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Type: Article
Published: 2021-08-17
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A reappraisal of Bufo levicristatus Boettger, 1885 (Anura: Bufonidae), a long forgotten toad from Paraguay

Laboratorio de Genética Evolutiva “Claudio Juan Bidau”, Instituto de Biología Subtropical (CONICET-UNaM), Facultad de Ciencias Exactas Químicas y Naturales, Universidad Nacional de Misiones, Félix de Azara 1552, Posadas, Misiones, Argentina
Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, 199034, St. Petersburg, Russia
Instituto de Investigación Biológica del Paraguay, Del Escudo 1607, 1429 Asunción, Paraguay
Sección Herpetología, Museo Nacional de Historia Natural, Montevideo
Laboratorio de Genética Evolutiva “Claudio Juan Bidau”, Instituto de Biología Subtropical (CONICET-UNaM), Facultad de Ciencias Exactas Químicas y Naturales, Universidad Nacional de Misiones, Félix de Azara 1552, Posadas, Misiones, Argentina
Amphibia Bufo ornatus Bufo scitulus nomen oblitum nomen protectum species inquirenda

Abstract

In this work, we show that Bufo levicristatus Boettger, 1885 is a senior synonym of Bufo scitulus Caramaschi & Niemeyer, 2003, and not a junior synonym of Bufo ornatus Spix, 1824, as previously considered. In addition, we present evidence that Bufo scitulus Caramaschi & Niemeyer, 2003 complies with the International Code of Zoological Nomenclature requirements for a reversal of precedence over the name Bufo levicristatus Boettger, 1885, and so the name Bufo scitulus is to be maintained for this species.

 

References

  1. Aguayo, R., Lavilla, E.O., Vera Candioti, M.F. & Camacho, T. (2009) Living in fast-flowing water: Morphology of the gastromyzophorous tadpole of the bufonid Rhinella quechua (R. veraguensis group). Journal of Morphology, 270 (12), 1431–1442. https://doi.org/10.1002/jmor.10768
    Anonymous. (1885) Original Mittheilungen aus der ethnologischen Abtheilung der Königlichen Museen zu Berlin. Verlag von W. Spemann, Berlin, 56 pp.
    Ávila, R.W., Pansonato, A. & Strüssmann, C. (2010) A new species of the Rhinella margaritifera group (Anura: Bufonidae) from Brazilian Pantanal. Zootaxa, 2339 (1), 57–68. https://doi.org/10.11646/zootaxa.2339.1.3
    Ávila, R.W., Morais, D.H., Perez, R., Pansonato, A., de Carvalho, V.T., Rojas, R.R., Gordo, M. & Farias, I.P. (2020) A new species of the Rhinella margaritifera (Laurenti 1768) species group (Anura, Bufonidae) from southern Brazilian Amazonia. Zootaxa, 4868 (3), 368–388. https://doi.org/10.11646/zootaxa.4868.3.3
    Baldissera, F.A., Caramaschi, U. & Haddad, C.F.B. (2004) Review of the Bufo crucifer species group, with descriptions of two new related species (Amphibia, Anura, Bufonidae). Arquivos do Museu Nacional do Rio de Janeiro, 62 (3), 255–282.
    Blotto, B., Pereyra, M.O. & Baldo, D. (2014) The tadpole of Rhinella azarai (Gallardo, 1965) with comments on the larval morphology in the Rhinella granulosa species group (Anura: Bufonidae). Journal of Herpetology, 48 (3), 434–438. https://doi.org/10.1670/12-121
    Boettger, O. (1885a) Liste von Reptilien und Amphibien aus Paraguay. Zeitschrift fur Naturforschung, 58, 213–248.
    Boettger, O. (1885b) Berichtigung der Liste von Reptilien und Amphibien am Paraguay von Dr. O. Boettcher. Zeitschrift für Naturwissenschaften, Halle, 58, 436–437.
    Boulenger, G.A. (1886) A synopsis of the reptiles and batrachians of the Province Rio Grande do Sul, Brazil. Annals and Magazines of Natural History, London, 18, 423–445.
    Brandão, R.A., Maciel, N.M. & Sebben, A. (2007) A new species of Chaunus from central Brazil (Anura; Bufonidae). Journal of Herpetology, 41 (2), 309–316. https://doi.org/10.1670/0022-1511(2007)41[309:ANSOCF]2.0.CO;2
    Brandão, R.A., Fenker, J., Lopes, B.E.P.D.C., de Sena, V.M.D.A. & Vasconcelos, B.D. (2020) Diet of terrestrial anurans in an ephemeral and simplified habitat during the dry season in the Brazilian Cerrado. Ethology Ecology & Evolution, 32 (6), 1–24. https://doi.org/10.1080/03949370.2020.1755373
    Brusquetti, F. & Lavilla, E.O. (2006) Lista comentada de los anfibios de Paraguay. Cuadernos de Herpetología, 20 (2), 3–79.
    Borges da Rocha, S., Sugai, J.L.M.M., Martins, F.I. & Sabino, J. (2015) Anfíbios. Guia do Mato Grosso do Sul. Natureza em foco, Campo Grande, 45 pp.
    Cabral, H., Casagranda, M.D., Brusquetti, F., Netto, F., Ferreira, V. & Lavilla, E.O. (2020) Multiscale endemism analysis for amphibians of Paraguay. Herpetological Journal, 30 (1), 35–46. https://doi.org/10.33256/hj30.1.3546
    Caldwell, J.P. (1991) A new species of toad in the genus Bufo from Pará, Brazil, with an unusual breeding site. Papéis Avulsos de Zoologia, 37 (26), 389–400.
    Camera, B., Krinski, D. & Calvo, I. (2014) Diet of the Neotropical frog Leptodactylus mystaceus (Anura: Leptodactylidae). Herpetology Notes, 7, 31–36.
    Caramaschi, U. & Niemeyer, H. (2003) Nova espécie do complexo de Bufo margaritifer (Laurenti, 1768) do estado do Mato Grosso do Sul. Brasil (Amphibia, Anura, Bufonidae). Boletim do Museo Nacional, 501, 1–16.
    Caramaschi, U. & Pombal Jr., J.P. (2006) A new species of Rhinella Fitzinger, 1826 from the Atlantic rain forest, eastern Brazil (Amphibia, Anura, Bufonidae). Papéis Avulsos de Zoologia, 46, 251–259. https://doi.org/10.1590/S0031-10492006002300001
    Cei, J.M. (1962) Batracios de Chile. Universidad de Chile, Santiago, 124 pp.
    Chaparro, J.C., Pramuk, J.B. & Gluesenkamp, A.G. (2007) A new species of arboreal Rhinella (Anura: Bufonidae) from cloud forest of southeastern Peru. Herpetologica, 63 (2), 203–212. https://doi.org/10.1655/0018-0831(2007)63[203:ANSOAR]2.0.CO;2
    Cochran, D.M. (1955) Frogs of southeastern Brazil. Bulletin of the United States National Museum, 206, 1–423.
    de Oliveira, J.E. (2007) Os primeiros passos em direção a uma arqueologia pantaneira: de Max Schmidt e Branka Susink a outras interpretaçoes sobre povos indígenas nas terras baixas do Pantanal. Revista de Arqueologia, 20 (1), 83–115.
    Dinerstein, E., Olson, D.M., Graham, D.J., Webster, A.L., Primm, S.A., Bookbinder, M.P. & Ledec, G. (1995) A conservation assessment of the terrestrial ecoregions of Latin America and the Caribbean. World Bank Group, Washington, D.C., 129 pp.
    dos Santos, S.P., Ibáñez, R. & Ron, S.R. (2015) Systematics of the Rhinella margaritifera complex (Anura, Bufonidae) from western Ecuador and Panama with insights in the biogeography of Rhinella alata. ZooKeys, 501, 109–145. https://doi.org/10.3897/zookeys.501.8604
    Duellman, W.E. (1970) The hylid frogs of Middle America. Monograph of the Museum of Natural History, University of Kansas, 1970, 1–753.
    Duellman, W.E. & Schulte, R. (1992) Description of a new species of Bufo from northern Peru with comments on phenetic groups of South American toads (Anura: Bufonidae). Copeia, 1992, 162–172. https://doi.org/10.2307/1446548
    Duméril, A.M.C. & Bibron, G. (1841) Erpétologie genérale ou Histoire naturelle complète des reptiles. Vol. 8. Librarie Enclyclopedique de Roret, Paris, 780 pp.
    Ferrão, M., Lima, A.P., Ron, S.R., dos Santos, S.P.L. & Hanken, J. (2020) New species of leaf-litter toad of the Rhinella margaritifera species group (Anura: Bufonidae) from Amazonia. Copeia, 108 (4), 967–986. https://doi.org/10.1643/CH2020043
    Fouquet, A., Gaucher, P., Blanc, M. & Vélez–Rodriguez, C.M. (2007) Description of two new species of Rhinella (Anura: Bufonidae) from the lowlands of the Guiana Shield. Zootaxa, 1663 (1), 17–32. https://doi.org/10.11646/zootaxa.1663.1.2
    Günther, A. (1858) Neue Batrachier in der Sammlung des britischen Museums. Archiv für Naturgeschichte, 24 (1), 319–328.
    Haad, M.B., Vera Candioti, F. & Baldo, D. (2011). Shape variation in lentic and lotic tadpoles of Melanophryniscus (Anura: Bufonidae). Studies on Neotropical Fauna and Environment, 46 (2), 91–99. https://doi.org/10.1080/01650521.2011.593124
    Hoogmoed, M.S. (1986) Biosystematic studies of the Bufo “typhonius” group. A preliminary progress report. In: Roček, Z. (Ed.), Studies in Herpetology. Charles University, Prague, pp. 147–150.
    Hoogmoed, M.S. (1990) Biosystematics of South American Bufonidae, with special reference to the Bufo “typhonius” group. In: Peters, G. & Hutterer, R. (Eds.), Vertebrates in the tropics. Museum Alexander Koening, Bonn, pp. 113–123.
    ICZN [International Commission on Zoological Nomenclature] (1999) International Code of Zoological Nomenclature. Fourth edition. International Trust for Zoological Nomenclature, London, 306 pp.
    Jiménez de la Espada, M. (1875) Vertebrados del viaje al Pacífico verificado de 1862 a 1865 por una comisión de naturalistas enviada por el gobierno español. Batracios. Imprenta de Miguel Ginesta, Madrid, 208 pp.
    Kelehear, C., Brown, G.P. & Shine, R. (2012) Size and sex matter: infection dynamics of an invading parasite (the pentastome Raillietiella frenatus) in an invading host (the cane toad Rhinella marina). Parasitology, 139 (12), 1596–1604. https://doi.org/10.1017/S0031182012000832
    Köhler, J. (2000) Amphibian diversity in Bolivia: A study with special reference to montane forest regions. Bonner Zoologische Monographien, 48, 1–243.
    Kolenc, F., Borteiro, C., Cotichelli, L., Baldo, D., Martínez Debat, C. & Vera Candioti, F. (2013) The tadpole and karyotype of Rhinella achavali (Anura: Bufonidae). Journal of Herpetology, 47 (4), 599–606. https://doi.org/10.1670/13-028
    Laurenti, J.N. (1768) Specimen medicum, exhibens synopsis reptilium emendatam, cum experimentis circa venena et antidote reptilium austriacorum, Wien, Austria. Zeitschrift für Feldherpetologie. Joan Thomae nob. de Trattnern, Viennae, 214 pp.
    Lavilla, E.O. & Brusquetti, F. (2018) On the identity of Bufo diptychus Cope, 1862 (Anura: Bufonidae). Zootaxa, 4442 (1), 161–170. https://doi.org/10.11646/zootaxa.4442.1.9
    Lavilla, E.O., Caramaschi, U., Langone, J.A., Pombal Jr., J.P. & de Sá, R.O. (2013) The Identity of Rana margaritifera Laurenti, 1768 (Anura, Bufonidae). Zootaxa, 3646 (3), 251–264. https://doi.org/10.11646/zootaxa.3646.3.4
    Leão, A.T. & Cochran, D.M. (1952) Revalidation and re-description of Bufo ocellatus Günther, 1858 (Anura: Bufonidae). Memórias do Instituto Butantan, 24 (2), 271–280.
    Lima, A.P., Menin, M. & Araújo, M.C. (2007) A new species of Rhinella (Anura: Bufonidae) from Brazilian Amazon. Zootaxa, 1663 (1), 1–15. https://doi.org/10.11646/zootaxa.1663.1.1
    Lötters, S. & Köhler, J. (2000) A new toad of the Bufo typhonius complex from humid montane forests of Bolivia. Spixiana, 23 (3), 293–303.
    Maia-Carneiro, T., Kiefer, M.C., Van Sluys, M. & Duarte Rocha, C.F. (2013) Feeding habits, microhabitat use, and daily activity period of Rhinella ornata (Anura, Bufonidae) from three Atlantic rainforest remnants in southeastern Brazil. North-Western Journal of Zoology, 9 (1), 157–165.
    Maragno, F.P. & Souza, F.L. (2011) Dieta de Rhinella scitula (Anura, Bufonidae): la importancia de la variación estacional y la talla corporal. Revista Mexicana de Biodiversidad, 82 (3), 879–886.
    Matavelli, R., Campos, A.M., Silva, G.R. & Andrade, G.V. (2014) First record of Rhinella ocellata (Günther, 1858) (Bufonidae) for the State of Maranhão, northeastern Brazil. Check List, 10 (2), 432–433. https://doi.org/10.15560/10.2.432
    Melin, D. (1941) Contribution to the knowledge of the Amphibia of South America. Göteborgs Kungl Vetenskaps-och Vitterhets-samhälles Handlingar Serien B, Matematiska och Naturvetenskapliga Skrifter, 1, 1–71.
    Menin, M., Rodrigues, D.J. & Lima, A.P. (2006) The tadpole of Rhinella proboscidea (Anura: Bufonidae) with notes on adult reproductive behavior. Zootaxa, 1258 (1), 47–56. https://doi.org/10.11646/zootaxa.1258.1.4
    Mercês, E.A., Juncá, F.A. & Casal, F.S.C. (2009) Girinos de três espécies do gênero Rhinella Fitzinger, 1826 (Anura–Bufonidae) ocorrentes no estado da Bahia, Brasil. Sitientibus Série Ciências Biologicas, 9, 133–138. https://doi.org/10.13140/2.1.4860.7363
    Mijares-Urrutia, A. & Arends, A. (2001) A new toad of the Bufo margaritifer complex (Amphibia: Bufonidae) from northwestern Venezuela. Herpetologica, 57 (4), 523–531.
    Milto, K.D. (2010) On the identity of Bufo levicristatus Boettger, 1885. Russian Journal of Herpetology, 17 (4), 257–258. https://doi.org/10.30906/1026-2296-2010-17-4-257-258
    Milto, K.D. & Barabanov, A.V. (2011) An annotated catalogue of the amphibian types in the collection of the Zoological Institute, Russian Academy of Sciences, St. Petersburg. Russian Journal of Herpetology, 18 (2), 137–153. https://doi.org/10.30906/1026-2296-2011-18-2-137-153
    Moravec, J., Lehr, E., Cusi, J.C., Córdova, J.H. & Gvoždík, V. (2014) A new species of the Rhinella margaritifera species group (Anura, Bufonidae) from the montane forest of the Selva Central, Peru. ZooKeys, 371, 35–56. https://doi.org/10.3897/zookeys.371.6580
    Motte, M., Núñez, K., Cacciali Sosa, P., Brusquetti, F., Scott, N.J. & Aquino, A.L. (2009) Categorización del estado de conservación de los anfibios y reptiles de Paraguay. Cuadernos de Herpetología, 23, 5–18.
    Motte, M., Zaracho, V., Caballero-Gini, A., Ferreira-Riveros, M., Nardelli, L.R., Coronel-Bejarano, D., Netto, F., Carosini, A., Rojas, V., Bueno, D., Cabral, H. & Martínez, N. (2019) Estado de conservación y lista roja de los anfibios del Paraguay. Boletín del Museo Nacional de Historia Natural del Paraguay, 23 (1), 1–66.
    Myers, G.S. & Carvalho, A.L. (1945). Notes on some new or little-known Brazilian amphibians, with an examination of the history of the Plata salamander, Ensatina platensis. Boletim do Museu Nacional, 35, 1–24.
    Narvaes, P. & Rodrigues, M.T. (2009) Taxonomic revision of Rhinella granulosa species group (Amphibia, Anura, Bufonidae), with a description of a new species. Arquivos de Zoologia, 40 (1), 1–73. https://doi.org/10.11606/issn.2176-7793.v40i1p1-73
    Oliveira, M.I.R.R., Weber, L.N. & Napoli, M.F. (2014) Chondrocranial and hyobranchial morphology in larvae of the genus Rhinella Fitzinger, 1826 (Amphibia, Anura, Bufonidae). Herpetological Journal, 24 (4), 229–237.
    Pereyra, M.O., Vera Candioti, M.F., Faivovich, J. & Baldo, D. (2015) Egg clutch structure of Rhinella rumbolli (Anura: Bufonidae) from Argentina with a review of reproductive diversity in Rhinella. Salamandra, 51 (2), 161–170.
    Pereyra, M.O., Baldo, D., Blotto, B.L., Iglesias, P.P., Thomé, M.T., Haddad, C.F.B., Barrio-Amorós, C., Ibañez, R. & Faivovich, J. (2016) Phylogenetic relationships of toads of the Rhinella granulosa group (Anura: Bufonidae): A molecular perspective with comments on hybridization and introgression. Cladistics, 32 (1), 36–53. https://doi.org/10.1111/cla.12110
    Pereyra, M.O., Blotto, B.L., Baldo, D., Chaparro, J.C., Ron, S., Elias-Costa, A., Iglesias, P.P., Venegas, P., Thomé, M.T.C., Ospina-Sarria, J.J., Maciel, N., Rada, M., Kolenc, F., Borteiro, C., Rivera-Correa, M., Rojas-Runjaic, F., Moravec, J., de la Riva, I., Wheeler, W.C., Castroviejo, S., Grant, T., Haddad, C.F.B. & Faivovich, J. (2021) Evolution in the genus Rhinella: a total evidence phylogenetic analysis of neotropical true toads (Anura: Bufonidae). Bulletin of the American Museum of Natural History, 447 (1), 1–156. https://doi.org/10.1206/0003-0090.447.1.1
    Pramuk, J.B. (2006) Phylogeny of South American Bufo (Anura: Bufonidae) inferred from combined evidence. Zoological Journal of the Linnean Society, 146 (3), 407–452. https://doi.org/10.1111/j.1096-3642.2006.00212.x
    Rezende Oliveira, S., Silva, D., Fachi, M. & Morais, A.R. (2017) Predation on Rhinella mirandaribeiroi (Gallardo, 1965) (Anura; Bufonidae) by a snake from Central Brazil. Herpetology Notes, 10, 151–155.
    Roberto, I.J., Brito, L. & Cascon, P. (2011) Temporal and spatial patterns of reproductive activity in Rhinella hoogmoedi (Anura: Bufonidae) from a tropical rainforest in northeastern brazil, with the description of it’s advertisement call. South American Journal of Herpetology, 6 (2), 87–97. https://doi.org/10.2994/057.006.0207
    Santos, S., Ibáñez, R. & Ron, S. (2015) Systematics of the Rhinella margaritifera complex (Anura, Bufonidae) from western Ecuador and Panama with insights in the biogeography of Rhinella alata. ZooKeys, 501, 109–145. https://doi.org/10.3897/zookeys.501.8604
    Schmeda-Hirschmann, G., Gomez, C.V., de Arias, A.R., Burgos-Edwards, A., Alfonso, J., Rolon, M., Brusquetti, F., Netto, F., Urra, F.A. & Cárdenas, C. (2017) The Paraguayan Rhinella toad venom: Implications in the traditional medicine and proliferation of breast cancer cells. Journal of Ethnopharmacology, 199, 106–118. https://doi.org/10.1016/j.jep.2017.01.047
    Segalla, M.V., Caramaschi, U., Cruz, C.A.G., Grant, T., Haddad, C.F.B., Langone, J.A. & Garcia, P.C.A. (2016) Brazilian amphibians: list of species. Herpetologia Brasileira, 5 (2), 34–46.
    Segalla, M.V., Berneck, B., Canedo, C., Caramaschi, U., Cruz, C.A.G., Garcia, P.C.A., Grant, T., Haddad, C.F.B., Lourenço, A.C.C., Mângia, S., Mott, T., Nascimento, L.B., Toledo, L.F., Werneck, F.P. & Langone, J.A. (2021) List of brazilian amphibians. Herpetologia Brasileira, 10 (1), 118–208. https://doi.org/10.5281/zenodo.4716176
    Severo-Neto, F. & Sugai, J.L.M. (2014) Rhinella scitula (Cope’s Toad). Predation. Herpetological Review, 45, 683.
    Smith, P., Atkinson, K., Brouard, J.P. & Pheasey, H. (2016) Reserva Natural Laguna Blanca, departamento San Pedro: Paraguay’s first Important Area for the Conservation of Amphibians and Reptiles? Russian Journal of Herpetology, 23 (1), 25–34.
    Smith, P., Cacciali, P., Atkinson, K., Pheasey, H. & Motte, M. (2012) New distributional records of amphibians for departamento San Pedro, Paraguay (Amphibia). Check List, 8 (5), 903–907. https://doi.org/10.15560/8.5.903
    Souza, F.L., Prado, C.P., Sugai, J.L.M.M., Ferreira, V.L., Terra, J., Aoki, C., Landgref-Filho, P., Strüssmann, C., Ávila, R.W., Rodrigues, D.J., Albuquerque, N.R. Terra, J., Uetanabaro, M., Béda, A.F., Piatti, L., Kawashita-Ribeiro, R.A., Delatorre, M., Faggioni, G.P., Demczuk, S.D.B., Duleba, S. & de Carvalho, L.A. (2019) Diversidade de anfíbios do estado de Mato Grosso do Sul, Brasil. Iheringia, Série Zoologia, 107 (Supplement), e2017152. [2017]
    Spix, J.B. (1824) Animalia nova sive species novae Testudinum et Ranarum quas in itinere per Brasiliam annis MDCCCXVII–MDCCCXX jussu et auspiciis Maximiliani Josephi I. Bavariae Regis. F.S. Hübschmann, München, 53 pp.
    Stynoski, J.L., Trama, F.A., Patrón, F.L.R., Tapia, E. & Hoke, K.L. (2020) Reproductive ecology of the peruvian earless toad Rhinella yunga (Amphibia, Bufonidae) with descriptions of calls, tadpole, and female competition. South American Journal of Herpetology, 2020 (15), 85–96. https://doi.org/10.2994/SAJH-D-18-00030.1
    Sugai, J.L.M.M., Souza, F.L., Landgref Filho, P. & Sczesny-Moraes, E.A. (2014) Rhinella scitula (Caramaschi & Niemeyer, 2003) (Amphibia: Anura: Bufonidae): New distribution records. Check List, 10, 694–696. https://doi.org/10.15560/10.3.694
    Thominot, A. (1884) Note sur un batracien d’espèce nouvelle provenant de Panama. Bulletin de la Société Philomathique de Paris, 8, 151–152.
    Trueb, L. (1971) Phylogenetic relationships of certain neotropical toads with the description of a new genus (Anura: Bufonidae). Bulletin of Los Angeles County Museum of Natural History, 216, 1–40.
    Uetanabaro, M., Souza, F.L., Landgref, P., Beda, A.F. & Brandão, R.A. (2007) Anfíbios e répteis do Parque Nacional da Serra da Bodoquena, Mato Grosso do Sul, Brasil. Biota Neotropica, 7, 279–289.
    Vaz-Silva, W., Valdujo, P.H. & Pombal Jr., J.P. (2012) New species of the Rhinella crucifer group (Anura, Bufonidae) from the brazilian Cerrado. Zootaxa, 3265 (1), 57–65. https://doi.org/10.11646/zootaxa.3265.1.4
    Vaz-Silva, W., Maciel, N.M., Bastos, R.P. & Pombal, J.P. (2015) Revealing two new species of the Rhinella margaritifera species group (Anura, Bufonidae): an enigmatic taxonomic group of Neotropical toads. Herpetologica, 71 (3), 212–222. https://doi.org/10.1655/HERPETOLOGICA-D-14-00039
    Vélez-Rodríguez, C.M. & Ruiz-Carranza, P.M. (2002) A new species of Bufo (Anura: Bufonidae) from Colombia. Herpetologica, 58 (4), 453–462.
    Vellard, J. 1959. Estudios sobre batracios andinos. V. El género Bufo. Memorias del Museo de Historia Natural “Javier Prado”, 8, 1–48.
    Vera Candioti, F., Grosso, J., Haad, B., Pereyra, M.O., Bornschein, M.R., Borteiro, C., Costa, P., Kolenc, F., Pie, M.R., Proaño, B., Ron, S., Stanescu, F. & Baldo, D. (2016) Structural and Heterochronic Variations during the early ontogeny in toads (Anura: Bufonidae). Herpetological Monographs, 30, 79–118. https://doi.org/10.1655/0018-0831(2002)058[0453:ANSOBA]2.0.CO;2
    Weiler, A., Nuñez, K., Airaldi, K., Lavilla, E.O., Peris, S. & Baldo, D. (2013) Anfibios del Paraguay. Facultad de Ciencias Exactas y Naturales. Universidad Nacional de Asunción, Universidad de Salamanca, San Lorenz, 126 pp.
    Wied, M.A.P. (1821) Reise nach Brasilien in den Jahren 1815 bis 1817. Vol. 2. Henrich Ludwig Brönner, Franfurt a. M., 345 pp.