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Brandão, R. A., & W. R. Heyer.  2005.  The complex calls of Leptodactylus pustulatus (Amphibia, Anura, Leptodactylidae).  Amphibia-Reptilia, 26(4):566-570.

Camargo, A., R. O. de Sá, & W. R. Heyer. 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.

Camargo, A., W. R. Heyer, and R. O. de Sá.  2009.  Phylogeography of the frog Leptodactylus validus (Amphibia: Anura): Patterns and timing of colonization events in the Lesser Antilles.  Molecular Phylogenetics and Evolution, 53:571-579.

Caramaschi, U., R. O. de Sá, & W. R. Heyer.  2005.  Common names for the frog genus Leptodactylus (Amphibia, Anura, Leptodactylidae).  Herpetological Review, 36(2):119-120.

Cardoso, A. J., & W. R. Heyer.  1995.  Advertisement, aggressive, and possible seismic signals of the frog Leptodactylus syphax (Amphibia, Leptodactylidae).  Alytes, 13:67-76.`

Crombie, R. I., & W. R. Heyer.  1983.  Leptodactylus longirostris (Anura: Leptodactylidae):  Advertisement call, tadpole, ecological and distributional notes.  Revista Brasileira de Biologia, 43:291-296.

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de Sá, R. O., & A. Gehrau.  1983.  Observaciones sobre la biología de Phyllomedusa iheringi Boulenger, 1885, (Anura, Hylidae).  Boletín de la Sociedad Zoológica del Uruguay, Segunda Época, 1:44-49.

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de Sá, R. O., W. R. Heyer, & A. Camargo.  2006 ("2005"). A phylogenetic analysis of Vanzolinius Heyer, 1974 (Amphibia, Anura, Leptodactylidae):  Taxonomic and life history implications.  Arquivos do Museu Nacional, Rio de Janeiro, 63(4):707-726.

Dixon, J. R., & W. R. Heyer. 1968. Anuran succession in a temporary pond in Colima, Mexico.  Bulletin of the Southern California Academy of Sciences, 67:129-137.

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

Fernández, M., D. Cole, W. R. Heyer, S. Reichle, & R. O. de Sá.  2009.  Predicting Leptodactylus (Amphibia, Anura, Leptodactylidae) distributions:  Broad-ranging versus patchily distributed species using a presence-only environmental niche modeling technique.  South American Journal of Herpetology, 4(2):103-116.

Frost, D. R., T. Grant, J. Faivovich, R. H. Bain, A. Haas, C. F. B. Haddad, R. O. de Sá, A. Channing, M. Wilkinson, S. C. Donnellan, C. J. Raxworthy, J. A. Campbell, B. L. Blotto, P. Moler, R. C. Drewes, R. A. Nussbaum, J. D. Lynch, D. M. Green, & W. C. Wheeler.  2006.  The amphibian tree of life.  Bulletin of the American Museum of Natural History, (297):1-37 + insert.

Furness, A. I., R. W. McDiarmid, W. R. Heyer, & G. R. Zug.  2010.  Oviduct modifications in foam-nesting frogs, with emphasis on the genus Leptodactylus (Amphibia, Leptodactylidae).  South American Journal of Herpetology, 5(1):13-29.

García Pérez, L., & W. R. Heyer.  1993.  Description of the advertisement call and resolution of the systematic status of Leptodactylus gracilis delattini Müller, 1968 (Amphibia: Leptodactylidae).  Proceedings of the Biological Society of Washington, 106:51-56.

Hayek, L. C., & W. R. Heyer. 2005. Determining sexual dimorphism in frog measurement data: Integration of statistical significance, measurement error, effect size and biological significance. Anais da Academia Brasileira de Ciências, 77:45-76.

Hayek, L. C., W. R. Heyer, & C. Gascon.  2001.  Frog morphometrics:  A cautionary tale.  Alytes, 18:153-177.

Heyer, M. M., W. R. Heyer, & R. O. de Sá.  2006.  Leptodactylus fragilis.  Catalogue of American Amphibians and Reptiles, (830):1-26.

Heyer, M. H., W. R. Heyer, & R. O. de Sá.  2009.  Bibliography of the frogs of the Leptodactylus Clade - Adenomera, Hydrolaetare, Leptodactylus, Lithodytes (Amphibia, Anura, Leptodactylidae).  Volume 1.  References.  Smithsonian Herpetological Information Service, Special Publication 1:1-677.

Heyer, M. H., W. R. Heyer, & R. O. de Sá.  2009.  Bibliography of the frogs of the Leptodactylus Clade - Adenomera, Hydrolaetare, Leptodactylus, Lithodytes (Amphibia, Anura, Leptodactylidae).  Volume 2.  Indices.  Smithsonian Herpetological Information Service, Special Publication 2:1-167.

Heyer, M. M., W. R. Heyer, S. Spear, & R. O. de Sá.  2003.  Leptodactylus mystacinus.  Catalogue of American Amphibians and Reptiles, (767):1-11.

Heyer, W. R. 1967. A herpetofaunal study of an ecological transect through the Cordilla de Tilarán, Costa Rica.  Copeia, 1967:259-271.

Heyer, W. R. 1968. The proper name for the type-species of the genus Leptodactylus.  Copeia, 1968:160-162.

Heyer, W. R. 1969. Studies on the genus Leptodactylus (Amphibia, Leptodactylidae).  III.  A redefinition of the genus Leptodactylus and a description of a new genus of leptodactylid frogs.  Contributions in Science, Los Angeles County Museum of Natural History, (155):1-14.

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.

Heyer, W. R.  1988.  On frog distribution patterns East of the Andes.  Pages 245-273 in P. E. Vanzolini & W. R. Heyer (editors), Proceedings of a Workshop on Neotropical Distribution Patterns Held 12-16 January 1987.  Academia Brasileira de Ciências.

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., & C. L. Barrio-Amorós.  2009.  The advertisement calls of two sympatric frogs, Leptodactylus lithonaetes (Amphibia: Anura: Leptodactylidae) and Pristimantis vilarsi (Amphibia: Anura: Strabomantidae).  Proceedings of the Biological Society of Washington, 122(3):282-291.

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.

Heyer, W. R., R. O. de Sá, & S. Muller. 2005. The enigmatic distribution of the Honduran endemic Leptodactylus silvanimbus (Amphibia: Anura: Leptodactylidae). Pages 81-101 in M. A. Donnelly, B. I. Crother, C. Guyer, M. H. Wake, & M. E. White (editors), Ecology and Evolution in the Tropics: A Herpetological Perspective. University of Chicago Press: Chicago & London. xv + 675 pages.

Heyer, W. R., R. O. de Sá, & A. Rettig.  2005.  Sibling species, advertisement calls, and reproductive isolation in frogs of the Leptodactylus pentadactylus species cluster (Amphibia, Leptodactylidae).  In N. Ananjeva & O. Tsinenko (editors), Herpetologica Petropolitana.  Proceedings of the 12th Ordinary General Meeting of the Societas Europaea Herpetologica, August 12-16, 2003, St. Petersburg, Russia.  Russian Journal of Herpetology, 12(Supplement):35-39.

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

Heyer, W. R., J. M. García-López, & A. J. Cardoso.  1996.  Advertisement call variation in the Leptodactylus mystaceus species complex (Amphibia: Leptodactylidae) with a description of a new sibling species.  Amphibia-Reptilia, 17:7-31.

Heyer, W. R., and A. A. Giaretta.  2009.  Advertisement calls, notes on natural history, and distribution of Leptodactylus chaquensis (Amphibia: Anura: Leptodactylidae) in Brasil.  Proceedings of the Biological Society of Washington, 122(3):292-305.

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

Heyer, W. R., & M. M. Heyer.  2002.   Leptodactylus elenae.  Catalogue of American Amphibians and Reptiles, (742):1-5.

Heyer, W. R., & M. M. Heyer.  2004.  Leptodactylus furnarius.  Catalogue of American Amphibians and Reptiles, (785):1-5.

Heyer, W. R., & M. M. Heyer. 2006. Leptodactylus knudseni. Catalogue of
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Heyer, W. R., M. M. Heyer, & R. O. de Sá. 2008. Leptodactylus cunicularius.  Catalogue of American Amphibians and Reptiles, (845):1-5.

Heyer, W. R., M. M. Heyer, & R. O. de Sá. 2010. Leptodactylus savagei..Catalogue of American Amphibians and Reptiles, (867):1-19.

Heyer, W. R., M. M. Heyer, & R. O. de Sá. 2010. Leptodactylus syphax. Catalogue of American Amphibians and Reptiles, (868):1-9.

Heyer, W. R., & F. A. Juncá.  2003.  Leptodactylus caatingae, a new species of frog from eastern Brazil (Amphibia: Anura: Leptodactylidae).  Proceedings of the Biological Society of Washington, 116(2):317-329.

Heyer, W. R., R. W. McDiarmid, & D. L. Weigmann.  1975.  Tadpoles, predation and pond habitats in the tropics.  Biotropica, 7:100-111.

Heyer, W. R., & L. R. Maxson.  1982.  Distributions, relationships, and zoogeography of lowland frogs:  The Leptodactylus complex in South America, with special reference to Amazonia.  Pages 375-388 in Ghillean T. Prance (ed.), Biological Diversification in the Tropics.  Columbia University Press.

Heyer, W. R., & L. R. Maxson.  1982.  Neotropical frog biogeography:  Paradigms and problems.  American Zoologist, 22:397-410.

Heyer, W. R., & V. R. Morales.  1995.  The advertisement call of the leptodactylid frog Leptodactylus griseigularis.  Amphibia-Reptilia, 16:91-92.

Heyer, W. R., & J. A. Peters.  1971.  The frog genus Leptodactylus in Ecuador.  Proceedings of the Biological Society of Washington, 84:163-170.

Heyer, W. R., & W. F. Pyburn.  1983.  Leptodactylus riveroi, a new frog species from Amazonia, South America (Anura: Leptodactylidae).  Proceedings of the Biological Society of Washington, 96:560-566.

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.

Heyer. W. R., A. S. Rand, C. A. G. Cruz, O. L. Peixoto, & C. E. Nelson. 1990.  Frogs of Boracéia.  Arquivos de Zoologia, 31:231-410. 

Heyer, W. R., & Y. R. Reid. 2003. Does advertisement call variation coincide with genetic variation in the genetically diverse frog taxon currently known as Leptodactylus fuscus (Amphibia: Leptodactylidae)? Anais da Academia Brasileira de Ciências, 75:39-54.

Heyer, W. R., & N. J. Scott, Jr.  2006.  The advertisement call of Leptodactylus laticeps (Amphibia, Anura, Leptodactylidae):  Predatory aural luring?  Herpetological Natural History, 9(2):189-194.

Heyer, W. R., & P. A. Silverstone.  1969.  The larva of Leptodactylus hylaedactylus (Amphibia, Leptodactylidae).  Fieldiana: Zoology, 51:141-145.

Heyer, W. R., & A. S. Thompson.  2000.  Leptodactylus rugosus.  Catalogue of American Amphibians and Reptiles, (708):1-5.

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Larson, P. M., & R. O. de Sá. 1998.  Chondrocranial morphology of Leptodactylus larvae (Leptodactylidae: Leptodactylinae):  Its utility in phylogenetic reconstruction.  Journal of Morphology, 238:287-305.

Larson, P. M., R. O. de Sá, & D. Arrieta.  2003.  Chondrocranial, hyobranchial and internal oral morphology in larvae of the basal bufonid genus Melanophryniscus (Amphibia: Anura).  Acta Zoologica (Stockholm), 84:145-154.

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Pyburn, W. F., & W. R. Heyer.  1975.  Identity and call of the frog, Leptodactylus stenodema.  Copeia, 1975(3):585-587.

Radice, G., M. K. Boggiano, M. DeSantis, P. M. Larson, J. Oppong, M. Smetanick, T. Stevens, J. Tripp, R. Weber, M. Kerckhove, & R. O. de Sá.  1999.  Three-dimensional reconstructions of tadpole chondrocrania from histological sections.  Virginia Journal of Science, 50:227-235.

Straughan, I. R., & W. R. Heyer.  1976.  A functional analysis of the mating calls of the neotropical frog genera of the Leptodactylus complex (Amphibia, Leptodactylidae).  Papéis Avulsos de Zoologia, 19(23):221-245.

Vanzolini, P. E., & W. R. Heyer.  1985.  The American herpetofauna and the interchange.  Pages 475-487 in F. G. Stehli & D. S. Webb (eds.), The Great American Biotic Interchange.  Plenum Press.

Vaz-Ferreira, R., R. O. de Sá, F. Achaval, & A. Gehrau.  1984.  Leptodactylus podicipinus (Cope, 1862) y Leptodactylus chaquensis Cei, 1950 (Anura, Leptodactylidae), en el Uruguay.  Boletín de la Sociedad Zoológica del Uruguay, Segunda Época, 2:72-77.

Wassersug, R. J., & W. R. Heyer.  1988.  A survey of internal oral features of leptodactylid larvae (Amphibia: Anura).  Smithsonian Contributions to Zoology, (457):1-99.  [This article is broken up into four pdf files.  pdf file 1.  pdf file 2.  pdf file 3.  pdf file 4.]

Wynn, A., & W. R. Heyer.  2001.  Do geographically widespread species of tropical amphibians exist?  An estimate of genetic relationships within the Neotropical frog Leptodactylus fuscus (Schneider 1799) (Anura Leptodactylidae).  Tropical Zoology, 14:255-285.

Yanek, K., W. R. Heyer, & R. O. de Sá.  2006. Genetic resolution of the enigmatic Lesser Antillean distribution of the frog Leptodactylus validus (Anura, Leptodactylidae).  South American Journal of Herpetology, 1(3):192-201.