Abstract
Frogs of the subfamily Mantellinae (Amphibia: Anura: Mantellidae) are a species-rich and diverse lineage endemic to the Madagascan region. The major synapomorphy of this clade is a derived reproductive mode including an unusual mating behaviour (loss of strong mating amplexus, egg deposition outside of water) and associated morphological adaptations (evolution of femoral glands, loss of nuptial pads). However, the evolutionary steps towards this unique character complex remain obscure. We here describe a recently discovered new frog, Tsingymantis antitra gen. nov., sp. nov. from the moderately dry karstic massif Tsingy de Ankarana in northern Madagascar. The new species is not referable to any existing genus or species groups. A phylogenetic analysis, based on DNA sequences of four mitochondrial genes (12S and 16S rRNA, tRNAVal, cytochrome b) and one nuclear gene (rhodopsin) placed Tsingymantis without significant support as sister taxon of the Mantellinae which was found to be a well-defined monophyletic group (100% Bayesian and 99% bootstrap support). The position of Tsingymantis as the most basal clade of the Mantellinae is in agreement with several morphological and osteological characters, suggesting that this subfamily including Tsingymantis may be a monophyletic group whereas the Boophinae could represent the most basal clade of the Mantellidae. We therefore include Tsingymantis in the Mantellinae in a preliminary way, pending further study. In contrast to the large majority of recent mantellid species which are adapted to humid rainforests, the most basal clades of the three subfamilies show adaptations to relatively dry conditions, indicating that the climate during the early radiation of mantellids (probably in the Eocene) may have been drier than in recent times.References
Andreone, F., Cadle, J.E., Cox, N., Glaw, F., Nussbaum, R.A., Raxworthy, C.J., Stuart, S., Vallan, D. & Vences, M. (2005) Species review of amphibian extinction risks in Madagascar: conclusions from the Global Amphibian Assessment. Conservation Biology, 19(6), 1790–1802.
Biju, S.D. & Bossuyt, F. (2003) New frog family from India reveals an ancient biogeographical link with the Seychelles. Nature, 425, 711–714.
Blommers-Schlösser, R.M.A. (1979) Biosystematics of the Malagasy frogs. I. Mantellinae (Ranidae). Beaufortia, 352(2), 1–77.
Blommers-Schlösser, R.M.A. & Blanc, C.P. (1991) Amphibiens (première partie). Faune de Madagascar, 75(1), 1–379.
Bossuyt, F. & Milinkovitch, M. C. (2000) Convergent adaptive radiations in Madagascan and Asian ranid frogs reveal covariation between larval and adult traits. Proceedings of the National Academy of Sciences of the USA, 97, 6585–6590.
Dubois, A. (1992) Notes sur la Classification des Ranidae (Amphibiens, Anoures). Bulletin de la Societé Linnéenne de Lyon, 61, 305–352.
Dubois, A. (2005) Amphibia Mundi. 1.1. An ergotaxonomy of recent amphibians. Alytes, 23(1–2), 1–24.
Frost, D.R. (2004) Amphibian Species of the World: an Online Reference. Version 3.0 (22 August, 2004). Available from: http://research.amnh.org/herpetology/amphibia/index.html (accessed 29 April 2006)
Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., de Sá, R.O., Channing, A., Wilkinson, M., Donnellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P., Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D.M. & Wheeler, W.C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1–370.
Glaw, F., & Vences, M. (1994) A fieldguide to the amphibians and reptiles of Madagascar. second edition, Köln, Vences & Glaw, 480 pp.
Glaw, F. & Vences, M. (2006). Phylogeny and genus-level classification of mantellid frogs (Amphibia, Anura). Organisms, Diversity and Evolution, 6 (3), 236–253.
Glaw, F., Vences, M. & Böhme, W. (1998) Systematic revision of the genus Aglyptodactylus Boulenger, 1919 (Amphibia: Ranidae), and analysis of its phylogenetic relationships to other Madagascan ranid genera (Tomopterna, Boophis, Mantidactylus, and Mantella). Journal of Zoological Systematics and Evolutionary Research, 36, 17–37.
Guibé, J. (1974) Batraciens nouveaux de Madagascar. Bulletin du Muséum National d'Histoire Naturelle, ser. 3, 171, 1169–1192.
Guindon, S. & Gascuel, O. (2003) A simple, fast, and accurate algorithm to estimate large phylogenies by maximum likelihood. Systematic Biology, 52, 696–704.
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.
Hawkins, A.F.A., Chapman, P., Ganzhorn, J.U., Bloxam, Q.M.C., Barlow, S.C. & Tonge, S.J. (1990) Vertebrate conservation in Ankarana Special Reserve, northern Madagascar. Biological Conservation, 54, 83–110.
Huelsenbeck, J.P., & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics, 17, 754–755.
Keane, T.M., Naughton, T.J. & McInerney, J.O. (2004) ModelGenerator: amino acid and nucleotide substitution model selection. National University of Ireland, Maynooth, Ireland.
Lang, M. (1990) Phylogenetic analysis of the genus group Tracheloptychus-Zonosaurus (Reptilia: Gerrhosauridae) with the hypothesis of biogeographical unit relationships in Madagascar. In: Peters, G. & Hutterer, R. (Eds.) Vertebrates in the Tropics. Museum Alexander Koenig, Bonn, 261–274.
Min, M.S.,Yang, S.Y., Bonett, R.M., Vieites, D.R., Brandon, R.A. & Wake, D.B. (2005) Discovery of the first Asian plethodontid salamander. Nature, 435, 87–90.
Richards, C.M., Nussbaum, R.A. & Raxworthy, C.J. (2000) Phylogenetic relationships within the Madagascan boophids and mantellids as elucidated by mitochondrial ribosomal genes. African Journal of Herpetology, 49, 23–32.
Ronquist, F., & Huelsenbeck, J. P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.
Sket, B. (1997) Distribution of Proteus (Amphibia: Urodela: Proteidae) and its possible explanation. Journal of Biogeography, 24, 263–280.
Thompson, J.D., Higgins, D.G. & Gibson, T.J. (1994) Clustal W.: improving the sensivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Research, 22, 4673–4680.
Van der Meijden, A., Vences, M., Hoegg, S. & Meyer, A. (2005) A previously unrecognized radiation of ranid frogs in southern Africa revealed by nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 37, 674–685.
Vences, M., Andreone, F., Glaw, F. & Mattioli, F. (2002a) New dwarf species of Mantidactylus from northwestern Madagascar (Anura: Mantellidae). Copeia, 2002 (4), 1057–1062.
Vences, M., Glaw, F., Andreone, F., Jesu, R. & Schimmenti, G. (2002b) Systematic revision of the enigmatic Malagasy broad-headed frogs (Laurentomantis Dubois, 1980) and their phylogenetic position within the endemic mantellid radiation of Madagascar. Contributions to Zoology, 70 (4), 191–212.
Vences, M. & Glaw, F. (2001) When molecules claim for taxonomic changes: New proposals on the classification of Old World treefrogs. Spixiana, 24, 85–92.
Vences, M., Glaw, F. & Böhme, W. (1999) A review of the genus Mantella (Anura, Ranidae, Mantellinae): taxonomy, distribution and conservation of Malagasy poison frogs. Alytes, 17, 3–72.
Vences, M., Glaw, F., Kosuch, J., Das, I. & Veith, M. (2000) Polyphyly of Tomopterna (Amphibia: Ranidae) based on sequences of the mitochondrial 16S and 12S rRNA genes, and ecological biogeography of Malagasy relict amphibian groups. In: Lourenço, W.R. & Goodman, S.M. (Eds.), Diversité et endémisme a Madagascar. Mémoires de la Société de Biogéographie, Paris, pp. 229–242.
Vences, M., Vieites, D.R., Glaw, F., Brinkmann, H., Kosuch, J., Veith, M. & Meyer, A. (2003) Multiple overseas dispersal in amphibians. Proceedings of the Royal Society of London B, 270, 2435–2442.
Wells, N.A. (2003) Some hypotheses on the Mesozoic and Cenozoic paleoenvironmental history of Madagascar, In: Goodman, S.M. & Benstead, J.P. (Eds.), The Natural History of Madagascar. The University of Chicago Press, Chicago and London, pp. 16–34.