Heyer, W. Ronald, and Rafael O. de Sá. Variation, Systematics, and Relationships of the Leptodactylus bolivianus Complex (Amphibia: Anura: Leptodactylidae). Smithsonian Contributions to Zoology, number 635, viii + 58 pages, 21 figures, 20 tables, 2011.—The Leptodactylus bolivianuscomplex has been considered to consist of one or two species, L. bolivianus alone or L. bolivianus and L. insularum. Detailed morphological analyses were undertaken to evaluate variation in the complex, which ranges from Costa Rica through Panama, across northern South America in the river valleys draining to the Caribbean, and throughout much of the Amazon basin with southern limits in Bolivia. Members of the complex also occur on several islands off Nicaragua, Panama, and Colombia. Analyses of morphological and advertisement call data indicate that there are either two or three species comprising the complex. Analysis of molecular data strongly supports recognition of three species, one of which is described as a new species, Leptodactylus guianensis. The three species comprising the L. bolivianus clade are most closely related to the L. ocellatus clade within the genus Leptodactylus.
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Heyer, W.R. and R. O. de Sá. 2011. Variation, Systematics, and Relationships of the Leptodactylus bolivianus Complex (Amphibia: Anura: Leptodactylidae). Smithsonian Contributions to Zoology, 635:1-58. https://doi.org/10.5479/si.00810282.635.1