Wüster, W. (1992). Cobras and other herps in South–east Asia. British Herpetological Society Bulletin, 39, 19–25.

    Wüster, W & Thorpe R.S. (1992) Asiatic cobras: population systematics of the Naja naja species complex (Serpentes: Elapidae) in India and Central Asia. Herpetologica, 48 (1), 69–85.