Organisms Diversity & Evolution

, Volume 13, Issue 1, pp 87–95 | Cite as

Molecules and morphology suggest cryptic species diversity and an overall complex taxonomy of fish scale geckos, genus Geckolepis

  • Inga Lemme
  • Martina Erbacher
  • Nathalie Kaffenberger
  • Miguel Vences
  • Jörn Köhler
Original Article


The current classification of the Malagasy-Comoroan geckos of the genus Geckolepis recognizes three valid species and is based on morphological differences only. Species allocation of individuals is difficult, mainly because of the insufficiently known degree of variation and the frequent partial loss of the integument in preserved specimens. Here we study Geckolepis specimens from almost their entire known range. We combine molecular and morphological data to estimate species richness in the genus, and to assess the taxonomic validity of the morphological differences proposed. Analyses of mitochondrial (12S, ND4) and nuclear (RAG1, CMOS) gene sequences support three major clades in addition to the morphologically distinct G. polylepis (included for 12S only), and the presence of various divergent lineages within these clades, some occurring in sympatry. Among lineages, the external morphology seems to be comparatively conservative, and differences are faint in many cases. For each major lineage, we summarize the most diagnostic morphological characters that distinguish it from other lineages. The combined molecular and morphological data strongly indicate the presence of highly divergent lineages, three of which are tentatively referred to the names Geckolepis maculata, G. polylepis and G. typica, whereas three other major lineages are considered Confirmed Candidate Species. Among the remaining lineages, differentiation is shallower, and these may qualify for Deep Conspecific Lineages. In conclusion, our analysis provides evidence for an underestimation of species richness in the genus and a complex evolutionary history, not reflected by current Geckolepis species-level taxonomy.


Squamata Gekkonidae Geckolepis Madagascar Molecular genetics Morphology Systematics Species diversity 



We are grateful to Franco Andreone, Parfait Bora, Neil D’Cruze, Hildegard Enting, Michael Franzen, Frank Glaw, Kathrin Glaw, Angelika Knoll, Aurelien Miralles, Zoltan Nagy, Jasmin E. Randrianirina, Fanomezana M. Ratsoavina, Angelin and Angeluc Razafimanantsoa, and Augustin Sarovy for help during fieldwork and/or collection of crucial samples. Wolfgang Böhme (ZFMK), Frank Glaw (ZSM), Gunther Köhler (SMF) and Mark-Oliver Rödel (ZMB) allowed examination of specimens held in their care. Special thanks to Frank Glaw for sharing data and photos. Fieldwork was carried out in the framework of collaboration accords with the Département de Biologie Animale, Université d'Antananarivo, Madagascar. We are grateful to the Malagasy authorities for collection and export permits.


  1. Angel, F. (1942). Les lézards de Madagascar. Memoires de l'Académie Malgache, 36, 1–139.Google Scholar
  2. Arévalo, E., Davis, S. K., & Sites, J. (1994). Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in Central Mexico. Systematic Biology, 43, 387–418.Google Scholar
  3. Avise, J. C., & Ball, R. M. (1990). Principles of genealogical concordance in species concepts and biological taxonomy. In D. Futuyma & J. Antonovics (Eds.), Surveys in Evolutionary Biology ((pp, Vol. 7, pp. 45–67). Oxford: Oxford University Press.Google Scholar
  4. Bauer, A., Glaw, F., Gehring, P.-S., & Vences, M. (2011). New species of Blaesodactylus (Squamata: Gekkonidae) from Ankarafantsika National Park in north-western Madagascar. Zootaxa, 2942, 57–68.Google Scholar
  5. Brygoo, E. R. (1987). L'endémisme des reptiles de Madagascar. Bulletin de la Societé Zoologique de France, 112, 5–38.Google Scholar
  6. Clement, D. M., Posada, D., & Crandall, K. A. (2000). TCS: a computer program to estimate gene genealogies. Molecular Ecology, 9, 1657–1659.PubMedCrossRefGoogle Scholar
  7. Crottini, A., Gehring, P.-S., Glaw, F., Harris, D. J., Lima, A., & Vences, M. (2011). Deciphering the cryptic species diversity of dull-coloured day geckos Phelsuma (Squamata: Gekkonidae) from Madagascar, with description of a new species. Zootaxa, 2982, 340–348.Google Scholar
  8. Glaw, F., & Vences, M. (1994). A Fieldguide to the Amphibians and Reptiles of Madagascar, 2nd ed. (pp. 480) Köln: Vences & Glaw Verlag.Google Scholar
  9. Glaw, F., & Vences, M. (2007). A Field Guide to the Amphibians and Reptiles of Madagascar, 3rd ed. (pp. 496). Köln: Vences & Glaw Verlag.Google Scholar
  10. Greenbaum, E., Bauer, A. M., Jackman, T. R., Vences, M., & Glaw, F. (2007). A phylogeny of the enigmatic Madagascan geckos of the genus Uroplatus (Squamata: Gekkonidae). Zootaxa, 1493, 41–51.Google Scholar
  11. Greenbaum, E., Jackman, T., & Bauer, A. M. (2007). Homopholis and Blaesodactylus (Squamata: Gekkonidae) revisited: new insights from a molecular phylogeny. African Journal of Herpetology, 56, 101–114.CrossRefGoogle Scholar
  12. Han, D., Zhou, K., & Bauer, A. M. (2004). Phylogenetic relationships among gekkotan lizards inferred from Cmos nuclear DNA sequences and a new classification of the Gekkota. Biological Journal of the Linnean Society, 83, 353–368.CrossRefGoogle Scholar
  13. Jackman, T. R., Bauer, A. M., Greenbaum, E., Glaw, F., & Vences, M. (2008). Molecular phylogenetic relationships among species of the Malagasy-Comoran gecko genus Paroedura (Squamata: Gekkonidae). Molecular Phylogenetics and Evolution, 46, 74–81.PubMedCrossRefGoogle Scholar
  14. Köhler, G., Diethert, H.-H., Nussbaum, R. A., & Raxworthy, C. J. (2009). A revision of the fish scale geckos, genus Geckolepis Grandidier (Squamata, Gekkonidae) from Madagascar and the Comoros. Herpetologica, 65, 419–435.CrossRefGoogle Scholar
  15. Librado, P., & Rozas, J. (2009). DnaSP v5: A software for comprehensive analysis of DNA polymorphism data. Bioinformatics, 25, 1451–1452.PubMedCrossRefGoogle Scholar
  16. Nagy, Z. T., Sonet, G., Glaw, F., & Vences, M. (2012). First large-scale DNA barcoding assessment of reptiles in the biodiversity hotspot of Madagascar, based on newly designed COI primers. PLoS ONE, 7, e34506.PubMedCrossRefGoogle Scholar
  17. Nylander, J. A. A. (2004). MrModeltest version 2.3. Evolutionary Biology Centre. Uppsala: Uppsala University.Google Scholar
  18. Padial, J. M., Castroviejo-Fisher, S., Köhler, J., Vilà, C., Chaparro, J. C., & De la Riva, I. (2009). Deciphering the products of evolution at the species level: the need for an integrative taxonomy. Zoologica Scripta, 38, 431–447.CrossRefGoogle Scholar
  19. Padial, J. M., Miralles, A., De la Riva, I., & Vences, M. (2010). The integrative future of taxonomy. Frontiers in Zoology, 7, article 16.Google Scholar
  20. Palumbi, S., Martin, A., Romano, S., McMillan, W. O., Stice, L., & Grabowski, G. (1991). The simple fool’s guide to PCR Version 2. Hawai: Honululu.Google Scholar
  21. Ronquist, F., & Huelsenbeck, J. P. (2003). MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.PubMedCrossRefGoogle Scholar
  22. Schubert, C., & Christophers, E. (1985). "Dermolytische Schreckhäutung", ein besonderes Autotomieverhalten von Geckolepis typica (Reptilia, Gekkonidae). Zoologischer Anzeiger, 214, 129–141.Google Scholar
  23. Schubert, C., Steffen, T., & Christophers, E. (1990). Weitere Beobachtungen zur "dermolytischen Schreckhäutung" bei Geckolepis typica (Reptilia, Gekkonidae). Zoologischer Anzeiger, 224, 175–192.Google Scholar
  24. Stephens, M., Smith, N. J., & Donnelly, P. (2001). A new statistical method for haplotype reconstruction from population data. American Journal of Human Genetics, 68, 978–989.PubMedCrossRefGoogle Scholar
  25. Templeton, A. R., Crandall, K. A., & Sing, C. F. (1992). A cladistic analysis of phenotypic associations with haplotypes inferred from restriction endonuclease mapping and DNA sequence data. Cladogram estimation. Genetics, 132, 619–633.PubMedGoogle Scholar
  26. Vieites, D. R., Wollenberg, K. C., Andreone, F., Köhler, J., Glaw, F., & Vences, M. (2009). Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Science of the U.S.A., 106, 8267–8272.CrossRefGoogle Scholar
  27. Wermuth, H. (1965). Liste der rezenten Amphibien und Reptilien. Gekkonidae, Pygopodidae, Xantusiidae. Das Tierreich, 80, 1–246.Google Scholar

Copyright information

© Gesellschaft für Biologische Systematik 2012

Authors and Affiliations

  • Inga Lemme
    • 1
    • 2
  • Martina Erbacher
    • 4
  • Nathalie Kaffenberger
    • 1
    • 3
    • 4
  • Miguel Vences
    • 1
  • Jörn Köhler
    • 4
  1. 1.Division of Evolutionary Biology, Zoological InstituteTechnical University of BraunschweigBraunschweigGermany
  2. 2.Alfred Wegener InstituteBremerhavenGermany
  3. 3.Senckenberg Natural History CollectionsDresdenGermany
  4. 4.Hessisches Landesmuseum DarmstadtDarmstadtGermany

Personalised recommendations