Dr. Rafael O. de Sa
Professor of Biology

The overall research framework in our lab is on evolutionary biology of anurans. Within this framework we have active research projects on molecular and morphological phylogenetic, systematics, taxonomy, and development. Our lab maintains active collaborations that involved reciprocal research visits with colleagues in the US as well as with several labs in South America, Europe, and Africa. These collaborations provide unique academic as well as cultural exchange opportunities.

In addition to the anuran research program, we are also collaborating with colleagues at the University of Uruguay on understanding the evolutionary biology of annual fishes.

Current Lab Members

Drew Braet
Travis Cuddy
Hanna Moreland

Grants and Fellowships
Grants: National Science Council of Spain, "Genetic and Bioacoustic Characterization of Endemic Threatened Species in a Biodiversity Hotspot: The Amphibians of the Tropical Andes."

Revision of the poorly known frog genus Chiasmocleis (Anura, Microhylidae). National Science Foundation.  Initiated in 2012.

Using sequencing in undergraduate research. 2004. LI-Cor Genomics Education Matching Funds Program.

Diversity beyond morphology: A revision of the frog genus Leptodactylus (Amphibia, Anura, Leptodactylidae). 2004-2008. National Science Foundation.

Bringing Evolution into the curriculum for non-science majors at the University of Richmond. 2004. Association of Colleges of the South.

Evolutionary relationships within the Neotropical genus Leptodactylus. 2000-2004 National Science Foundation.

A molecular phylogeny of the frog genus Leptodactylus. 1998-2000. Thomas F. Jeffress and Kate Miller Jeffress Memorial Trust.

Timing of musculoskeletal development: A 3-dimensional approach to the analysis of form. 1995-1998 National Science Foundation.

Molecular Evolution: Integration of Theory and Practice. 1994-1996 National Science Foundation.
Distinguished Visiting Professor. National Museum of Natural History, Paris. 2007
Fulbright Distinguished Chair. University of Sao Paulo, Sao Paulo, Brazil. 2013-2014.

Building conservation capacity for a changing Amazonia.” PI: David Salisbury (UR), CO-PIs: M. Finley-Brook (UR), R.O. de Sá (UR), and P. Smallwood (UR). 2013-2015. HED-USAID

Distinguished Educator, University of Richmond. 1998
Selected Publications

* denotes student author

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

de Sá, R. O. and C. Swart*. 1999. Development of the suprarostral plate of pipoid frogs. Journal of Morphology 240:143-153.

Camargo*, A., de Sá, R. O. and 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.

de Sá, R., W.R. Heyer, and A. Camargo*. 2005. A phylogenetic analysis of Vanzolinius (Amphibia, Anura, Leptodactylidae): Taxonomic and life histroy implications. Arquivos do Museu de Historia Natural do Rio de Janeiro, 63(4):1-20.

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, and W. C. Wheeler. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History. 297:1-370, +71 figs, 5 tbls, 7 Append.

Yanek*, K., W.R. Heyer, and 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.

Loader, S. P., G. J. Measey, R. O. de Sá, and P. K. Malonza. 2009. A new brevicipitid species (Brevicipitidae: Callulina) from the fragmented forests of the Taita Hills, Kenya. Zootaxa, 2123:55-68.

Heyer, W. R, M. Heyer, and R. O. de Sá. 2009. Leptodactylus syphax. Catalogue of American Amphibians and Reptiles, 868:1-9.

do Nascimento*, Filipe A,. C. M. G. de Lima*, G. O. Skuk, and R. O. de Sá. 2009. The tadpole of Hypsiboas atlanticus (Anura: Hylidae) from Northeastern Brazil. Iheringia, 99(4):431-436.

Heyer, W. R., M. M. Heyer, & R. O. de Sá. Leptodactylus savagei. 2009. Catalogue of American Amphibians and Reptiles, 867:1-19.

Lavilla, E. O., J. J. Nunez, F. E. Rabanal, J. A Langone, and R. O. de Sá. 2010. The identity of Zachaenus roseus Cope, 1890 (Anura:species inquirenda). Zootaxa, 2561:49-58.

Lavilla, E.O., J.A. Langone, J.M. Padial, and R.O. de Sá. 2010. The identity of the crackling, luminescent frog of Suriname (Rana typhonia Linn., 1758) (Amphibia, Anura). Zootaxa, 2671:17-30.

do Nascimento*, Filipe A. C., B. S. Lisboa*, G. O. Skuk, and R. O. de Sá. 2010. Description of the tadpole of Proceratoprhys renalis (Miranda-Ribeiro, 1920) (Anura: Cycloramphidae). South American Journal of Herpetology, 5(3):241-248.

Greenbaum, E., E.N. Smith, R.O. de Sá. 2011. Molecular systematics of the Middle American genus Hypopachus (Anura: Microhylidae). Molecular Phylogenetics and Evolution, 61:265-277.

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.

Heyer, M. M., Heyer, W. R. and R. O. de Sá. 2012. Leptodactylus pentadactylus. Catalogue of American Amphibians and Reptiles, 887: 1-48.

Streicher, J. W., Cox, C. L., Campbell, J.A., Smith, E. N., and de Sá, R. O. 2012 Rapid range expansion in the Great Plains narrow-mouthed toad (Gastrophryne olivacea) and a revised taxonomy for North American microhylids. Molecular Phylogenetics and Evolution, 64(2012):645-653.

Gower, D., Doherty-Bone, T. M., Aberra, R. K., Mengistu, A., Schwaller, S., Menegon, M., de Sá, R., Saber, S. A., Cunningham, A. A., and Loader, S. P. 2012. High prevalence of the amphibian chytrid fungus (Batrachochytrium dendrobatidis) across multiple taxa and localities in the highlands of Ethiopia. Herpetological Journal, 22: 225–233.

Fabrezi M., Quinzio S. I., Goldberg, J., and de Sá, R. O. 2012. The development of Dermatonotus muelleri (Anura: Gastrophryninae). Journal of Herpetology, 46(3):363-380.

de Sá, R.O., Streicher, J.W., Sekonyela R*., Forlani, M.C., Loader, S.P., Greenbaum, E., Richards, S., and Haddad, CFB. 2012. Molecular phylogeny of microhylid frogs (Anura: Microhylidae) with emphasis on relationships among New World genera. BMC Evolutionary Biology, 2012 (241):1-40.

Gower, D., Aberra, R. K., Mengistu, A., Schwaller, S., Menegon, M., de Sá, R. O, Saber, S. A., Cunningham, A. A., and Loader, S. P. 2013. Long-term data for endemic frog genera reveal potential conservation crisis in the Bale Mountains, Ethiopia. Fauna & Flora International, Oryx, 47(1), 59–69.

Brandão, R., Alvarez, G., and de Sá, R.O. 2013. The advertisement call of the poorly known Leptodactylus tapiti (Anura, Leptodactylidae)Zootaxa, 3616(3):284-286.

Lavilla, E.O., Caramaschi, U., Langone, J. A., Pombal Jr., J., and de Sá, R.O. 2013. The identity of Rana margaritifer Laurenti, 1768 (Anura: Bufonidae). Zootaxa, 3646(3):251-264.

Fouquet, A., Maronna, M.M., Verdade, V.K., de Sá, R.O., and Trefaut-Rodrigues, M., 2013. Unexpected phylogenetic positions of the genera Rupirana and Crossodactylodes reveal insights into the biogeography and reproductive evolution of Leptodactylidae. Molecular Phylogenetics and Evolution, 67(2):445-457.

Loader, S.P., Ceccarelli, S., Wilkinson, M., Menegon, M., Mariaux J., de Sá, R.O, Howell, K.M., and Gower, D.J. 2013. Species boundaries and biogeography in the East African Torrent Frogs (Petropedetes: Petropeditidae). African Journal of Herpetology, 62(1):40-48.

Forlani*, M.C., Vinicius, C., Dias, I.R., Ruas, D., Tonini*, J.F.R., de Sá, R.O.  2013. The advertisement calls and distribution records of two sympatric species of Chiasmocleis (Méhely 1904) (Anura, Microhylidae) from the Atlantic forest. South American Journal of Herpetology, 8(1):46-51.

Minelli, A.,   Ohler, A., Aescht, E.,  Bauer, A.  Bonato, L., Bour, R., De Carvalho, M.R. De Sá, R.O., Foissner, W., Fusco, G.,  González-Oreja, J.A., Grandcolas, P., Iverson, J., Leschen, R.,  Löbl, I., Maruzzo, D., Nel, A.,  Nemésio, A., Riff, D., Dubois, A. 2013. Pushing Taxonomy to Extinction? Science, 339(6118):413-416.

Do Nascimento*,.F. A. C., Mott, T., Langone, J. A., Davies, C. A., and de Sá, R. O. 2013. Chondrocranium and tadpole oral cavity of four Odontophrynus species (Anura, Odontophrynidae) with a review of the larval external morphology of the genus. Zootaxa, 3700 (1):140-158.

Magalhaes*, F. M, Garda, A. A., Amado, T., and de Sá, R. O. 2013. The tadpole of Leptodactylus caatingae Heyer & Junca 2003 (Anura: Leptodactylidae): external morphology, internal anatomy, and natural history. South American Journal of Herpetology, 8(3):203-210.

Siu-Ting,  K., Gower J.D, Pisani  D., Kassahun  R., Gebresenbet  F., Menegon  M., Mengistu A. A., Saber A S., de Sá  R., Wilkinson  M., Loader P S. 2014. Evolutionary relationships of the Critically Endangered frog Ericabatrachus baleensis Largen, 1991 with notes on incorporating previously unsampled taxa into large-scale phylogenetic analyses. BMC Evolutionary Biology, 2014, 14:1-44.

Laufer, G., Gobel, N., Mauntone, J. M., Galán, M., and de Sá, R. O. 2014. First report of satellite males during breeding in Leptodactylus latrans (Amphibia: Anura)Cuadernos Herpetologia, 28(1):37-38.

Tonini, J. F. R., Forlani, M. C., and de Sá, R. O. 2014. A new species of Chiasmocleis (Microhylidae, Gastrophryninae) from the Atlantic Forest of Espirito Santo State, Brazil. ZooKeys, 428:109-113.

Berois, N., Arezo, J. M., and de Sá, R. O. 2014. The Neotropical genus Austrolebias: An emerging model of annual killifishes. Cell and Developmental Biology, 3(2):1-9.

Loader, P. S., Ceccarelli, F. S., Menegon, M., Howell, K.M., Kassahun, R.,  Mengistu, A. A., Saber, S. A, Gebresenbet, F., de Sa R. O., Davenport, T.B.D., Larson, J., Müller, H., Wilkinson, M., and Gower, D.J.  2014. Persistence and stability of Eastern Afromontane forests: evidence from brevicipitid frogs. Journal of Biogeography, 41(9):1781-1792.

de Sá,  R.O., Grant, T., Camargo, A., Heyer,W. R., Ponssa,  M. L.,  and Stanley, E*.  2014. Systematics of the Neotropical genus Leptodactylus Fitzinger, 1826 (Anura: Leptodactylidae): Phylogeny, the Relevance of Non-Molecular Evidence, and Species accounts. South American Journal of Herpetology, 9 (Special Issue 1):1-128.

Additional Publications

Heyer, M. H., W. R. Heyer, & R. O. de Sá. 2009. Bibliography of frogs of the Leptodactylus clade (Amphibia, Anura, Leptodactylidae – Adenomera, Hydrolaetare, Leptodactylus, Lithodytes). I. Bibliography. Smithsonian Herpetological Information Series, 677 pgs.

Heyer, M. H., W. R. Heyer, & R. O. de Sá. 2009. Bibliography of frogs of the Leptodactylus clade (Amphibia, Anura, Leptodactylidae – Adenomera, Hydrolaetare, Leptodactylus, Lithodytes). II. Indices. Smithsonian Herpetological Information Series, 167 pgs.

Ph.D., University of Texas-Austin 1991
M.Sc., The University of Kansas 1987
B.Sc., University of Uruguay
Contact Information
B115 Gottwald Science Center
(804) 289-8542
(804) 289-8233 (Fax)
Areas of Expertise
Evolutionary Biology and Biodiversity
Phylogenetic, Systematics, and Taxonomy
Developmental Biology