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Australian Journal of Zoology Australian Journal of Zoology Society
Evolutionary, molecular and comparative zoology
RESEARCH ARTICLE

Refining the polytypic species concept of mangrove monitors (Squamata: Varanus indicus group): a new cryptic species from the Talaud Islands, Indonesia, reveals the underestimated diversity of Indo-Australian monitor lizards

André Koch A E , Evy Arida B , Andreas Schmitz C , Wolfgang Böhme A and Thomas Ziegler D
+ Author Affiliations
- Author Affiliations

A Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany.

B Museum Zoologicum Bogoriense, Jl. Raya Bogor km 46, 16911 Cibinong, Indonesia. Current address: Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany.

C Muséum d’histoire naturelle, C.P. 6434, CH-1211 Geneva 6, Switzerland.

D AG Zoologischer Garten Köln, Riehler Straße 173, D-50735 Köln, Germany.

E Corresponding author. Email: a.koch.zfmk@uni-bonn.de

Australian Journal of Zoology 57(1) 29-40 https://doi.org/10.1071/ZO08072
Submitted: 3 September 2008  Accepted: 16 March 2009   Published: 6 May 2009

Abstract

The description of a new cryptic member of the mangrove monitor (Varanus indicus) (Daudin, 1802) group from Indonesia presents a refinement of the systematic concept for a polytypic complex of closely related species. The new species is recognised on the basis of morphological and molecular evidence. It is so far known only from the type locality, the remote Talaud Islands, located between Sulawesi, Halmahera and Mindanao at the outer boundary of the Wallacea transition zone between the Oriental and Australian faunal regions. While the descriptions of several sibling and mostly sympatric species within the last 15 years have not affected the range of V. indicus, here, we demonstrate that morphologically distinct island populations represent independent evolutionary lineages and warrant specific recognition as distinct species within an evolving superspecies complex. In addition, some basic natural history information is provided for the new species based on observations at the type locality. These systematic and zoogeographic investigations reveal shortcomings in the current knowledge concerning the diversity and endemism of Indo-Australian monitor lizards, particularly within the Wallacean region.


Acknowledgements

The authors are grateful to the Indonesian Institute of Sciences (LIPI) for research permit (2767/SU.3/KS/2005 to AK) in support of their biodiversity sampling on Sulawesi and adjacent islands. The generous support of Pak Dedy Darnaedy (Research Center for Biology, LIPI, Indonesia), and Pak Mulyadi (MZB, Indonesia) is acknowledged also. Further thanks are due to Mumpuni (MZB) for allowing AK to examine monitor specimens of the MZB collections and to Henry (Salibabu Island) and to Sahid Kampi and his family (Sangihe Island) for hospitality and manifold support. The authors also thank Ronald Vonk (ZMA) for the loan of material under his care and for the exchange of one paratype of the newly described monitor species for deposition in the ZFMK collection. Bernhard Huber (ZFMK), Peter Jäger (Naturmuseum Senckenberg, Frankfurt, Germany) and Siegfried Ingrisch (Bad Karlshafen, Germany) identified the arthropod remains. Kai Philipp (Baden-Baden, Germany) and local staff of Conservation International Indonesia (Sangihe Island) kindly provided photographs of Varanus lirungensis. Finally, an earlier version of the manuscript benefited from critical comments by Robert Neal (Brisbane, Australia), Johan van Rooijen (Voorburg, The Netherlands), and two anonymous reviewers. Field work in Sulawesi and the Talaud Islands was made possible by a Ph.D. grant (AK) from the Evangelisches Studienwerk e.V. Villigst. This is publication No. 9 of the cooperation project about the herpetofaunal diversity of Sulawesi and adjacent islands between the herpetological sections of the Zoologisches Forschungsmuseum Alexander Koenig Bonn (ZFMK), Germany, and the Museum Zoologicum Bogoriense (MZB), Indonesia.


References

Ast, J. C. (2001). Mitochondrial DNA evidence and evolution in Varanoidea (Squamata). Cladistics 17, 211–226.
Crossref | GoogleScholarGoogle Scholar | Brandenburg T. (1983). Monitors of the Indo-Australian Archipelago. M.Sc. Thesis, University of Leyden, The Netherlands.

Brown, W. C. (1956). The distribution of terrestrial reptiles in the islands of the Pacific Basin. Proceedings of the 8th Pacific Science Congress 3A, 1479–1491.
De Queiroz K. (1998). The general lineage concept of species, species criteria, and the process of speciation: a conceptual unification and terminological recommendations. In ‘Endless Forms: Species and Speciation’. (Eds D. J. Howard and S. H. Berlocher.) pp. 57–75. (Oxford University Press: Oxford.)

De Queiroz, K. (2005). A unified concept of species and its consequences for the future of taxonomy. Proceedings of the California Academy of Sciences 56(Suppl. 1), 196–215.
De Rooij N. (1915). ‘The Reptiles of the Indo-Australian Archipelago.’ (E. J. Brill Ltd.: Leiden.)

Devitt, T. J. , LaDuc, T. J. , and McGuire, J. A. (2008). The Trimorphodon biscutatus (Squamata: Colubridae) species complex revisited: a multivariate statistical analysis of geographic variation. Copeia 2008, 370–387.
Crossref | GoogleScholarGoogle Scholar | Dryden G. , and Ziegler T. (2004). Varanus indicus. In ‘Varanoid Lizards of the World’. (Eds E. R. Pianka and D. R. King.) pp. 184–188. (Indiana University Press: Bloomington, IN.)

Eidenmüller B. , and Philippen H.-D. (2008). ‘Varanoid Lizards – Warane und Krustenechsen. Terralog 6.’ (Chimaira: Frankfurt a. M.)

Hammer, Ø. , Harper, D. A. T. , and Ryan, P. D. (2001). PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica 4, 9.
International Union for Conservation of Nature (2001). ‘IUCN Red List Categories and Criteria, Version 3.1.’ (IUCN Species Survival Commission: Gland, Switzerland & Cambridge, UK.)

Jennings W. B. , and Pianka E. R. (2004). The tempo and timing of the Australian Varanus radiation. In ‘Varanoid Lizards of the World’. (Eds E. R. Pianka and D. R. King.) pp. 77–87. (Indiana University Press: Bloomington, IN.)

Koch, A. , Auliya, M. , Schmitz, A. , Kuch, U. , and Böhme, W. (2007). Morphological studies on the systematics of south east Asian water monitors (Varanus salvator ssp. Complex): nominotypic populations and taxonomic overview. Mertensiella 16, 109–180.
Lever C. (2003). ‘Naturalized Reptiles and Amphibians of the World.’ (Oxford University Press: Oxford.)

Mayr E. (1942). ‘Systematics and the Origin of Species.’ (Columbia University Press: New York.)

McCoid, M. J. (1993). Reproductive output in captive and wild mangrove monitors (Varanus indicus). Varanews 3, 4–5.
McGarigal K. , Cushman S. , and Stafford S. G. (2000). ‘Multivariate Statistics for Wildlife and Ecology Research.’ (Springer: New York.)

McGuire, J. A. , Brown, R. M. , Mumpuni, , Riyanto, A. , and Anayani, N. (2007). The flying lizards of the Draco lineatus group (Squamata: Iguana: Agamidae): a taxonomic revision with descriptions of two new species. Herpetological Monographs 21, 179–212.
Crossref | GoogleScholarGoogle Scholar |

Mertens, R. (1926). Über die Rassen einiger indo-australischer Reptilien. Senckenbergiana 8, 272–279.


Mertens, R. (1942a). Die Familie der Warane. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 462, 1–116.


Mertens, R. (1942b). Die Familie der Warane. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 465, 117–234.


Mertens, R. (1942c). Die Familie der Warane. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 466, 235–391.


Mertens, R. (1963). Liste der rezenten Amphibien und Reptilien: Helodermatidae, Varanidae, Lanthanotidae. Das Tierreich 79, 1–26.


Philipp, K. (1999). Niche partitioning of Varanus doreanus, V. indicus and V. jobiensis in Irian Jaya: preliminary results. Mertensiella 11, 307–316.


Philipp, K. , Böhme, W. , and Ziegler, T. (1999). The identity of Varanus indicus: redefinition and description of a sibling species coexisting at the type locality. Spixiana 22, 273–287.


Rastegar-Pouyani, N. (2005). A multivariate analysis of geographic variation in the Trapelus agilis complex (Sauria: Agamidae). Amphibia-Reptilia 26, 159–173.
Crossref | GoogleScholarGoogle Scholar |

Van Rooijen, J. , and Vogel, G. (2008). An investigation into the taxonomy of Dendrelaphis tristis (Daudin, 1803): revalidation of Dipsas schokari (Kuhl, 1820) (Serpentes: Colubridae). Contributions to Zoology (Amsterdam, Netherlands) 77, 33–43.


Van Steenis-Kruseman, M. J. (1950). Malaysian plant collectors and collections. Flora Malesiana 1, 1–639.


Vogel, G. , and van Rooijen, J. (2008). Contributions to a review of the Dendrelaphis pictus (Gmelin, 1789) complex – 2. The eastern forms. (Serpentes: Colubridae). Herpetozoa (Wien) 21, 3–29.


Wikramanayake, E. D. , and Dryden, G. L. (1988). The reproductive ecology of Varanus indicus on Guam. Herpetologica 44, 338–344.


Ziegler, T. , and Böhme, W. (1997). Genitalstrukturen und Paarungsbiologie bei squamaten Reptilien, speziell den Platynota, mit Bemerkungen zur Systematik. Mertensiella 8, 3–207.


Ziegler, T. , and Böhme, W. (1999). Genital morphology and systematics of two recently described monitor lizards of the Varanus (Euprepiosaurus) indicus group. Mertensiella 11, 121–128.


Ziegler, T. , Böhme, W. , and 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 (Wien) 12, 45–56.


Ziegler, T. , Philipp, K. M. , and Böhme, W. (1999b). Zum Artstatus und zur Genitalmorphologie von Varanus finschi Böhme, Horn & Ziegler, 1994, mit neuen Verbreitungsangaben für V. finschi und V. doreanus (Meyer, 1874) (Reptilia: Squamata: Varanidae). Zoologische Abhandlungen. Staatliches Museum für Tierkunde 50, 267–279.


Ziegler, T. , Böhme, W. , Eidenmüller, B. , and 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 Beitrage 50, 27–30.


Ziegler, T. , Böhme, W. , and Schmitz, A. (2007a). A new species of the Varanus indicus group (Squamata, Varanidae) from Halmahera Island, Moluccas: morphological and molecular evidence. Mitteilungen des Museum für Naturkunde Berlin. Zoologische Reihe 83(Suppl.), 109–119.
Crossref | GoogleScholarGoogle Scholar |

Ziegler, T. , Schmitz, A. , Koch, A. , and Böhme, W. (2007b). A review of the subgenus Euprepiosaurus of Varanus (Squamata: Varanidae): morphological and molecular phylogeny, distribution and zoogeography, with an identification key for the members of the V. indicus and the V. prasinus species groups. Zootaxa 1472, 1–28.





Appendix 1.  Additional specimens of the Varanus indicus species complex included in this study
Specimens that have been examined for the first time are indicated by an asterisk
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