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The status of Elachistocleis pearsei, E. surinamensis, E. sikuani and E. tinigua Elachistocleissurinamensis and E. pearsei aretheoldest available names in this group. The former was named by Daudin (1802) as Bufo surinamensis, and the latter was named by Ruthven (1914) as Hypopachus pearsei. Both were considered synonyms of Elachistocleis ovalis by Parker (1934). Dunn (1944; 1949) commented on H. pearsei under the new combination Elachistocleis pearsei. Carvalho (1954) placed it into the new genus Relictivomer Carvalho, 1954 based on the presence of the posterior part of the prevomer and, recently, de Sá et al. (2012) reallocated it into Elachistocleis based on DNA data. Kenny (1969) redescribed E. surinamensis from specimens from Trinidad and cited a personal communication from A. Grandison, who had found no differences between specimens of E. surinamensis from Trinidad and a paratype of E. pearsei. Duellman (1997) also compared specimens of E. pearsei from Panama and specimens of Elachistocleis sp. from Gran Sabana, south-eastern Venezuela (which he did not assign to any species, but which falls in the range of occurrence of E. surinamensis; see Fig. 3) and did not find any noticeable difference in the external morphology of adults or in their advertisement calls. Jowers et al. (2021) recently redescribed and designated a neotype for E. surinamensis from Trinidad but, unfortunately, provided no comparisons of the species with E. pearsei. While the present paper was in review, two new species from Colombia were described and named: Elachistocleis sikuani and E. tinigua (AcostaGalvis et al., 2022). Although we did not access the vouchers and did not include the new sequences generated therein in our analyses, we confirmed that two lineages already represented in our tree correspond to them (based on genetic similarity, 99.8%). The authors made a brief comparison of the new species with E. pearsei and E. surinamensis, mentioning some differences in body size, quantity and colour of the blotches in live specimens, degree of conspicuousness of the post-commissural glands and presence/absence of the mid-dorsal white line (Acosta-Galvis et al., 2022). The latter feature was reported as absent in E. sikuani and E. tinigua but present (‘evident’) in E. surinamensis. However, as mentioned above in discussion about the variation of E. surumu, the presence of a mid-dorsal white line in E. surinamensis was not reported in its original description (see: Daudin, 1802) or in its redescription and neotype designation (see: Jowers et al., 2021). On the contrary, Jowers et al. (2021) used the absence of the mid-dorsal line in E. surinamensis to distinguish it from E. nigrogularis. We also have not observed a mid-dorsal white line in any examined specimens of E. surinamensis (Fig. 1). Acosta-Galvis et al. (2022) did not provide a comparison between E. pearsei and E. surinamensis. Instead, they added more pieces to the puzzle by giving new names to two allopatric lineages. They followed the prevailing tradition in the taxonomy of Elachistocleis of creating more and more nomina rather than conducting a thorough review addressing the validity and application of available names. We reiterate that a comprehensive morphological revision of Elachistocleis is needed and should take priority over the naming of additional new Elachistocleis species. Unfortunately, our sampling of specimens of those species is limited, and a reassessment of the taxonomic status of those species is beyond the scope of the present work. Therefore, for time being, we recognize E. pearsei, E. surinamensis, E. sikuani and E. tinigua as valid species., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on page 560, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481, {"references":["Daudin FM. 1802. Histoire naturelle des rainettes, des grenouilles et des crapauds. Paris: Levrault.","Ruthven AG. 1914. Description of a new engystomatid frog of the genus Hypopachus. Proceedings of the Biological Society of Washington 27: 77 - 80.","Parker H. 1934. A monograph of the frogs of the family Microhylidae. London: Trustees of the British Museum.","Dunn ER. 1944. Los generos de anfibios y reptiles de Colombia, 1. Primera parte: anfibios. Caldasia 2: 497 - 529.","Dunn ER. 1949. Notes on South American frogs of the family Microhylidae. American Museum NoVitates 1419: 1 - 21.","de Carvalho AL. 1954. A preliminary synopsis of the genera of American microhylid frogs. Occasional Papers of the Museum of Zoology. UniVersity of Michigan 555: 1 - 22.","de Sa RO, Streicher JW, Sekonyela R, Forlani MC, Loader SP, Greenbaum E, Richards S, Haddad CFB. 2012. Molecular phylogeny of microhylid frogs (Anura: Microhylidae) with emphasis on relationships among New World genera. BMC EVolutionary Biology 12: 241.","Kenny JS. 1969. The amphibia of Trinidad. Studies on the fauna of Curacao and other Caribbean islands 108: 1 - 78.","Duellman WE. 1997. Amphibians of La Escalera region, southeastern Venezuela: taxonomy, ecology, and biogeography. Scientific Papers Natural History Museum the UniVersity of Kansas 2: 1 - 52.","Jowers MJ, Othman SN, Borzee A, Rivas GA, Sanchez- Ramirez S, Auguste RJ, Downie JR, Read M, Murphy JC. 2021. Unraveling unique island colonization events in Elachistocleis frogs: phylogeography, cryptic divergence, and taxonomical implications. Organisms DiVersity & EVolution 21: 189 - 206.","Acosta-Galvis AR, Tonini JFR, de Sa RO. 2022. Two new species of Elachistocleis Parker, 1927 (Anura: Microhylidae: Gastrophryninae) from Colombia. Zootaxa 5099: 527 - 548."]}

Elachistocleis surumu Caramaschi, 2010 as a junior subjective synonym of Elachistocleis surinamensis (Daudin, 1802) Although we do not have samples from the type locality of E. surumu (Vila Surumu, Municipality of Pacaraima, State of Roraima, Brazil), we sampled many specimens from nearby localities surrounding the type locality (Fig. 3) and all of them are recovered in a single lineage that is widespread in the eastern portion of the Guiana Shield (Figs 1, 3). The proposed morphological differences between E. surumu and E. surinamensis are subtle. Caramaschi (2010) differentiated E. surumu from E. surinamensis based on the alleged presence of a light vertebral stripe in E. surinamensis (absent in E. surumu) and minor details in the ventral colour patterns. However, neither the original description of E. surinamensis (Daudin, 1802) nor the recent species redescription and neotype designation (Jowers et al., 2021) stated that E. surinamensis has any kind of dorsal stripe or line. Caramaschi (2010) did not list any voucher specimens of E. surinamensis in his list of examined material. Ventral colour patterns of E. surumu are contained in the variation found herein for specimens of E. surinamensis (see also: Jowers et al., 2021). Therefore, we consider Elachistocleis surumu Caramaschi, 2010 as a junior subjective synonym of Elachistocleis surinamensis (Daudin, 1802)., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on page 558, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481, {"references":["Caramaschi U. 2010. Notes on the taxonomic status of Elachistocleis oValis (Schneider, 1799) and description of five new species of Elachistocleis Parker, 1927 (Amphibia, Anura, Microhylidae). Boletim do Museu Nacional 527: 1 - 30.","Daudin FM. 1802. Histoire naturelle des rainettes, des grenouilles et des crapauds. Paris: Levrault.","Jowers MJ, Othman SN, Borzee A, Rivas GA, Sanchez- Ramirez S, Auguste RJ, Downie JR, Read M, Murphy JC. 2021. Unraveling unique island colonization events in Elachistocleis frogs: phylogeography, cryptic divergence, and taxonomical implications. Organisms DiVersity & EVolution 21: 189 - 206."]}

THE ELACHISTOCLEIS BICOLOR GROUP We recognize four nominal species in the E. bicolor group: E. bicolor, E. haroi, E. helianneae and E. nigrogularis. Except for E. nigrogularis, species of this group predominantly have an immaculate belly, but see comments below., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on page 557, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481

Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill, Peloso, Pedro (2023): A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis). Zoological Journal of the Linnean Society 197: 545-568, DOI: 10.1093/zoolinnean/zlac057

The (unsettling) status of Elachistocleis ovalis There has been a long controversy regarding the validity and applicability of this nomen, especially regarding whether it should be applied to immaculate or to maculate-bellied specimens of Elachistocleis [see Lavilla et al. (2003) and Caramaschi (2010) for lengthy discussions on the issue]. While some authors (e.g. Parker, 1927; Dunn, 1949; Kenny, 1969; De la Riva et al., 2000) applied this name to maculate-bellied specimens – thus conveniently distinguishing them from the immaculate-bellied E. bicolor – others did not. In a meeting abstract of his unpublished dissertation, Carcerelli (1992) regarded E. bicolor a junior synonym of E. ovalis; a view that was followed by a few other authors (e.g. Klappenbach & Langone, 1992; Olmos & Achaval, 1997; Kwet & Di-Bernardo, 1998). That decision was mainly grounded on the fact that Schneider (1799) stated that Rana ovalis had an ‘ inferne flavidus ’ (yellow venter). Lavilla et al. (2003) also remarked that, considering Schneider’s description, E. ovalis should be applied to the specimens with ‘immaculate, yellow, ventral colouration’; but they did not adhere to the idea of synonymizing E. ovalis and E. bicolor. Instead, they regarded E. ovalis as restricted to the ‘northern portion of the generic range’, whereas E. bicolor would be restricted to the ‘southern portion of the generic range’. Nevertheless, ventral patterns have been recorded idiosyncratically in the literature and ‘uniform’, ‘yellow’ and ‘immaculate’ bellies have often been treated as synonyms, although this is not necessarily the case. Some individuals of typically maculatebellied species can present the venter almost entirely covered by yellow blotches (e.g. Fig. 5G, J, L) and thus could also fit into the ‘ inferne flavidus ’ description. Thus, assuming that the type of E. ovalis had an immaculate venter, because Schneider (1799) stated that it has ‘ inferne flavidus ’, is not solid reasoning in itself. Moreover, as described above for our samples from Cuiabá (AAG-UFU 5953 and AAG-UFU 5954), sympatric specimens may present different ventral patterns and still be genetically similar, increasing the complexity of this puzzle. The identity of E. ovalis is still a conundrum, especially difficult to solve due to the lack of a type locality and the unknown whereabouts of the type specimen. For these reasons, among others, Caramaschi (2010) proposed an operational solution: he considered Rana ovalis Schneider, 1799 and the combination Elachistocleis ovalis a nomen dubium associated with a species inquirenda; a decision that was followed by Jowers et al. (2021). Since we could not find any new evidence on the contrary, we also adhere to that decision by Caramaschi (2010). AmphibiaWeb (2022) and Frost (2022), the two largest amphibian taxonomic catalogues, have essentially also followed this suggestion and, although both still include this species in their list of 22 recognized Elachistocleis species, the following notes are added for E. ovalis: ‘Nominally, Elachistocleis ovalis is a nomen inquirenda (see comment), not applied to a biological population. But, pending revision, this name is applied to populations from Bolivia, although a great deal of confusion surrounds the identification of specimens mentioned from other countries’ (Frost, 2022); ‘This ancient name (1799) lacks type material and there is no type locality. Despite wide usage it should be considered an invalid name and not used [...] Pending revision, AmphibiaWeb continues to use the name E. ovalis ’ (AmphibiaWeb, 2022)., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on pages 561-562, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481, {"references":["Lavilla EO, Vaira M, Ferrari L. 2003. A new species of Elachistocleis (Anura: Microhylidae) from the Andean Yungas of Argentina, with comments on the Elachistocleis oValis - E. bicolor controversy. Amphibia-Reptilia 24: 269 - 284.","Caramaschi U. 2010. Notes on the taxonomic status of Elachistocleis oValis (Schneider, 1799) and description of five new species of Elachistocleis Parker, 1927 (Amphibia, Anura, Microhylidae). Boletim do Museu Nacional 527: 1 - 30.","Parker HW. 1927. The brevicipitid frogs allied to the genus Gastrophryne. Occasional Papers of the Museum of Zoology. UniVersity of Michigan 187: 1 - 6.","Dunn ER. 1949. Notes on South American frogs of the family Microhylidae. American Museum NoVitates 1419: 1 - 21.","Kenny JS. 1969. The amphibia of Trinidad. Studies on the fauna of Curacao and other Caribbean islands 108: 1 - 78.","De la Riva I, Kohler J, Lotters S, Reichle S. 2000. Ten years of research on Bolivian amphibians: updated checklist, distribution, taxonomic problems, literature and iconography. ReVista Espanola de Herpetologia 14: 19 - 164.","Klappenbach MA, Langone JA. 1992. Lista sistematica y sinonimica de los anfibios del Uruguay con comentarios y notas sobre su distribucion. Anales del Museo Nacional de Historia Natural de MonteVideo (2. a Serie) 8: 163 - 222.","Olmos A, Achaval F. 1997. Anfibios y reptiles del Uruguay. Montevideo: Barreiro y Ramos.","Kwet A, Di-Bernardo M. 1998. Elachistocleis erythrogaster, a new microhylid species from Rio Grande do Sul, Brazil. Studies on Neotropical Fauna and EnVironment 33: 7 - 18.","Schneider JG. 1799. Historia amphibiorum naturalis et literarariae. Fasciculus primus. Continens ranas, calamitas, bufones, salamandras et hydros in genera et species descriptos notisque suis distinctos. Jena: Friederici Frommanni.","Jowers MJ, Othman SN, Borzee A, Rivas GA, Sanchez- Ramirez S, Auguste RJ, Downie JR, Read M, Murphy JC. 2021. Unraveling unique island colonization events in Elachistocleis frogs: phylogeography, cryptic divergence, and taxonomical implications. Organisms DiVersity & EVolution 21: 189 - 206.","Frost DR. 2022. Amphibian species of the aeorld: an online reference, V. 6.1. New York: American Museum of Natural History. Doi: 10.5531 / db. vz. 0001. Electronic database available at https: // amphibiansoftheworld. amnh. org / index. php (accessed 2 April 2022)."]}

The status of Elachistocleis cesarii, E. piauiensis and E. magna Miranda-Ribeiro (1920) described two subspecies of Engystoma ovale based on maculate-bellied specimens from the state of São Paulo, Brazil, which were later synonymized by Parker (1934) with Elachistocleis ovalis. Toledo et al. (2010) recently resurrected and redescribed them under the new combination Elachistocleis cesarii. Herein we recognize E. cesarii as a species distributed in south-eastern and central-western Brazil (Fig. 3) and closely related to E. piauiensis and E. magna (Fig. 1). Elachistocleis piauiensis was described and named by Caramaschi & Jim (1983) from Picos, Piauí. We found four lineages distributed in north-eastern Brazil, but we could not confidently assign any to E. piauiensis, because we did not have access to data from the type locality. Specimens assigned to each of these four lineages are morphologically similar to each other and to the original species description of E. piauiensis. The ventral colour pattern varies widely and similarly along the clade composed of E. cesarii, E. magna and the four candidate species E. cf. piauiensis 1–4, with no clear difference between them. The presence/absence of the mid-dorsal line also varies, except for E. cf. piauiensis 3, in which all examined vouchers lack the line. The holotype of E. piauiensis also lacks the line, and this could be evidence to assign the name to this specific clade. However, we cannot rule out that the consistent absence in E. cf. piauiensis 3 is not an effect of our sampling, given the high variability of this character state in several of our delimited species (Fig. 1). Elachistocleis cesarii was distinguished from E. piauiensis by differences in the advertisement call, size of the post-commissural gland and details of the inguinal region colouration (Toledo et al., 2010) – phenotypic traits that we have not explored for the present contribution. On the other hand, E. magna was distinguished from E. cesarii and E. piauiensis solely by its size (male SVL above 31 mm in E. magna, against SVL below 29 mm in E. cesarii and E. piauiensis; Toledo, 2010). Among examined specimens, we observed a consistent difference in size between E. magna and E. cf. piauiensis 1–4, but not between E. magna and E. cesarii. Males of E. cesarii can have SVL up to 31 mm, whereas some males of E. magna have SVL below 29 mm. Therefore, a thorough evaluation of phenotypic data allied with a denser molecular sampling is needed to verify if the differential diagnoses among E. cesarii, E. piauiensis and E. magna will hold true with additional data., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on pages 560-561, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481, {"references":["Miranda-Ribeiro A. 1920. Os engystomatideos do Museu Paulista (com um genero e tres especies novos). ReVista do Museu Paulista. Sao Paulo 12: 281 - 288.","Parker H. 1934. A monograph of the frogs of the family Microhylidae. London: Trustees of the British Museum.","Toledo LF, Loebmann D, Haddad CFB. 2010. Revalidation and redescription of Elachistocleis cesarii (Miranda-Ribeiro, 1920) (Anura: Microhylidae). Zootaxa 2418: 50 - 60.","Caramaschi U, Jim J. 1983. A new microhylid frog, genus Elachistocleis (Amphibia, Anura), from northeastern Brasil. Herpetologica 39: 390 - 394."]}

The (mysterious) status of Elachistocleis corumbaensis Unfortunately, we have no samples confidently assignable to E. corumbaensis. We have two samples from two localities near (around 20 km and 80 km) but not from the type locality of the species (Parque Municipal de Piraputangas, Corumbá municipality; Piva et al., 2017) (Fig. 3). For one of these tissue samples (CFBHT 00078), we were unable to find the voucher specimen. The other (MAPT 1363) was identified by the collectors as E. matogrosso (see: Koroiva et al., 2020). So we assume, albeit tentatively, that it has an immaculate venter. Both samples fall with E. bicolor in our species delimitation (Fig. 1). However, E. corumbaensis was described as having a maculate belly (Piva et al., 2017). The closest locality that we have a sample of a maculate-bellied species (E. cesarii) is Campo Grande, state of Mato Grosso do Sul, Brazil, 357 km from the type locality of E. corumbaensis. It is also worth remarking that E. corumbaensis has several similarities with E. haroi, particularly the dorsal pattern with a dark mark in the shape of a pine tree (Pereyra et al., 2013; Piva et al., 2017). Our closest sample of E. haroi is from Puerto Carmelo Peralta, Department of Alto Paraguay (Paraguay), 275 km south of the type locality of E. corumbaensis. For the moment, we consider E. corumbaensis a valid species, although it is unclear if this species belongs to the E. bicolor group or to the E. surinamensis group. The (unresolved) status of Elachistocleis erythrogaster This species is apparently restricted to the Centre of Research and Nature Conservation Pró-Mata, state of Rio Grande do Sul, above 900 m a.s.l. (Kwet & Di-Bernardo, 1998). It was distinguished from E. bicolor [treated as E. ovalis in Kwet & Di-Bernardo (1998)], which occurs in sympatry, by the unique red belly, larger size and other bioacoustic and reproductive characteristics (Kwet & Di-Bernardo, 1998). We have sampled some individuals from localities close to the type locality – although from the lowland – and they all fall with E. bicolor. This species has not been collected for nearly 20 years and is considered to be rare (Kwet et al., 2010)., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on page 561, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481, {"references":["Piva A, Caramaschi U, Albuquerque NR. 2017. A new species of Elachistocleis (Anura: Microhylidae) from the Brazilian Pantanal. Phyllomedusa. Journal of Herpetology 16: 143 - 154.","Koroiva R, Rodrigues LRR, Santana DJ. 2020. DNA barcoding for identification of anuran species in the central region of South America. PeerJ 8: e 10189.","Pereyra LC, Akmentins MS, Laufer G, Vaira M. 2013. A new species of Elachistocleis (Anura: Microhylidae) from north-western Argentina. Zootaxa 3694: 525 - 544.","Kwet A, Di-Bernardo M. 1998. Elachistocleis erythrogaster, a new microhylid species from Rio Grande do Sul, Brazil. Studies on Neotropical Fauna and EnVironment 33: 7 - 18.","Kwet A, Lingnau R, Di-Bernardo M. 2010. Pro-Mata: anfibios da Serra Gaucha, sul do Brasil - Amphibien der Serra Gaucha, Sudbrasilien - amphibians of the Serra Gaucha, south of Brazil. Tubingen and Porto Alegre: Brasilien-Zentrum de Universitat Tubingen and EDIPUCRS."]}

Elachistocleis bumbameuboi Caramaschi, 2010 and Elachistocleis carvalhoi Caramaschi, 2010 as junior subjective synonyms of Elachistocleis magna Toledo, 2010 Caramaschi (2010) distinguished E. bumbameuboi and E. carvalhoi from E. magna based on subtle differences in the dorsal and ventral colour patterns, as follows: regarding the dorsal pattern, E. magna has ‘dorsum uniform dark greyish with scarce minute brighter dots in the outer boundaries’, whereas E. bumbameuboi and E. carvalhoi have, respectively, ‘dorsum uniformly dark grey or black without marks’ and ‘dorsum uniformly brown or dark grey without marks’; regarding the ventral pattern, E. magna has ‘venter grey with minute scattered white spots, mainly on the belly and ventral surfaces of legs’, whereas E. bumbameuboi and E. carvalhoi have, respectively, ‘venter grey with minute anastomosed whitish spots, producing a salt-and-pepper pattern’ and ‘venter greyish with large anastomosed yellow or whitish yellow blotches, producing a coarse marbled pattern, mainly in the chest area. Photographs of the holotypes of E. carvalhoi (Caramaschi, 2010: fig. 3) and E. bumbameuboi (Caramaschi, 2010: fig. 4) and a photograph of a paratype of E. bumbameuboi (Nunes et al., 2010: fig. 1) depict specimens with dorsa scattered with minute white dots. Thus, our examination of specimens of E. bumbameuboi, E. carvalhoi and E. magna show a broad overlap among the various shades of dorsal colour and the size and distribution pattern of ventral markings (e.g. see Fig. 5). Based on our phylogenetic hypothesis, on the broad geographic sampling, which included a paratype of E. bumbameuboi, topotypes of E. carvalhoi and E. magna, and on the examination of several voucher specimens, we consider that E. bumbameuboi Caramaschi, 2010 and E. carvalhoi Caramaschi, 2010 are junior subjective synonyms of E. magna Toledo, 2010., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on pages 558-560, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481, {"references":["Caramaschi U. 2010. Notes on the taxonomic status of Elachistocleis oValis (Schneider, 1799) and description of five new species of Elachistocleis Parker, 1927 (Amphibia, Anura, Microhylidae). Boletim do Museu Nacional 527: 1 - 30.","Nunes I, Canedo C, Carvalho RR. 2010. Advertisement call and geographic distribution of Elachistocleis piauiensis Caramaschi & Jim, 1983 (Amphibia, Microhylidae), with notes on the presence of post-commissural gland in the genus. South American Journal of Herpetology 5: 30 - 34."]}

TAXONOMY OF ELACHISTOCLEIS – A COMPLEX MATTER Parker (1927) erected Elachistocleis to allocate Rana ovalis Schneider, 1799 and Engystoma ovale bicolor (Guérin-Méneville, 1838). As usual at the time, the descriptions of both species were brief and did not designate type specimens. Lavilla et al. (2003) suggested that these names apply to complexes of species, because species of this genus are morphologically similar, with few obvious external characters that can be used as reliable diagnostic characters (i.e. non-overlapping and fixed). Ultimately, as mentioned throughout our contribution, diagnoses in this genus rely heavily on the ventral colour pattern, head proportions and the presence/absence and extension of body lines, such as a mid-dorsal white line and lines on the hidden surface of hindlimbs (Parker, 1927; Lavilla et al., 2003; Caramaschi, 2010; Pereyra et al., 2013). Caramaschi (2010) and Piva et al. (2017) suggested the ventral colour pattern as a diagnostic character for two internal species groups: a group with immaculate bellies (uniformly clear, free of markings) and another with maculate bellies. We found that the groups proposed by the above-cited authors are not monophyletic. For example, Elachistocleis muiraquitan (immaculate-bellied) is not related to the remaining immaculate-bellied species; instead, it was recovered in the E. surinamensis species group, a clade composed of species with predominantly maculate bellies. Furthermore, whereas E. panamensis has a maculate belly (Dunn et al., 1948; Nelson, 1972), E. araios has an immaculate belly (Sánchez-Nivicela et al., 2020). Given their phylogenetic position as successive sistertaxa to the remaining species of the genus (but see our comments on the position of E. araios), the description of E. araios already rendered the immaculate-bellied species group as paraphyletic. Also, it implies that the optimization of the ventral pattern of the Elachistocleis ancestor is ambiguous, given our topology and that of Sánchez-Nivicela et al. (2020). More importantly, we found intraspecific variation of the ventral pattern in some species, such as E. bicolor and E. nigrogularis (Figs 1, 4). Ontogenetic variation of the ventral pattern has also been recently reported for E. haroi (Bueno-Villafañe et al., 2020). Therefore, the available phylogenetic and morphological evidence emphatically rejects the existence of groups diagnosed solely by the colour of their bellies. It is noteworthy that this discussion seems far from settled. Ventral patterns have been recorded idiosyncratically in the literature. Sánchez-Nivicela et al. (2020) cite E. cesarii as ‘having uniform, immaculate, ventral colouration’, although Toledo et al. (2010) when resurrecting the species clearly state that it has ‘ventral colouration white or yellow with grey marks and reticulations’, and their figure 3d depicts a specimen with a maculate belly. Jowers et al. (2021) state that E. cesarii has a ‘uniform’ belly. This same was said of E. erythrogaster and E. bicolor. While E. cesarii and E. erythrogaster are maculate (the latter comparatively less maculate), E. bicolor bellies usually present no markings whatsoever (but see discussion on E. bicolor below). Given the historical use of the ventral pattern in the taxonomy of Elachistocleis, we attempted to verify as many specimens as possible for this trait. We have not extensively checked our vouchers for other morphological diagnostic traits (male throat colour; presence and shape of the femoral stripe; post-commissural gland). Nevertheless, our preliminary data suggest that none of the above-cited morphological traits is as reliable as suggested in the literature [see also comments in Nunes et al. (2010) and Marinho et al. (2018)]. Data from other sources are scarce. Tadpoles of few species are described (e.g. Rossa-Feres & Nomura, 2006; Pereyra et al., 2013; Schulze et al., 2015; Ferreira & Weber, 2021), but it is difficult to confidently assign these descriptions to lineages recovered because most lack an association with available DNA sequences. Moreover, there is no comprehensive understanding of variation, although Gómez & Kehr (2012) reported some variability in larval morphology related to chemical cues in the presence of predators. Advertisement calls of several Elachistocleis have been described (e.g. Nelson, 1972; Duellman, 1997; Kwet & Di-Bernardo, 1998; Lavilla et al., 2003; Nunes et al., 2010; Toledo, 2010; Marinho et al., 2018; Pansonato et al., 2018; Jowers et al., 2021). Some of these are associated with vouchers present in our phylogeny (see: Marinho et al., 2018; Jowers et al., 2021). These are not adequately distributed in our topology and most are described from a few specimens and localities. Published data have already shown that even slight increases in sample size may affect recorded variation (see: Marinho et al., 2018). Finally, some localities present at least two sympatric species of Elachistocleis. For instance, both E. nigrogularis and E. surinamensis are syntopically found at the type locality of E. nigrogularis (Jowers et al., 2021), and both E. bicolor and E. erythrogaster are found at the type locality of E. erythrogaster (Kwet & Di-Bernardo, 1998) (also see Fig. 3). This makes it even more difficult to associate larvae with adults and advertisement calls of unvouchered specimens to named species without molecular data. A fine line A white or light yellow mid-dorsal line is present in many individuals of Elachistocleis, and other genera of Gastrophryninae, such as Chiasmocleis Méhelÿ, 1904 (e.g. Peloso et al., 2014), Ctenophryne Mocquard, 1904 (e.g. Duellman, 1978), Hamptophryne A.L. Carvalho, 1954 (e.g. Parker, 1927; Duellman, 1978), Hypopachus [e.g. Cope (1889); see also figure 5 in Greenbaum et al. (2011)], Dasypops Miranda-Ribeiro, 1924 and Stereocyclops Cope, 1870 (PP, personal observation). Many authors have noticed important intraspecific or intrapopulational variation (i.e. being present in some individuals and absent in others) in this character in other genera (e.g. Chiasmocleis, Peloso et al., 2014; Ctenophryne, Zweifel & Myers, 1989; Stereocyclops, GNF, personal observation) and in Elachistocleis (e.g. Nelson, 1972; Toledo, 2010; Toledo et al., 2010; Marinho et al., 2018). Notwithstanding, the occurrence of this line and the variation of its extension when present (e.g. from snout to vent or from post-cephalic fold to vent) were used in diagnoses of several Elachistocleis species (e.g. Caramaschi & Jim, 1983; Caramaschi, 2010; Nunes-de-Almeida & Toledo, 2012; Piva et al., 2017). We found a remarkable variation in the occurrence of the mid-dorsal white line in and among species (Fig. 1), similar to the variation reported for Chiasmocleis (Peloso et al., 2014). This variation challenges the reliability of the trait as a diagnostic feature and weakens the known diagnoses of several currently recognized species (see below). It was impossible to confidently evaluate the extension of the mid-dorsal line, thus we only scored the presence or the absence of the line. Hence, we recommend caution when using the mid-dorsal line in the systematics of Elachistocleis, especially for diagnostic purposes., Published as part of Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill & Peloso, Pedro, 2023, A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis), pp. 545-568 in Zoological Journal of the Linnean Society 197 on pages 554-555, DOI: 10.1093/zoolinnean/zlac057, http://zenodo.org/record/7695481, {"references":["Parker HW. 1927. The brevicipitid frogs allied to the genus Gastrophryne. Occasional Papers of the Museum of Zoology. UniVersity of Michigan 187: 1 - 6.","Schneider JG. 1799. Historia amphibiorum naturalis et literarariae. Fasciculus primus. Continens ranas, calamitas, bufones, salamandras et hydros in genera et species descriptos notisque suis distinctos. Jena: Friederici Frommanni.","Guerin-Meneville F-E. 1838. Iconographie du regne Animal de G. CuVier ou representation d'apres nature de l'une des especes les plus remarquables et souVent non enVore figurees, de chaque genre d'animaux, aVec un texte descriptiff mis au courant de la science, Vol. 3 (Part - Reptiles). Paris: J. B. Balliere.","Lavilla EO, Vaira M, Ferrari L. 2003. A new species of Elachistocleis (Anura: Microhylidae) from the Andean Yungas of Argentina, with comments on the Elachistocleis oValis - E. bicolor controversy. Amphibia-Reptilia 24: 269 - 284.","Caramaschi U. 2010. Notes on the taxonomic status of Elachistocleis oValis (Schneider, 1799) and description of five new species of Elachistocleis Parker, 1927 (Amphibia, Anura, Microhylidae). Boletim do Museu Nacional 527: 1 - 30.","Pereyra LC, Akmentins MS, Laufer G, Vaira M. 2013. A new species of Elachistocleis (Anura: Microhylidae) from north-western Argentina. Zootaxa 3694: 525 - 544.","Piva A, Caramaschi U, Albuquerque NR. 2017. A new species of Elachistocleis (Anura: Microhylidae) from the Brazilian Pantanal. Phyllomedusa. Journal of Herpetology 16: 143 - 154.","Dunn ER, Trapido H, Evans H. 1948. A new species of the microhylid frog genus Chiasmocleis from Panama. American Museum NoVitates 1376: 1 - 8.","Nelson CE. 1972. Distribution and biology of Chiasmocleis panamensis (Amphibia: Microhylidae). Copeia 1972: 895 - 898.","Sanchez-Nivicela JC, Peloso PLV, Urgiles VL, Yanez- Munoz MH, Sagredo Y, Paez N, Ron S. 2020. Description and phylogenetic relationships of a new trans-Andean species of Elachistocleis Parker 1927 (Amphibia, Anura, Microhylidae). Zootaxa 4779: 323 - 340.","Bueno-Villafane D, Caballlero-Gini A, Ferreira M, Netto F, Fernandez Rios D, Brusquetti F. 2020. Ontogenetic changes in the ventral colouration of post metamorphic Elachistocleis haroi Pereyra, Akmentins, Laufer, Vaira, 2013 (Anura: Microhylidae). Amphibia-Reptilia 41: 191 - 200.","Toledo LF, Loebmann D, Haddad CFB. 2010. Revalidation and redescription of Elachistocleis cesarii (Miranda-Ribeiro, 1920) (Anura: Microhylidae). Zootaxa 2418: 50 - 60.","Jowers MJ, Othman SN, Borzee A, Rivas GA, Sanchez- Ramirez S, Auguste RJ, Downie JR, Read M, Murphy JC. 2021. Unraveling unique island colonization events in Elachistocleis frogs: phylogeography, cryptic divergence, and taxonomical implications. Organisms DiVersity & EVolution 21: 189 - 206.","Nunes I, Canedo C, Carvalho RR. 2010. Advertisement call and geographic distribution of Elachistocleis piauiensis Caramaschi & Jim, 1983 (Amphibia, Microhylidae), with notes on the presence of post-commissural gland in the genus. South American Journal of Herpetology 5: 30 - 34.","Marinho P, Carvalho TR, Bang DL, Teixeira BFDV, Azarak PA, Campos CEC, Giaretta AA. 2018. Advertisement calls, intraspecific variation and species diagnosis of six Brazilian species of Elachistocleis (Anura: Microhylidae: Gastrophryninae). Zootaxa 4521: 357 - 375.","Rossa-Feres D de C, Nomura F. 2006. Characterization and taxonomic key for tadpoles (Amphibia: Anura) from the northwestern region of Sao Paulo State, Brazil. Biota Neotropica 6: 1 - 26.","Schulze A, Jansen M, Kohler G. 2015. Tadpole diversity of Bolivia's lowland anuran communities: molecular identification, morphological characterisation, and ecological assignment. Zootaxa 4016: 1 - 111.","Ferreira JS, Weber LN. 2021. A survey of the external morphology, internal oral morphology, chondrocranium and hyobranchial apparatus of Elachistocleis larvae Parker, 1927 (Anura, Microhylidae). Journal of Morphology 282: 472 - 484.","Gomez VI, Kehr AI. 2012. The effect of chemical signal of predatory fish and water bug on the morphology and development of Elachistocleis bicolor tadpoles (Anura: Microhylidae). Biologia 67: 1001 - 1006.","Duellman WE. 1997. Amphibians of La Escalera region, southeastern Venezuela: taxonomy, ecology, and biogeography. Scientific Papers Natural History Museum the UniVersity of Kansas 2: 1 - 52.","Kwet A, Di-Bernardo M. 1998. Elachistocleis erythrogaster, a new microhylid species from Rio Grande do Sul, Brazil. Studies on Neotropical Fauna and EnVironment 33: 7 - 18.","Pansonato A, Mudrek JR, Nunes CA, Strussmann C. 2018. Advertisement calls of topotypes of Elachistocleis matogrosso (Anura: Microhylidae). Salamandra 54: 92 - 96.","Peloso PLV, Sturaro MJ, Forlani MC, Gaucher P, Motta AP, Wheeler WC. 2014. Phylogeny, taxonomic revision, and character evolution of the genera Chiasmocleis and Syncope (Anura, Microhylidae) in Amazonia, with descriptions of three new species. Bulletin of the American Museum of Natural History 386: 1 - 112.","Duellman WE. 1978. The Biology of an Equatorial Herpetofauna in Amazonian Ecuador. Lawrence: University of Kansas.","de Carvalho AL. 1954. A preliminary synopsis of the genera of American microhylid frogs. Occasional Papers of the Museum of Zoology. UniVersity of Michigan 555: 1 - 22.","Cope ED. 1889. Batrachia of North America. Bulletin of the United States National Museum 34: 5 - 525.","Greenbaum E, Smith EN, de Sa RO. 2011. Molecular systematics of the Middle American genus Hypopachus (Anura: Microhylidae). Molecular Phylogenetics and EVolution 61: 265 - 277.","Zweifel RG, Myers CW. 1989. A New Frog of the Genus Ctenophryne (Microhylidae) from the Pacific Lowlands of Northwestern South America. American Museum NoVitates 2947: 1 - 16.","Caramaschi U, Jim J. 1983. A new microhylid frog, genus Elachistocleis (Amphibia, Anura), from northeastern Brasil. Herpetologica 39: 390 - 394.","Nunes-de-Almeida CHL, Toledo LF. 2012. A new species of Elachistocleis Parker (Anura, Microhylidae) from the state of Acre, northern Brazil. Zootaxa 3424: 43 - 50."]}

Chiasmocleis supercilialba Morales and McDiarmid, 2009 Figure 35, plate 8 Chiasmocleis supercilialbus (Morales and McDiarmid, 2009). Incorrect spelling in the original description (see Peloso et al., 2013). Syncope supercilialbus (de Sá et al., 2012). HOLOTYPE: MHNSM 16174; examined solely from a photograph, in life (not shown). TYPE LOCALITY: Pakitza, Reserve Zone, Manu National Park, ca. 57 km northwestern from the mouth of Rio Manu (11 ° 569470S / 71 ° 179000W), on Rio Manu, Departamento Madre de Dios, Peru. DIAGNOSIS: A medium-sized species for the genus; SVL in males 18.4–18.7 mm (Morales and McDiarmid, 2009). Females unknown. Body ovoid to elongate; head triangular, snout rounded in dorsal and lateral views. Four distinctive fingers; FI well developed, subarticular tubercle may be absent or present; subarticular tubercles present on all remaining fingers, sometimes hardly visible; adpressed FI never extends past the distal margin of subarticular tubercle of FII; adpressed FIV does not reach distal tubercle of FIII; palmar tubercles present, not divided; relative finger lengths I,II, IV,III. Five distinctive toes present, first may be much reduced; toes may be slightly fringed; toes not webbed; TI lacks tubercle; adpressed TI does not touch subarticular tubercle of TII; adpressed TV does not touch middle subarticular tubercle of TIV; TII–IV with terminal discs, usually more developed in females, but also present in males; relative toe lengths I,II,V,III,IV. An inguinal blotch of varied shape is always present. Venter (belly and under surfaces of thighs) usually white or cream with black or dark brown irregular spots. VARIATION: Some variation in color pattern was observed from photos of live specimens (pl. 8) and examination of a paratype (USNM 342862: fig. 35). The dorsum can be dark brown, to almost uniformly whitish (silver) and may show reddish blotches and white spots (when not entirely whitish). A wide white stripe is almost invariably present from the snout to past the eyelids, sometimes extending to the inguinal region (forming the split stripe). A very narrow dorsal white line commonly present, extending from the occipital region to the cloacal region. Dorsal surfaces of forearm yellowish, orange or red, while dorsal surfaces of hind limbs usually follow the general dorsal pattern. As reported by Morales and McDiarmid (2009) the shape and number of inguinal and lateral blotches (or spots) is variable, but commonly dark brown. Throat region with intense brown reticulation over a white (or cream-colored) background. Chest, belly, and undersurfaces of limbs with large, dark brown spots or irregular blotches (fig. 35, pl. 8D, F, H). CALL AND TADPOLE: The advertisement call of C. supercilialba was described by Morales and McDiarmid (2009). Call consists of a fast series of multipulsed notes (mean note duration 31.0 ms; mean interval between notes 37.0 ms) with dominant frequency between 2985.8–3205.0 Hz. Number of pulses per note not given. Tadpoles are unknown. REMARKS: A population from Rio Formoso, state of Rondônia, Brazil (OMNH 37204–37206, 37308–37309, 37319), present dermal spines on dorsum, upper lips, cloacal region, fingers, and toes. The specimens also show well-developed fringes on the fingers and toes, and the toes are basally webbed. These specimens, therefore, agree (in part) with the diagnosis of Chiasmocleis supercilialba (see Morales and McDiarmid 2009, and text above). We tentatively assign the Rio Formoso specimens to C. supercilialba. DISTRIBUTION (fig. 36): Southern Peru (Madre de Dios) and western Brazil (Acre and Rondônia)., Published as part of Peloso, Pedro L. V., Sturaro, Marcelo José, Forlani, Mauricio C., Gaucher, Philippe, Motta, Ana Paula & Wheeler, Ward C., 2014, Phylogeny, Taxonomic Revision, And Character Evolution Of The Genera Chiasmocleis And Syncope (Anura, Microhylidae) In Amazonia, With Descriptions Of Three New Species, pp. 1 in Bulletin of the American Museum of Natural History 2014 (386) on pages 55-57, DOI: 10.1206/834.1, http://zenodo.org/record/5378601, {"references":["Morales, V. R., and R. W. McDiarmid. 2009. A new species of Chiasmocleis (Anura: Microhylidae) from southern Amazonian Peru with comments on some other michrohylids. Biotempo 9: 71 - 76.","Peloso, P. L. V., R. W. McDiarmid, and U. Caramaschi. 2013. The gender of '' cleis '': correct spelling of Chiasmocleis supercilialbus Morales and McDiarmid, 2009, and Elachistocleis magnus Toledo, 2010 (Anura, Microhylidae). Zootaxa 3718: 297 - 298.","de Sa, R. O., et al. 2012. Molecular phylogeny of microhylid frogs (Anura: Microhylidae) with emphasis on relationships among New World genera. BMC Evolutionary Biology 12: 1 - 21."]}

Osteopilus Fitzinger, 1843 TYPE SPECIES: Trachycephalus marmoratus Duméril and Bibron, 1841 (5 Hyla septentrionalis Duméril and Bibron, 1841). Calyptahyla Trueb and Tyler, 1974. Type species: Trachycephalus lichenatus Gosse, 1851 (5 Hyla crucialis Harlan, 1826), by original designation. DIAGNOSIS: This genus is diagnosed by 43 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. No morphological synapomorphies are known for this genus. CONTENTS: Eight species. Osteopilus brunneus (Gosse, 1851); Osteopilus crucialis (Harlan, 1826); Osteopilus dominicensis (Tschudi, 1838); Osteopilus marianae (Dunn, 1926); Osteopilus pulchrilineatus (Cope ‘‘1869’’ [1870]); Osteopilus septentrionalis (Duméril and Bibron, 1841); Osteopilus vastus (Cope, 1871); Osteopilus wilderi (Dunn, 1925)., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 109, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Itapotihyla, new genus TYPE SPECIES: Hyla langsdorffii Duméril and Bibron, 1841. DIAGNOSIS: Molecular autapomorphies include 122 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these transformations. A possible morphological autapomorphy is the presence of a prominent subcloacal flap. ETYMOLOGY: From Itapoti 1 ­ Hyla. The generic name is an allusion to the resemblance of the unique known species of this genus with lichens and mosses. Itapoti is a Tupi­Guarani term, a composition of ‘‘ita´’’ (5 rock) with ‘‘poti’’ (5 flower or to flourish), which means lichen or moss. CONTENTS: Monotypic. Itapotihyla langsdorffii (Duméril and Bibron, 1841), new comb.

Nyctimantis Boulenger, 1882 TYPE SPECIES: Nyctimantis rugiceps Boulenger, 1882, by monotypy. DIAGNOSIS: Molecular autapomorphies include 139 transformations in mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. Possible morphological autapomorphies are the development of an irregular orbital flange in the frontoparietal, and the sphenethmoid almost completely concealed dorsally by the frontoparietals and nasals (Duellman and Trueb, 1976). CONTENTS: Monotypic. Nyctimantis rugiceps Boulenger, 1882., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 109, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Plectrohyla bistincta Group DIAGNOSIS: Exemplars of this species group in our analysis are diagnosed by 16 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these transformations. We are not aware of any morphological synapomorphy supporting this group. CONTENTS: Twenty­one species. Plectrohyla ameibothalame (Canseco­Márquez, Mendelson, and Gutiérrez­Mayén et al., 2002), new comb.; Plectrohyla arborescandens (Taylor, ‘‘1938’’[1939]), new comb.; Plectrohyla bistincta (Cope, 1877), new comb.; Plectrohyla calthula (Ustach, Mendelson, McDiarmid, and Campbell, 2000), new comb.; Plectrohyla calvicollina (Toal, 1994), new comb.; Plectrohyla celata (Toal and Mendelson, 1995), new comb.; Plectrohyla cembra (Caldwell, 1974), new comb.; Plectrohyla charadricola (Duellman, 1964), new comb.; Plectrohyla chryses (Adler, 1965), new comb.; Plectrohyla crassa (Brocchi, 1877), new comb.; Plectrohyla cyanomma (Caldwell, 1974), new comb.; Plectrohyla cyclada (Campbell and Duellman, 2000) new comb.; Plectrohyla hazelae (Taylor, 1940), new comb.; Plectrohyla labedactyla (Mendelson and Toal, 1996), new comb.; Plectrohyla mykter (Adler and Dennis, 1972), new comb.; Plectrohyla pachyderma, (Taylor, 1942), new comb.; Plectrohyla pentheter (Adler, 1965), new comb.; Plectrohyla psarosema (Campbell and Duellman, 2000), new comb.; Plectrohyla robertsorum (Taylor, 1940), new comb.; Plectrohyla sabrina (Caldwell, 1974), new comb.; Plectrohyla siopela, (Duellman, 1968), new comb.; Plectrohyla thorectes (Adler, 1965), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on pages 104-105, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Osteocephalus Steindachner, 1862 TYPE SPECIES: Osteocephalus taurinus Steindachner, 1862, by subsequent designation of Kellogg (1932). DIAGNOSIS: This genus is diagnosed by 34 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy supporting this genus. CONTENTS: Seventeen species. Osteocephalus buckleyi (Boulenger, 1882); Osteocephalus cabrerai (Cochran and Goin, 1970); Osteocephalus deridens Jungfer, Ron, Seipp, and Almendáriz, 2000; Osteocephalus elkejungingerae (Henle, 1981); Osteocephalus exophthalmus Smith and Noonan, 2001; Osteocephalus fuscifacies Jungfer, Ron, Seipp, and Almendáriz, 2000; Osteocephalus heyeri Lynch, 2002; Osteocephalus leoniae Jungfer and Lehr, 2001; Osteocephalus leprieurii (Duméril and Bibron, 1841); Osteocephalus mutabor Jungfer and Hödl, 2002; Osteocephalus oophagus Jungfer and Schiesari, 1995; Osteocephalus pearsoni (Gaige, 1929); Osteocephalus planiceps Cope, 1874; Osteocephalus subtilis Martins and Cardoso, 1987; Osteocephalus taurinus Steindachner, 1862; Osteocephalus verruciger (Werner, 1901); Osteocephalus yasuni Ron and Pramuk, 1999., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 109, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Tlalocohyla, new genus TYPE SPECIES: Hyla smithii Boulenger, 1902. DIAGNOSIS: This genus is diagnosed by 92 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy. ETYMOLOGY: From Tlaloc, the Olmec God of the rain, 1 connecting ­ o 1 Hyla. The gender is feminine. COMMENTS: The inclusion of Hyla godmani and H. loquax is tentative and based on its association with H. picta and H. smithii in the former H. godmani group by Duellman (2001). The larvae of H. loquax and H. smithii share a reduction in the length of the third posterior tooth row (Caldwell, 1986; Lee, 1996). This feature is not present in the larvae of H. godmani as described by Duellman (1970). CONTENTS: Four species. Tlalocohyla godmani (Günther, 1901), new comb.; Tlalocohyla loquax (Gaige and Stuart, 1934), new comb.; Tlalocohyla picta (Günther, 1901), new comb.; Tlalocohyla smithii (Boulenger, 1902), new comb.

Plectrohyla bistincta Group DIAGNOSIS: Exemplars of this species group in our analysis are diagnosed by 16 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these transformations. We are not aware of any morphological synapomorphy supporting this group. CONTENTS: Twenty­one species. Plectrohyla ameibothalame (Canseco­Márquez, Mendelson, and Gutiérrez­Mayén et al., 2002), new comb.; Plectrohyla arborescandens (Taylor, ‘‘1938’’[1939]), new comb.; Plectrohyla bistincta (Cope, 1877), new comb.; Plectrohyla calthula (Ustach, Mendelson, McDiarmid, and Campbell, 2000), new comb.; Plectrohyla calvicollina (Toal, 1994), new comb.; Plectrohyla celata (Toal and Mendelson, 1995), new comb.; Plectrohyla cembra (Caldwell, 1974), new comb.; Plectrohyla charadricola (Duellman, 1964), new comb.; Plectrohyla chryses (Adler, 1965), new comb.; Plectrohyla crassa (Brocchi, 1877), new comb.; Plectrohyla cyanomma (Caldwell, 1974), new comb.; Plectrohyla cyclada (Campbell and Duellman, 2000) new comb.; Plectrohyla hazelae (Taylor, 1940), new comb.; Plectrohyla labedactyla (Mendelson and Toal, 1996), new comb.; Plectrohyla mykter (Adler and Dennis, 1972), new comb.; Plectrohyla pachyderma, (Taylor, 1942), new comb.; Plectrohyla pentheter (Adler, 1965), new comb.; Plectrohyla psarosema (Campbell and Duellman, 2000), new comb.; Plectrohyla robertsorum (Taylor, 1940), new comb.; Plectrohyla sabrina (Caldwell, 1974), new comb.; Plectrohyla siopela, (Duellman, 1968), new comb.; Plectrohyla thorectes (Adler, 1965), new comb.

Bokermannohyla circumdata Group DIAGNOSIS: This species group is diagnosed by 52 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. A putative morphological synapomorphy of this group is the presence of (usually thin) dark vertical stripes on the posterior surface of the thigh (Heyer, 1985). CONTENTS: Fifteen species. Bokermannohyla ahenea (Napoli and Caramaschi, 2004), new comb.; Bokermannohyla astartea (Bokermann, 1967), new comb.; Bokermannohyla caramaschii (Napoli, 2005) new comb.; Bokermannohyla carvalhoi (Peixoto, 1981), new comb.; Bokermannohyla circumdata (Cope, ‘‘1870’’ [1871]), new comb.; Bokermannohyla feioi (Napoli and Caramaschi, 2004), new comb.; Bokermannohyla gouveai (Peixoto and Cruz, 1992), new comb.; Bokermannohyla hylax (Heyer, 1985), new comb.; Bokermannohyla ibitipoca (Caramaschi and Feio, 1990), new comb.; Bokermannohyla izeckshoni (Jim and Caramaschi, 1979), new comb.; Bokermannohyla lucianae (Napoli and Pimenta, 2003), new comb.; Bokermannohyla luctuosa (Pombal and Haddad, 1993), new comb.; Bokermannohyla nanuzae (Bokermann and Sazima, 1973), new comb.; Bokermannohyla ravida (Caramaschi, Napoli and Bernardes, 2001), new comb.; Bokermannohyla sazimai (Cardoso and Andrade, ‘‘1982’’ [1983]), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on pages 82-83, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Scinax ruber Clade DIAGNOSIS: This clade is supported by 53 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. A morphological synapomorphy suggested for this clade by Faivovich (2002) is the proctodeal tube positioned above the margin of the lower fin. COMMENTS: Faivovich (2002) was skeptical about the monophyly of the S. ruber clade; however, the present analysis recovers it as monophyletic, with a considerable number of transformations supporting its monophyly. As in the case of several of the synapomorphies suggested by Faivovich’s (2002) analysis for Scinax, we are unsure as to whether the suggested morphological synapomorphies are optimized identically in our analysis. In particular, we do not know the taxonomic distribution within Dendropsophini for two other synapomorphies proposed for this clade (Faivovich, 2002): the arytenoids with a dorsal prominence developed over the pharyngeal margin, and absence of the lateral m. extensor brevis distalis digiti V (pes). Preliminary observation on the larvae of some species of Sphaenorhynchus (Sphaenorhynchus bromelicola, S. orophilus, S. pauloalvini, and S. prasinus; Faivovich, personal obs.) indicate that their proctodeal tubes are attached to the free margin of the lower fin, similar to the S. catharinae clade, instead of having the characteristic position seen in larvae of the S. ruber clade. Scinax megapodius and S. trachythorax are considered here to be junior synonyms of S. fuscovarius for reasons discussed in appendix 4. There are two species, Hyla dolloi and H. karenanneae, that upon examination of their type series we consider to be species of Scinax (see appendix 4 for further comments on them). CONTENTS: Fifty­six species. Eleven assigned to two groups, 43 unassigned to any group., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 96, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Phyllomedusa perinesos Group DIAGNOSIS: A possible synapomorphy of this group is the purple coloration on the hands, feet, flanks, and concealed surfaces, as well as the purple venter with white granules (Cannatella, 1982). COMMENTS: We did not include any exemplar of this group in the analysis. Its monophyly is tentatively assumed following Cannatella (1982) and is based on the evidence mentioned above. CONTENTS: Four species. Phyllomedusa baltea Duellman and Toft, 1979; Phyllomedusa duellmani Cannatella, 1982; Phyllomedusa ecuatoriana Cannatella, 1982; Phyllomedusa perinesos Duellman, 1973., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 117, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Hyloscirtus armatus Group DIAGNOSIS: This species group is diagnosed by 103 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. Duellman et al. (1997) suggested four synapomorphies of the H. armatus group: the presence of keratincovered bony spines on the proximal ventral surface of the humerus, on the expanded distal element of the prepollex, and on the first metacarpal; forearms hypertrophied; tadpole tail long with low fins and bluntly rounded tip; and the presence of a ‘‘shelf’’ on the larval upper jaw sheath. COMMENTS: Our observations of breeding males of the two species of this group indicate the presence of darkly pigmented, keratinized spicules in the dorsum, head (particularly lips), forelimbs, undersides of forelimbs, and pectoral and abdominal region. As the breeding biology of this and the other two species groups of the genus becomes better known, it will be possible to understand if the presence of these spicules are a putative synapomorphy of the H. armatus group. CONTENTS: Two species. Hyloscirtus armatus (Boulenger, 1902), new comb.; Hyloscirtus charazani (Vellard, 1970), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 84, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Hyla arborea Group DIAGNOSIS: This species group is diagnosed by 37 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy supporting this group. COMMENTS: The contents of the Hyla arborea group are restricted to avoid its paraphyly. The inclusion of the species that were not included in the present analysis and do not show the NOR in chromosome 6 is tentative, because no evidence, other than the molecular data presented here, is known to support its monophyly. CONTENTS: Fourteen species. Hyla annectans (Jerdon, 1870); Hyla arborea (Linnaeus, 1758); Hyla chinensis Günther, 1858; Hyla hallowellii Thomson, 1912; Hyla immaculata Boettger, 1888; Hyla intermedia Boulenger, 1882; Hyla meridionalis Boettger, 1874; Hyla sanchiangensis Pope, 1929; Hyla sarda (De Betta, 1853); Hyla savignyi Audouin, 1827; Hyla simplex Boettger, 1901; Hyla tsinlingensis Liu and Hu, 1966; Hyla ussuriensis Nikolsky, 1918; Hyla zhaopingensis Tang and Zhang, 1984., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on pages 101-102, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Aplastodiscus albosignatus Group DIAGNOSIS: This species group is diagnosed by 42 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. A possible morphological synapomorphy of this group is the presence of elaborate tubercles and ornamentation around the cloacal region (Cruz and Peixoto, ‘‘1985’’ [1987]). CONTENTS: Seven species. Aplastodiscus albosignatus (A. Lutz and B. Lutz, 1938), new comb.; Aplastodiscus callipygius (Cruz and Peixoto, ‘‘1984’’ [1985]), new comb.; Aplastodiscus cavicola (Cruz and Peixoto, ‘‘1984’’ [1985]), new comb.; Aplastodiscus flumineus (Cruz and Peixoto, ‘‘1984’’ [1985]), new comb.; Aplastodiscus ibirapitanga (Cruz, Pimenta, and Silvano, 2003), new comb.; Aplastodiscus leucopygius (Cruz and Peixoto, ‘‘1984’’ [1985]), new comb.; Aplastodiscus sibilatus (Cruz, Pimenta, and Silvano, 2003), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 82, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Hyla cinerea Group DIAGNOSIS: This species group is diagnosed by 35 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy supporting this group. COMMENTS: Hyla femoralis is excluded from the H. cinerea group to avoid the paraphyly of the group. CONTENTS: Three species. Hyla cinerea (Schneider, 1799); Hyla gratiosa LeConte, ‘‘1856’’ [1857]; Hyla squirella Bosc, 1800., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 102, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Ecnomiohyla, new genus TYPE SPECIES: Hypsiboas miliarius Cope, 1886. DIAGNOSIS: This genus is diagnosed by 37 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy supporting this genus. ETYMOLOGY: From the Greek, ecnomios, meaning marvelous, unusual; an obvious reference to the incredible frogs of the Hyla tuberculosa group. The gender is feminine. COMMENTS: This new genus contains the Hyla tuberculosa group, excluding H. dendrophasma, and including one species of the H. miotympanum group as well. Erecting a new genus for this clade is the only way of being consistent with the new monophyletic taxonomy that is proposed for hylids. Although naming the former H. tuberculosa group as a genus constitutes a testable claim of monophyly, we expect that it will ultimately be found to be two or three different clades, with one of these being the one named here. CONTENTS: Ten species. Ecnomiohyla echinata (Duellman, 1962), new comb.; Ecnomiohyla fimbrimembra (Taylor, 1948), new comb.; Ecnomiohyla miliaria (Cope, 1886), new comb.; Ecnomiohyla minera (Wilson, McCranie, and Williams, 1985), new comb.; Ecnomiohyla miotympanum (Cope, 1863), new comb.; Ecnomiohyla phantasmagoria (Dunn, 1943); Ecnomiohyla salvaje (Wilson, McCranie, and Williams, 1985), new comb.; Ecnomiohyla thysanota (Duellman, 1966), new comb.; Ecnomiohyla tuberculosa (Boulenger, 1882), new comb.; Ecnomiohyla valancifer (Firschein and Smith, 1956), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 100, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Phyllomedusa burmeisteri Group DIAGNOSIS: We are not aware of any synapomorphy of this group. COMMENTS: We included only a single species of this group in our analysis, and as such we did not test its monophyly, but we recognize it following Pombal and Haddad (1992), pending a rigorous test of its monophyly. CONTENTS: Four species. Phyllomedusa burmeisteri Boulenger, 1882; Phyllomedusa distincta B. Lutz, 1950; Phyllomedusa iheringii Boulenger, 1885; Phyllomedusa tetraploidea Pombal and Haddad, 1992., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 117, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Bokermannohyla pseudopseudis Group DIAGNOSIS: This group is diagnosed by 48 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy for this group. COMMENTS: Considering our results, which show the undescribed species of the Hyla pseudopseudis group (Hyla sp. 6) to be the sister taxon of an undescribed species similar with H. alvarengai (Hyla sp. 9), we are tentatively including the H. alvarengai in this species group. Eterovick and Brandão (2001) characterized this group on the basis of the presence of short, lateral irregular tooth rows and for having more tooth rows (between six and eight rows) in the oral discs of the larvae than do those of the Bokermannohyla circumdata group. However, the tadpole of H. ibitiguara, included in this group by Caramaschi et al. (2001), has a labial tooth formula of 2/4 (Cardoso, 1983) and seems to lack the short, lateral irregular tooth rows, as do tadpoles of H. alvarengai (Sazima and Bokermann, 1977). CONTENTS: Four species. Bokermannohyla alvarengai (Bokermann, 1956), new comb.; Bokermannohyla ibitiguara (Cardoso, 1983), new comb.; Bokermannohyla pseudopseudis (Miranda­Ribeiro, 1937), new comb.; Bokermannohyla saxicola (Bokermann, 1964), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on pages 83-84, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Bokermannohyla claresignata Group DIAGNOSIS: We are not aware of any synapomorphy supporting the monophyly of this group; see below. COMMENTS: We did not include any exemplar of this group, and as such we did not test its monophyly. See the earlier discussion regarding its position in the South American I clade. Considering the topology of Cophomantini, synapomorphies suggested for the Bokermannohyla claresignata group can only be maintained if assumed to be reversals (i.e., a enlarged larval oral disc, complete marginal papillae, and large number of labial tooth rows are also present in Myersiohyla new genus and the Hyloscirtus armatus group; complete marginal papillae and large number of labial tooth rows are also present in the H. bogotensis and H. larinopygion groups; the marginal papillae are also complete or with an extremely reduced gap in other species of Bokermannohyla). This would be unproblematic if it were a result of the analysis, but we prefer not to assume it a priori. While these character states cannot be considered at this stage to support the monophyly of the B. claresignata group, considering that its two species are barely distinguishable from each other, we think its nonmonophyly is unlikely and we continue to recognize the group as a hypothesis to be tested. CONTENTS: Two species. Bokermannohyla claresignata (A. Lutz and B. Lutz, 1939), new comb.; Bokermannohyla clepsydra (A. Lutz, 1925), new comb.

Tlalocohyla, new genus TYPE SPECIES: Hyla smithii Boulenger, 1902. DIAGNOSIS: This genus is diagnosed by 92 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy. ETYMOLOGY: From Tlaloc, the Olmec God of the rain, 1 connecting ­ o 1 Hyla. The gender is feminine. COMMENTS: The inclusion of Hyla godmani and H. loquax is tentative and based on its association with H. picta and H. smithii in the former H. godmani group by Duellman (2001). The larvae of H. loquax and H. smithii share a reduction in the length of the third posterior tooth row (Caldwell, 1986; Lee, 1996). This feature is not present in the larvae of H. godmani as described by Duellman (1970). CONTENTS: Four species. Tlalocohyla godmani (Günther, 1901), new comb.; Tlalocohyla loquax (Gaige and Stuart, 1934), new comb.; Tlalocohyla picta (Günther, 1901), new comb.; Tlalocohyla smithii (Boulenger, 1902), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 107, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Aplastodiscus perviridis Group DIAGNOSIS: This species group is diagnosed by 58 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. Apparent morphological synapomorphies of this group include the bicolored iris and the absence of webbing between toes I and II (known instances of homoplasy within hylids occur in some Scinax and in various groups of Lophiohylini) and reduction of webbing between the other toes (Garcia et al., 2001). CONTENTS: Two species. Aplastodiscus cochranae (Mertens, 1952); Aplastodiscus perviridis A. Lutz in B. Lutz, 1950. Bokermannohyla, new genus TYPE SPECIES: Hyla circumdata Cope, ‘‘1870’’ [1871]. DIAGNOSIS: This genus is diagnosed by 65 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy. ETYMOLOGY: This genus is dedicated to Werner Carlos Augusto Bokermann (1929– 1995), as homage to his contribution to the knowledge of Brazilian anurans. He also described several species now included in the new genus. The name derives from Bokermann 1 connecting ­ o 1 Hyla. We are adopting the ending ­ hyla for several of the new genera described here, most of which contain species groups formerly placed in Hyla. The gender is feminine. COMMENTS: Bokermannohyla includes all species previously allocated in the Hyla circumdata, H. martinsi, and H. pseudopseudis groups. We include tentatively the H. claresignata group pending the inclusion of its species in the analysis, because it was associated to the H. circumdata group by Bokermann (1972) and Jim and Caramaschi (1979). Hyla alvarengai is also included because our analysis shows that Hyla sp. 9 (aff. H. alvarengai) is nested within this new genus. These species groups should be maintained within Bokermannohyla until their monophyly is rigorously tested. CONTENTS: Twenty­three species, placed in four species groups., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 82, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Phyllomedusa tarsius Group DIAGNOSIS: We are not aware of any synapomorphy supporting the monophyly of this group. COMMENTS: We included a single species of this group, and as such we did not test its monophyly, but we continue to recognize it following De la Riva (1999) until its monophyly is rigorously tested. CONTENTS: Four species. Phyllomedusa boliviana Boulenger, 1902; Phyllomedusa camba De la Riva, 2000; Phyllomedusa sauvagii Boulenger, 1882; Phyllomedusa tarsius (Cope, 1868). Species of Phyllomedusa Unassigned to Group There are several species that are currently not assigned to any group. These are: Phyllomedusa atelopoides Duellman, Cadle, and Cannatella, 1988; Phyllomedusa bicolor (Boddaert, 1772); Phyllomedusa coelestis (Cope, 1874); Phyllomedusa palliata Peters, ‘‘1872’’ [1873]; Phyllomedusa tomopterna (Cope, 1868); Phyllomedusa trinitatis Mertens, 1926; Phyllomedusa vaillanti Boulenger, 1882; and Phyllomedusa venusta Duellman and Trueb, 1967., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on pages 117-118, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Bokermannohyla pseudopseudis Group DIAGNOSIS: This group is diagnosed by 48 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy for this group. COMMENTS: Considering our results, which show the undescribed species of the Hyla pseudopseudis group (Hyla sp. 6) to be the sister taxon of an undescribed species similar with H. alvarengai (Hyla sp. 9), we are tentatively including the H. alvarengai in this species group. Eterovick and Brandão (2001) characterized this group on the basis of the presence of short, lateral irregular tooth rows and for having more tooth rows (between six and eight rows) in the oral discs of the larvae than do those of the Bokermannohyla circumdata group. However, the tadpole of H. ibitiguara, included in this group by Caramaschi et al. (2001), has a labial tooth formula of 2/4 (Cardoso, 1983) and seems to lack the short, lateral irregular tooth rows, as do tadpoles of H. alvarengai (Sazima and Bokermann, 1977). CONTENTS: Four species. Bokermannohyla alvarengai (Bokermann, 1956), new comb.; Bokermannohyla ibitiguara (Cardoso, 1983), new comb.; Bokermannohyla pseudopseudis (Miranda­Ribeiro, 1937), new comb.; Bokermannohyla saxicola (Bokermann, 1964), new comb.

Dendropsophus columbianus Group DIAGNOSIS: The only morphological synapomorphy suggested for this group is the presence of two close, triangular lateral spaces between the cricoid and arytenoids at the posterior part of the larynx (Kaplan, 1999). COMMENTS: We included a single exemplar of this group, and as such we did not test its monophyly, but following Kaplan (1999) we recognize it on the basis of the evidence mentioned above. CONTENTS: Three species. Dendropsophus bogerti (Cochran and Goin, 1970), new comb.; Dendropsophus carnifex (Duellman, 1969), new comb.; Dendropsophus columbianus (Boettger, 1892), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 90, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Dendropsophus labialis Group DIAGNOSIS: We are not aware of any synapomorphy for this group. COMMENTS: We included a single exemplar of this group in the analysis, and as such we did not test its monophyly. Following Duellman and Trueb (1983) and Duellman (1989), we continue to recognize the group pending a rigorous test of its monophyly. CONTENTS: Three species. Dendropsophus labialis (Peters, 1863), new comb.; Dendropsophus meridensis (Rivero, 1961) new comb.; Dendropsophus pelidna (Duellman, 1989), new comb.

Itapotihyla, new genus TYPE SPECIES: Hyla langsdorffii Duméril and Bibron, 1841. DIAGNOSIS: Molecular autapomorphies include 122 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these transformations. A possible morphological autapomorphy is the presence of a prominent subcloacal flap. ETYMOLOGY: From Itapoti 1 ­ Hyla. The generic name is an allusion to the resemblance of the unique known species of this genus with lichens and mosses. Itapoti is a Tupi­Guarani term, a composition of ‘‘ita´’’ (5 rock) with ‘‘poti’’ (5 flower or to flourish), which means lichen or moss. CONTENTS: Monotypic. Itapotihyla langsdorffii (Duméril and Bibron, 1841), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 109, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Scinax catharinae Clade DIAGNOSIS: This clade is diagnosed by 90 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. Morphological synapomorphies suggested for this clade by Faivovich (2002) are absence of the anterior process of the suprascapula, internal vocal sac, distal division of the middle branch of the m. extensor digitorum comunis longus, and insertion of the medial side of this branch on the tendon of the m. extensor brevis medius digiti IV. COMMENTS: Regardless of problems imposed by the present results to interpretation of the possible synapomorphies of Scinax resulting from Faivovich’s (2002) analysis, the sparse available knowledge on the taxonomic distribution of the transformations supporting the monophyly of the very distinctive S. catharinae clade suggests that most of them still hold in the present analysis. An exception is the m. depressor mandibulae without an origin from the dorsal fascia at the level of the m. dorsalis scapulae, which also occurs in Hyla uruguaya, rendering its optimization ambiguous in our analysis., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 95, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Hypsiboas pellucens Group DIAGNOSIS: This species group is diagnosed by 115 transformations in mitochondrial ribosomal genes. See appendix 5 for a complete list of these transformations. We are not aware of any morphological synapomorphy for the group. COMMENTS: We recognize this new species group to include the clade composed of the fragment of the former Hyla albomarginata complex that includes H. pellucens and H. rufitela. The inclusion of H. rubracyla is tentative, based on its previous association with H. pellucens. CONTENTS: Three species. Hypsiboas pellucens (Werner, 1901), new comb.; Hypsiboas rubracylus (Cochran and Goin, 1970), new comb.; Hypsiboas rufitelus (Fouquette, 1958), new comb.

Hypsiboas punctatus Group DIAGNOSIS: This species group is diagnosed by 30 transformations in mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy for the group. COMMENTS: We do not see any reason to keep the former Hyla granosa and H. punctata groups separated, as our analysis shows that the two nominal species form a monophyletic group and they are phenotypically similar, so we include all the species in the Hypsiboas punctatus group. The inclusion of Hyla alemani, H. atlantica, H. hobbsi, and H. ornatissima is tentative, based on their previous association with the former H. granosa and H. punctata groups. The inclusion of Hyla picturata is based on our analysis. CONTENTS: Eight species. Hypsiboas alemani (Rivero, 1964), new comb.; Hypsiboas atlanticus (Caramaschi and Velosa, 1996), new comb.; Hypsiboas granosus (Boulenger, 1882), new comb.; Hypsiboas hobbsi (Cochran and Goin, 1970), new comb.; Hypsiboas ornatissimus (Noble, 1923), new comb.; Hypsiboas picturatus (Boulenger, 1882), new comb.; Hypsiboas punctatus (Schneider, 1799), new comb.; Hypsiboas sibleszi (Rivero, 1971), new comb.

Hypsiboas pellucens Group DIAGNOSIS: This species group is diagnosed by 115 transformations in mitochondrial ribosomal genes. See appendix 5 for a complete list of these transformations. We are not aware of any morphological synapomorphy for the group. COMMENTS: We recognize this new species group to include the clade composed of the fragment of the former Hyla albomarginata complex that includes H. pellucens and H. rufitela. The inclusion of H. rubracyla is tentative, based on its previous association with H. pellucens. CONTENTS: Three species. Hypsiboas pellucens (Werner, 1901), new comb.; Hypsiboas rubracylus (Cochran and Goin, 1970), new comb.; Hypsiboas rufitelus (Fouquette, 1958), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 87, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Dendropsophus labialis Group DIAGNOSIS: We are not aware of any synapomorphy for this group. COMMENTS: We included a single exemplar of this group in the analysis, and as such we did not test its monophyly. Following Duellman and Trueb (1983) and Duellman (1989), we continue to recognize the group pending a rigorous test of its monophyly. CONTENTS: Three species. Dendropsophus labialis (Peters, 1863), new comb.; Dendropsophus meridensis (Rivero, 1961) new comb.; Dendropsophus pelidna (Duellman, 1989), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 91, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Dendropsophus parviceps Group DIAGNOSIS: This species group is diagnosed by 27 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy supporting the monophyly of this group. COMMENTS: We expressed our scepticism regarding the monophyly of this group, as currently defined, but found no evidence to reject monophyly. We recognize the group pending a rigorous test of its monophyly. CONTENTS: Fifteen species. Dendropsophus allenorum (Duellman and Trueb, 1989), new comb.; Dendropsophus bokermanni (Goin, 1960), new comb.; Dendropsophus brevifrons (Duellman and Crump, 1974), new comb.; Dendropsophus gaucheri (Lescure and Marty, 2001), new comb.; Dendropsophus giesleri (Mertens, 1950), new comb.; Dendropsophus grandisonae (Goin, 1966), new comb.; Dendropsophus koechlini (Duellman and Trueb, 1989), new comb.; Dendropsophus luteoocellatus (Roux, 1927), new comb.; Dendropsophus microps (Peters, 1872), new comb.; Dendropsophus parviceps (Boulenger, 1882), new comb.; Dendropsophus pauiniensis (Heyer, 1977), new comb.; Dendropsophus ruschii (Weygoldt and Peixoto, 1987), new comb.; Dendropsophus schubarti (Bokermann, 1963), new comb.; Dendropsophus subocularis (Dunn, 1934), new comb.; Dendropsophus timbeba (Martins and Cardoso, 1987), new comb. Species of Dendropsophus Unassigned to Group There are several species of Dendropsophus that have not been associated with any group. These are: Dendropsophus amicorum (Mijares­Urrutia, 1998), new comb.; Dendropsophus battersbyi (Rivero, 1961), new comb.; Dendropsophus haraldschultzi (Bokermann, 1962), new comb.; Dendropsophus stingi (Kaplan, 1994), new comb.; Dendropsophus tintinnabulum (Melin, 1941), new comb.; Dendropsophus yaracuyanus (Mijares­Urrutia and Rivero, 2000), new comb.

Dendropsophus columbianus Group DIAGNOSIS: The only morphological synapomorphy suggested for this group is the presence of two close, triangular lateral spaces between the cricoid and arytenoids at the posterior part of the larynx (Kaplan, 1999). COMMENTS: We included a single exemplar of this group, and as such we did not test its monophyly, but following Kaplan (1999) we recognize it on the basis of the evidence mentioned above. CONTENTS: Three species. Dendropsophus bogerti (Cochran and Goin, 1970), new comb.; Dendropsophus carnifex (Duellman, 1969), new comb.; Dendropsophus columbianus (Boettger, 1892), new comb.

Hypsiboas punctatus Group DIAGNOSIS: This species group is diagnosed by 30 transformations in mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. We are not aware of any morphological synapomorphy for the group. COMMENTS: We do not see any reason to keep the former Hyla granosa and H. punctata groups separated, as our analysis shows that the two nominal species form a monophyletic group and they are phenotypically similar, so we include all the species in the Hypsiboas punctatus group. The inclusion of Hyla alemani, H. atlantica, H. hobbsi, and H. ornatissima is tentative, based on their previous association with the former H. granosa and H. punctata groups. The inclusion of Hyla picturata is based on our analysis. CONTENTS: Eight species. Hypsiboas alemani (Rivero, 1964), new comb.; Hypsiboas atlanticus (Caramaschi and Velosa, 1996), new comb.; Hypsiboas granosus (Boulenger, 1882), new comb.; Hypsiboas hobbsi (Cochran and Goin, 1970), new comb.; Hypsiboas ornatissimus (Noble, 1923), new comb.; Hypsiboas picturatus (Boulenger, 1882), new comb.; Hypsiboas punctatus (Schneider, 1799), new comb.; Hypsiboas sibleszi (Rivero, 1971), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 88, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Corythomantis Boulenger, 1896 TYPE SPECIES: Corythomantis greeningi Boulenger, 1896, by monotypy. DIAGNOSIS: Molecular autapomorphies include 132 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these transformations. Morphological autapomorphies of this monotypic genus include the absence of palatines, and nasals that conceal the alary processes of premaxillaries (Trueb, 1970a). CONTENTS: Monotypic. Corythomantis greeningi Boulenger, 1896., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 108, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Dendropsophus minimus Group DIAGNOSIS: No synapomorphy is known for this group. COMMENTS: We included a single species of this group in the analisis, and as such we did not test its monophyly and we are not aware of any evidence supporting it. Following Duellman (1982), we continue to recognize it pending a rigorous test of its monophyly. CONTENTS: Four species. Dendropsophus aperomeus (Duellman, 1982), new comb.; Dendropsophus minimus (Ahl, 1933), new comb.; Dendropsophus miyatai (Vigle and Goberdhan­Vigle, 1990), new comb.; Dendropsophus riveroi (Cochran and Goin, 1970), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 92, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Pseudacris regilla Clade DIAGNOSIS: This clade is diagnosed by molecular data presented by Moriarty and Cannatella (2004). CONTENTS: Two species. Pseudacris cadaverina (Cope, 1866); Pseudacris regilla (Baird and Girard, 1852)., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 106, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Acris Duméril and Bibron, 1841 TYPE SPECIES: Rana gryllus LeConte, 1825, by subsequent designation of Fitzinger (1843). DIAGNOSIS: This genus is diagnosed by 138 transformations in nuclear and mitochondrial proteins and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. Other apparent synapomorphies include the differentiation of the m. intermandibularis into an apical supplementary element (Tyler, 1971), and diploid chromosome number of 22 (Bushnell et al., 1939; Cole, 1966; Duellman, 1970). CONTENTS: Two species. Acris crepitans Baird, 1854; Acris gryllus (LeConte, 1825)., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 99, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Scinax perpusillus Group DIAGNOSIS: Presumed synapomorphies of this group are the oviposition in bromeliads and the extreme reduction of webbing between toes II and III (Peixoto, 1987; Faivovich, 2002). COMMENTS: The monophyly of this group was not tested by Faivovich (2002) because only one species of the group was available for his analysis, where it obtained as the sister taxon of all exemplars of the S. catharinae clade. For these two reasons, we continue recognizing this group until its monophyly is rigorously tested. CONTENTS: Seven species. Scinax alcatraz (B. Lutz, 1973); Scinax arduous Peixoto, 2002; Scinax atratus (Peixoto, 1989); Scinax littoreus (Peixoto, 1988); Scinax melloi (Peixoto, 1989); Scinax perpusillus (A. Lutz and B. Lutz, 1939); Scinax v ­signatus (B. Lutz, 1968)., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 96, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Bokermannohyla claresignata Group DIAGNOSIS: We are not aware of any synapomorphy supporting the monophyly of this group; see below. COMMENTS: We did not include any exemplar of this group, and as such we did not test its monophyly. See the earlier discussion regarding its position in the South American I clade. Considering the topology of Cophomantini, synapomorphies suggested for the Bokermannohyla claresignata group can only be maintained if assumed to be reversals (i.e., a enlarged larval oral disc, complete marginal papillae, and large number of labial tooth rows are also present in Myersiohyla new genus and the Hyloscirtus armatus group; complete marginal papillae and large number of labial tooth rows are also present in the H. bogotensis and H. larinopygion groups; the marginal papillae are also complete or with an extremely reduced gap in other species of Bokermannohyla). This would be unproblematic if it were a result of the analysis, but we prefer not to assume it a priori. While these character states cannot be considered at this stage to support the monophyly of the B. claresignata group, considering that its two species are barely distinguishable from each other, we think its nonmonophyly is unlikely and we continue to recognize the group as a hypothesis to be tested. CONTENTS: Two species. Bokermannohyla claresignata (A. Lutz and B. Lutz, 1939), new comb.; Bokermannohyla clepsydra (A. Lutz, 1925), new comb., Published as part of FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, pp. 1-240 in Bulletin of the American Museum of Natural History 2005 (294) on page 83, DOI: 10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2, http://zenodo.org/record/5363218

Dendropsophus leucophyllatus Group DIAGNOSIS: This species group is diagnosed by 35 transformations in mitochondrial ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. COMMENTS: On the basis of our molecular results, we are including Hyla anceps in this group. With the exception of this species, all other members of the group share the presence of pectoral glands in males and females (Duellman, 1970). CONTENTS: Eight species. Dendropsophus anceps (A. Lutz, 1929), new comb.; Dendropsophus bifurcus (Andersson, 1945), new comb.; Dendropsophus ebraccatus (Cope, 1874), new comb.; Dendropsophus elegans (Wied­Neuwied, 1824), new comb.; Dendropsophus leucophyllatus (Beireis, 1783), new comb.; Dendropsophus rossalleni (Goin, 1959), new comb.; Dendropsophus sarayacuensis (Shreve, 1935), new comb.; Dendropsophus triangulum (Günther, ‘‘1868’’ [1869]), new comb.