{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,31]],"date-time":"2026-10-31T22:31:58Z","timestamp":1793485918195,"version":"4.5.0"},"reference-count":1,"publisher":"Magnolia Press","issue":"2","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Zootaxa"],"abstract":"<jats:p>Brachycephalus bufonoides was described as a \u201cvariety\u201d of B. ephippium based on two specimens which 90 years later was considered full species. Besides its brief original description, nothing else is known for this species. Herein we report the rediscovery of the pumpkin-toadlet Brachycephalus bufonoides from Nova Friburgo, State of Rio de Janeiro, the second most populous area within the Atlantic Forest in Brazil. A detailed osteological description of this species was also provided, including skull, hyolaryngeal skeleton and postcranium skeleton. The laryngeal skeleton of Brachycephalus genus was depicted for the first time. We conducted a molecular phylogenetic analysis of Brachycephalus using DNA sequences comprising two fragments of mitochondrial gene (16S). Both analysis with Bayesian inference and maximum parsimony supported the recognition of B. bufonoides as an exclusive lineage, allocated within the B. ephippium species group in B. vertebralis lineage. We improved the diagnosis and variation of the species, including more collected specimens, coloration in vivo and advertisement call description. Compared with its congeners, B. bufonoides has skin on head and dorsum with dermal hyperossification; skull with hyperossification of postorbital crests; a pair of hyperossified bulges about equidistant between postorbital crests; fourth presacral vertebra with transverse process hyperossified, ornamented and sacral diapophyses hyperossified, which can be seen externally (lineage of B. vertebralis sensu Condez et al. 2020); presence of dermal ossification as separated bulges of each vertebrae; general background color orange with different intensities of dark orange blotches on dorsum, including bordering of sacral region; absence of osteoderms and presence of warts on the dorsolateral surface of body; medium body size (SVL of adults: 12.0\u201314.5 mm for males and 14.7\u201316.3 mm for females; Table 1); rough dorsum; advertisement calls with 13 to 17 pulses; presence of pulse period modulation; and advertisement calls with notes longer than 0.2 s (0.22 to 0.31 s). Herein an important contribution for the taxonomy and systematics of this genus is provided, including a large amount of novel information for B. bufonoides from different sources (i.e., molecular, morphological variation, bioacoustic), allowing it to be included in future studies of species delimitation and relationships within Brachycephalus. Also, the discovery of this species reiterates the importance of Nova Friburgo for the conservation of the Atlantic Forest biodiversity.<\/jats:p>","DOI":"10.11646\/zootaxa.4819.2.3","type":"journal-article","created":{"date-parts":[[2020,7,23]],"date-time":"2020-07-23T00:54:28Z","timestamp":1595465668000},"source":"Crossref","is-referenced-by-count":9,"title":["Rediscovery of the toadlet Brachycephalus bufonoides Miranda-Ribeiro, 1920 (Anura: Brachycephalidae) with osteological and acoustic descriptions"],"prefix":"10.11646","volume":"4819","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5353-3906","authenticated-orcid":false,"given":"MANUELLA","family":"FOLLY","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5411-5522","authenticated-orcid":false,"given":"LUCAS COUTINHO","family":"AMARAL","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7558-4926","authenticated-orcid":false,"given":"SERGIO POTSCH DE","family":"CARVALHO-E-SILVA","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4900-630X","authenticated-orcid":false,"given":"JOS\u00c9 P.","family":"POMBAL JR.","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"4913","published-online":{"date-parts":[[2020,7,23]]},"reference":[{"key":"49264","unstructured":"<p class=\"Reference\">Alberch, P. &amp; Gale, E.A. (1985) A developmental analysis of an evolutionary trend: digital reduction in amphibians. <em>Evolution<\/em>, 39, 8\u201323.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1111\/j.1558-5646.1985.tb04076.x<\/p><p class=\"Reference\">Almeida-Santos, M. Siqueira, C.C., Van Sluys, M. &amp; Rocha, C.F.D. (2011) Ecology of the Brazilian flea frog <em>Brachycephalus didactylus<\/em> (Terrarana: Brachycephalidae). <em>Journal of Herpetology<\/em>, 45 (2), 251\u2013255.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1670\/10-015.1<\/p><p class=\"Reference\">Altschul, S.F., Madden, T.L., Sch\u00e4ffer, A.A., Zhang, J., Zhang, Z., Miller, W. &amp; Lipman, D.J. (1997) Gapped BLAST and PSI-BLAST: a new generation of protein database search programs. <em>Nucleic Acids Research<\/em>, 25 (17), 3389\u20133402.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1093\/nar\/25.17.3389<\/p><p class=\"Reference\">Alves, A.C.R., Ribeiro, L.F., Haddad, C.F.B. &amp; Reis, S.F. (2006) Two new species of <em>Brachycephalus<\/em> (Anura: Brachycephalidae) from the Atlantic forest in Paran\u00e1 State, southern Brazil. <em>Herpetologica<\/em>, 62 (2), 221\u2013233.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1655\/05-41.1<\/p><p class=\"Reference\">Alves, A.C.R., Sawaya, R., Reis, S.F. &amp; Haddad, C.B.F. (2009) A new species of <em>Brachycephalus<\/em> (Anura: Brachycephalidae) from the Atlantic Rain Forest in S\u00e3o Paulo State, Southeastern Brazil. <em>Journal of Herpetology<\/em>, 43 (2), 212\u2013219.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1670\/0022-1511-43.2.212<\/p><p class=\"Reference\">Andersen, M.L. (1978) <em>The comparative myology and osteology of the carpus and tarsus of selected anurans<\/em>. Ph.D. dissertation, University of Kansas, Lawrence, Kansas, 295 pp.<\/p><p class=\"Reference\">Angulo, A. &amp; Reichle, S. (2008) Acoustic signals, species diagnosis, and species concepts: the case of a new cryptic species of <em>Leptodactylus<\/em> (Amphibia, Anura, Leptodactylidae) from the Chapare region, Bolivia. \u200e<em>Zoological Journal of the Linnean Society<\/em>, 152 (1), 59\u201377.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1111\/j.1096-3642.2007.00338.x<\/p><p class=\"Reference\">Ara\u00fajo, C.B., Guerra, T.J., Amatuzzi, M.C.O. &amp; Campos, L.A. (2012) Advertisement and territorial calls of <em>Brachycephalus pitanga<\/em> (Anura: Brachycephalidae). <em>Zootaxa<\/em>, 3302 (1), 66\u201367.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.11646\/zootaxa.3302.1.5<\/p><p class=\"Reference\">Bokermann, W.C.A. (1966) <em>Lista anotada das localidades tipo de anf\u00edbios brasileiros<\/em>. Servi\u00e7o de Documenta\u00e7\u00e3o, S\u00e3o Paulo, 183 pp.<\/p><p class=\"Reference\">Bornschein, M.R., Ribeiro, L.F., Blackburn, D.C., Stanley, E.L. &amp; Pie, M.R. (2016) A new species of <em>Brachycephalus<\/em> (Anura: Brachycephalidae) from Santa Catarina, southern Brazil. <em>PeerJ<\/em>, 4 (e2629), 1\u201319.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.7717\/peerj.2629<\/p><p class=\"Reference\">Bornschein, M.R., Ribeiro, L.F., Rollo Jr., M.M., Confetti, A.E. &amp; Pie, M.R. (2017) Advertisement call of <em>Brachycephalus albolineatus<\/em> (Anura: Brachycephalidae). <em>PeerJ<\/em>, 6 (e5273), 1\u201318.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.7287\/peerj.preprints.3461v1<\/p><p class=\"Reference\">Bornschein, M.R., Pie, M.R. &amp; Teixeira, L. (2019a) Conservation Status of <em>Brachycephalus<\/em> Toadlets (Anura: Brachycephalidae) from the Brazilian Atlantic Rainforest. <em>Diversity<\/em>,<em> <\/em>11, 150, 1\u201329.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.3390\/d11090150<\/p><p class=\"Reference\">Bornschein, M.R., Rollo, M.M., Pie, M.R., Confetti, A.E. &amp; Ribeiro, L.F. (2019b) Redescription of the advertisement call of <em>Brachycephalus tridactylus<\/em> (Anura: Brachycephalidae). <em>Phyllomedusa<\/em>, 18 (1), 3\u201312.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.11606\/issn.2316-9079.v18i1p3-12<\/p><p class=\"Reference\">Brown, J.H. (1978) The theory of insular biogeography and the distribution of boreal birds and mammals. <em>Great Basin Naturalist Memoirs<\/em>, 2, 209\u2013227.<\/p><p class=\"Reference\">Campos, L.A., Silva, H.R. &amp; Sebben, A. (2010) Morphology and development of additional bony elements in the genus <em>Brachycephalus<\/em> (Anura: Brachycephalidae). <em>The Biological Journal of the Linnean Society<\/em>, 99 (4), 752\u2013767.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1111\/j.1095-8312.2010.01375.x<\/p><p class=\"Reference\">Canedo, C. &amp; Pombal, J.P. Jr. (2007) Two new species of torrent frog of the genus <em>Hylodes<\/em> (Anura, Hylodidae) with nuptial thumb tubercles. <em>Herpetologica<\/em>, 63 (2), 224\u2013235.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1655\/0018-0831(2007)63[224:TNSOTF]2.0.CO;2<\/p><p class=\"Reference\">Carvalho, T.R. &amp; Giaretta, A.A. (2013) Taxonomic circumscription of <em>Adenomera martinezi<\/em> (Bokermann, 1956) (Anura: Leptodactylidae: Leptodactylinae) with the recognition of a new cryptic taxon through a bioacoustic approach. <em>Zootaxa<\/em>, 3701 (2), 207\u2013237.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.11646\/zootaxa.3701.2.5<\/p><p class=\"Reference\">Cei, J.M. (1980) <em>Amphibians of Argentina. Monitore Zoologico Italiano. New Series. Monograph 2<\/em>. Instituto de Biologia Animal, Facultad de Ciencias Agrarias, Universidad Nacional de Cuyo, Mendoz, 609 pp.<\/p><p class=\"Reference\">Clemente-Carvalho, R.B., Antoniazzi, M.M., Jared, C., Haddad, C.F., Alves, A.C., Rocha, H.S., Pereira, G.R., Oliveira, D.F., Lopes, R.T. &amp; dos Reis, S.F. (2009) Hyperossification in miniaturized toadlets of the genus <em>Brachycephalus<\/em> (Amphibia: Anura: Brachycephalidae): microscopic structure and macroscopic patterns of variation. <em>Journal of Morphology<\/em>, 270 (11), 1285\u20131295.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1002\/jmor.10755<\/p><p class=\"Reference\">Clemente-Carvalho, R.B.G., Klaczko, J., Perez, S.I., Alves, A.C.R., Haddad, C.F.B. &amp; dos Reis, S.F. (2011) Molecular phylogenetic relationships and phenotypic diversity in miniaturized toadlets, genus <em>Brachycephalus <\/em>(Amphibia: Anura: Brachycephalidae). <em>Molecular Phylogenetics and Evolution<\/em>, 61 (1), 79\u201389.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1016\/j.ympev.2011.05.017<\/p><p class=\"Reference\">Clemente-Carvalho, R.B.G., Giaretta, A.A., Condez, T.H., Haddad, C.F.B. &amp; dos Reis, S.F. (2012) A new species of miniaturized toadlet, genus <em>Brachycephalus <\/em>(Anura: Brachycephalidae), from the Atlantic forest of Southeastern Brazil. <em>Herpetologica<\/em>, 68 (3), 365\u2013374.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1655\/HERPETOLOGICA-D-11-00085.1<\/p><p class=\"Reference\">Cochran, D.M. (1955) Frogs of southeastern Brazil. <em>Bulletin of the United States National Museum<\/em>, 206, 1\u2013423.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.5479\/si.03629236.206.1<\/p><p class=\"Reference\">Condez, T.H., Clemente-Carvalho, R.B.G., Haddad, C.F.B. &amp; dos Reis, S.F. (2014) A new species of <em>Brachycephalus<\/em> (Anura: Brachycephalidae) from the highlands of the Atlantic Forest, Southeastern Brazil. <em>Herpetologica<\/em>, 70 (1), 89\u201399.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1655\/HERPETOLOGICA-D-13-00044<\/p><p class=\"Reference\">Condez, T.H., Monteiro, J.P.C., Comitti, E.J., Garcia, P.C.A., Amaral, I.B. &amp; Haddad, C.F.B. (2016) A new species of flea-toad (Anura: Brachycephalidae) from southern Atlantic Forest, Brazil. <em>Zootaxa<\/em>, 4083 (1), 40\u201356.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.11646\/zootaxa.4083.1.2<\/p><p class=\"Reference\">Condez, T.H., Haddad, C.F.B. &amp; Zamudio, K.R. (2020) Historical biogeography and multi-trait evolution in miniature toadlets of the genus <em>Brachycephalus <\/em>(Anura: Brachycephalidae), <em>Biological Journal of the Linnean Society<\/em>, 129 (3), 664\u2013686.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1093\/biolinnean\/blz200<\/p><p class=\"Reference\">Dirnb\u00f6ck, T., Essl, F. &amp; Rabitsch, W. (2011) Disproportional risk for habitat loss of high-altitude endemic species under climate change. <em>Global Change Biology<\/em>, 17 (2), 990\u2013996.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1111\/j.1365-2486.2010.02266.x<\/p><p class=\"Reference\">Dorigo, T.A., Siqueira, C.C., Vrcibradic, D., Maia-Carneiro, T., Almeida-Santos, M. &amp; Rocha, C.F.D. (2012) Ecological aspects of the pumpkin toadlet, <em>Brachycephalus garbeanus<\/em> Miranda-Ribeiro, 1920 (Anura: Neobatrachia: Brachycephalidae), in a highland forest of southeastern Brazil. <em>Journal of Natural History<\/em>, 46 (39\u201340), 2497\u20132507. https:\/\/doi.org\/10.1080\/00222933.2012.713525<\/p><p class=\"Reference\">Duellman, W.E. (1970) <em>The hylids frogs of Middle America<\/em>. Monograph of the Museum of Natural History, University of Kansas, Lawrence, Kansas, pp.<\/p><p class=\"Reference\">Duellman, W.E. (1999) Global distribution of amphibians: patterns, conservation, and future challenges. <em>In<\/em>: Duellman, W.E. (Ed.), <em>Patterns of distribution of amphibians: a global perspective<\/em>. Johns Hopkins University Press, Baltimore, Maryland, pp. 1\u201330<\/p><p class=\"Reference\">Fabrezi, M. (1992) El carpo de los anuros. <em>Alytes<\/em>, 10, 1\u201336.<\/p><p class=\"Reference\">Fabrezi, M. (1993) The anuran tarsus. <em>Alytes<\/em>, 11, 47\u201363.<\/p><p class=\"Reference\">Fabrezi, M. (2001) A survey of prepollex and prehallux variation in anuran limbs. <em>Zoological Journal of the Linnean Society<\/em>, 131 (2), 227\u2013248.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1111\/j.1096-3642.2001.tb01316.x<\/p><p class=\"Reference\">Felsenstein, J. (1985) Confidence limits on phylogenies, an approach using bootstrap. <em>Evolution<\/em>, 39 (4), 783\u2013791.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1111\/j.1558-5646.1985.tb00420.x<\/p><p class=\"Reference\">Firkowski, C.R., Bornschein, M.R., Ribeiro, L.F. &amp; Pie, M.R. (2016) Species delimitation, phylogeny and evolutionary demography of co-distributed, montane frogs in the southern Brazilian Atlantic Forest. <em>Molecular Phylogenetics and Evolution<\/em>, 100, 345\u2013360.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1016\/j.ympev.2016.04.023<\/p><p class=\"Reference\">Folly, M., Hepp, F., Carvalho-e-Silva, S.P. &amp; Duellman, W.E. (2014) Taxonomic status and redescription of <em>Flectonotus ulei<\/em> (Anura: Hemiphractidae), with a key for the species of <em>Fritziana<\/em>. <em>Zoologia<\/em>, 31 (4), 393\u2013399. https:\/\/doi.org\/10.1590\/S1984-46702014000400011<\/p><p class=\"Reference\">Folly, M., Hepp, F. &amp; Carvalho-e-Silva, S.P. (2018) A new bromeligenous species of <em>Fritziana<\/em> Mello-Leit\u00e3o, 1937 (Amphibia: Anura: Hemiphractidae) from high elevations in the Serra dos \u00d3rg\u00e3os, Rio de Janeiro, Brazil. <em>Herpetologica<\/em>, 74 (1), 58\u201372.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1655\/Herpetologica-D-16-00035<\/p><p class=\"Reference\">Frost, D.R. (2019) Amphibian Species of the World: An Online Reference. American Museum of Natural History, New York, USA. 21 May 2019. Available from: http:\/\/research.amnh.org\/herpetology\/amphibia\/index.html (accessed 21 December 2019)<\/p><p class=\"Reference\">Garey, M.V., Lima, A.M.X., Hartmann, M.T. &amp; Haddad, C.F.B. (2012) A new species of miniaturized toadlet, genus <em>Brachycephalus<\/em> (Anura: Brachycephalidae), from southern Brazil. <em>Herpetologica<\/em>, 68 (2), 266\u2013271.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1655\/HERPETOLOGICA-D-11-00074.1<\/p><p class=\"Reference\">Giaretta, A.A. &amp; Sawaya, R.J. (1998) Second species of <em>Psyllophryne<\/em> (Anura: Brachycephalidae). <em>Copeia<\/em>, 1998 (4), 985\u2013987.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.2307\/1447345<\/p><p class=\"Reference\">Goloboff, P.A., Farris, J.S. &amp; Nixon, K.C. (2003) T.N.T.: Tree analysis using new technology. Program and documentation. Available from: http:www.zmuc.dk\/public\/phylogeny (accessed 21 December 2019)<\/p><p class=\"Reference\">Goutte, S., Mason, M.J., Christensen-Dalsgaard, J., Montealegre-Z, F., Chivers, B.D., Sarria-S, F.A., Antoniazzi, M.M., Jared, C., Sato, L.A. &amp; Toledo, L.F. (2017) Evidence of auditory insensitivity to vocalization frequencies in two frogs. <em>Scientific Reports<\/em>, 7 (12121), 1\u20139.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1038\/s41598-017-12145-5<\/p><p class=\"Reference\">Guimar\u00e3es, C. da S., Luz, S., Rocha, P.C. &amp; Feio, R.N. (2017) The dark side of pumpkin toadlet: a new species of <em>Brachycephalus<\/em> (Anura: Brachycephalidae) from Serra do Brigadeiro, southeastern Brazil. <em>Zootaxa<\/em>, 4258 (4), 327\u2013344.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.11646\/zootaxa.4258.4.2<\/p><p class=\"Reference\">Haddad, C.F.B., Alves, A.C.R., Clemente-Carvalho, R.B.G. &amp; dos Reis, S.F. (2010) A new species of <em>Brachycephalus<\/em> from the Atlantic Rain Forest in S\u00e3o Paulo State, southeastern Brazil (Amphibia: Anura: Brachycephalidae). <em>Copeia<\/em>, 2010 (3), 410\u2013420.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1643\/CH-09-102<\/p><p class=\"Reference\">Haddad, C.F.B. &amp; Pombal, J.P. Jr. (1998) Redescription of <em>Physalaemus spiniger<\/em> (Anura: Leptodactylidae) and description of two new reproductive modes. <em>Journal of Herpetolology<\/em>, 32 (4), 557\u2013565.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.2307\/1565210<\/p><p class=\"Reference\">Hedges, S.B., Duellman, W.E. &amp; Heinicke, M.P. (2008) New World direct-development frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation. <em>Zootaxa<\/em>, 1737 (1), 1\u2013182.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.11646\/zootaxa.1737.1.1<\/p><p class=\"Reference\">Hedges, S.B. (1994) Molecular evidence for the origin of birds. <em>Proceedings of the National Academy of Sciences of the United States of America<\/em>, 91 (7), 2621\u20132624.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1073\/pnas.91.7.2621<\/p><p class=\"Reference\">Heyer, W.R., Rand, A.S. Cruz, C.A.G., Peixoto, O.L. &amp; Nelson, C.E. (1990) Frogs of Borac\u00e9ia. <em>Arquivos de Zoologia<\/em>, 31, 231\u2013410.<\/p><p class=\"Reference\">Hepp, F. &amp; Pombal, J.P. Jr. (2019) Naming structures and qualifying properties of anuran bioacoustical signals: a call for homology-based nomenclature and equality for quantitative data. <em>Anais da Acad\u00eamia Brasileira de Ci\u00eancias<\/em>, 91 (4), e20190965.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1590\/0001-3765201920190965<\/p><p class=\"Reference\">Ihering, H.V. &amp; Ihering, R.V. (1911) O Museu Paulista nos anos de 1906 a 1909. <em>Revista do Museu Paulista<\/em>, 8, 1\u201322.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.5962\/bhl.part.20998<\/p><p class=\"Reference\">Instituto Brasileiro de Geografia e Estat\u00edstica\u2014IBGE (1959) <em>Enciclop\u00e9dia dos munic\u00edpios brasileiros<\/em>. IBGE, Rio de Janeiro, 467 pp.<\/p><p class=\"Reference\">Izecksohn, E. (1971) Novo g\u00eanero e nova esp\u00e9cie de Brachycephalidae do Estado do Rio de Janeiro, Brasil (Amphibia, Anura). <em>Boletim do Museu Nacional, <\/em>Boletim do Museu Nacional (N.S.), Zoologia, New Series, Zoologia, 280, 1\u201312.<\/p><p class=\"Reference\">Katoh, K. &amp; Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. <em>Molecular Biology and Evolution<\/em>, 30 (4), 772\u2013780.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1093\/molbev\/mst010<\/p><p class=\"Reference\">Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Meintjes, P. &amp; Drummond, A. (2012) Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. <em>Bioinformatics<\/em>, 28 (12), 1647\u20131649.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1093\/bioinformatics\/bts199<\/p><p class=\"Reference\">K\u00f6hler, J., Jansen, M., Rodr\u00edgues, A., Kok, P.J.R., Toledo, L.F., Emmrich, M., Glaw, F., Haddad, C.F.B., R\u00f6del, M.O. &amp; Vences, M. (2017) The use of bioacoustics in anuran taxonomy: theory, terminology, methods and recommendations form best practice. <em>Zootaxa<\/em>, 4251 (1), 1\u2013124.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.11646\/zootaxa.4251.1.1<\/p><p class=\"Reference\">Koichiro, T., Stecher, G., Peterson, D., Filipski, A. &amp; Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. <em>Molecular Biology and Evolution<\/em>, 30 (12), 2725\u20132729.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1093\/molbev\/mst197<\/p><p class=\"Reference\">Littlejohn, M.J. (2001) Patterns of differentiation in temporal properties of acoustic signals of anurans. <em>In<\/em>: Ryan, M.J. (Ed.), <em>Anuran Communication<\/em>. Smithsonian Institution Press, Washington, pp. 102\u2013120.<\/p><p class=\"Reference\">McDonald, K. &amp; Brown, J.H. (1992) Using montane mammals to model extinctions due to global change. <em>Conservation Biology<\/em>, 6 (3), 409\u2013415.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1046\/j.1523-1739.1992.06030409.x<\/p><p class=\"Reference\">Merckx, V.S.F.T., Hendriks, K.P., Beentjes, K.K., Mennes, C.B., Becking, L.E., Peijnenburg, K.T.C.A., Afendy, A., Arumugam, N., de Boer, H., Biun, A., Buang, M.M., Chen, P., Chung, A.Y.C., Dow, R., Feijen, F.A.A., Feijen, H., Soest, C.F., Geml, J., Geurts, R., Gravendeel, B., Hovenkamp, P., Imbun, P., Ipor, I., Janssens, S.B., Jocqu\u00e9, M., Kappes, H., Khoo, E., Koomen, P., Lens, F., Majapun, R.J., Morgado, L.N., Neupane, S., Nieser, N., Pereira, J.T., Rahman, H., Sabran, S., Sawang, A., Schwallier, R.M., Shim, P., Smit, H., Sol, N., Spait, M., Stech, M., Stokvis, F., Sugau, J.B., Suleiman, M., Sumail, S., Thomas, D.C., van Tol, J., Tuh, F.Y.Y., Yahya, B.E., Nais, J., Repin, R., Lakim, M. &amp; Schilthuizen, M. (2015) Evolution of endemism on a young tropical mountain. <em>Nature<\/em>, 524, 347\u2013350.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1038\/nature14949<\/p><p class=\"Reference\">Miranda-Ribeiro, A. (1920) Os Brachycephalideos do Museu Paulista (com tr\u00eas especes novas). <em>Revista do Museu Paulista<\/em>, 12, 307\u2013316.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.5962\/bhl.part.22255<\/p><p class=\"Reference\">Monteiro, J.P.C., Condez, T.H., Garcia, P.C.A., Comitti, E.J., Amaral, I.B. &amp; Haddad, C.F.B. (2018a) A new species of <em>Brachycephalus<\/em> (Anura, Brachycephalidae) from the coast of Santa Catarina State, southern Atlantic Forest, Brazil. <em>Zootaxa<\/em>, 4407 (4), 483\u2013505.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.11646\/zootaxa.4407.4.2<\/p><p class=\"Reference\">Monteiro, J.P.C., Condez, T.H., Garcia, P.C.A. &amp; Haddad, C.F.B. (2018b) The advertisement calls of two species of <em>Brachycephalus<\/em> (Anura: Brachycephalidae) from southern forest Brazil. <em>Zootaxa<\/em>, 4415 (1), 183\u2013188.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.11646\/zootaxa.4415.1.10<\/p><p class=\"Reference\">Napoli, M.F., Caramaschi, U., Cruz, C.A.G. &amp; Dias, I.R. (2011) A new species of flea-toad, genus <em>Brachycephalus<\/em> Fitzinger (Amphibia: Anura: Brachycephalidae), from the Atlantic rainforest of southern Bahia, Brazil. <em>Zootaxa<\/em>, 2739 (1), 33\u201340.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.11646\/zootaxa.2739.1.3<\/p><p class=\"Reference\">Oliveira, E.G. &amp; Haddad, C.F.B. (2015) Diet Seasonality and Feeding Preferences of <em>Brachycephalus pitanga<\/em> (Anura: Brachycephalidae). <em>Journal of Herpetology<\/em>, 49 (2), 252\u2013256.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1670\/13-211<\/p><p class=\"Reference\">Padial, J.M., Grant, T. &amp; Frost, D.R. (2014) Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria. <em>Zootaxa<\/em>, 3825 (1), 1\u2013132.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.11646\/zootaxa.3825.1.1<\/p><p class=\"Reference\">Palumbi, S.R., Martin, A., McMillan, W.O., Stice, L. &amp; Grabowski, G. (1991) <em>The simple fool\u2019s guide to PCR<\/em>. Version 2.0. Privately published, compiled by Palumbi S., Department of Zoology, University of Hawaii, Honolulu, 47 pp.<\/p><p class=\"Reference\">Pie, M.R., Meyer, A.L.S., Firkowski, C.R., Ribeiro, L.F. &amp; Bornschein, M.R. (2013) Understanding the mechanisms underlying the distribution of microendemic montane frogs (<em>Brachycephalus<\/em> spp., Terrarana: Brachycephalidae) in the Brazilian Atlantic Rainforest. <em>Ecological Modelling<\/em>, 250, 165\u2013176.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1016\/j.ecolmodel.2012.10.019<\/p><p class=\"Reference\">Pie, M.R. &amp; Ribeiro, L.F. (2015) A new species of <em>Brachycephalus<\/em> (Anura: Brachycephalidae) from the Quiriri mountain range of southern Brazil. <em>PeerJ<\/em>, 3 (e1179), 1\u20139.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.7717\/peerj.1179<\/p><p class=\"Reference\">Pie, M.R., Ribeiro, L.F., Confetti, A.E., Nadaline, M.J. &amp; Bornschein, M.R. (2018) A new species of <em>Brachycephalus<\/em> (Anura: Brachycephalidae) from southern Brazil. <em>PeerJ<\/em>, 6 (e5683), 1\u201327.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.7717\/peerj.5683<\/p><p class=\"Reference\">Pombal, J.P. Jr. (1999) Oviposi\u00e7\u00e3o e desenvolvimento de <em>Brachycephalus ephippium<\/em> (Spix) (Anura: Brachycephalidae). <em>Revista Brasileira de Zoologia<\/em>, 16 (4), 947\u2013956.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1590\/S0101-81751999000400004<\/p><p class=\"Reference\">Pombal, J.P. Jr. (2010) A posi\u00e7\u00e3o taxon\u00f4mica das variedades de <em>Brachycephalus ephippium<\/em> (Spix, 1824) descritas por Miranda-Ribeiro, 1920 (Amphibia, Anura, Brachycephalidae). <em>Boletim do Museu Nacional (N.S.), Zoologia,<\/em> 526, 1\u201312.<\/p><p class=\"Reference\">Pombal, J.P. Jr. &amp; Gasparini, J.L. (2006) A new <em>Brachycephalus<\/em> (Anura: Brachycephalidae) from the Atlantic Rainforest of Esp\u00edrito Santo, southeastern Brazil. <em>South American Journal of Herpetology<\/em>, 1 (2), 87\u201393.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.2994\/1808-9798(2006)1[87:ANBABF]2.0.CO;2<\/p><p class=\"Reference\">Pombal, J.P. Jr. &amp; Izecksohn, E. (2011) Uma nova esp\u00e9cie de <em>Brachycephalus<\/em> (Anura: Brachycephalidae) do estado do Rio de Janeiro. <em>Pap\u00e9is Avulsos de Zoologia<\/em>, 51 (28), 443\u2013451.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1590\/S0031-10492011002800001<\/p><p class=\"Reference\">Pombal, J.P. Jr., Sazima, I. &amp; Haddad, C.F.B. (1994) Breeding behavior of the pumpkin toadlet, <em>Brachycephalus ephippium<\/em> (Brachycephalidae). <em>Journal of Herpetology<\/em>, 28 (4), 516\u2013519.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.2307\/1564972<\/p><p class=\"Reference\">Pombal, J.P. Jr., Wistuba, E.M. &amp; Bornschein, M.R. (1998) A new species of brachycephalid (Anura) from the Atlantic Rain Forest of Brazil. <em>Journal of Herpetology<\/em>, 32 (1), 70\u201374.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.2307\/1565481<\/p><p class=\"Reference\">Pombal, J.P. Jr., Siqueira, C.C., Dorigo, T.A., Vrcibradic, D. &amp; Rocha, C.F.D. (2008) A third species of the rare frog genus <em>Holoaden<\/em> (Terrarana, Strabomantidae) from a montane rainforest area of southeastern Brazil. <em>Zootaxa<\/em>, 1938 (1), 61\u201368.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.11646\/zootaxa.1938.1.4<\/p><p class=\"Reference\">Posada, D. (2008) jModelTest: Phylogenetic Model Averaging. <em>Molecular Biology and Evolution<\/em>, 25 (7), 1253\u20131256.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1093\/molbev\/msn083<\/p><p class=\"Reference\">Pounds, J.A., Bustamante, M.R., Coloma, L.A., Consuegra, J.A., Fodgen, M.P.L., Foster, P.N., La Marca, E., Masters, K.L., Merino-Viteri, A., Puschendorf, R., Ron, S.R., S\u00e1nchez-Azofeifa, G.A., Still, C.J. &amp; Young, B.E. (2006) Widespread amphibian extinctions from epidemic disease driven by global warming. <em>Nature<\/em>, 439, 161\u2013167.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1038\/nature04246<\/p><p class=\"Reference\">Pugener, L.A. &amp; Maglia, A.M. (1997) Osteology and skeletal development of <em>Discoglossus sardus<\/em> (Anura: Discoglossidae). <em>Journal of Morphology<\/em>, 233 (3), 267\u2013286.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1002\/(SICI)1097-4687(199709)233:3&lt;267::AID-JMOR6&gt;3.0.CO;2-0<\/p><p class=\"Reference\">R Core Team R (2016) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from: http:\/\/www.r-project.org\/ (accessed 24 December 2019)<\/p><p class=\"Reference\">Rambaut, A. &amp; Drummond, A. (2007) TRACER. MCMC trace analysis tool. Version 1.4. University of Oxford, Oxford. Available from: http:\/\/tree.bio.ed.ac.uk\/software\/tracer (accessed 24 December 2019)<\/p><p class=\"Reference\">Ribeiro, L.F., Alves, A.C.R., Haddad, C.F.B. &amp; Reis, S.F. dos (2005) Two new species of <em>Brachycephalus <\/em>G\u00fcnther, 1858 from the State of Paran\u00e1, southern Brazil (Amphibia, Anura, Brachyephalidae). <em>Boletim do Museu Nacional<\/em>, New Series,<em> <\/em>Zoologia, 519, 1\u201318.<\/p><p class=\"Reference\">Ribeiro, L.F., Bornschein, M.R., Belmonte-Lopes, R., Firkowski, C.R., Morato, S.A.A. &amp; Pie, M.R. (2015) Seven new microendemic species of <em>Brachycephalus <\/em>(Anura: Brachycephalidae) from southern Brazil. <em>PeerJ<\/em>, 3, e1011.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.7717\/peerj.1011<\/p><p class=\"Reference\">Ribeiro, L.F., Blackburn, D.C., Stanley, E.L., Pie, M.R. &amp; Bornschein, M.R. (2017) Two new species of the <em>Brachycephalus pernix<\/em> group (Anura: Brachycephalidae) from the state of Paran\u00e1, southern Brazil. <em>PeerJ<\/em>, 5, e3603.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.7717\/peerj.3603<\/p><p class=\"Reference\">Ronquist, F., Teslenko, M., Mark, P. van der, Ayres, D.L., Darling, A., H\u00f6hna, S., Larget, B., Liu, L., Suchard, M.A. &amp; Huelsenbeck, J.P. (2012) MRBAYES 3.2: Efficient Bayesian phylogenetic inference and model selection across a large model space. <em>Systematic Biology<\/em>, 61 (3), 539\u2013542.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1093\/sysbio\/sys029<\/p><p class=\"Reference\">Sabaj, M.H. (2016) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an online reference. Version 6.5. American Society of Ichthyologists and Herpetologists, Washington, D.C. Available from: <a href=\"http:\/\/www.asih.org\/\">http:\/\/www.asih.org\/<\/a> (accessed 16 August 2016)<\/p><p class=\"Reference\">Siqueira, C.C., Vrcibradic, D., Nogueira-Costa, P., Martins, A.R., Dantas, L., Gomes, V.L.R., Bergallo, H.G. &amp; Rocha, C.F.D. (2014) Environmental parameters affecting the structure of leaf-litter frog (Amphibia: Anura) communities in tropical forests: a case study from an Atlantic Rainforest area in Southeastern Brazil. <em>Iheringia<\/em>, Zoologia, 31 (2), 147\u2013152.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1590\/S1984-46702014000200005<\/p><p class=\"Reference\">Spehn, E.M., Rudmann-Maurer, K., K\u00f6rner, C. &amp; Maselli, D. (2010) <em>Mountain biodiversity and global change.<\/em> GMBA-DIVERSITAS, University of Basel, Basel, 59 pp.<\/p><p class=\"Reference\">Tandel, M.C.F.F., Loibel, S., Oliveira, E.G. &amp; Haddad, C.F.B. (2014) Diferencia\u00e7\u00e3o de 3 tipos de vocaliza\u00e7\u00f5es (cantos) na esp\u00e9cie <em>Brachycephalus pitanga<\/em>. <em>Revista da Estat\u00edstica da Universidade Federal de Ouro Preto<\/em>, 3, 374\u2013386.<\/p><p class=\"Reference\">Taylor, W.R. &amp; Van Dyke, G.C. (1985) Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study. <em>Cybium<\/em>, 9, 107\u2013119.<\/p><p class=\"Reference\">Trewavas, E. (1933) The hyoid and larynx of the Anura. <em>Philosophical transactions of the Royal Society of London<\/em>, Series B, 222, 410\u2013527.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1098\/rstb.1932.0020<\/p><p class=\"Reference\">Trueb, L. (1973) Bones, frogs, and evolution. <em>In<\/em>: Vial, J.L. (Ed.), <em>Evolutionary biology of anurans: contemporary research on major problems<\/em>. University of Missouri Press, Columbia, Missouri, pp. 65\u2013132.<\/p><p class=\"Reference\">Trueb, L. (2015) Osteology. <em>In<\/em>: Duellman, W.E. (Ed.), <em>Marsupial frog: Gastrotheca and allied genera<\/em>. Johns Hopkins University Press, Baltimore, Maryland, pp. 31\u201351.<\/p><p class=\"Reference\">Verdade, V.K., Rodrigues, M.T., Cassimiro, J., Pavan, D., Liou, N. &amp; Lange, M.C. (2008) Advertisement call, vocal activity, and geographic distribution of <em>Brachycephalus hermogenesi<\/em> (Giaretta and Sawaya, 1998) (Anura: Brachycephalidae). <em>Journal of Herpetology<\/em>, 42 (3), 542\u2013549.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1670\/07-287.1<\/p><p class=\"Reference\">Weber, L.N., Verdade, V.K., Salles, R.O.L., Fouquet, A. &amp; Carvalho-e-Silva, S.P. de (2008) A new species of <em>Cycloramphus<\/em> Tschudi (Anura: Cycloramphidae) from the Parque Nacional da Serra dos \u00d3rg\u00e3os, southeastern Brazil. <em>Zootaxa<\/em>, 2737 (1), 19\u201333.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.11646\/zootaxa.2737.1.2<\/p><p class=\"Reference\">Wilkinson, J.A., Matsui, M. &amp; Terachi, T. (1996) Geographic variation in a Japanese tree frog (<em>Rhacophorus arboreus<\/em>) revealed by PCR-aided restriction site analysis of mtDNA. <em>Journal of Herpetology<\/em>, 30 (3), 418\u2013423.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.2307\/1565184<\/p><p class=\"Reference\">Wistuba, E.M. (1998) <em>Hist\u00f3ria natural de Brachycephalus pernix Pombal, Wistuba and Bornschein, 1998 (Anura) no Morro Anhangava, Munic\u00edpio de Quatro Barras, Estado do Paran\u00e1<\/em>. M.Sc. dissertation, Universidade Federal do Paran\u00e1, Curitiba, 89 pp.<\/p><p class=\"Reference\">Yeh, J. (2002) The effect of miniaturized body size on skeletal morphology in frogs. <em>Evolutio<\/em>n, 56 (3), 628\u2013641.<\/p><p class=\"Reference\"> https:\/\/doi.org\/10.1111\/j.0014-3820.2002.tb01372.x<\/p><p class=\"Reference\">Zeileis, A. &amp; Grothendieck, G. (2005) Zoo: s3 Infrastructure for Regular and Irregular Time Series. <em>Journal of Statistical Software<\/em>, 14 (6), 1\u20137.<\/p><p> https:\/\/doi.org\/10.18637\/jss.v014.i06<\/p>"}],"container-title":["Zootaxa"],"original-title":[],"link":[{"URL":"https:\/\/www.mapress.com\/zt\/article\/download\/zootaxa.4819.2.3\/40666","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.mapress.com\/zt\/article\/download\/zootaxa.4819.2.3\/40666","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,2,23]],"date-time":"2023-02-23T03:08:26Z","timestamp":1677121706000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mapress.com\/zt\/article\/view\/zootaxa.4819.2.3"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,7,23]]},"references-count":1,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2020,7,23]]}},"URL":"https:\/\/doi.org\/10.11646\/zootaxa.4819.2.3","relation":{},"ISSN":["1175-5334","1175-5326"],"issn-type":[{"value":"1175-5334","type":"electronic"},{"value":"1175-5326","type":"print"}],"subject":[],"published":{"date-parts":[[2020,7,23]]}}}