Abstract
The Leptodactylus melanonotus group consists of 15 species, but references to skeletal characters are available for only three species: L. leptodactyloides, L. melanonotus, and L. diedrus. Leptodactylus nesiotus is a member of the melanonotus group known only from the type locality, Bonasse swamp, on the Southwestern peninsula of Trinidad, Trinidad and Tobago. This species has been categorized as vulnerable given its restricted distribution. Herein, we report the adult osteology of L. nesiotus, the skeletal characters are compared with the available data from other Leptodactylus species. A phylogenetic analysis recovers a paraphyletic L. melanonotus group relative to the L. latrans group. A monophyletic “latrans-melanonotus” clade is supported by five synapomorphies. L. nesiotus is recovered as the sister species of L. validus, a relationship supported by two synapomorphies: T-shaped terminal phalanges and a dark-colored stripe on the outer surface of arm. In addition, we report on the ecology of this poorly known species.
References
Camargo, A., de Sá, R.O. & Heyer, W.R. (2006) Phylogenetic analyses of mtDNA sequences reveal three cryptic lineages in the widespread neotropical frog Leptodactylus fuscus (Schneider, 1799) (Anura, Leptodactylidae). Biological Journal of the Linnean Society, 87, 325–341.
de Sá, R.O., Heyer, W.R. & Camargo, A. (2005) Are Leptodactylus didymus and L. mystaceus phylogenetically sibling species (Amphibia, Anura, Leptodactylidae)? Herpetologia Petropolitana, 90–92.
Fabrezi, M. (1992) El carpo de anuros. Alytes, 10, 1–36.
Fabrezi, M. & Alberch, P. (1996) The carpal elements of anurans. Herpetologica, 52, 188–204.
Farris, J.S. (1969) A successive approximations approach to character weighting. Systematic Zoology, 18, 374–385.
Farris, J.S. (1988) Hennig86, program and documentation. Published by the author, Port Jefferson, New York.
Fitzinger, L.J.F.J. (1826) Neue Classification der Reptilien nach ihren natu¨rlichen Verwandtschaften nebst einer Verwandtschafts-Tafel und einem Verzeichnisse der Reptilien-Sammlung des k.k. zoologisch Museum’s zu Wien. Wien: J.G. Heubner.
Frost, D.R. (2009) Amphibian Species of the World: an Online Reference. Version 5.2 (15 July, 2008). Electronic Database available from: http://research.amnh.org/ herpetology/amphibia/index.php. American Museum of Natural History, New York, USA (accessed March 2 2010).
Goloboff, P.A. (1993) Estimating character weigths during tree search. Cladistics, 9, 83–91.
Goloboff, P.A. (1997) Self-weighted optimization: tree searches and character states reconstruction under implied transformation costs. Cladistics, 13, 225–245.
Goloboff, P.A. & Farris, J.S. (2001) Methods for quick consensus estimation. Cladistics, 17, 26–34.
Goloboff, P., Farris, J. & Nixon, K. (2003a) T.N.T.: tree analysis using new technology. Program and documentation available from the authors and http://www.zmuc.dk/public/ phylogeny (accessed March 2 2010).
Goloboff, P.A., Farris, J.S., Källersjö, M., Oxelman, B., Ramírez, M.J. & Szumik, C.A. (2003b) Improvements to resampling measures of group support. Cladistics, 19, 324–332.
Heyer, W.R. (1969a) The adaptive ecology of the species groups o the genus Leptodactylus (Amphibia, Leptodactylidae). Evolution, 23, 421–428.
Heyer, W.R. (1969b) Studies on the genus Leptodactylus (Amphibia, Leptodactylidae) III. A redefinition of the genus Leptodactylus and a description of a new genus of Leptodactylid frogs. Contribution in Science, Natural History Museum Los Angeles County, 155, 1–14.
Heyer, W.R. (1974) Relationships of the marmoratus species group (Amphibia, Leptodactylidae) within the subfamily Leptodactylinae. Contributions in Science of the Natural History Museum of the Los Angeles County, 253, 1–46.
Heyer, W.R. (1994) Variation within then Leptodactylus podicipinus-wagneri complex of frogs (Amphibia: Leptodactylidae). Smithsonian Contributions to Zoology, 546, 1–124.
Heyer, W.R. (1998) The relationships of Leptodactylus diedrus (Anura, Leptodactylidae). Alytes, 16, 1–24.
Heyer, W.R. & Crombie, R.I. (2005) Leptodactylus lauramiriamae, a distinctive new species of frog (Amphibia: Anura: Leptodactylidae) from Rondônia, Brazil. Preceedings of the Biological Society of Washington, 118, 590–595.
IUCN, Conservation International, and Nature Serve. (2006) Global Amphibian Assessment. www.globalamphibians.org. (accessed 12 November 2008)
Larson, P.M. & de Sá, R.O. (1998) Chondrocranial morphology of Leptodactylus larvae (Leptodactylidae: Leptodactylinae): its utility in phylogenetic reconstruction. Journal of Morphology, 238, 287–305.
Lavilla, E.O., Langone, J.A., Caramaschi, U., Heyer, W.R. & de Sá, R.O. (2010) The identification of Rana ocellata Linnaeus, 1758. Nomenclatrual impact on the species currently known as Leptodactylus ocellatus (Leptodactylidae) and Osteopilus brunneus (Gosse, 1851) (Hylidae). Zootaxa, 2346, 1–16.
Lipscomb, D.L. (1992) Parsimony, homology and the analysis of multistate characters. Cladistics, 8, 45–65.
Lynch, J.D. (1971) Evolutionary relationships, osteology, and zoogeography of leptodactyloid frogs. Miscellaneous Publications Museum of Natural History University of Kansas, 53, 1–238.
Maglia, A.M., Pugener, L.A. & Mueller, J.M. (2007) Skeletal morphology and postmetamorphic ontogeny of Acris crepitans (Anura: Hylidae): a case of miniaturization in frogs. Journal of Morphology, 268, 194–223.
Maxson, L.R. & Heyer, W.R. (1988) Molecular systematics of the frogs genus Leptodactylus (Amphibia: Leptodactylidae). Fieldana: Zoology, 41, 1–13.
Perotti, M.G. (2001) Skeletal development of Leptodactylus chaquensis (Anura: Leptodactylidae). Herpetologica, 57, 318–335.
Ponssa, M.L. (2006) On the osteology of a distinctive species of the genus Leptodactylus: Leptodactylus laticeps (Boulenger, 1917) (Anura, Leptodactylidae). Zootaxa, 1188, 23–36.
Ponssa, M.L. (2008) Cladistic analysis and osteological descriptions of the species of the L. fuscus species group of the genus Leptodactylus (Anura, Leptodactylidae). Journal of Zoological Systematics and Evolutionary Research, 46, 249–266.
Ponssa, M.L., Goldberg, J. & Abdala, V. (2010) Sesamoids in anurans: new data, old issues. Anatomical Record, 293, 1646–1668.
Ponssa, M.L. & Heyer, W.R. (2007) Osteological characterization of four putative species of the genus Adenomera (Anura: Leptodactylidae), with comments on intra- and interspecific variation. Zootaxa, 1403, 37–54.
Ponssa, M.L. & Lavilla, E.O. (1998) Osteology of Leptodactylus latinasus (Anura: Leptodactylidae) and the validity of its subspecies. Bulletin of the Maryland Herpetological Society, 34, 57–63.
Pugener, L.A. & Maglia, A.M. (2007) Skeletal morphology and development of the olfactory region of Spea (Anura: Scaphiopodidae). Journal of Anatomy, 211, 754–768.
Sebben, A., Maciel, N.M., Campos, L.A., Kokubum, M.N.C. & Da Silva, H.R. (2007) Occurrence of a calcified pseudodontoid in Leptodactylus troglodytes (Anura: Leptodactylidae). Journal of Herpetology, 41, 337–340.
Trewavas, E. (1933) The hyoid and larynx of the Anura. Philosophical Transactions of the Royal Society of London, 222, 401–527.
Trueb, L. (1973) Bones, frogs and evolution. In: Vial, J. L. (Ed.), Evolutionary biology of the anurans: contemporary research on major problems. The University Missouri Press, Columbia, pp. 65–132.
Trueb, L. (1993) Patterns of cranial diversity among the Lissamphibia. In: Hanken, J. & B. K. Hall (Eds.), The Skull 2. The University of Chicago Press, pp. 255–343.
Trueb, L., Pugener, L.A. & Maglia, A.M. (2000) Ontogeny of the bizarre: an osteological description of Pipa pipa (Anura: Pipidae), with an account of skeletal development in the species. Journal of Morphology, 243, 75–104.
Wassersug, R.J. (1976) A procedure for differential staining of cartilage and bone in hole formalin fixed vertebrates. Staining Technology, 51, 131–134.