Unravelling the underestimated diversity of Philippine water monitor lizards (Squamata: Varanus salvator complex), with the description of two new species and a new subspecies

• Main Authors: Koch, André, Gaulke, Maren, Böhme, Wolfgang
• Format: Article in Journal/Newspaper
• Language: unknown
• Published: Zenodo 2010
• Subjects: Biodiversity; Taxonomy; Animalia; Chordata; Reptilia; Squamata; Varanidae; Cramer; Custodio; Dickerson; Fairbanks; Fuller; Gonzalez; Harper; Huxley; Krebs; Lorius; Maren; Mayhew; Meier; Myers; Nickerson; Pacific; Sunda; Visser; Antarc*; Antarctica; ice core
• Online Access: https://dx.doi.org/10.5281/zenodo.195067; https://zenodo.org/record/195067

Koch, André, Gaulke, Maren, Böhme, Wolfgang (2010): Unravelling the underestimated diversity of Philippine water monitor lizards (Squamata: Varanus salvator complex), with the description of two new species and a new subspecies. Zootaxa 2446: 1-54, DOI: 10.5281/zenodo.195067 : {"references": ["Alcala, A.C. (1957) Philippine notes of the ecology of the giant marine toad. Silliman Journal, 4(2), 90-96.", "Alcala, A.C. (1986) Guide to Philippine flora and fauna - Amphibians and Reptiles. JMC Press, Quezon City, Philippines, 195 pp.", "Ast, J.C. (2001) Mitochondrial DNA evidence and evolution in Varanoidea (Squamata). Cladistics, 17, 211-226.", "Auffenberg, W. (1988) Gray 's monitor lizard. University Press of Florida, 419 pp.", "Bauer, A.M. & Adler, K. (2001) The dating and correct citation of A. F. A. Wiegmann's \"Amphibien\" section of Meyen's Reise um die Erde, with a bibliography of Wiegmann's herpetological publications. Archives of Natural History, 28(3), 313-326.", "Baverstock, P.R., King, D., King, M., Birrell, J., & Krieg, M. (1993) The evolution of species of the Varanidae: Microcomplement fixation analysis of serum albumins. Australian Journal of Zoology, 41, 621-638.", "Bayless, M.K. & Adragna, J.A. (1997) Monitor lizards in the Philippine Islands: a historical perspective (Sauria: Varanidae). Asia Life Sciences, 6(1/2), 39-50.", "Bayless, M.K. & Adragna, J.A. (2002) John G. Veitch and the Negros water monitor lizard (Varanus salvator nuchalis Gunther, 1872) of the Philippine Islands. Dragon News, 3.", "Becker, H.O., Bohme, W. & Perry, S.F. (1989) Die Lungenmorphologie der Warane (Reptilia: Varanidae) und ihre systematisch-stammesgeschichtliche Bedeutung. Bonner Zoologische Beitrage, 40, 27-56.", "Bennett, D. (1998) Monitor lizards: natural history, biology and husbandry. Edition Chimaira, Frankfurt a.M., 354 pp.", "Bennett, D. (2000) Notes on Varanus salvator marmoratus on Polillo Island, Philippines. In: Bennett, D. (Ed), Wildlife of Polillo Island, Philippines. Viper Press, Aberdeen. pp. 29-32.", "Bohme, W. (1988) Zur Genitalmorphologie der Sauria: Funktionelle und stammesgeschichtliche Aspekte. Bonner Zoologische Monographien, 27, 3-176.", "Bohme, W. (2003) Checklist of the living monitor lizards of the world (family Varanidae). Zoologische Verhandelingen, 341, 1-43.", "Bohme, W. & Koch, A. (2010) On the type selection and re-typification of two monitor lizard taxa (Squamata: Varanidae): Monitor bivittatus celebensis Schlegel, 1844 and Monitor kordensis Meyer, 1874; with some comments and corrections on other name-bearing type specimens. Zootaxa.", "Bohme, W. & Ziegler, T. (2009) A review of iguanian and anguimorph lizard genitalia (Squamata: Chamaeleonidae; Varanoidea, Shinisauridae, Xenosauridae, Anguidae) and their phylogenetic significance: comparisons with molecular data sets. Journal of Zoological Systematics and Evolutionary Research, 47, 189-202.", "Bohme, W., Horn, H.-G. & Ziegler, T. (1994) Zur Taxonomie der Pazifikwarane (Varanus indicus -Komplex): Revalidierung von Varanus doreanus (A. B. Meyer, 1874) mit Beschreibung einer neuen Unterart. Salamandra, 30, 119-142.", "Borden, R. (2007) Varanus salvator (Asian Water Monitor) Migration. Biawak, 1, 84.", "Boulenger, G.A. (1885) Catalogue of the lizards in the British Museum (Natural History). Second edition. Vol. 2. Iguanidae, Xenosauridae, Zonuridae, Anguidae, Anniellidae, Helodermatidae, Varanidae, Xantusiidae, Teeidae, Amphisbaenidae. Taylor and Francis, London, 436 pp.", "Boulenger, G.A. (1894) On the herpetological Fauna of Palawan and Balabac. Annals and Magazine of Natural History, 14, 81-90.", "Brandenburg, T. (1983) Monitors in the Indo-Australian Archipelago. Unpubl. Thesis, University of Leiden, Leiden, 98 pp.", "Brown, R.M. & Diesmos, A. C. (2001) Application of lineage-based species concepts to oceanic island frog populations: the effects of differing taxonomic philosophies on the estimation of Philippine biodiversity. Silliman Journal, 42, 133-162.", "Brown, R.M. & Diesmos, A. (2009) Philippines, Biology. In: Gillespie, R. & Clague, D. (Eds), Encyclopedia of Islands. University of California Press, Berkeley. pp. 723-732.", "Brown, R.M., Diesmos, A.C. & Alcala, A.C. (2001) The state of Philippine Herpetology and the Challenges of the next decade. Silliman Journal, 42, 18-87.", "Brown, R.M., Ferner, J.W., Sison, R.V., Gonzales, P.C. & Kennedy, R.S. (1996) Amphibians and reptiles of the Zambales Mountains of Luzon Island, Republic of the Philippines. Herpetological Natural History, 4(1), 1-22.", "Brown, R.M., Siler, C.D., Diesmos, A.C., & Alcala, A.C. (2009) Philippine frogs of the genus Leptobrachium (Anura; Megophryidae): phylogeny-based species delimitation, taxonomic review, and description of three new species. Herpetological Monographs, 23, 1-44.", "Brown, W.C. & Alcala, A.C. (1970) The zoogeography of the herpetofauna of the Philippine Islands, a fringing archipelago. Proceedings of the California Academy of Sciences, 38, 105-130.", "Brown, W.C. & Alcala, A.C. (1980) Philippine lizards of the family Scincidae. Silliman University Natural Sciences Monographs, Dumaguete City, pp. 264.", "Brygoo, E.-R. (1987) Les types de varanides (reptiles, sauriens) du Museum national d'Histoire naturelle. Catalogue critique. Bulletin du Museum national d'Histoire naturelle, Paris, 9(2) Suppl., 21-38.", "Censky, E.J., Hodge, K. & Dudley, J. (1998) Over-water dispersal of lizards due to hurricanes. Nature, 395, 556.", "Cramer, D. (2003) Advanced quantitative data analysis. Open University Press, Philadelphia, 254 pp.", "Cuvier, G. (1829) Le Regne animal distribue d'apres son organisation, pour servir de base a l'Histoire Naturelle des animaux et d'introduction a l'anatomiecomparee, Ed. 2, Vol. 2, Deterville et Crochard, Paris.", "Das, I. & Kueh, B.H. (2008) Herpetofauna of the Pulau Banggi Group of islands off northeastern Borneo. Herpetological Review, 39, 296-298.", "de Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation: a conceptual unification and terminological recommendations. In: Howard, D.J. & Berlocher, S.H. (Eds), Endless forms: species and speciation. Oxford University Press, Oxford. pp. 57-75.", "de Queiroz, K. (2005) A unified concept of species and its consequences for the future of taxonomy. Proceedings of the California Academy of Science, 56, 196-215.", "Evans, B.J., Brown, R.M., McGuire, J.A., Supriatna, J., Andayani, N., Diesmos, A., Iskandar, D., Melnick, D.J. & Cannatella, D.C. (2003) Phylogenetics of fanged frogs: testing biogeographical hypotheses at the interface of the Asian and Australian faunal Zones. Systematic Biology, 52, 794-819.", "Dickerson, R.E. (1928) Distribution of life in the Philippines. Philippine Bureau of Sciences Monographs, 21, 322 pp.", "Diesmos, A. & Gaulke, M. (2007) Varanus nuchalis. In: IUCN 2009. IUCN red list of threatened species. version 2009.2, www.iucnredlist.org, downloaded on 17 February 2010.", "Esselstyn, J.A., Widmann, P. & Heaney, L.R. (2004) The mammals of Palawan Island, Philippines. Proceedings of the Biological Society of Washington, 117, 271-302.", "Everett, A.H. (1889) Remarks on the zoo-geographical relationships of the island of Palawan and some adjacent Islands. Proceedings of the Royal Society of London, 1889, 220-228.", "Fairbanks, R.G. (1989) A 17,000-year glacio-eustatic sea level record: influence of glacial melting rates on the Younger Dryas event and deep-ocean circulation. Nature, 342, 637-642.", "Ferner, J.W., Brown, R.M., Sison, R.V. & Kennedy, R.S. (2001) The amphibians and reptiles of Panay Island, Philippines. Asiatic Herpetological Research, 9, 34-70.", "Fuller, S., Baverstock, P. &King D. (1998) Biogeographic origins of goannas (Varanidae): A molecular perspective.", "Gaulke, M. (1986) Uber die Situation des Bindenwarans (Varanus salvator nuchalis) auf Negros, Philippinen. Herpetofauna, 8(44), 16-18.", "Gaulke, M. (1989) Zur Biologie des Bindenwaranes, unter Berucksichtigung der palaogeographischen Verbreitung und der phylogetischen Entwicklung der Varanidae. Courier Forschungsinstitut Senckenberg, 112, 1-242.", "Gaulke, M. (1991a) Systematic relationships of the Philippine water monitors as compared with Varanus s. salvator, with a discussion of dispersal routes. In: Bohme, W. & Horn, H.-G. (Eds), Advances in Monitor research. Mertensiella, 2, pp. 154-167.", "Gaulke, M. (1991b) On the diet of the water monitor, Varanus salvator, in the Philippines. In: Bohme, W. & Horn, H.-G. (Eds), Advances in monitor research, Mertensiella, 2, pp. 143-153.", "Gaulke, M. (1992a) Taxonomy and biology of Philippine water monitors (Varanus salvator). Philippine Journal of Science, 121(4), 345-381.", "Gaulke, M. (1992b) Distribution, population density, and exploitation of the water monitor (Varanus salvator) in the Philippines. Hamadryad, 17, 21-27.", "Gaulke, M. (1994) Notes on the herpetofauna of Panaon and Samar, east Visayans, Philippines. Hamadryad, 19, 1-10.", "Gaulke, M. (1995) Der Sulu-Archipel - Besiedlungsgeschichte, Geologie und Herpetofauna. Natur und Museum, 125(7), 217-226.", "Gaulke, M. (1996) Die Herpetofauna von Sibutu Island (Philippinen), unter Berucksichtigung zoogeographischer und okologischer Aspekte. Senckenbergiana Biologica, 75(1/2), 45-56.", "Gaulke, M. (1998) Utilization and conservation of lizards and snakes in the Philippines. In: Erdelen, W. (Ed), Conservation, trade and sustainable use of lizards and snakes in Indonesia, Mertensiella, 9, pp. 137-142.", "Gaulke, M. (2001) Philippinen-Naturreisefuhrer. Natur und Tier-Verlag, 398 pp.", "Gaulke, M. (2008) Unterwegs im Suden der Philippinen. Terraria, 10, 69-80.", "Gaulke, M. & Curio, E. (2001) A new monitor lizard from Panay Island, Philippines. Spixiana, 24(3), 275-286.", "Gaulke, M. & Reiter, J. (2001) Varanus salvator nuchalis, eine wenig bekannte Unterart des Bindenwarans. Draco, 7, 42-49.", "Gaulke, M., Sy, E., Afuang, L, Duya, M.R., Diesmos, M. & Gonzalez, J.C. (2007) Varanus marmoratus. In: IUCN 2009. IUCN red list of threatened species. version 2009.2, www.iucnredlist.org, downloaded on 17 February 2010.", "Good, D.A., Bauer, A.M. & Gunther, R. (1993) An annotated type Catalogue of the anguimorph lizards (Squamata: Anguidae, Helodermatidae, Varanidae, Xenosauridae) in the Zoological Museum, Berlin. Mitteilungen des Zoologischen Museum Berlin, 69(1), 45-56.", "Gray, J. (1845) Catalogue of the specimens of lizards in the collection of the British Museum. British Museum, London, 289 pp.", "Guibe, J. (1954) Catalogue des types des lezards dans la collection du Museum National d'Histoire Naturelle, Paris. Colas, Bayeux, Paris.", "Gunther, A. (1872) On two species of Hydrosaurus from the Philippine Islands. Proceedings of the Zoological Society London, 1872, 145-146.", "Hall, R. (2002) Cenozoic geological and plate tectonic evolution of SE Asia and the SW Pacific: computer-based reconstructions, model and animations. Journal of Asian Earth Sciences, 20, 353-431.", "Hammer, O., Harper, D.A.T. & Ryan, P.D. (2001) PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica, 4, 1-9.", "Heaney, L.R. (1985) Zoogeographic evidence for middle and late Pleistocene land bridges to the Philippine Islands. Modern Quaternary Research in Southeast Asia, 9, 127-165.", "Heaney, L.R. & Regalado Jr., J.C. (1998) Vanishing treasures of the Philippine rain forest. Field Museum, Chicago, 88 pp.", "Heaney, L.R., Balete, D.S., Dolar, M.L., Alcala, A.C., Dans, A.T.L., Gonzales, P.C. , Ingle, N.R., Lepiten, M.V., Oliver, W.L.R., Ong, P.S., Rickart, E.A., Tabaranza Jr., B.R. & Utzurrum, R.C.B. (1998) A synonpsis of the mammalian fauna of the Philippine Islands. Fieldiana: Zoology, 88, 1-61.", "Horn, H.-G. (1994) Der Ritualkampf von Varanus salvator cumingi. Herpetofauna, 16(92), 27-30.", "Horn, H.-G. & Visser, G.J. (1997) Review of reproduction of monitor lizards Va ra nu s spp. in captivity. II. International Zoo Yearbook, 35, 227-246.", "Horn, H.-G. & Gaulke M. (2004) Varanus salvator (Subspecies). In: Pianka, E.R. & King, D.R. (Eds), Varanoid lizards of the world. Bloomington. Indiana University Press, Indianapolis. pp. 258-271.", "Horn, H.-G, Gaulke, M. & Bohme, W. (1994) New data on ritualized combats in monitor lizards (Sauria: Varanidae) with remarks on their function and phylogenetic implications. Zoologischer Garten N. F., 64(5), 265-280.", "Huxley, T.H. (1868) On the classification and the distribution of the Alectoromorphae and Heteromorphae. Proceedings of the Zoological Society of London, 6, 249-319.", "Inger, R.F. (1954) Systematics and zoogeography of Philippine amphibia. Fieldiana: Zoology, 33, 181-531.", "Jolliffe, I.T. (1986) Principal component analysis. Springer, Berlin, Germany.", "Jones, A.W. & Kennedy, R.S. (2008) Evolution in a tropical archipelago: comparative phylogeography of Philippine fauna and flora reveals complex patterns of colonization and diversification. Biological Journal of the Linnean Society, 95, 620-639.", "Khan, M. (1969) A preliminary study of the water monitor, Varanus salvator. Malay Naturalist Journal, 22(2), 64-68.", "King, M., & King, D. (1975) Chromosomal evolution in the lizard genus Varanus (Reptilia). Australian Journal of Biological Sciences, 28, 89-108.", "Koch, A., Auliya, M., Schmitz, A., Kuch, U. & Bohme, W. (2007) Morphological studies on the systematics of South East Asian water monitors (Varanus salvator Complex): nominotypic populations and taxonomic overview. In: Horn, H-G, Bohme, W. & Krebs, U. (Eds), Advances in Monitor Research III. Mertensiella, 16, pp. 109-180.", "Lambeck, K., Esat, T.M., Potter, E.-K. (2002) Links between climate and sea levels for the past three million years. Nature, 419, 199-206.", "Leviton, A.E., McDiarmid, R., Moody, S., Nickerson, M., Rosado, J., Sokol, O. & Voris, H. (1980) Museum acronyms - second edition. Herpetological Review, 11(4), 93-102.", "Luxmoore, R. & Groombridge, B. (1989) Asian monitor lizards: a review of distribution, status, exploitation and trade in four selected species. Report to the CITES Secretariat, World Conservation Monitoring Centre, Cambridge.", "Mallet, J. (2005) Hybridization as an invasion of the genome. Trends in Ecology and Evolution, 20, 229-237.", "Mallet, J. (2007) Hybrid speciation. Nature, 446, 279-283.", "Martin, W. (1838) On two specimens of saurian reptiles sent to the society by Mr. Cuming. Proceedings of the Zoological Society of London, 1838, 68-70.", "Mavarez, J. & Linares, M. (2008) Homoploid hybrid speciation in animals. Molecular Ecology, 17, 4181-4185.", "Mayhew, P.J., Jenkins, G.B. & Benton, T.G. (2008) A long-term association between global temperature and biodiversity, origination and extinction in the fossil record. Proceedings of the Royal Society of London B, 275, 47-53.", "Mayr, E. (2000) The biological species concept. In: Wheeler Q.D. & Meier, R. (Eds), Species concepts and phylogenetic theory. A debate. Columbia University Press, New York. pp. 17-29.", "Mayr, E. & Ashlock, P.D. (1991) Principles of systematic zoology. Second edition. McGraw-Hill Book & Co., Singapore, 475 pp.", "McGuire, J.A. & Kiew, B.H. (2001) Phylogenetic systematics of Southeast Asian flying lizards (Iguania: Agamidae: Draco) as inferred from mitochondrial DNA sequence data. Biological Journal of the Linnean Society, 72, 203-229.", "McGuire, J.A., Brown, R.M., Mumpuni, Riyanto, A. & Andayani, N. (2007) The flying lizards of the Draco lineatus group (Squamata: Iguania: Agamidae): a taxonomic revision with description of two new species. Herpetological Monographs, 21, 179-212.", "Mertens, R. (1942) Die Familie der Warane (Varanidae). Abhandlungen der Senckenbergischen naturforschenden Gesellschaft, 462, 465, 466, 1-391.", "Mertens, R. (1959) Liste der Warane Asiens und der Indoaustralischen Inselwelt, mit systematischen Bemerkungen. Senckenbergiana Biologica, 40(5/6), 221-240.", "Mertens, R. (1963) Liste der rezenten Amphibien und Reptilien: Helodermatidae, Varanidae, Lanthanotidae. In: Mertens, R. & Hennig, W. (Eds), Das Tierreich. Walter de Gruyter & Co., Berlin. pp. i-x, 26.", "Mittermeier, R.A., Gil, P.R., Hoffmann, M., Pilgrim, J., Brooks, T., Goettsch, Mittermeier, C.G., Lamoreux, J. & da Fonseca, G.A.B. (2004) Hotspots revisited. CEMEX, Mexico City, 390 pp.", "Moschny, D. (2009) Haltung & Zucht des Mindanao-Bindenwarans. Varanus, 1(1), 8-20.", "Mulcahy, D.G. (2008) Phylogeography and species boundaries of the western North American Nightsnake (Hypsiglena torquata): revisiting the subspecies concept. Molecular Phylogenetics and Evolution, 46, 1095-1115.", "Myers, N., Mittermeier, R.A., Mittermeier, C.G., da Fonseca, G.A.B. & Kent, J. (2000) Biodiversity hotspots for conservation priorities. Nature, 403, 853-858.", "Pearson, K. (1901) On lines and planes of closest fit to systems of points in space. Philosophy Magazine, 2, 59-572.", "Peterson, A.T., Ball, L.G. & Brady, K.W. (2000) Distribution of the birds of the Philippines: biogeography and conservation priorities. Bird Conservation International, 10, 149-167.", "Petit, R.J., Jouzel, J., Raynaud, D., Barkov, N.I., Barnola, J.-M., Basile, I., Bender, M., Chappellaz, J., Davis, M., Delaygue, G., Delmotte, M., Kotlyakov, V.M., Lipenkov, V.Y., Lorius, C., Pepin, L., Ritz, C., Saltzman, E. & Stievenard, M. (1999) Climate and atmospheric history of the past 420,000 years from the Vostok ice core, Antarctica. Nature, 399, 429-436.", "Pianka, E.R. & King, D.R. (2004) Varanoid lizards of the world. Indiana University Press, Bloomington, Indianapolis, 588 pp.", "Piper, P.J., Ochoa, J., Lewis, H., Paz, V. & Ronquillo, W.P. (2008) The first evidence for the past presence of the tiger Panthera tigris (L.) on the island of Palawan, Philippines: extinction in an island population. Palaeogeography, Palaeoclimatology, Palaeoecology, 264, 123-127.", "Rabineau, M., Berne, S., Aslanian, J.-L.O.D., Guillocheau, F. & Joseph, P. (2006) Paleo sea levels reconsidered from direct observation of paleoshoreline position during Glacial Maxima (for the last 500,000 yr). Earth and Planetary Science Letters, 252, 119-137.", "Rawlinson, P.A., Widjoya, A.H.T., Hutchinson, M.N., Brown, G.W. (1990) The terrestrial vertebrates of Krakatau Islands, Sunda Strait, 1883-1986. Philosophical Transaction Royal Society London B, 328, 2-28.", "Reis, K.R. & Garong, A.M. (2001) Late Quaternary terrestrial vertebrates from Palawan Island, Philippines. Palaeogeography, Palaeoclimatology, Palaeoecology, 171, 409-421.", "Ross, C.A. & Gonzales, P.C. (1992) Amphibians and reptiles of Catanduanes Island, Philippines. National Museum Papers, Manila, 2(2), 50-76.", "Shi, G.R. (1993) Multivariate data analysis in palaeontology and palaeobiogeography - a review. Palaeogeography, Palaeoclimatology, Palaeoecology, 105, 199-234.", "Shine, R., Harlow, P.S., Keogh, J.S. & Boeadi (1996) Commercial harvesting of giant lizards: the biology of water monitors Varanus salvator in southern Sumatra. Biological Conservation, 77, 125-134.", "Sy, E., Diesmos, A., Jakosalem, P.G., Gonzalez, J.C., Paguntalan, L.M., Demegillo, A., Custodio, C., Delima, E., Tampos, G., Gaulke, M. & Jose, R. (2007) Varanus cumingi. In: IUCN (2009). IUCN Red List of Threatened Species. Version 2009.2. www.iucnredlist.org, downloaded on 17 February 2010.", "Taylor, E.H. (1918) Reptiles of the Sulu Archipelago. Philippine Journal of Science, 13(5), 233-265.", "Taylor, E.H. (1922) The lizards of the Philippine Islands. Philippine Journal of Science, 17, 1-269.", "van Rooijen, J. & Vogel, G. (2008) Contributions to a review of the Dendrelaphis pictus complex (Serpentes: Colubridae) - 1. Description of a sympatric species. Amphibia-Reptilia, 29, 101-115.", "Villamor, C.I. (1993) Morphometry and conservation status of water monitor lizard (Varanus salvator) in the Philippines. Asia Life Science, 2, 113-120.", "Vogel, D. (1994) Erstmalige Nachzucht des Mindanao Bindenwarans (Varanus [Varanus] salvator cumingi) im Zoologischen Garten Frankfurt. Monitor, 3, 41.", "Vogel, G. & van Rooijen, J. (2008) Contributions to a review of the Dendrelaphis pictus (Gmelin, 1789) complex - 2. the eastern forms (Serpentes: Colubridae). Herpe