Abstract
We review the intrageneric systematics of day geckos, genus Phelsuma (Squamata: Sauria: Gekkonidae) in the light of a recently published multigene phylogeny, and provide an overview of 19 selected chromatic, morphological and behavioural characters for all currently recognized species and subspecies in the genus. To replace previous attempts of dividing the genus into phenetic species assemblages, we propose the following monophyletic species groups and summarize the morphological characters that partly support this grouping: Phelsuma barbouri group (P. barbouri, P. pronki); P. dubia group (P. berghofi, P. dubia, P. flavigularis, P. hielscheri, P. malamakibo, P. modesta, P. nigristriata, P. ravenala); P. mutabilis group (P. borai, P. breviceps, P. mutabilis); P. cepediana group (P. borbonica, P. cepediana, P. guentheri, P. guimbeaui, P. inexpectata, P. ornata, P. rosagularis as well as the extinct species P. edwardnewtonii and P. gigas); P. astriata group (P. astriata, P. sundbergi); P. madagascariensis group (P. abbotti, P. grandis, P. guttata, P. kochi, P. madagascariensis, P. parkeri, P. seippi and probably P. masohoala); P. lineata group (P. antanosy, P. comorensis, P. kely, P. lineata, P. pusilla, P. quadriocellata and probably P. hoeschi); P. laticauda group (P. laticauda, P. pasteuri, P. robertmertensi, P. v-nigra); P. klemmeri group (P. klemmeri); P. andamanense group (P. andamanense) and P. standingi group (P. standingi). Due to their equivocal placements based on different molecular markers, we refrain from assigning P. serraticauda and P. vanheygeni to any of these species groups at this time. We complement the molecular data with new trees based on partial sequences of the 16S rRNA gene for the P. madagascariensis- and P. lineata groups and propose to elevate P. lineata dorsivittata and P. quadriocellata parva to full species rank as P. dorsivittata and P. parva.References
Akaike, H. (1974) A new look at the statistical model identification. IEEE Transactions on automatic Control, 19, 716–723.
Ali, J.R. & Huber, M. (2010) Mammalian biodiversity on Madagascar controlled by ocean currents. Nature, 653–656.
Austin, J.J., Arnold, E.N. & Jones, C.G. (2004) Reconstructing an island radiation using ancient and recent DNA: the extinct and living day geckos (Phelsuma) of the Mascarene islands. Molecular Phylogenetics and Evolution, 31, 102–109.
Bauer, A.M. (2003) Gekkonidae, Geckos. Pp. 973–977 In: The Natural History of Madagascar. Goodman, S.M. and J.P. Benstead (Eds.). The University of Chicago Press, Chicago, Illinois, USA.
Berghof, H.P. & Trautmann, G. (2009) Eine neue Art der Gattung Phelsuma Gray, 1825 (Sauria: Gekkonidae) von der Ostküste Madagaskars. Sauria, 31, 5–14.
Blanc, C.P. (1972) Les Reptiles de Madagascar et des Iles Voisines. In: Battisttini, R. & Richard-Vindard, G. (Eds.), Biogeography and Ecology in Madagascar. W Junk, The Hague, pp. 501–614.
Böhme, W. & Meier, H. (1981) Eine neue Form der madagascariensis-Gruppe der Gattung Phelsuma von den Seychellen (Reptilia: Sauria: Gekkonidae). Salamandra, 17, 12–19.
Börner, A.R. (1972) Revision der Gekkonidengattung Phelsuma Gray 1825. Saurologica [published privately], 1, 1-145.
Börner, A.R. & Minuth, W. (1984) On the taxonomy of the Indian Ocean lizards of the Phelsuma madagascariensis group (Reptilia: Gekkonidae). Journal of the Bombay Natural History Society, 81, 243–281.
Boulenger, G.A. (1885) Catalogue of the lizards in the British Museum (Natural History). Volume I. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae. Trustees of the British Museum, London, I-XII, 436 pp.
Boumans, L., Vieites, D.R., Glaw, F. & Vences, M. (2007) Geographical patterns of deep mitochondrial differentiation in widespread Malagasy reptiles. Molecular Phylogenetics and Evolution, 45, 822–839.
Cheke, A.S. (1975) An undescribed gecko from Agalega, Phelsuma agalega sp. nov. Bulletin of the Mauritius Institute, 8, 33–48.
Cheke, A.S. (1981) What the book does not tell you about day geckos: Some comments on Oostveen's "Phelsuma's", on new species and on recent research. Herplite, 6, 4–10.
Cheke, A.S. (1982) Phelsuma Gray 1825 in the Seychelles and neighboring islands: A re-appraisal of their taxonomy and description of two new forms. Senckenbergiana biologica, 62, 181–198.
Checke, A.S. Hume, J. (2008) Lost land of the Dodo: The ecological history of Mauritius, Réunion and Rodrigues. A&C Black.
Daza, J. D. & Bauer, A. M. (2010 / Anatomical Record, 293, 402–413) The circumorbital bones of the Gekkota (Reptilia: Squamata). Anatomical Record.
D'Cruze, N., Sabel, J., Dawson, J. & Kumar, S. (2009) The influence of habitat type and structure on the abundance of Phelsuma madagascariensis grandis (Gekkonidae) in northern Madagascar. Herpetological Conservation and Biology, 4, 55–61.
Gardner, A.S. (1984) The evolutionary ecology and population systematics of day geckos (Phelsuma) in the Seychelles. Unpublished Ph.D Thesis, University of Aberdeen.
Gardner, A.S. (1986) Morphological evolution in the day gecko Phelsuma sundbergi in the Seychelles: A multivariate study. Biological Journal of the Linnean Society, 29, 223–244.
Gardner, A.S. (1987) The systematics of the Phelsuma madagascariensis species group of day geckos (Reptilia: Gekkonidae) in the Seychelles. Zoological Journal of the Linnean Society, 91, 93–105.
Glaw, F., Köhler. J. & Vences, M. (2009) A new species of cryptically coloured day gecko (Phelsuma) from the Tsingy de Bemaraha National Park in western Madagascar. Zootaxa, 2195, 61–68.
Glaw, F. & Vences, M. (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Second Edition. Vences and Glaw Verlag, Köln, 480 pp.
Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Third Edition. Vences and Glaw Verlag, Köln, 496 pp.
Glaw, F., Vences, M. & Ziegler, T. (1999) Bemerkungen zu Phelsuma dubia (Boettger, 1881): Wiederentdeckung des Holotypus, Verwandtschaftsbeziehungen und Daten zur Fortpflanzung. Salamandra, 35, 267–278.
Guayasamin, J. M., Castroviejo-Fisher, S., Trueb, L., Ayarzagüena, J., Rada, M. & Vilá, C. (2009) Phylogenetic systematics of Glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1–97.
Guindon S. & Gascuel O. (2003) A simple, fast, and accurate algorithm to estimate large phylogenies by maximum likelihood. Systematic Biology, 52, 696–704.
Hallmann, G., Krüger, J. & Trautmann, G. (2008, eds.) Faszinierende Taggeckos. Die Gattung Phelsuma. Second Edition. Natur und Tier-Verlag, Münster.
Harmon, L., Melville, J., Larson, A. & Losos, J.B. (2008) The role of geography and ecological opportunity in the diversification of day geckos (Phelsuma). Systematic Biology, 57, 562–573.
Hedges, S. B., Duellman, W. E. & Heinicke, M. P. (2008) New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1–182.
Huelsenbeck, J.P., Ronquist, F. (2001) MrBayes: Bayesian inference of phylogenetic trees. Bioinformatics, 17, 754–755.
Ikeuchi, I., Mori, A. & Hasegawa, M. (2005) Natural history of Phelsuma madagascariensis kochi from a dry forest in Madagascar. Amphibia-Reptilia 26, 475–483.
Katoh, K. & Toh, H. (2008) Recent developments in the MAFFT multiple sequence alignment program. Briefings in Bioinformatics, 9, 286–298.
Kluge, A.R. (1991) Checklist of gekkonid lizards. Smithsonian Herpetological Information Service, 85, 1–35.
Krüger, J. (1993) Beschreibung einer neuen Unterart von Phelsuma quadriocellata aus dem Nord-Osten Madagaskars. Salamandra, 29, 133–139.
Krüger, J. (1996a) Untersuchungen zur Validität einiger Taxa der Gattung Phelsuma. Herpetofauna, 18 (102), 5–11.
Krüger, J. (1996b) Beschreibung einer neuen Art aus der Gattung Phelsuma aus dem Süd-Osten Madagaskars. Herpetofauna, 18 (105), 14–18.
Krüger, J. (1996c) Zur Nomenklatur der lineata-Gruppe in der Gattung Phelsuma (Reptilia: Sauria: Gekkonidae). Sauria, 18 (3), 37–42.
Krysko, K. L., Nichole, A. N., Sheehy, C. M. (2003) The Madagascar giant gecko, Phelsuma madagascariensis grandis Gray 1870 (Sauria: Gekkonidae): A new established species in Florida. Florida Scientist, 66, 222–225.
Lerner, A. (2004) A new taxonomically isolated species of the genus Phelsuma Gray, 1825 from the Ampasindava Peninsula, Madagascar. Phelsuma, 12, 91–98.
Lever, C. (2003) Naturalized Reptiles and Amphibians of the World. Oxford University Press, Oxford.
Losos, J. (1986) Island biogeography of day geckos (Phelsuma) in the Indian Ocean. Oecologia, 68, 338–343.
Loveridge, A. (1941) New geckos (Phelsuma & Lygodactylus), snake (Leptotyphlops), and frog (Phrynobatrachus) from Pemba Island, East Africa. Proceedings of the Biological Society of Washington, 54, 175–176.
Loveridge, A. (1942) Revision of the Afro-Oriental geckos of the genus Phelsuma. Bulletin of the Museum Comparative Zoology, Harvard, 89, 439–482.
Meier, H. (1977) Beobachtungen an Phelsuma standingi (Reptilia: Sauria: Gekkonidae). Salamandra, 13, 1–12.
Meier, H. (1981) Zur Taxonomie und Ökologie der Gattung Phelsuma (Reptilia: Sauria: Gekkonidae) auf den Komoren, mit Beschreibung einer neuen Art. Bonner Zoologische Beiträge, 31 [1980], 323–331.
Meier, H. (1982) Ergebnisse zur Taxonomie und Ökologie einiger Arten und Unterarten der Gattung Phelsuma auf Madagaskar, gesammelt in den Jahren 1972 bis 1981, mit Beschreibung einer neuen Form (Reptilia: Sauria: Gekkonidae). Salamandra, 18, 168–190.
Meier, H. (1983) Neue Ergebnisse über Phelsuma lineata pusilla Mertens, 1964, Phelsuma bimaculata Kaudern, 1922 und Phelsuma quadriocellata (Peters, 1883), mit Beschreibung von zwei neuen Unterarten (Sauria: Gekkonidae). Salamandra, 19, 108–122.
Meier, H. (1984) Zwei neue Formen der Gattung Phelsuma von den Komoren (Sauria: Gekkonidae). Salamandra, 20, 32–38.
Meier, H. (1986) Der Formenkreis von Phelsuma v-nigra (Boettger, 1913) (Sauria: Gekkonidae) auf den Komoren: Beschreibung von zwei neuen Unterarten. Salamandra, 22, 11–20.
Meier, H. (1987) Vorläufige Beschreibung einer neuen Art der Gattung Phelsuma von Madagaskar (Sauria: Gekkonidae). Salamandra, 23, 204–211.
Meier, H. (1989) Eine neue Form aus der lineata-Gruppe der Gattung Phelsuma auf Madagaskar. Salamandra, 25, 230–236.
Meier, H. (1993) Neues über einige Taxa der Gattung Phelsuma auf Madagaskar, mit Beschreibung zweier neuer Formen. Salamandra, 29, 119–132.
Meier, H. & Böhme, W. (1991) Zur Arealkunde von Phelsuma madagascariensis (Gray, 1831) anhand der Museumssammlungen A. Koenig und Senckenberg, mit Bemerkungen zur Variabilität von P. m. kochi Mertens, 1954. Salamandra, 27, 143–151.
Meier, H. & Böhme, W. (1996) Zum taxonomischen Status des Formenkreises von Phelsuma abbotti Stejneger, 1893, mit Bemerkungen über P. masohoala Raxworthy & Nussbaum, 1994. Salamandra, 32, 85–98.
Mertens, R. (1954) Studien über die Reptilienfauna Madagaskars II. Ein neue Rasse von Phelsuma madagascariensis. Senckenbergiana biologica, 35, 13–16.
Mertens, R. (1962) Studien über die Reptilienfauna Madagaskars. III. Die Arten und Unterarten der Geckonengattung Phelsuma. Senckenbergiana biologica, 43, 81–127.
Mertens, R. (1963a) Studien über die Reptilienfauna Madagascars IV. Zwei neue Arten der Geckonengattung Phelsuma. Senckenbergiana biologica,.44, 349–356.
Mertens, R. (1963b) The geckos of the genus Phelsuma on Mauritius and adjacent islands. Bulletin of the Mauritius Institute, 5, 299–304.
Mertens, R. (1964) Studien über die Reptilienfauna Madagaskars V. Fünf neue Rassen der Geckonengattung Phelsuma. Senckenbergiana biologica, 45, 99–112.
Mertens, R. (1966) Die nichtmadegassischen Arten und Unterarten der Geckonengattung Phelsuma. Senckenbergiana biologica, 47, 85–110.
Mertens, R. (1970) Neues über einige Taxa der Geckonengattung Phelsuma. Senckenbergiana biologica, 51, 1–13.
Mertens, R. (1973) Studien über die Reptilienfauna Madagaskars. VII. Eine neue Unterart des Taggeckos Phelsuma lineata. Senckenbergiana biologica, 54, 299–301.
Nussbaum, R.A., Raxworthy, C.J., Raselimanana, A.P. & Ramanamanjato, J.B. (2000) New species of Day Gecko, Phelsuma Gray (Reptilia: Squamata: Gekkonidae), from the Réserve Naturelle Intégrale d’Andohahela, southern Madagascar. Copeia, 2000, 763–770.
Nylander, J.A., Wilgenbusch, J.C., Warren, D.L., Swofford, D.L. (2008) Awty (are we there yet?): A system for graphical exploration of mcmc convergence in Bayesian phylogenetics. Bioinformatics, 24, 581–583.
Ota, H., Ineich, I. (2006) Colonization of the gold dust day gecko, Phelsuma laticauda (Reptilia: Gekkonidae), in Moorea Archipelago, French Polynesia. Current Herpetology, 25, 97–99.
Palumbi, S.R., Martin, A., Romano, S., McMillan, W.O., Stice, L. & Grabowski, G. (1991) The Simple Fool’s Guide to PCR, Version 2.0. Privately published, Hawaii.
Posada, D. (2008) jModelTest: Phylogenetic Model Averaging. Molecular Biology and Evolution, 25, 1253–1256.
Posada, D. & Crandall, K.A. (1998) MODELTEST: testing the model of DNA substitution. Bioinformatics, 14, 817–818.
Puente, M., Glaw, F., Vieites, D. R. & Vences, M. (2009) Review of the systematics, morphology and distribution of Malagasy dwarf geckos, genera Lygodactylus and Microscalabotes (Squamata: Gekkonidae). Zootaxa, 2103, 1–76.
Radtkey, R. (1996) Adaptive radiation of day-geckos (Phelsuma) in the Seychelles Archipelago: A phylogenetic analysis. Evolution, 50, 604–623.
Raxworthy, C.J. & Nussbaum, R.A. (1993) A new Madagascan Phelsuma, with a review of Phelsuma trilineata and comments on Phelsuma cepediana in Madagascar (Squamata: Gekkonidae). Herpetologica, 49, 342–349.
Raxworthy, C. J. & Nussbaum, R.A. (1994) A partial systematic revision of the day geckos, Phelsuma Gray, of Madagascar (Reptilia: Squamata: Gekkonidae). Zoological Journal of the Linnean Society, 112, 321–335.
Raxworthy, C.J., Ingram, C.M., Rabibisoa, N. & Pearson, R.G. (2007) Applications of ecological niche modelling for species delimitation: A review and empirical evaluation using day geckos (Phelsuma) from Madagascar. Systematic Biology, 56, 907–923.
Rendahl, H. (1939) Zur Herpetofauna der Seychellen. I. Reptilien. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere, 72, 157–328.
Rocha S., Posada D., Carretero M.A., Harris D.J., 2007. Phylogenetic affinities of Comoroan and East African day geckos (genus Phelsuma): multiple natural colonisations, introductions and island radiations. Molecular Phylogenetics and Evolution, 43, 685–692.
Rocha S., Vences M., Glaw F., Posada D., Harris D.J., 2009. Multigene phylogeny of Malagasy day geckos of the genus Phelsuma. Molecular Phylogenetics and Evolution, 52, 530–537.
Rösler, H. (1998) Zur Taxonomie und Verbreitung von Phelsuma madagascariensis (Gray, 1831) in Nordost-Madagaskar (Sauria: Gekkonidae). Gekkota, 1, 21–32.
Rösler, H. (2001) Zur Rumpfbeschuppung der Gattung Phelsuma Gray, 1825: zoogeographische Aspekte (Sauria: Gekkonidae). Gekkota, 3, 47–73.
Rösler, H., Obst, F.J. & Seipp, R. (2001) Eine neue Taggecko-Art von Westmadagaskar: Phelsuma hielscheri sp.n. (Reptilia: Sauria: Gekkonidae). Zoologische Abhandlungen Staatliches Museum für Tierkunde in Dresden, 51, 51–60.
Sambrook, J., Fritsh, E.F. & Maniatis, T. (1999) Molecular Cloning: A Laboratory Manual, second ed. Cold Spring Harbor Press, New York.
Schönecker, P. (2008) Geckos of Madagascar, the Seychelles, Comoros and Mascarene Islands. Terralog. Edition Chimaira, Frankfurt a. M., 144 pp.
Schönecker, P., Bach, S. & Glaw, F. (2004) Eine neue Taggecko-Art der Gattung Phelsuma aus Ost-Madagaskar (Reptilia: Squamata: Gekkonidae). Salamandra, 40, 105–112.
Seipp, R. (1990) Eine neue Art der Gattung Phelsuma Gray, 1825 von Madagaskar (Reptilia: Sauria: Gekkonidae). Senckenbergiana biologica, 71, 11–14.
Sound, P., Kosuch, J., Vences, M., Seitz, A. & Veith, M. (2006) Preliminary molecular relationships of Comoran day geckos (Phelsuma). Proceedings of the 13th Congress Societas Europaea Herpetologica, 175–179
Swofford, D.L. (2002) PAUP*. Phylogenetic analysis using parsimony (*and other methods). Version 4. Sinauer Associates, Sunderland, MA.
Taggart, J.B., Hynes, R.A., Prodoul, P.A., Ferguson, A. (1992) A simplified protocol for routine total DNA isolation from salmonid fishes. Journal of Fish Biology, 40, 963–965.
Van Heygen, E. (2004) Remarks on the Phelsuma barbouri- and Phelsuma klemmeri-phenetic groups. Phelsuma, 12, 153–154.
Vinson, J. & Vinson, J.M. (1969) The saurian fauna of the Mascarene Islands. Mauritius Institute Bulletin, 4, 203–320.
Zwickl, D.J. (2006) Genetic algorithm approaches for the phylogenetic analysis of large biological sequence datasets under the maximum likelihood criterion. Ph.D.dissertation, The University of Texas at Austin. Available from http://www.bio.utexas.edu/faculty/antisense/garli/Garli.html