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Dubois, A., & W. R. Heyer.  1992.  Leptodactylus labialis, the valid name for the American white-lipped frog (Amphibia: Leptodactylidae).  Copeia, 1992:584-585.

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Heyer, W. R. 1969. Studies on frogs of the genus Leptodactylus (Amphibia, Leptodactylidae).  V.  Taxonomic notes on L. latinasus, rhodonotus, romani, and wuchereri.  Herpetologica, 25:1-8.

Heyer, W. R. 1969. The adaptive ecology of the species groups of the frog genus Leptodactylus (Amphibia, Leptodactylidae). Evolution, 23:421-428.

Heyer, W. R.  1970.  Studies on frogs of the genus Leptodactylus (Amphibia, Leptodactylidae).  VI.  Biosystematics of the Melanonotus group.  Contributions in Science, Los Angeles County Museum of Natural History, (191):1-48.

Heyer, W. R. 1970.  Studies on the genus Leptodactylus (Amphibia, Leptodactylidae).  II.  Diagnosis and distribution of the Leptodactylus of Costa Rica.  Revista de Biología Tropical, 16:171-205.

Heyer, W. R.  1971.  Leptodactylus labialis.  Catalogue of American Amphibians and Reptiles, (104):1-3.

Heyer, W. R.  1972.  The status of Leptodactylus pumilio Boulenger (Amphibia, Leptodactylidae) and the description of a new species of Leptodactylus from Ecuador.  Contributions in Science, Los Angeles County Museum of Natural History, (231):1-8.

Heyer, W. R.  1973.  Systematics of the Marmoratus group of the frog genus Leptodactylus (Amphibia, Leptodactylidae).  Contributions in Science, Los Angeles County Museum of Natural History, (251):1-50.

Heyer, W. R. 1974. Relationships of the marmoratus species group (Amphibia, Leptodactylidae) within the Subfamily Leptodactylinae. Contributions in Science, Los Angeles County Museum of Natural History, (253):1-46.

Heyer, W. R.  1974.  Vanzolinius, a new genus proposed for Leptodactylus discodactylus (Amphibia, Leptodactylidae).  Proceedings of the Biological Society of Washington, 87:81-90.

Heyer, W. R.  1975.  A preliminary analysis of the intergeneric relationships of the frog family Leptodactylidae.  Smithsonian Contributions to Zoology, (199):1-55. [This article is broken up into two pdf files.  pdf file 1.  pdf file 2.]

Heyer, W. R.  1975.  Adenomera lutzi (Amphibia: Leptodactylidae), a new species of frog from Guyana.  Proceedings of the Biological Society of Washington, 88(28):315-318.

Heyer, W. R.  1976.  Notes on the frog fauna of the Amazon Basin.  Acta Amazonica, 6:369-378.

Heyer, W. R.  1977.  A discriminant function analysis of the frogs of the genus Adenomera (Amphibia: Leptodactylidae).  Proceedings of the Biological Society of Washington, 89:581-592.

Heyer, W. R.  1977.  Taxonomic notes on frogs from the Madeira and Purus rivers, Brasil.  Papéis Avulsos de Zoologia, 31:141-162.

Heyer, W. R.  1978.  Systematics of the fuscus group of the frog genus Leptodactylus (Amphibia, Leptodactylidae).  Natural History Museum of Los Angeles County Science Bulletin, (29):1-85.  [This article is broken up into two pdf files.  pdf file 1.  pdf file 2.]

Heyer, W. R. 1979. Systematics of the pentadactylus species group of the frog genus Leptodactylus (Amphibia: Leptodactylidae). Smithsonian Contributions to Zoology, (301):1-43.

Heyer, W. R.  1983.  Clarification of the names Rana mystacea Spix, 1824, Leptodactylus amazonicus Heyer, 1978 and a description of a new species, Leptodactylus spixi (Amphibia: Leptodactylidae).  Proceedings of the Biological Society of Washington, 96:270-272.

Heyer, W. R.  1984.  The systematic status of Adenomera griseigularis Henle, with comments on systematic problems in the genus Adenomera (Amphibia: Leptodactylidae).  Amphibia-Reptilia, 5:97-100.

Heyer, W. R. 1998. The relationships of Leptodactylus diedrus (Anura, Leptodactylidae). Alytes, 16:1-24.

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Heyer, W. R.  1994.  Variation within the Leptodactylus podicipinus-wagneri complex of frogs (Amphibia: Leptodactylidae).  Smithsonian Contributions to Zoology, (546):1-124.

Heyer, W. R.  1995.  South American rocky habitat Leptodactylus (Amphibia: Anura: Leptodactylidae) with description of two new species.  Proceedings of the Biological Society of Washington, 108:695-716.

Heyer, W. R.  1997.  Geographic variation in the frog genus Vanzolinius (Anura: Leptodactylidae).  Proceedings of the Biological Society of Washington, 110:338-365.

Heyer, W. R.  2000.  The lectotype of Cystignathus hylodes Reinhardt and Lütken, 1862.  Journal of Herpetology, 34:150-153.

Heyer, W. R.  2002.  Leptodactylus fragilis, the valid name for the Middle American and northern South American white-lipped frog (Amphibia: Leptodactylidae).  Proceedings of the Biological Society of Washington, 115:321-322.

Heyer, W. R.  2005.  Leptodactylus fragilis Brocchi, 1877.  White-lipped frog.  Pages 500-501 in M. J. Lannoo (editor), Amphibian Declines.  The Conservation Status of United States Species.  University of California Press:  Berkeley & Los Angeles.  xxi+1094 pages.

Heyer, W. R.  2005.  Variation and taxonomic clarification of the large species of the Leptodactylus pentadactylus species group (Amphibia: Leptodactylidae) from Middle America, northern  South America, and Amazonia.  Arquivos de Zoologia, 37(3):269-348.

Heyer, W. R., & M. S. Bellin.  1973.  Ecological notes on five sympatric Leptodactylus (Amphibia, Leptodactylidae) from Ecuador.  Herpetologica, 29:66-72.

Heyer, W. R., & K. A. Berven.  1973.  Species diversities of herpetofaunal samples from similar habitats at two tropical sites.  Ecology, 54:642-645.

Heyer, W. R., U. Caramaschi, & R. O. de Sá.  2006.  Rana ocellata Linnaeus, 1758 (currently Leptodactylus ocellatus; Amphibia, Anura:  Proposed conservation of usage of the specific name by the designation of a neotype.  Bulletin of Zoological Nomenclature, 63(3):184-186.

Heyer, W. R., & C. M. Carvalho.  2000.  Calls and calling behavior of the frog Leptodactylus natalensis (Amphibia: Anura: Leptodactylidae).  Proceedings of the Biological Society of Washington, 113:284-290.

Heyer, W. R., J. Coddington, W. J. Kress, P. Acevedo, D. Cole, T. L. Erwin, B. J. Meggers, M. G. Pogue, R. W. Thorington, R. P. Vari, M. J. Weitzman, & S. H. Weitzman. 2000 ("1999").  Amazonian biotic data and conservation decisions.  Ciência e Cultura, Journal of the Brazilian Association for the Advancement of Science, 51:372-385.

Heyer, W. R., & R. I. Crombie.  2005.  Leptodactylus lauramiriamae, a distinctive new species of frog (Amphibia: Anura: Leptodactylidae) from Rondônia, Brazil.  Proceedings of the Biological Society of Washington, 118(3):590-595.

Heyer, W. R., R. O. de Sá, J. R. McCranie, & L. D. Wilson.  1996.  Leptodactylus silvanimbus (Amphibia: Anura: Leptodactylidae):  Natural history notes, advertisement call, and relationships.  Herpetological Natural History, 4:169-174.

Heyer, W. R., R. O. de Sá, & S. Muller.  2002.   Leptodactylus silvanimbus.  Catalogue of American Amphibians and Reptiles, (742):1-5.

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Heyer, W. R., & M. J. Diment.  1974.  The karyotype of Vanzolinius discodactylus (Amphibia, Leptodactylidae).  Proceedings of the Biological Society of Washington, 87:327-336.

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Heyer, W. R., & M. M. Heyer.  2001.  Leptodactylus lithonaetes.  Catalogue of American Amphibians and Reptiles, (723):1-3.

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Heyer, W. R., & A. S. Rand. 1977. Foam nest construction in the leptodactylid frogs Leptodactylus pentadactylus and Physalaemus pustulosus (Amphibia, Anura, Leptodactylidae). Journal of Herpetology, 11:225-228.

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