Abstract
We provide a synopsis of the currently recognized taxa within the subgenus Euprepiosaurus of Varanus, consisting of the V. indicus species group (in chronological order, V. indicus, V. doreanus, V. jobiensis, V. finschi, V. melinus, V. yuwonoi, V. caerulivirens, V. cerambonensis, V. juxtindicus, V. zugorum) and the V. prasinus species group (accordingly, V. prasinus, V. beccarii, V. kordensis, V. bogerti, V. keithhornei, V. telenesetes, V. macraei, V. boehmei, V. reisingeri). We summarize the taxonomic history of the species groups and highlight the morphology and distribution of the species in detail. Molecular genetic analyses confirm Euprepiosaurus and also the two contained species groups as monophyla. Our molecular (mitochondrial 16S rRNA gene) data further reinforce that V. beccarii, V. boehmei, V. keithhornei, and V. macraei are distinct species within the V. prasinus group. V. kordensis consistently proves to be a sister species to all remaining members of the V. prasinus species group studied by us. Comparatively low genetic distances argue for relatively recent speciation processes within the V. indicus group. The species status of V. caerulivirens and V. finschi is again corroborated. The analyses consistently place V. cerambonensis and V. melinus as sister species. It is further evident that both species groups within Euprepiosaurus still contain distinct unrecognized taxa. Finally, we discuss the phylogeny and zoogeography of Euprepiosaurus in the light of our data and provide an identification key for the species of this subgenus.References
Ahl, E. (1932) Eine neue Eidechse und zwei neue Frösche von der Insel Jobi. Mitt. Zool. Mus. Berlin, 17, 892–894.
Allison, A. (1996) Zoogeography of amphibians and reptiles of New Guinea and the Pacific region. In: Keast, A. & Miller, S.E. (Ed), The Origin and Evolution of Pacific island biotas, New Guinea to eastern Polynesia: patterns and processes. SPB Academic Publishing, Amstersam, 407–436.
Ast, J.C. (2001) Mitochondrial DNA evidence and evolution in Varanoidea (Squamata). Cladistics, 17, 211–226.
Balouet, J.C. (1991) The fossil vertebrate record of New Caledonia. In: Rich, P.-V., Monaghan, J.M., Baird, R.F., Rich, T.H., Thompson, E.M. & Williams, C. (Ed), Vertebrate palaeontology of Australia. Melbourne, Pioneer Design Studio and Monash University Publications Committee, 1383–1409.
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. Austral. J. Zool., 41, 621–638.
Böhme, W. (1988) Zur Genitalmorphologie der Sauria: funktionelle und stammesgeschichtliche Aspekte. Bonn. zool. Monogr., 27, 1–176.
Böhme, W. (1991) New findings on the hemipenial morphology of monitor lizards and their systematic implications. In: Böhme, W. & Horn, H.-G. (Ed), Advances in monitor research. Mertensiella, 2, 42–49.
Böhme, W. (1997) Robert Mertens’ Systematik und Klassifikation der Warane: Aktualisierung seiner 1942er Monographie und eine revidierte Checkliste. Anhang zum Nachdruck von Mertens, R., 1942, Die Familie der Warane (Varanidae). Erster bis dritter Teil. Abh. senckenberg. naturf. Ges., Frankfurt/M. 462, 465, 466, 1–391. Edition Chimaira, Frankfurt am Main, I–XXIII pp.
Böhme, W. (2003) Checklist of the living monitor lizards of the world (family Varanidae). Zool. Verh., Leiden 341, 3–43.
Böhme, 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.
Böhme, W. & Jacobs, H.J. (2001) Varanus macraei sp. n., eine neue Waranart der V. prasinus-Gruppe aus West Irian, Indonesien. Herpetofauna, 23, 5–10.
Böhme, W., Philipp, K. & Ziegler, T. (2002) Another new member of the Varanus (Euprepiosaurus) indicus group (Sauria, Varanidae): an undescribed species from Rennell Island, Solomon Islands. Salamandra, 38, 15–26.
Böhme, W. & Ziegler, T. (1997) Varanus melinus sp. n., ein neuer Waran aus der V. indicus-Gruppe von den Molukken, Indonesien. Herpetofauna, 19, 26–34.
Böhme, W. & Ziegler, T. (2005) A new monitor lizard from Halmahera, Moluccas, Indonesia (Reptilia: Squamata: Varanidae). Salamandra, 41, 51–59.
Böhme, W. & Ziegler, T. (accepted) Notes on the distribution, diet, hemipenis morphology and systematics of Varanus spinulosus Mertens, 1941. In: Horn, H.-G., Böhme, W., Krebs, U. (Ed), Advances in Monitor Research III. Mertensiella, Rheinbach.
Boulenger, G.A. (1885) Catalogue of the lizards in the British Museum (Natural History). II. Iguanidae, Xenosauridae, Zonuridae, Anguidae, Anniellidae, Helodermatidae, Varanidae, Xantusidae, Teiidae, Amphisbaenidae. London, British Museum.
Branch, W.R. (1982) Hemipenial morphology of platynotan lizards. J. Herpetol., 16, 16–38.
Brandenburg, T. (1983) Monitors in the Indo-Australian archipelago. Univ. Leiden (Unpubl. thesis).
Card, W. & Kluge, A.G. (1995) Hemipeneal skeleton and varanid lizard systematics. J. Herpetol., 29, 275–280.
Daudin, F.M. (1802) Histoire naturelle générale et particulière des reptiles. III. Paris.
Diamond, J.M. (1987) Did Komodo dragons evolve to eat pygmy elephants? Nature, 326, 832.
Doria, G. (1874) Enumerazione dei rettili raccolti dal Dott. O. Beccari in Amboina, alle isole Aru ed alle isole Kei durante gli anni 1872–73. Ann. Mus. civ. Stor. nat. Genova, 6, 325–357.
Dryden, G. (1965) The food and feeding of Varanus indicus on Guam. Micronesia, 2, 72–76.
Dryden, G. & Ziegler, T. (2004) Varanus indicus. In: Pianka, E.R., King, D.R., King, R. A. (Ed), Varanoid lizards of the world. Indiana University Press, Bloomington, 184–188.
Eidenmüller, B. & Wicker, R. (2005) Eine weitere neue Waranart aus dem Varanus prasinus-Komplex von der Insel Misol, Indonesien. Sauria, 27, 3–8.
Fairbanks, R.G. (1989) A 17000-year glacio-eustatic sea level record: influence of glacial melting rates on the Younger Dryas event and deep ocean circulaton. Nature, 342, 637–642.
Fitch, A.J., Goodman, A.E. & Donnellan, S.C. (2006) A molecular phylogeny of the Australian monitor lizards (Squamata: Varanidae) inferred from mitochondrial DNA sequences. Austral. J. Zool., 54, 253–269.
Fuller, S., Baverstock, P. & King, D. (1998) Biogeographic origin of goannas (Varanidae): a molecular perspective. Mol. Phylogeny & Evol., 9, 294–307.
Godthelp, H. (2001) The Australian rodent fauna, flotilla’s, flotsam or just fleet footed? In: Metcalfe, I., Smith, J.M.B., Morwood, M. & Davidson, I. (Ed), Faunal and Floral Migration and Evolution in SE Asia-Australasia. Balkema Publishers, Lisse, 319–322.
Greene, H.W. (1986) Diet and Arboreality in the Emerald Monitor, Varanus prasinus, with Comments on the Study of Adaptation. Fieldiana Zoology, 31, 1–12.
Greene, H.W. (2004) Varanus prasinus. In: Pianka, E.R., King, D.R., King, R. A. (Ed), Varanoid lizards of the world. Indiana University Press, Bloomington, 225–229.
Hall, R. (2001) Cenozoic reconstruction of SE Asia and the SW Pacific: changing patterns of land and sea. In: Metcalfe, I., Smith, J.M.B., Morwood, M. & Davidson, I. (Ed), Faunal and Floral Migration and Evolution in SE Asia-Australasia. Balkema Publishers, Lisse, 35–56.
Hall, R. (2002) Cenozoic geological and plate tectonic evolution of SE Asia and the SW Pacific: computer-based reconstructions, model and animations. J. Asian Earth Sciences, 20, 353–431.
Hall, T.A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98.
Harvey, M.B. & Barker, D.G. (1998) A new species of blue-tailed monitor lizard (genus Varanus) from Halmahera island, Indonesia. Herpetologica, 54, 34–44.
Heaney, L.R. (1991) A synopsis of climatic and vegetational change in Southeast Asia. Climatic Change, 19, 53–61.
Hooijer, D.A. (1975) Quaternary mammals west and east of Wallace's Line. Neth. J. Zool., 25, 46–56.
Hope, G. (1996) Quaternary change and the historical biogeography of Pacific islands. In: Keast, A. & Miller, S. E (Ed), The Origin and Evolution of Pacific island biotas, New Guinea to eastern Polynesia: patterns and processes. SPB Academic Publishing, Amstersam, 165–190.
How, R.A. & Kitchener, D.J. (1997) Biogeography of Indonesian snakes. J. Biogeography, 24, 725–735.
Huelsenbeck, J.P. & Ronquist, F. 2001. MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics 17: 754–755.
Irwin, S. (2004) Varanus keithhornei. In: Pianka, E.R., King, D.R., King, R. A. (Ed), Varanoid lizards of the world. Indiana University Press, Bloomington, 401–405.
Iskandar, D.T. & Tjan, K.N. (1996) The Amphibians and Reptiles of Sulawesi, with notes on the Distribution and Chromosomal Number of Frogs. In: Kitchener, D.J. & Suyanto, A. (Ed), First international conference on eastern Indonesian-Australian vertebrate fauna. Western Australian Museum, 39–46.
Jacobs, H.J. (2002) Zur morphologischen Variabilität der nominellen Smaragdwaran-Taxa Varanus prasinus (H. Schlegel, 1839) und V. kordensis (A. B. Meyer, 1874), mit Bemerkungen zur Erstzucht des letzteren. Herpetofauna, 24, 21–34.
Jacobs, H.J. (2003) A further new emerald tree monitor lizard of the Varanus prasinus species group from Waigeo, West Irian (Squamata: Sauria: Varanidae). Salamandra, 39, 65–74.
Jacobs, H.J. (2004) Varanus kordensis. In: Pianka, E.R., King, D.R., King, R.A. (Ed), Varanoid lizards of the world. Indiana University Press, Bloomington, 205–207.
Koch, A. & Böhme, W. (2005) Die Herpetofauna Sulawesis unter besonderer Berücksichtigung der Gattung Varanus: phylogeographische Beziehungen zu angrenzenden Gebieten. - Das Projekt stellt sich vor. Elaphe, 13, 43–46.
Kuch, U., Keogh, J.S., Weigel, J., Smith, L.A. & Mebs, D. (2005) Phylogeography of Australia’s king brown snake (Pseudechis australis) reveals Pleistocene divergence and Pleistocene dispersal of a top predator. Naturwissenschaften, 92, 121–127.
McGuigan, K., Zhu, D., Allen, G.R. & Moritz, C. (2000) phylogenetic relationships and historical biogeography of melanotaeiid fishes in Australia and New Guinea. Mar. Freshwater Res., 51, 713–723.
Mertens, R. (1934) Die Inselreptilien. Ihre Ausbreitung, Variation und Artbildung. - E. Schweizerbart'sche Verlagsbuchhandlung, Stuttgart, 209 pp.
Mertens, R. (1941) Zwei neue Warane des australischen Faunengebietes. Senckenbergiana, Frankfurt/Main, 23, 266–272.
Mertens, R. (1942) Die Familie der Warane (Varanidae). Dritter Teil: Taxonomie. Abh. senckenberg. naturf. Ges., Frankfurt a. M., 466, 235–391.
Mertens, R. (1950) Notes on some Indo-Australian monitors (Sauria: Varanidae). Amer. Mus. Nov., 1456, 1–7.
Mertens, R. (1951) A new lizard of the genus Varanus from New Guinea. Fieldiana Zool., Chicago, 31, 467–471.
Meyer, A.B. (1874) Übersicht über die von mir auf Neu-Guinea und den Inseln Jobi, Mysore und Mafoor im Jahre 1873 gesammelten Amphibien. MBer. K. Akad. Wiss., Berlin, 1874, 128–138.
Molnar, R.E. (2004) The long and honorable History of Monitors and their kin. In: Pianka, E.R., King, D.R. & King, R.A. (Ed), Varanoid lizards of the World. Indiana University Press, Bloomington, 10–67.
Molnar, R.E. & Pianka, E.R. (2004) Biogeography and Phylogeny of Varanoids. In: Pianka, E.R., King, D.R. & King, R.A. (Ed), Varanoid lizards of the World. Indiana University Press, Bloomington, 68–76.
Nylander, J.A.A. (2005) MrModeltest 1.1b, Uppsala, Sweden.
Palumbi, S.R., Martin, A., Romano, S., McMillan, W.O., Stice, L. & Grabowski, G. (1991) The simple fool's guide to PCR. Department of Zoology and Kewalo Marine Laboratory, Hawai: 47 pp.
Philipp, K.M. (1999) Niche partitioning of Varanus doreanus, V. indicus and V. jobiensis in Irian Jaya: preliminary results. In: Horn, H.-G. & Böhme, W. (Ed), Advances in monitor research II. Mertensiella, 11, 307–316.
Philipp, K.M., Böhme, W. & Ziegler, T. (1999) The identity of Varanus indicus: Redefinition and description of a sibling species coexisting at the type locality (Sauria: Varanidae: Varanus indicus group). Spixiana, 22, 273–287.
Philipp, K., Ziegler, T. & Böhme, W. (2004a) Varanus cerambonensis. In: Pianka, E.R., King, D.R., King, R. A. (Ed), Varanoid lizards of the world. Indiana University Press, Bloomington, 165–167.
Philipp, K., Ziegler, T. & Böhme, W. (2004b) Varanus jobiensis. In: Pianka, E.R., King, D.R., King, R. A. (Ed), Varanoid lizards of the world. Indiana University Press, Bloomington, 189–192.
Philipp, K., Ziegler, T. & Böhme, W. (2004c) Varanus finschi. In: Pianka, E.R., King, D.R., King, R. A. (Ed), Varanoid lizards of the world. Indiana University Press, Bloomington, 176–178.
Philipp, K.M., Ziegler, T. & Böhme, W. (accepted) Preliminary investigations of the natural diet of six monitor lizard species of the Varanus (Euprepiosaurus) indicus group. In: Horn, H.-G., Böhme, W. & Krebs, U. (Ed), Advances in monitor research III. Mertensiella, Rheinbach.
Posada, D. & Crandall, K.A. (1998) Modeltest: testing the model of DNA substitution. Bioinformatics, 14, 817–818.
Rawlings, L. H., Barker, D. & S. C. Donnellan (2004) Phylogenetic relationships of the Australo-Papuan Liasis pythons (Reptilia: Macrostomata), based on mitochondrial DNA. Austr. J. Zool. 52, 215–227.
Rawlings, L.H. & Donnellan, S.C. (2003) Phylogeographic analysis of the green python, Morelia viridis, reveals cryptic diversity. Mol. Phyl. Evol., 27, 36–44.
Reeder, T.W. (2003) A phylogeny of the Australian Sphenomorphus group (Scincidae: Squamata) and the phylogenetic placement of the crocodile skinks (Tribolonotus): Bayesian approaches to assessing congruence and obtaining confidence in maximum likelihood inferred relationships. Mol. Phyl. Evol., 27, 384–397.
Rodda, G.H., Fritts, T.H. & Reichel, J.D. (1991) The distributional patterns of reptiles and amphibians in the Mariana Islands. Micronesia, 24(2), 195–210.
Schlegel, H. (1839) Abbildungen neuer oder unvollständig bekannter Amphibien, nach der Natur oder dem Leben entworfen und mit einem erläuternden Texte begleitet. Düsseldorf.
Schmitz, A. (2003) Taxonomic and phylogenetic studies on scincid lizards (Reptilia : Scincidae). Unpubl. PhD thesis. University of Bonn, 262 pp.
Schmitz, A., Ineich, I. & Chirio, L. (2005) Molecular review of the genus Panaspis sensu lato in Cameroon, with special reference to the status of the proposed subgenera. Zootaxa, 863, 1–28.
Schulte II, J.A., Melville, J. & Larson, A. (2003) Molecular phylogenetic evidence of ancient divergence of lizard taxa on either side of Wallace’s Line. Proc. R. Soc. Lond. B, 270, 597–603.
Sprackland, R.G. (1991) Taxonomic review of the Varanus prasinus group with description of two new species. Mem. Qld. Mus., 30, 561–576.
Sprackland, R.G. (1993) Rediscovery of a Solomon Islands monitor lizard (Varanus indicus spinulosus) Mertens, 1941. The Vivarium, Neptune City 4(5), 25–27.
Sprackland, R.G. (1994) Rediscovery and taxonomic review of Varanus indicus spinulosus Mertens, 1941. Herpetofauna 24(2), 33–39.
Sprackland, R.G. (1997) Mangrove monitor lizards. Reptiles, 5, 48–63.
Sweet, S.S. & Pianka, E.R. (in press.) Monitors, Mammals and Wallace’s Line. In: Horn, H.-G., Böhme, W. & Krebs, D. (Ed), Mertensiella, Rheinbach.
Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4.0b10. Sinauer Associates, Sunderland, Massachusetts, USA.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin F. & Higgins, D.G. (1997) The ClustalX windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 24, 4876–4882.
Uchida, T. (1967) Observations on the monitor lizard as a rat control agent on Ifaluk, Western Caroline Islands. Micronesia, 3(1), 17–18.
Voris, H.K. (2000) Maps of Pleistocene sea levels in Southeast Asia: shorelines, river systems and time durations. J. Biogeogr., 27, 1153–1167.
Walsh, P.S., Metzger, D.A. & Higuchi, R. (1991) Chelex 100 as a medium for simple extraction of DNA for PCR-based typing from forensic material. BioTechniques, 10(4), 506–513.
Wells, R.W. & Wellington, C.R. (1985) A classification of the Amphibia and Reptilia of Australia. Austr. J. Herpetol., Suppl. Ser., (1), 1–61.
Yang, Z.H., Goldman, N. & Friday, A. (1994) Comparison of Models for Nucleotide Substitution Used in Maximum-Likelihood Phylogenetic Estimation. Molecular Biology and Evolution, 11, 316–324.
Ziegler, T. & Böhme, W. (1997) Genitalstrukturen und Paarungsbiologie bei squamaten Reptilien, speziell den Platynota, mit Bemerkungen zur Systematik. Mertensiella, 8, 1–207.
Ziegler, T. & Böhme, W. (1998) Comment on the proposed conservation of the specific name of Varanus teriae Sprackland, 1991 (Reptilia, Squamata). Bulletin of Zoological Nomenclature, London, 55, 111–113.
Ziegler, T. & Böhme, W. (1999) Genital morphology and systematics of two recently described monitor lizards of the Varanus (Euprepiosaurus) indicus group. In: Horn, H.-G., Böhme, W. (Ed), Advances in monitor research II. Mertensiella, 11, Rheinbach, 121–128.
Ziegler, T. & Böhme, W. (2004) Varanus melinus. In: Pianka, E.R., King, D.R., King, R. A. (Ed), Varanoid lizards of the world. Indiana University Press, Bloomington, 215–219.
Ziegler, T., Böhme, W., Eidenmüller, B. & Philipp, K.M. (2001) A note on the coexistence of three species of Pacific monitor lizards in Australia (Sauria, Varanidae, Varanus indicus group). Bonner Zoologische Beiträge, Bonn, 50, 27–30.
Ziegler, T., Böhme, W. & Philipp, K.M. (1999a) Varanus caerulivirens sp. n., a new monitor lizard of the V. indicus group from Halmahera, Moluccas, Indonesia (Squamata: Sauria: Varanidae). Herpetozoa, 12, 45–56.
Ziegler, T., Böhme, W. & Schmitz, A.(in press) A new species of the Varanus indicus group (Squamata, Varanidae) from Halmahera Island, Moluccas: morphological and molecular evidence. Mitt. Mus. Nat.kd. Berlin, Zool. Reihe.
Ziegler, T., Philipp, K.M. & Böhme, W. (1999b) Zum Artstatus und zur Genitalmorphologie von Varanus finschi Böhme, Horn et Ziegler, 1994, mit neuen Verbreitungsangaben für V. finschi und V. doreanus (Meyer, 1874) (Reptilia: Squamata: Varanidae). Zoologische Abhandlungen, Staatliches Museum für Tierkunde, Dresden, 50, 267–279.