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Family Microhylidae Günther, 1858 The only species of the genus Elachistocleis known to occur in the state of Amapá is Elachistocleis helianneae Caramaschi, 2010 (Jowers et al. 2021). We considered the record of Elachistocleis sp. from Pereira-Júnior et al. (2013) to be E. helianneae based on the distribution and the overall morphology of the specimens (Costa-Campos, pers. obs.) and calls (Marinho et al. 2018). Fouquet et al. (2021b) showed in their integrative study that Otophryne pyburni is probably restricted to the western portion of the Guiana Shield, and that the populations of Otophryne inhabiting French Guiana and the Brazilian states of Amapá and Pará belong to an unconfirmed candidate species related to Otophryne robusta Boulenger, 1900. Therefore, we chose to consider the records of O. pyburni in Amapá as O. cf. robusta until more studies arise to evaluate its taxonomic status, confirming or contradicting this candidate species. Synapturanus zombie Fouquet, Leblanc, Fabre, Rodrigues, Menin, Courtois, Dewynter, Hölting, Ernst, Peloso & Kok, 2021 was recently described from French Guiana with one population recorded in the municipality of Oiapoque, northern Amapá (locality 46, Fig. 10) and a second potential population in the upper Rio Calçoene (2.3734° N, 51.3782° W; Fouquet et al. 2021a). We did not add this locality to the map because of the uncertainty on the population identity according to the original publication. We have found records of Synapturanus mirandaribeiroi Nelson & Lescure, 1975 from three localities in the Tumucumaque Mountains National Park (localities 5–7; Lima 2008). The two species have previously been confused with each other and have close distribution ranges, with one of the records from Tumucumaque (locality 5) very close to the Mitaraka massif, French Guiana, one of the localities with known populations of S. mirandaribeiroi (Fouquet et al. 2021). We then consider the records from the Tumucumaque Park (Lima 2008) as S. mirandaribeiroi, but we are aware that some of these records may actually be S. zombie Fouquet, Leblanc, Fabre, Rodrigues, Menin, Courtois, Dewynter, Hölting, Ernst, Peloso & Kok, 2021. Finally, we have an unvouchered record of S. zombie from the municipality of Serra do Navio, with only a picture (Fig. 9G). Although the picture is slightly out of focus, it is possible to see one important diagnostic character distinguishing this species from S. mirandaribeiroi: the dorsum with numerous orange spots and blotches (dorsum with diffuse mottled pattern in S. mirandaribeiroi). Therefore, we consider the photograph-based record of S. zombie in Serra do Navio, extending the distribution of this species to the central portion of Amapá, more than 250 km southeast from its type locality (Itoupé, French Guiana; 3.0230° N, 53.0955° W)., Published as part of Taucce, Pedro P. G., Costa-Campos, Carlos Eduardo, Carvalho, Thiago R. & Michalski, Fernanda, 2022, Anurans (Amphibia: Anura) of the Brazilian state of Amapá, eastern Amazonia: species diversity and knowledge gaps, pp. 96-130 in European Journal of Taxonomy 836 on page 118, DOI: 10.5852/ejt.2022.836.1919, http://zenodo.org/record/7061668, {"references":["Jowers M. J., Othman S. N., Borzee A., Rivas G. A., Sanchez-Ramirez S., Auguste R. J., Downie J. R., Read M. & Murphy J. C. 2021. Unraveling unique island colonization events in Elachistocleis frogs: phylogeography, cryptic divergence, and taxonomical implications. Organisms Diversity and Evolution 21 (1): 189 - 206. https: // doi. org / 10.1007 / s 13127 - 021 - 00487 - y","Marinho P., Carvalho T. R., Bang D. L., Teixeira B. F. V., Azarak P. A., Costa-Campos C. E. & Giaretta A. A. 2018. Advertisement calls, intraspecific variation and species diagnosis of six Brazilian species of Elachistocleis (Anura: Microhylidae: Gastrophryninae). Zootaxa 4521 (3): 357 - 375. https: // doi. org / 10.11646 / zootaxa. 4521.3.3","Fouquet A., Leblanc K., Framit M., Rejaud A., Rodrigues M. T., Castroviejo-Fisher S., Pelos P. L. V., Prates I., Manzi S., Suescun U., Baroni S., Moraes L. J. C. L., Recoder R., Souza S. M., Dal Vecchio F., Camacho A., Ghellere J. M., Rojas-Runjaic F. J. M., Gagliardi-Urrutia G., De Carvalho V. T., Gordo M., Menin M., Kok P. J. R., Hrbek T., Werneck F. P., Crawford A. J., Ron S. R., Mueses-Cisneros J. J., Rojas Zamora R. R., Pavan D., Ivo Simoes P., Ernst R. & Fabre A. - C. 2021 b. Species diversity and biogeography of an ancient frog clade from the Guiana Shield (Anura: Microhylidae: Adelastes, Otophryne, Synapturanus) exhibiting spectacular phenotypic diversification. Biological Journal of the Linnean Society 132: 233 - 256. https: // doi. org / 10.1093 / biolinnean / blaa 204","Fouquet A., Leblanc K., Fabre A. C., Rodrigues M. T., Menin M., Courtois E. A., Dewynter M., Holting M., Ernst R., Peloso P. & Kok P. J. R. 2021 a. Comparative osteology of the fossorial frogs of the genus Synapturanus (Anura, Microhylidae) with the description of three new species from the Eastern Guiana Shield. Zoologischer Anzeiger 293: 46 - 73. https: // doi. org / 10.1016 / j. jcz. 2021.05.003","Lima J. D. 2008. A herpetofauna do Parque Nacional do Montanhas do Tumucumaque, Amapa, Brasil, Expedicoes I a V. In: Bernhard E. (ed.) Inventarios Biologicos Rapidos no Parque Nacional Montanhas do Tumucumaque, Amapa, Brasil.: 38 - 50."]}

Family Leptodactylidae Werner, 1896 Leptodactylus bolivianus Boulenger, 1898 is distributed in the western and central portions of the Amazon Basin in Bolivia, Brazil, Colombia, Peru and Venezuela (Heyer & de Sá 2011), whereas Leptodactylus guianensis Heyer & de Sá, 2011 is distributed across the Guiana Shield. These two species are morphologically quite similar to each other and, before the description of L. guianensis in 2011, it was commonly confused with L. bolivianus in the literature (Heyer & de Sá 2011). We consider the record of L. cf. bolivianus from Lima (2006b) to be L. guianensis. Gazoni et al. (2021) recently revisited the systematics of the group of L. melanonotus using an integrative approach. One of the taxonomic results was the revalidation of L. intermedius, previously a junior synonym of Leptodactylus petersii Steindachner, 1864. Besides L. petersii, there are records of two other species of the group of L. melanonotus in Amapá: L. wagneri (Lima 2006) and Leptodactylus podicipinus Cope, 1862 (PereiraJúnior et al. 2013; Araújo & Costa 2015; Lima 2018; Costa-Campos & Freire 2019). All records previously associated with L. podicipinus in Amapá should be assigned to L. intermedius. Leptodactylus wagneri is distributed in western Amazonia and the records in eastern Amazonia correspond either to L. intermedius (see Gazoni et al. 2021) or to unnamed species, treated here as Leptodactylus sp., endemic to the Eastern Guiana Shield (Carvalho et al. in press). Leptodactylus leptodactyloides (Andersson, 1945) is widely distributed in Amazonia but reported in Amapá for the first time in this study. The species was confirmed to occur at one location (Comunidade Ariri), within the municipality of Macapá, based on DNA-barcoded individuals (Carvalho et al. in press)., Published as part of Taucce, Pedro P. G., Costa-Campos, Carlos Eduardo, Carvalho, Thiago R. & Michalski, Fernanda, 2022, Anurans (Amphibia: Anura) of the Brazilian state of Amapá, eastern Amazonia: species diversity and knowledge gaps, pp. 96-130 in European Journal of Taxonomy 836 on page 118, DOI: 10.5852/ejt.2022.836.1919, http://zenodo.org/record/7061668, {"references":["Heyer W. R. & de Sa R. O. 2011. Variation, Systematics, and Relationships of the Leptodactylus bolivianus Complex (Amphibia: Anura: Leptodactylidae). Smithsonian Contributions to Zoology 635: 1 - 58. https: // doi. org / 10.5479 / si. 00810282.635.1","Lima J. D. 2006 b. Inventarios Biologicos da Herpetofauna na Floresta Nacional do Amapa - Expedicoes I e II. In: Bernard E. (ed.) Inventarios Biologicos na Floresta Nacional do Amapa - Expedicoes I e II: 57 - 77. Macapa, Amapa.","Gazoni T., Lyra M. L., Ron S. R., Strussmann C., Baldo D., Narimatsu H., Pansonato A., Schneider R. G., Giaretta A. A., Haddad C. F. B., Parise-Maltempi P. P. & Carvalho T. R. 2021. Revisiting the systematics of the Leptodactylus melanonotus group (Anura: Leptodactylidae): Redescription of L. petersii and revalidation of its junior synonyms. Zoologischer Anzeiger 290: 117 - 134. https: // doi. org / 10.1016 / j. jcz. 2020.12.002","Lima J. D. 2018. Relatorio final do programa de monitoramento de herpetofauna da area de influencia da UHE Cachoeira Caldeirao.","Costa-Campos C. E. & Freire E. M. X. 2019. Richness and composition of anuran assemblages from an Amazonian savanna. ZooKeys 2019 (843): 149 - 169. https: // doi. org / 10.3897 / zookeys. 843.33365"]}

Family Aromobatidae Grant, Frost, Caldwell, Gagliardo, Haddad, Kok, Mittel, Noonan, Schargel & Wheeler, 2006 Anomaloglossus baeobatrachus (Boistel & Massary, 1999) is found throughout French Guiana and the state of Amapá, and may be a species complex (Fouquet et al. 2019 a, 2019b). The specimens from Amapá are recovered in a different clade in comparison with the clade formed by specimens from the type locality (Saint Eugène, French Guiana). Moreover, what is called An. baeobatrachus in the literature has two markedly different phenotypes, one with endotrophic (as observed from topotypes) and the other one with exotrophic tadpoles, morphologically indistinguishable from each other (Fouquet et al. 2019 a, 2019b). The populations with exotrophic tadpoles have signs of past hybridization and more investigation is necessary to clarify their taxonomic status (Fouquet et al. 2019a). Although both phenotypes occur in Amapá (Fouquet et al. 2019a), we chose to consider all the records under the name An. baeobatrachus until the situation is clarified. Two problematic species appeared among the records, namely Anomaloglossus beebei (Noble, 1923) and Allobates marchesianus (Melin, 1941). Anomaloglossus beebei is an endangered species with a restricted distribution, known only from the Pakaraima region, western Guyana (Cole et al. 2013; IUCN 2022). There are records of An. beebei from seven localities in central and northeastern Amapá (localities 5, 7–8, 11–12, 14–15; Lima 2006a, 2006b, 2008; Queiroz et al. 2011). Allobates marchesianus is probably a species complex distributed in Colombia, Peru, Venezuela, and the Brazilian state of Amazonas (Caldwell et al. 2002). There are also records of Al. marchesianus from central and northeastern Amapá, from five localities (6–8, 14–15; Lima 2006a, 2006b, 2008). According to recent thorough molecular surveys (Fouquet et al. 2019b; Rejaud et al. 2020; Vacher et al. 2020) none of these species occur in the state of Amapá. Besides, we did not have access to any of the vouchers and, due to the complex taxonomic history of the two species (see Caldwell et al. 2002; Kok et al. 2006), we cannot assign these records undoubtedly to a single species and we chose to remove the records from the list. Nonetheless, it is important to note that An. beebei has long been mistaken for Allobates granti Kok, MacCulloch, Gaucher, Poelman, Bourne, Lathrop & Lenglet, 2006 (Kok et al. 2006) and, although there is no record of Al. granti in Amapá, the species is known to inhabit several localities throughout the western border of French Guiana. Thus, at least some of the records of An. beebei may actually correspond to Al. granti., Published as part of Taucce, Pedro P. G., Costa-Campos, Carlos Eduardo, Carvalho, Thiago R. & Michalski, Fernanda, 2022, Anurans (Amphibia: Anura) of the Brazilian state of Amapá, eastern Amazonia: species diversity and knowledge gaps, pp. 96-130 in European Journal of Taxonomy 836 on pages 100-101, DOI: 10.5852/ejt.2022.836.1919, http://zenodo.org/record/7061668, {"references":["Fouquet A., Ferrier B., Salmona J., Tirera S., Vacher J. - P., Courtois E. A., Gaucher P., Lima J. D., Nunes P. M. S., De Souza S. M., Rodrigues M. T., Noonan B. & De Thoisy B. 2019 a. Phenotypic and life-history diversification in Amazonian frogs despite past introgressions. Molecular Phylogenetics and Evolution 130: 169 - 180. https: // doi. org / 10.1016 / j. ympev. 2018.09.010","Fouquet A., Vacher J. - P., Courtois E. A., Deschamps C., Ouboter P., Jairam R., Gaucher P., Dubois A. & Kok P. J. R. 2019 b. A new species of Anomaloglossus (Anura: Aromobatidae) of the stepheni group with the redescription of A. baeobatrachus (Boistel and de Massary, 1999), and an amended definition of A. leopardus Ouboter and Jairam, 2012. Zootaxa 4576: 439 - 460. https: // doi. org / 10.11646 / zootaxa. 4576.3.2","Cole C. J., Townsend C. R., Reynolds R. P., MacCulloch R. D. & Lathrop A. 2013. Amphibians and reptiles of Guyana, South America: illustrated keys, annotated species accounts, and a biogeographic synopsis. Proceedings of the Biological Society of Washington 125: 317 - 578.","IUCN 2022. The IUCN red list of threatened species. Available from https: // www. iucnredlist. org [accessed 1 Sep. 2022].","Lima J. D. 2006 a. Projeto de Pesquisa para Inventarios da Fauna e Plantas Superiores da Reserva de Desenvolvimento Sustentavel - Rio Iratapuru, Amapa. Macapa, Amapa.","Lima J. D. 2006 b. Inventarios Biologicos da Herpetofauna na Floresta Nacional do Amapa - Expedicoes I e II. In: Bernard E. (ed.) Inventarios Biologicos na Floresta Nacional do Amapa - Expedicoes I e II: 57 - 77. Macapa, Amapa.","Lima J. D. 2008. A herpetofauna do Parque Nacional do Montanhas do Tumucumaque, Amapa, Brasil, Expedicoes I a V. In: Bernhard E. (ed.) Inventarios Biologicos Rapidos no Parque Nacional Montanhas do Tumucumaque, Amapa, Brasil.: 38 - 50.","Queiroz S. S., Silva A. R., Reis F. M., Lima J. D. & Lima J. R. F. 2011. Anfibios de uma area de castanhal da Reserva Extrativista do Rio Cajari, Amapa. Biota Amazonia 1 (1): 1 - 18. https: // doi. org / 10.18561 / 2179 - 5746 / biotaamazonia. v 1 n 1 p 1 - 18","Caldwell J. P., Lima A. P. & Keller C. 2002. Redescription of Colostethus marchesianus from its type locality. Copeia 2002: 157 - 165.","Vacher J. P., Chave J., Ficetola F. G., Sommeria-Klein G., Tao S., Thebaud C., Blanc M., Camacho A., Cassimiro J., Colston T. J., Dewynter M., Ernst R., Gaucher P., Gomes J. O., Jairam R., Kok P. J. R., Lima J. D., Martinez Q., Marty C., Noonan B. P., Nunes P. M. S., Ouboter P., Recoder R., Rodrigues M. T., Snyder A., Marques-Souza S. & Fouquet A. 2020. Large-scale DNA-based survey of frogs in Amazonia suggests a vast underestimation of species richness and endemism. Journal of Biogeography 47 (8): 1781 - 1791. https: // doi. org / 10.1111 / jbi. 13847","Kok P. J. R., Macculloch R. D., Gaucher P., Poelman E. H., Bourne G. R., Lathrop A. & Lenglet G. L. (2006). A new species of Colostethus (Anura, Dendrobatidae) from French Guiana with a redescription of Colostethus beebei (Noble, 1923) from its type locality. Phyllomedusa: Journal of Herpetology 5: 43 - 66. https: // doi. org / 10.11606 / issn. 2316 - 9079. v 5 i 1 p 43 - 66"]}

Family Craugastoridae Hedges, Duellman & Heinicke, 2008 Pristimantis marmoratus (Boulenger, 1900) is currently distributed in the western portion of the Guiana Shield, through Venezuela and Guyana, and its sister clade has a more eastern distribution, occurring in French Guiana and Amapá (Kok et al. 2018). Fouquet et al. (2022b) recently described this clade as Pristimantis crepitaculus Fouquet, Peloso, Jairam, Lima, Mônico, Ernst & Kok, 2022. We considered four out of our five records of P. marmoratus (Lima 2008; 2018; Queiroz et al. 2011; Benício & Lima 2017) as P. crepitaculus. Pristimantis ockendeni (Boulenger 1912) was once thought to occur all over the upper Amazon basin, from southern Peru to Colombia, but it is currently thought to be a species complex with the nominal P. ockendeni currently known only from the type locality, in Peru (Elmer et al. 2007; Elmer & Canatella 2008). Silva e Silva & Costa-Campos (2018) recorded a species with overall morphology similar to P. ockendeni which they identified as P. cf. ockendeni , but they also recorded P. marmoratus. Besides P. crepitaculus, Fouquet et al. (2022b) also mention P. grandoculis from the state of Amapá, a species they revalidated in their study. The main morphological difference between these two species is the tympanum, which is present in the first species and absent in the latter. Thus, we consider the records of Silva e Silva & Costa-Campos (2018) as P. crepitaculus (their P. cf. ockendeni) and P. grandoculis (their P. marmoratus).

Family Bufonidae Gray, 1825 A recent study revealed high levels of species diversity within the bufonid genus Amazophrynella (Rojas et al. 2018), and described a new species for Suriname, French Guiana, and Amapá: Amazophrynella teko Rojas, Fouquet, Ron, Hernández-Ruz, Melo-Sampaio, Chaparro, Vogt, Carvalho, Pinheiro, Ávila, Farias, Gordo & Hrbek, 2018. According to their study, it is the only species distributed in Amapá. Vacher et al. (2020) also found only one species of Amazophrynella in their molecular survey, but they identified it as A. manaos Rojas-Zamora, Carvalho, Gordo, Ávila, Farias & Hrbek, 2014 or A. aff. manaos. We assume that the entries A. manaos and A. aff. manaos in their supplemental material is likely an inconsistent nomenclatural update throughout their species list. Both applied names (A. manaos and A. aff. manaos) of Vacher et al. (2020) are undoubtedly conspecific with A. teko. In this context, we treated the records of A. minuta and Amazophrynella sp. 1 from the Amapá National Forest as conspecific with A. teko as well. For more than30 years, Atelopus hoogmoedi was considered a subspecies of Atelopus pulcher (Boulenger, 1882) (Lescure 1974, 1976) or Atelopus spumarius Cope, 1871 (Lescure et al. 1980; Lescure & Marty 2000; Lötters et al. 2002). However, Lötters & Schulte (2005) elevated the taxon to full species level and Noonan & Gaucher (2005) provided molecular support for their decision. The populations from Brazil distributed north of the Amazon River, including Amapá, are currently assigned to A. hoogmoedi (Costa-Campos & Carvalho 2018; Jorge et al. 2020; Silva et al. 2020). Thus, we considered the records of Atelopus barbotini Lescure, 1981 from Lima (2006a, 2006b) and A. spumarius from Queiroz et al. (2011) and Lima (2018) as A. hoogmoedi. According to Vacher et al. (2020), Atelopus flavescens Duméril & Bibron, 1841 is also expected to occur in Amapá. However, since the taxonomic status of the populations from Amapá remains partly ambiguous, we adopted only one of the names in our list (A. hoogmoedi). Although it is probable that A. flavescens occurs indeed in the state of Amapá, further taxonomic studies should clarify this matter. Regarding the genus Rhinella, there are two species of the group of R. granulosa registered in Amapá: Rhinella granulosa (Spix, 1824) (Pereira-Júnior et al. 2013; Ferreira-Lima et al. 2017; Missassi et al. 2017; Lima 2018) and R. major (Costa-Campos & Freire 2019). However, Amapá is contained only within the distribution range of R. major, thus we considered records of R. granulosa to be R. major. Within the group of R. margaritifera, Rhinella martyi Fouquet, Gaucher, Blanc & Vélez- Rodriguez, 2007 is currently considered a junior synonym of Rhinella margaritifera (Laurenti, 1768) (Pereyra et al. 2021), and we considered the records of R. martyi (Silva-e-Silva & Costa-Campos 2018; Pedroso-Santos et al. 2019) as R. margaritifera. We then considered the records of R. margaritifera as Rhinella dapsilis (Myers & Carvalho, 1945), following the same study. Pereyra et al. (2021) recovered Rhinella castaneotica (Caldwell, 1991) paraphyletic, with the specimen from French Guiana more related to Rhinella proboscidea Spix, 1824 than to the probable R. castaneotica sensu stricto from the Brazilian state of Pará. Ferrão et al. (2022) analyzed specimens from both French Guiana and Amapá, recovering them monophyletic and as the sister group of R. castaneotica and Rhinella teotoniensis Ferrão, Souza, Hanken & Lima, 2022. We agree with their conclusions and consider the species from Amapá an unnamed species, which we treat as R. aff. castaneotica., Published as part of Taucce, Pedro P. G., Costa-Campos, Carlos Eduardo, Carvalho, Thiago R. & Michalski, Fernanda, 2022, Anurans (Amphibia: Anura) of the Brazilian state of Amapá, eastern Amazonia: species diversity and knowledge gaps, pp. 96-130 in European Journal of Taxonomy 836 on pages 101-105, DOI: 10.5852/ejt.2022.836.1919, http://zenodo.org/record/7061668, {"references":["Rojas R. R., Fouquet A., Ron S. R., Hernandez-Ruz E. J., Melo-Sampaio P. R., Chaparro J. C., Vogt R. C., Carvalho V. T. de, Pinheiro L. C., Avila R. W., Farias I. P., Gordo M. & Hrbek T. 2018. A Pan-Amazonian species delimitation: High species diversity within the genus Amazophrynella (Anura: Bufonidae). PeerJ 2018 (7). https: // doi. org / 10.7717 / peerj. 4941","Vacher J. P., Chave J., Ficetola F. G., Sommeria-Klein G., Tao S., Thebaud C., Blanc M., Camacho A., Cassimiro J., Colston T. J., Dewynter M., Ernst R., Gaucher P., Gomes J. O., Jairam R., Kok P. J. R., Lima J. D., Martinez Q., Marty C., Noonan B. P., Nunes P. M. S., Ouboter P., Recoder R., Rodrigues M. T., Snyder A., Marques-Souza S. & Fouquet A. 2020. Large-scale DNA-based survey of frogs in Amazonia suggests a vast underestimation of species richness and endemism. Journal of Biogeography 47 (8): 1781 - 1791. https: // doi. org / 10.1111 / jbi. 13847","Lescure J. 1974. Presence d'une sous-espece d' Atelopus pulcher (Amphibien, Anoure) dans les Guyanes: Atelopus pulcher hoogmoedi. Bulletin du Museum national d'histoire naturelle, Paris. Serie 3, Zoologie 144: 997 - 1005.","Lescure J. 1976. Contribution a l'etude des amphibiens de Guyane Francaise. VI. Liste preliminaire des anoures. Bulletin du Museum national d'histoire naturelle, Paris. Serie 3, Zoologie 377: 475 - 524.","Lescure J., Grenand F. & Grenand P. 1980. Les Amphibiens dans l'univers wayapi. Journal d'Agriculture traditionnelle et de Botanique appliquee 27 (3): 247 - 261. https: // doi. org / 10.3406 / jatba. 1980.3826","Lescure J. & Marty C. 2000. Atlas des amphibiens de Guyane. Museum national d'histoire naturelle, Paris.","Lotters S., Haas W., Schick S. & Bohme W. 2002. On the systematics of the harlequin frogs (Amphibia: Bufonidae: Atelopus) from Amazonia. II: Redescription of Atelopus pulcher (Boulenger, 1882) from the eastern Andean versant in Peru. Salamandra 38 (3): 165 - 184.","Lotters S. & Schulte R. 2005. Atelopus pulcher. In: Rueda-Almonacid J. V., Rodriguez-Machecha J. V., Lotters S., la Marca E., Kahn T. R. & Angulo A. (eds) Ranas Arlequines: 104. Conservacion Internacional, Bogota.","Noonan B. P. & Gaucher P. 2005. Phylogeography and demography of Guianan harlequin toads (Atelopus): Diversification within a refuge. Molecular Ecology 14 (10): 3017 - 3031. https: // doi. org / 10.1111 / j. 1365 - 294 X. 2005.02624. x","Costa-Campos C. E. & Carvalho T. R. 2018. The advertisement call of the Hoogmoed's harlequin toad Atelopus hoogmoedi Lescure, 1974 from northern Brazil (Anura, Bufonidae). Zootaxa 4521 (1): 141 - 144. https: // doi. org / 10.11646 / zootaxa. 4521.1.11","Jorge R. F., Ferrao M. & Lima A. P. 2020. Out of bound: A new threatened Harlequin toad (Bufonidae, Atelopus) from the outer borders of the Guiana shield in Central Amazonia described through integrative taxonomy. Diversity 12 (8). https: // doi. org / 10.3390 / D 12080310","Silva G. W. B., Cornelio G. S., de Oliveira E. A., Trindade N. G. P., Franca I. & Ruz E. J. H. 2020. A candidate species currently classified as Atelopus hoogmoedi (Anura: Bufonidae) in the Eastern Amazon, Para, Brazil. Genetics and Molecular Research 19 (1): 1 - 10. https: // doi. org / 10.4238 / GMR 18392","Lima J. D. 2006 a. Projeto de Pesquisa para Inventarios da Fauna e Plantas Superiores da Reserva de Desenvolvimento Sustentavel - Rio Iratapuru, Amapa. Macapa, Amapa.","Lima J. D. 2006 b. Inventarios Biologicos da Herpetofauna na Floresta Nacional do Amapa - Expedicoes I e II. In: Bernard E. (ed.) Inventarios Biologicos na Floresta Nacional do Amapa - Expedicoes I e II: 57 - 77. Macapa, Amapa.","Queiroz S. S., Silva A. R., Reis F. M., Lima J. D. & Lima J. R. F. 2011. Anfibios de uma area de castanhal da Reserva Extrativista do Rio Cajari, Amapa. Biota Amazonia 1 (1): 1 - 18. https: // doi. org / 10.18561 / 2179 - 5746 / biotaamazonia. v 1 n 1 p 1 - 18","Lima J. D. 2018. Relatorio final do programa de monitoramento de herpetofauna da area de influencia da UHE Cachoeira Caldeirao.","Bokermann W. C. A. 1967. Nova especie de Hyla do Amapa. Revista Brasileira de Biologia 27: 109 - 111.","Fouquet A., Martinez Q., Zeidler L., Courtois E. A., Gaucher P., Blanc M., Lima J. D., Souza S. M., Rodrigues M. T. & Kok P. J. R. 2016. Cryptic diversity in the Hypsiboas semilineatus species group (Amphibia, Anura) with the description of a new species from the eastern Guiana Shield. Zootaxa 4084 (1): 79 - 104. https: // doi. org / 10.11646 / zootaxa. 4084.1.3","Heyer W. R. & de Sa R. O. 2011. Variation, Systematics, and Relationships of the Leptodactylus bolivianus Complex (Amphibia: Anura: Leptodactylidae). Smithsonian Contributions to Zoology 635: 1 - 58. https: // doi. org / 10.5479 / si. 00810282.635.1","Ferreira-Lima J. R., Lima J. D., Lima S. D., Silva R. B. L. & Andrade G. V. 2017. Amphibians found in the Amazonian Savanna of the Rio Curiau Environmental. Biota Neotropica, Inventory 17 (2): e 20160252. https: // doi. org / 10.1590 / 1676 - 0611 - BN- 2016 - 0252","Missassi A. F. R., Costa-Campos C. E., Lima J. D. & Lambertini C. 2017. Range extension of the poorly know tree frog Dendropsophus haraldschultzi (Bokermann, 1962) (Amphibia, Anura, Hylidae): first records for two Brazilian Amazon states. Herpetology Notes 10: 703 - 707.","Costa-Campos C. E. & Freire E. M. X. 2019. Richness and composition of anuran assemblages from an Amazonian savanna. ZooKeys 2019 (843): 149 - 169. https: // doi. org / 10.3897 / zookeys. 843.33365","Pereyra M. O., Blotto B. L., Baldo D., Chaparro J. C., Ron S. R., Elias-Costa A. J., Iglesias P. P., Venegas P. J., Maria M. T., Ospina-Sarria J. J., Maciel N. M., Rada M., Kolenc F., Borteiro C., Rivera-Correa M., Rojas-Runjaic F. J. M., Moravec J., de La Riva I., Wheeler W. C., Castroviejo-Fisher S., Grant T., Haddad C. F. B. & Faivovich J. 2021. Evolution in the Genus Rhinella: A total evidence phylogenetic analysis of Neotropical true toads (Anura: Bufonidae). Bulletin of the American Museum of Natural History 447 (1): 1 - 156. https: // doi. org / 10.1206 / 0003 - 0090.447.1.1","Pedroso-Santos F., Sanches P. R. & Costa-Campos C. E. 2019. Anurans and reptiles of the Reserva Extrativista Beija-Flor Brilho de Fogo, Amapa State, Eastern Amazon. Herpetology Notes 12 (July): 799 - 807.","Ferrao M., de Souza R. A., Colatreli O. P., Hanken J. & Lima A. P. 2022. Hidden in the litter: cryptic diversity of the leaf-litter toad Rhinella castaneotica - proboscidea complex revealed through integrative taxonomy, with description of a new species from south-western Amazonia. Systematics and Biodiversity 20: 1 - 24. https: // doi. org / 10.1080 / 14772000.2022.2039317"]}

Family Eleutherodactylidae Lutz, 1954 Silva e Silva & Costa-Campos (2018) and Pedroso-Santos et al. (2019) recorded Adelophryne gutturosa Hoogmoed & Lescure, 1984 at Cancão Municipal Natural Park, municipality of Serra do Navio, and Reserva Extrativista Beija-Flor-Brilho-de-Fogo, municipality of Pedra Branca do Amapari, respectively. These records belong to the recently described A. amapaensis., Published as part of Taucce, Pedro P. G., Costa-Campos, Carlos Eduardo, Carvalho, Thiago R. & Michalski, Fernanda, 2022, Anurans (Amphibia: Anura) of the Brazilian state of Amapá, eastern Amazonia: species diversity and knowledge gaps, pp. 96-130 in European Journal of Taxonomy 836 on page 112, DOI: 10.5852/ejt.2022.836.1919, http://zenodo.org/record/7061668, {"references":["Silva e Silva Y. B. & Costa-Campos C. E. 2018. Anuran species composition of Cancao municipal natural park, municipality of Serra do Navio, Amapa state, Brazil. ZooKeys 2018 (762): 131 - 148. https: // doi. org / 10.3897 / zookeys. 762.22634","Pedroso-Santos F., Sanches P. R. & Costa-Campos C. E. 2019. Anurans and reptiles of the Reserva Extrativista Beija-Flor Brilho de Fogo, Amapa State, Eastern Amazon. Herpetology Notes 12 (July): 799 - 807."]}

Family Dendrobatidae Cope, 1865 Ranitomeya populations in Amapá have been registered with two different names, Ranitomeya amazonica (Schulte, 1999) (Lima 2018) and Ranitomeya ventrimaculata (Shreve, 1935) (Lima 2006a, 2006b; Queiroz et al. 2011). However, R. amazonica is a species known for decades prior to its description, and has been erroneously identified as several other species, including R. ventrimaculata (for a complete taxonomic history see Brown et al. 2011). Currently, R. ventrimaculata is restricted to Colombia, Ecuador, Peru, and the western portion of the Brazilian state of Amazonas, and until recently the populations of Ranitomeya from eastern Amazonia were assigned to R. amazonica (Brown et al. 2011). Nevertheless, Muell et al. (2022) recovered R. amazonica nested within R. variabilis (Zimmermann & Zimmermann, 1988) and specifically populations from eastern Amazonia were recovered as R. variabilis. Thus, we agree with them and consider Amapá records to be R. variabilis., Published as part of Taucce, Pedro P. G., Costa-Campos, Carlos Eduardo, Carvalho, Thiago R. & Michalski, Fernanda, 2022, Anurans (Amphibia: Anura) of the Brazilian state of Amapá, eastern Amazonia: species diversity and knowledge gaps, pp. 96-130 in European Journal of Taxonomy 836 on page 108, DOI: 10.5852/ejt.2022.836.1919, http://zenodo.org/record/7061668, {"references":["Lima J. D. 2018. Relatorio final do programa de monitoramento de herpetofauna da area de influencia da UHE Cachoeira Caldeirao.","Shreve B. 1935. On a new teiid and Amphibia from Panama, Ecuador, and Paraguay. Occasional papers of the Boston Society of Natural History 8: 209 - 218.","Lima J. D. 2006 a. Projeto de Pesquisa para Inventarios da Fauna e Plantas Superiores da Reserva de Desenvolvimento Sustentavel - Rio Iratapuru, Amapa. Macapa, Amapa.","Lima J. D. 2006 b. Inventarios Biologicos da Herpetofauna na Floresta Nacional do Amapa - Expedicoes I e II. In: Bernard E. (ed.) Inventarios Biologicos na Floresta Nacional do Amapa - Expedicoes I e II: 57 - 77. Macapa, Amapa.","Queiroz S. S., Silva A. R., Reis F. M., Lima J. D. & Lima J. R. F. 2011. Anfibios de uma area de castanhal da Reserva Extrativista do Rio Cajari, Amapa. Biota Amazonia 1 (1): 1 - 18. https: // doi. org / 10.18561 / 2179 - 5746 / biotaamazonia. v 1 n 1 p 1 - 18","Brown J. L., Twomey E., Amezquita A., Souza M. B., Caldwell J. P., Lotters S., von May R., MeloSampaio P. R., Mejia-Vargas D., Perez-Pena P., Pepper M., Poelman E. H., Sanchez-Rodriguez M. & Summers K. 2011. A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae). Zootaxa 3083 (1): 1 - 120. https: // doi. org / 10.11646 / zootaxa. 3083.1.1","Muell M. R., Chavez G., Prates I., Guillory W. X., Kahn T. R., Twomey E. M., Rodrigues M. T. & Brown J. L. 2022. Phylogenomic analysis of evolutionary relationships in Ranitomeya poison frogs (Family Dendrobatidae) using ultraconserved elements. Molecular Phylogenetics and Evolution 168: 107389. https: // doi. org / 10.1016 / j. ympev. 2022.107389"]}

Family Hylidae Rafinesque, 1815 The population previously identified as Boana multifasciata (Günther, 1859) from the Guiana Shield is currently known to be an unnamed candidate species, inhabiting the Guianas and the Brazilian states of Roraima, Pará and Amapá (Fouquet et al. 2021c) based only on molecular data. We added the molecularbased records from Vacher et al. (2020) and Fouquet et al. (2021c) corresponding to this taxon, as well as the literature records with overall morphology more similar to B. multifasciata. We decided to keep using this name for Amapá populations until more studies clarify their taxonomic status. In the species group of B. semilineata, two species are known to occur in Amapá, Boana diabolica Fouquet, Martinez, Zeidler, Courtois, Gaucher, Blanc, Lima, Souza, Rodrigues & Kok, 2016 and an unnamed species, B. aff. semilineata (Fouquet et al. 2016). Besides these two names, records in Amapá have also been identified as Boana geographica (Spix, 1824) (Lima 2008; Pereira-Júnior et al. 2013; Benício & Lima 2017; Ferreira-Lima et al. 2017) and B. aff. geographica (Lima 2006b). These records could belong either to B. diabolica or B. aff. semilineata, so we chose to consider in our list only the molecular confirmed records (Fouquet et al. 2016; Vacher et al. 2020). Dendropsophus counani Fouquet, Orrico, Ernst, Blanc, Martinez, Vacher, Rodrigues, Ouboter, Jairam & Ron, 2015 has long been confused with Dendropsophus brevifrons (Duellman & Crump, 1974) (Fouquet et al. 2015), and we consider the record of D. brevifrons (Benício & Lima 2017) to be D. counani. Dendropsophus minusculus (Rivero, 1971) is morphologically similar to Dendropsophus branneri (Cochran, 1948), with which D. minusculus has been historically confused (Zina et al. 2014). Dendropsophus minusculus is distributed in northern Brazil and the Guiana Shield, mainly within Amazonia, with some sparse records in northeastern Brazil in the states of Maranhão, Piauí, Ceará, and an isolated record from Bahia, whereas D. branneri inhabits the Atlantic Forest of southeastern and northeastern Brazil (Zina et al. 2014; Frost 2022). Both species seem to have a contact zone in northeastern Brazil, but the state of Amapá is within the distribution range of D. minusculus. Thus, we consider the records of D. branneri (Lima 2006a, 2006b) to be D. minusculus. Orrico et al. (2021) has recently defined the populations with the overall morphology like D. minutus (Peters, 1872) in the Guiana Shield as D. amicorum, and we agreed with them regarding the records of Amapá. It is important to note that, even though the IUCN status of the species is Critically Endangered (CR, IUCN 2022), it was evaluated when it was known only from the type localiy, and the conservation status of D. amicorum is likely to change during next evaluations. Dendropsophus walfordi (Bokermann, 1962) and Dendropsophus nanus (Boulenger, 1889) are also very similar to each other, being even considered as representing a single species (Lutz 1973). Despite being currently considered as separate species (Langone & Basso 1987), these are morphologically and genetically closely related to each other, with D. walfordi being frequently recovered within D. nanus, leaving it paraphyletic (Fouquet et al. 2011; Medeiros et al. 2013; Orrico et al. 2021). Recently, Seger et al. (2021) evaluated the D. nanus - D. walfordi complex with a broad molecular study and defined that D. nanus is the lineage inhabiting Paraguay, northeastern Argentina, and southern Brazil, whereas D. walfordi is the single lineage inhabiting Amazonia. Thus, we decided to adopt the name D. walfordi to records from both species within the state of Amapá. Benício & Lima (2017) recorded Dendropsophus microcephalus (Cope, 1886) and Dendropsophus parviceps (Boulanger, 1882) in the Amapá National Forest. The first species is distributed from southern Mexico to northern South America (Frost 2022) but, although the specimens east of Venezuela have been called D. microcephalus due to morphological similarities, they probably correspond to other species, such as D. minusculus or even Dendropsophus gaucheri (Lescure & Marty, 2000) (V.G.D. Orrico, pers. comm.). Although Benício & Lima (2017) cited both D. minusculus and D. microcephalus, there are no voucher specimens cited in their study. Thus, we decided to remove the record of D. microcephalus from the list until more data from the Amapá National Forest is available to help clarifying the taxonomic status of this population. Moreover, we considered the record of D. microcephalus from Queiroz et al. (2011) as D. minusculus. Dendropsophus parviceps is currently known from Ecuador, Colombia, and the Brazilian state of Amazonas, with related populations from Brazilian states of Acre and Rondônia being recently described as Dendropsophus kamagarini Rivadeneira, Venegas & Ron, 2018 (Rivadeneira et al. 2018). Besides D. parviceps, Benício & Lima (2017) found another species from the group of D. parviceps (sensu Orrico et al. 2021) in the Amapá National Forest, D. brevifrons (that we consider D. counani, see above). As in the previous case concerning D. microcephalus and D. minusculus, we also decided to remove D. parviceps from our list and consider the record as D. counani until additional data is available. The record of Lysapsus laevis (Parker, 1935) from Araújo & Costa-Campos (2015) (as Pseudis laevis) is actually Lysapsus bolivianus (Gallardo, 1961) (Costa-Campos, pers. comm.). There are two species of the group of Osteocephalus buckleyi (sensu Jüngfer et al. 2013) cited for Amapá, O. buckleyi (Boulenger, 1882) (Lima 2008; Queiroz et al. 2011) and O. cabrerai (Cochran & Goin, 1970) (Lima 2006b, 2008; Pedroso-Santos et al. 2019). Dewynter et al. (2019) also found two species of the group of O. buckleyi in French Guiana, both reaching the border with Amapá: O. cabrerai and Osteocephalus helenae (Ruthven, 1919). Vacher et al. (2020) also adopted these two species names in their molecular survey for populations of Osteocephalus from Amapá and French Guiana. Even though the two species were recovered as a single taxonomic entity in their species delimitation analysis, they applied both names to the clade formed by two lineages of Osteocephalus that appear to be morphologically diagnosable to each other and coexist in the Eastern Guiana Shield (see Dewynter et al. 2016). Thus, we consider both O. cabrerai and O. helenae in our list. We did not have access to the voucher specimens of most studies and most of the known previous records do not provide photographs of species of Osteocephalus. As such, we could not be precise about the locations where each of the two species occurs in Amapá. One exception is at Reserva Extrativista Municipal Beija-Flor-Brilho-deFogo, central part of Amapá, from where Pedroso-Santos et al. provided a picture of what they identified as O. cabrerai (Pedroso-Santos et al. 2019: 801, fig. 1.11) and it corresponds to the same morphotype Dewynter et al. (2016) consider O. cabrerai. Scinax garbei (Miranda-Ribeiro, 1926) is a widespread species of the group of S. rostratus distributed throughout the middle and upper Amazon Basin in Venezuela, Ecuador, Colombia, Peru, Bolivia and Brazil (Faivovich et al. 2005; Frost 2022). Although there are records of S. garbei from Amapá (FerreiraLima et al. 2017; Lima 2018; Silva e Silva & Costa-Campos 2018), there is molecular evidence of only Scinax jolyi Lescure & Marty, 2000 inhabiting Amapá (Vacher et al. 2020). Due to the taxonomic complexity of S. garbei (Ron et al. 2018) and the overall morphological similarity between the two species, we consider more plausible that the records of S. garbei in Amapá are indeed S. jolyi. Scinax cruentomma (Duelmann, 1972) is mainly distributed in western Amazonia, but there are a few additional records in other Amazonian regions (Carvalho et al. 2015: fig. 3). Those authors conducted acoustic comparisons between populations from the type locality (Santa Cecilia, Ecuador), the upper Negro River (Amazonas, Brazil), and French Guiana (data derived from Lescure & Marty 2000) and found that the high variation in some of the analyzed call traits indicate that the French Guiana population could represent another, potentially unnamed, species. Therefore, we provisionally assign the Amapá population to S. cf. cruentomma until the taxonomic status of the populations of Scinax bearing a horizontal red streak on the iris from the Eastern Guiana Shield is addressed., Published as part of Taucce, Pedro P. G., Costa-Campos, Carlos Eduardo, Carvalho, Thiago R. & Michalski, Fernanda, 2022, Anurans (Amphibia: Anura) of the Brazilian state of Amapá, eastern Amazonia: species diversity and knowledge gaps, pp. 96-130 in European Journal of Taxonomy 836 on pages 112-114, DOI: 10.5852/ejt.2022.836.1919, http://zenodo.org/record/7061668, {"references":["Fouquet A., Marinho P., Rejaud A., Carvalho T. R., Caminer M. A., Jansen M., Rainha R. N., Rodrigues M. T., Werneck F. P., Lima A. P., Hrbek T., Giaretta A. A., Venegas P. J., Chavez G. & Ron S. 2021 c. Systematics and biogeography of the Boana albopunctata species group (Anura, Hylidae), with the description of two new species from Amazonia. Systematics and Biodiversity 19 (4): 375 - 399. https: // doi. org / 10.1080 / 14772000.2021.1873869","Vacher J. P., Chave J., Ficetola F. G., Sommeria-Klein G., Tao S., Thebaud C., Blanc M., Camacho A., Cassimiro J., Colston T. J., Dewynter M., Ernst R., Gaucher P., Gomes J. O., Jairam R., Kok P. J. R., Lima J. D., Martinez Q., Marty C., Noonan B. P., Nunes P. M. S., Ouboter P., Recoder R., Rodrigues M. T., Snyder A., Marques-Souza S. & Fouquet A. 2020. Large-scale DNA-based survey of frogs in Amazonia suggests a vast underestimation of species richness and endemism. Journal of Biogeography 47 (8): 1781 - 1791. https: // doi. org / 10.1111 / jbi. 13847","Fouquet A., Martinez Q., Zeidler L., Courtois E. A., Gaucher P., Blanc M., Lima J. D., Souza S. M., Rodrigues M. T. & Kok P. J. R. 2016. Cryptic diversity in the Hypsiboas semilineatus species group (Amphibia, Anura) with the description of a new species from the eastern Guiana Shield. Zootaxa 4084 (1): 79 - 104. https: // doi. org / 10.11646 / zootaxa. 4084.1.3","Lima J. D. 2008. A herpetofauna do Parque Nacional do Montanhas do Tumucumaque, Amapa, Brasil, Expedicoes I a V. In: Bernhard E. (ed.) Inventarios Biologicos Rapidos no Parque Nacional Montanhas do Tumucumaque, Amapa, Brasil.: 38 - 50.","Benicio R. A. & Lima J. D. 2017. Anurans of Amapa National Forest, Eastern Amazonia, Brazil. Herpetology Notes 10: 627 - 633.","Ferreira-Lima J. R., Lima J. D., Lima S. D., Silva R. B. L. & Andrade G. V. 2017. Amphibians found in the Amazonian Savanna of the Rio Curiau Environmental. Biota Neotropica, Inventory 17 (2): e 20160252. https: // doi. org / 10.1590 / 1676 - 0611 - BN- 2016 - 0252","Lima J. D. 2006 b. Inventarios Biologicos da Herpetofauna na Floresta Nacional do Amapa - Expedicoes I e II. In: Bernard E. (ed.) Inventarios Biologicos na Floresta Nacional do Amapa - Expedicoes I e II: 57 - 77. Macapa, Amapa.","Zina J., Silva G. R., Loebmann D. & Orrico V. G. D. 2014. The recognition of Dendropsophus minusculus (Rivero, 1971) (Hylidae, Dendropsophini) as a highly polymorphic, multi-domain distributed species. Brazilian Journal of Biology 74 (3): 146 - 153. https: // doi. org / 10.1590 / 1519 - 6984.22912","Frost D. R. 2022. Amphibian Species of the World: an Online Reference. Available from http: // research. amnh. org / herpetology / amphibia / index. html [accessed 11 Apr. 2022].","Lima J. D. 2006 a. Projeto de Pesquisa para Inventarios da Fauna e Plantas Superiores da Reserva de Desenvolvimento Sustentavel - Rio Iratapuru, Amapa. Macapa, Amapa.","Orrico V. G. D., Grant T., Faivovich J., Rivera-Correa M., Rada M. A., Lyra M. L., Cassini C. S., Valdujo P. H., Schargel W. E., Machado D. J., Wheeler W. C., Barrio-Amoros C., Loebmann D., Moravec J., Zina J., Sole M., Sturaro M. J., Peloso P. L. V., Suarez P. & Haddad C. F. B. 2021. The phylogeny of Dendropsophini (Anura: Hylidae: Hylinae). Cladistics 37 (1): 73 - 105. https: // doi. org / 10.1111 / cla. 12429","IUCN 2022. The IUCN red list of threatened species. Available from https: // www. iucnredlist. org [accessed 1 Sep. 2022].","Lutz B. 1973. Brazilian species of Hyla. University of Texas Press, Austin.","Langone J. A. & Basso N. G. 1987. Distribucion geografica y sinonimia de Hyla nana Boulenger, 1889 y de Hyla sanborni Schmidt, 1944 (Anura, Hylidae) y observaciones sobre formas afines. Comunicaciones Zoologicas del Museo de Historia Natural de Montevideo 11: 1 - 17.","Fouquet A., Noonan B. P., Michel Blanc & Dill Orrico V. G. 2011. Phylogenetic position of Dendropsophus gaucheri (Lescure & Marty, 2000) highlights the need for an in-depth investigation of the phylogenetic relationships of Dendropsophus (Anura: Hylidae). Zootaxa 3035: 59 - 67. https: // doi. org / 10.11646 / zootaxa. 3035.1.3","Medeiros L. R., Lourenco L. B., Rossa-Feres D. C., Lima A. P., Andrade G. V., Giaretta A. A., Egito G. T. B. T. & Recco-Pimentel S. M. 2013. Comparative cytogenetic analysis of some species of the Dendropsophus microcephalus group (Anura, Hylidae) in the light of phylogenetic inferences. BMC Genetics 14. https: // doi. org / 10.1186 / 1471 - 2156 - 14 - 59","Seger K. R., Teixeira B. F. V., Annibale F. S., Rossa-Feres D. D. C., Lima A. P., Andrade G. V., Giaretta A. A. & Lourenco L. B. 2021. Five independent lineages revealed by integrative taxonomy in the Dendropsophus nanus - Dendropsophus walfordi species complex. Diversity 13 (11): 522. https: // doi. org / 10.3390 / d 13110522","Lescure J. & Marty C. 2000. Atlas des amphibiens de Guyane. Museum national d'histoire naturelle, Paris.","Queiroz S. S., Silva A. R., Reis F. M., Lima J. D. & Lima J. R. F. 2011. Anfibios de uma area de castanhal da Reserva Extrativista do Rio Cajari, Amapa. Biota Amazonia 1 (1): 1 - 18. https: // doi. org / 10.18561 / 2179 - 5746 / biotaamazonia. v 1 n 1 p 1 - 18","Rivadeneira D., Venegas P. & Ron S. R. 2018. Species limits within the widespread Amazonian treefrog Dendropsophus parviceps with descriptions of two new species (Anura, Hylidae). ZooKeys 726: 25 - 77. https: // doi. org / 10.3897 / zookeys. 726.13864","Jungfer K. - H., Faivovich J., Padial J. M., Castroviejo-Fisher S., Lyra M. L., Berneck B. von M., Iglesias P. P., Kok P. J. R., MacCulloch R. D., Rodrigues M. T., Verdade V. K., Torres-Gastello C. P., Chaparro J. C., Valdujo P. H., Reichle S., Moravec J., Gvozdik V., Gagliardi-Urrutia L. A. G., Ernst R., De la Riva I., Means D. B., Lima A. P., Senaris J. C., Wheeler W. C., and Haddad C. F. B. 2013. Systematics of spinybacked treefrogs (Hylidae: Osteocephalus): an Amazonian puzzle. Zoologica Scripta 42: 351 - 380.","Pedroso-Santos F., Sanches P. R. & Costa-Campos C. E. 2019. Anurans and reptiles of the Reserva Extrativista Beija-Flor Brilho de Fogo, Amapa State, Eastern Amazon. Herpetology Notes 12 (July): 799 - 807.","Dewynter M. & Chaline O. 2016. Inventaire herpetologique de la Borne n ° 4. Les Cahiers scientifiques du Parc amazonien de Guyane 2 (1): 281 - 302.","Faivovich J., Haddad C. F. B., Garcia P. C. A., Frost D. R., Campbell J. A. & Wheeler W. C. 2005. Systematic review of the frog family Hylidae, with special reference to Hylinae: phylogenetic analysis and taxonomic revision. Bulletin of the American Museum of Natural History 294: 1 - 240.","Lima J. D. 2018. Relatorio final do programa de monitoramento de herpetofauna da area de influencia da UHE Cachoeira Caldeirao.","Silva e Silva Y. B. & Costa-Campos C. E. 2018. Anuran species composition of Cancao municipal natural park, municipality of Serra do Navio, Amapa state, Brazil. ZooKeys 2018 (762): 131 - 148. https: // doi. org / 10.3897 / zookeys. 762.22634","Ron S. R., Duellman W. E., Caminer M. A. & Pazmino D. 2018. Advertisement calls and DNA sequences reveal a new species of Scinax (Anura: Hylidae) on the Pacific lowlands of Ecuador. PLoS ONE 13: e 0203169. https: // doi. org / 10.1371 / journal. pone. 0203169","Carvalho T. R., Teixeira B. F. V., Duellman W. E. & Giaretta A. A. 2015. Scinax cruentommus (Anura: Hylidae) in the upper Rio Negro drainage, Amazonas state, Brazil, with the redescription of its advertisement call. Phyllomedusa 14 (2): 139 - 146. https: // doi. org / 10.11606 / issn. 2316 - 9079. v 14 i 2 p 139 - 146"]}

Family Centrolenidae Taylor, 1951 Vitreorana ritae (Lutz, 1952) is currently a senior synonym of V. oyampiensis (Lescure, 1975) (Cisneros- Heredita 2013). Despite some authors state that this synonymy remains dubious due to the lack of acoustic and molecular data from the type locality (headwaters of the Caiwama River, Amazonas, Colombia) (Fouquet et al. 2019c), we consider the records of V. oyampiensis (as Cochranella oyampiensis; Lima 2006b; 2008) equivalent to V. ritae., Published as part of Taucce, Pedro P. G., Costa-Campos, Carlos Eduardo, Carvalho, Thiago R. & Michalski, Fernanda, 2022, Anurans (Amphibia: Anura) of the Brazilian state of Amapá, eastern Amazonia: species diversity and knowledge gaps, pp. 96-130 in European Journal of Taxonomy 836 on page 108, DOI: 10.5852/ejt.2022.836.1919, http://zenodo.org/record/7061668, {"references":["Fouquet A., Vidal N. & Dewynter M. 2019 c. The amphibians of the Mitaraka massif, French Guiana. Zoosystema 41 (sp 1): 359 - 374. https: // doi. org / 10.5252 / zoosystema 2019 v 41 a 19","Lima J. D. 2006 b. Inventarios Biologicos da Herpetofauna na Floresta Nacional do Amapa - Expedicoes I e II. In: Bernard E. (ed.) Inventarios Biologicos na Floresta Nacional do Amapa - Expedicoes I e II: 57 - 77. Macapa, Amapa.","Lima J. D. 2008. A herpetofauna do Parque Nacional do Montanhas do Tumucumaque, Amapa, Brasil, Expedicoes I a V. In: Bernhard E. (ed.) Inventarios Biologicos Rapidos no Parque Nacional Montanhas do Tumucumaque, Amapa, Brasil.: 38 - 50."]}

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

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."]}

The outstanding species richness of Amazonia has fascinated biologists for centuries. However, the records of actual numbers and distribution of species forming its ecosystems are so incomplete that the understanding of the historical causes and regional determinants of this diversity remain speculative. Anuran clades have repeatedly been documented to harbour many unnamed species in this region, notably the Boana albopunctata species group. Considering the documented distribution and the ecology of the species of that group, we hypothesized that it diversified via successive trans-riverine dispersals during the late Miocene and Pliocene, after the formation of the modern Amazon watershed. To test this hypothesis, we gathered an extensive dataset of 16S rDNA sequences sampled throughout Amazonia and a mitogenomic dataset representative of the diversity of the clade to (1) re-evaluate species boundaries and distributions, and (2) infer the spatio-temporal history of diversification within Amazonia. We delimited 14 Operational Taxonomic Units (OTUs) in an Amazonian clade, i.e., 75% higher than currently recognized (14 OTUs for eight described species). Combining molecular data with morphological and acoustic data, two new species, Boana courtoisae sp. nov. from the eastern Guiana Shield and Boana eucharis sp. nov. from Southern Amazonia, are described herein. These species belong to a clade that diversified throughout Amazonia during the last 10 Ma, thus more recently than co-distributed small terrestrial anurans but concomitantly with other more vagile vertebrates. Our time-scaled phylogeny and biogeographic analyses suggest an initial east-west divergence and confirm reciprocal trans-riverine dispersals during the last 5 Ma. The geomorphological evolution of the region and species-specific dispersal ability largely explain these distinct spatio-temporal patterns across anurans. http://www.zoobank.org/zoobank.org:act:4F8ACA9F-F6F1-4605-BD6C-6D4650AACCBE http://www.zoobank.org/zoobank.org:act:51CC7B40-2D6B-4A9E-AF50-AB34D4CE1042

Leptodactylus kilombo sp. nov. (Figs. 2, 5d, 6f, 8) [http://zoobank.org/ urn:lsid:zoobank.org:act: E9291B2E-D63E-49AE-9CDD-122560774016] Leptodactylus sp. (aff. mystaceus): Silva et al. (2020) Holotype. CHUFPB 28204, an adult male from Bezerra stream (12.897899°S, 46.810419°W; 626 m above sea level [a.s.l.]), in Arraias, Tocantins State, northern Brazil, collected by L. A. da Silva on 7 November 2018. Paratypes. One adult male (CHUFPB 28205) and one adult female (CHUFPB 30409) collected with the holotype. Referred material. One genetic voucher (UFBA 7694) from Bom Jesus da Lapa, Bahia State (Fig. 1), originally assigned to L. mystacinus by Dória et al. (2018). Etymology. The Kimbundu word kilombo (quilombo in Portuguese), used as a noun in apposition, literally means war camp. Quilombos are hinterland settlements founded by people of African origin including the quilombolas (inhabitants of quilombos), maroons, and some other names and variations by which these communities recognize themselves. Most of the original inhabitants were runaway slaves. There are hundreds of recognized quilombos throughout Brazil and also several out of record by the federal government. Such settlements carry plenty of historical, cultural, and anthropological history back into the Brazilian Imperial Age (colonial Brazil), especially in the mid-17th century, related to the slavery in the country. The type locality of Leptodactylus kilombo is located nearby an ancient quilombola settlement named Chapada dos Negros in Arraias, in the south of Tocantins State, north central Brazil (Gualberto 2017). Diagnosis. Leptodactylus kilombo is characterized by the following combination of character states: (1) small to moderate size, male SVL = 37.6–41.3 mm, female SVL = 41.8 mm; (2) advertisement call composed of single, nonpulsed notes; (3) note duration varying from 30–56 ms; (4) note dominant frequency varying from 1098–1572 Hz; (5) frequency upsweep varying from 388–775 Hz. Comparisons with members of the L. mystaceus complex. Leptodactylus kilombo can be distinguished from other species by having the sole of the foot covered with few, non-obvious tubercles (prominent tubercles in the other species; Heyer 1978, 1983; Heyer et al. 1996), except Leptodactylus notoaktites. The new species (male SVL = 37.6–41.3 mm, mean = 40.8; female SVL = 41.8 mm; Table 2) can be further distinguished from the other species of the L. mystaceus complex, except L. elenae and L. spixi, by a smaller size: L. didymus (male SVL = 45.9–52.2 mm, mean = 49.3; female SVL = 45.8–53.5 mm, mean = 49.8; Heyer et al. 1996), L. mystaceus (male SVL = 42.4–52.2 mm, mean = 47.4; female SVL = 44.8–56.1 mm, mean = 50.0; Heyer et al. 1996), L. notoaktites (male SVL = 42.5–54.2 mm, mean = 47.5; female SVL = 43.4–55.8 mm, mean = 48.0; de Sá et al. 2014). Members of the L. mystaceus complex are mainly distinguished from each other by their distinct advertisement calls (Fig. 6). Acoustic comparisons in the L. fuscus group. The advertisement call of Leptodactylus kilombo is composed of single notes (Fig. 6f), differing from the trill calls of L. cunicularius, L. cupreus, L. oreomantis, and L. plaumanni (Fig. 7; Carvalho et al. 2013). The new species is distinguished from species with pulsed calls (L. caatingae, L. fragilis, L. labrosus, L. mystaceus, and L. cf. mystaceus; Heyer 1978; Heyer & Juncá 2003; Heyer et al. 2006; Carvalho & Ron 2011) by having a nonpulsed call. From the remaining species of the L. fuscus group also having single-note, nonpulsed calls, L. kilombo is distinguished by a shorter note duration (30–56 ms) from L. bufonius, L. camaquara, L. didymus, L. elenae, L. laticeps, and L. spixi (> 100 ms; Table 3; Heyer & Scott 2006; Carvalho et al. 2013). The call of L. kilombo has a modest frequency upsweep (frequency modulation, FM = 388–775 Hz), whereas L. albilabris, L. fuscus, L. gracilis, L. jolyi, L. longirostris, L. notoaktites, L. sertanejo, and L. syphax have a pronounced frequency upsweep in their calls (FM> 1000 Hz; Heyer 1978; Giaretta & Costa 2007; Heyer et al. 2010), while that of L. poecilochilus has a negligible/slight frequency upsweep (FM Leptodactylus kilombo is distinguished by a lower dominant frequency (1098–1572 Hz) from L. furnarius, L. latinasus, L. marambaiae, L. mystacinus, L. tapiti, and L. troglodytes (> 2000 Hz) (Heyer 1978; Sazima & Bokermann 1978; Brandão et al. 2013). Description of holotype. Body robust. Snout sub-elliptical in dorsal and ventral views (Fig. 8 a–b), acuminate in lateral view (Fig. 8e). Canthus rostralis rounded; loreal region flat; tympanic annulus well-defined, circular (TD = 60.4% ED); tympanic membrane translucent (brownish); supratympanic fold from the posterior corner of the eye, passing over the dorsal edge of the tympanic annulus and ending at the base of the arm; vocal sac subgular; vocal slits present; vomerine teeth in two nearly straight rows medial and posterior to choanae and almost parallel to sagittal plane. Tongue rounded, free at its posterior third. Relative finger lengths IV Colors. In life, dorsal surfaces (body and limbs) reddish; snout whitish. Light brown blotches scattered on the dorsum (Fig. 5d). Upper portion of the loreal region dark brown with green specks, lower portion reddish on a cream-colored background, lips nonpigmented with light-colored spots and stains. Flank light gray; tubercles reddish and light cream. Throat white scattered with dark brown spots and dots. Chest and belly off-white; ventral surface of limbs cream-colored. Upper surface of thigh, tibia and tarsus brown with dark brown transversal bars. Posterior surface of thigh with brown spots and irregular longitudinal stripe light cream. In preservative, dorsal surfaces (body and limbs) brown with darker brown blotches (Fig. 8). The dorsal reddish coloration in life becomes brown or dark brown. Dark brown blotches and brown transversal bars on the upper surface of thigh and tibia are more conspicuous. Interorbital blotch poorly delimited posteriorly. Upper portion of the loreal region blackish brown, lower portion light brown, lips dark brown. Tympanic membrane light brown. Flank light gray. Throat white covered with dark brown spots and dots. Chest, belly, and ventral surface of limbs varying from cream-colored to off-white. Dark brown transversal bars on upper surface of thigh, tibia and tarsus. Posterior surface of thigh with scattered brown spots and irregular longitudinal stripe stripe light cream. Ventral surface of hand and foot brown interspersed with nonpigmented areas; Sole of foot with sparse white dots. Intraspecific variation. Lateral line of tubercles is complete in CHUFPB 28205. Posterior surface of tarsus can be smooth or covered with tiny tubercles (poorly visible in preservative). Tubercles on the sole of foot are more conspicuous in the examined female (CHUFPB 30409) than in male specimens. Advertisement call. Description based on 94 calls recorded from two males (Table 3). The call (Fig. 6f) consists of single, nonpulsed notes given at a rate of 109–122 per minute. Note duration varies from 30–56 ms. Rise time is 37–77% of note duration. The dominant frequency coincides with the fundamental harmonic at 1098–1572 Hz. Notes usually have a modest frequency upsweep throughout their duration, varying from 388–775 Hz. plex. Data are presented as mean SD (range). applicable to species with nonpulsed calls). Abbreviations: ND = note duration, RT = rise time, NR = note repetition rate, PP = pulses per note, PR = pulse repetition rate, DF = dominant frequency, and FM = frequency modulation. Habitat and natural history. At the type locality, we first found the species in breeding activity early in the rainy season (October 2018), when many calling males could be heard along a dried-up stream (Bezerra stream), which was humid in the sandy stream bed. On this occasion, most males were calling from underneath dense layer of leaf litter (see a calling male at: https://www.youtube.com/watch?v=SUyRQOiGReQ). During a second expedition to the type locality later in the rainy season (April 2019), we did not find or hear any specimen of the new species, suggesting that its breeding activity may be restricted to the early rainy season. Despite being located in a central region of the Cerrado domain, the type locality is more characterized by the presence of Seasonally Dry Tropical Forests (Werneck 2011) than typical savanna formations. Leptodactylus kilombo and L. cf. mystaceus are syntopic at the type locality (Fig. 1). Despite being quite similar to each other in overall morphology, these species can be easily distinguished by their advertisement calls (see “Comparisons with congeners” section). Besides L. cf. mystaceus, two other species of the L. fuscus group occur syntopically at the type locality (L. fuscus and L. troglodytes). All four species were observed calling simultaneously at the type locality. Distribution. Leptodactylus kilombo (Fig. 1) is known from two localities in the extreme southeast of Tocantins, the type locality (Arraias) and Conceição do Tocantins, from a single locality in northeastern Goiás (Flores de Goiás; see Appendix II) and a single locality in western Bahia (Bom Jesus da Lapa; see Appendix II). Remarks. Live specimens of Leptodactylus kilombo have a reddish-pink upper lip stripe. This color feature might be an additional diagnostic feature of the species in the L. mystaceus complex. Because of our small sample size of L. kilombo, further sampling is needed to confirm whether upper lip stripe coloration is an informative character in the diagnosis of this species. We examined three specimens collected from Conceição do Tocantins (field# RAB 1965–67), located approximately 85 km northwest of the type locality of L. kilombo. These specimens resemble L. kilombo in overall morphology and by having a reddish-pink upper lip stripe, thus assigned to L. kilombo based on morphological and color features from the preserved specimens and pictures in life, but acoustic and molecular data should also be obtained from this population.

Leptodactylus watu sp. nov. (Figs. 2, 4a, 4c, 5h, 6h, 10) [http://zoobank.org/ urn:lsid:zoobank.org:act: 5D79A081-C50C-463A-8101-04351E87AD23] Leptodactylus aff. spixi: Guimar��es et al. (2019) Holotype. UFMG 21332, an adult male from Parque Estadual do Rio Doce (19.753922��S, 42.633047��W, approximately 280 m a.s.l.), in Marli��ria, Minas Gerais State, southeastern Brazil, collected by H. Thomassen on 21 November 2017. Paratypes. Seven adult males (UFMG 21327���21331 and 21337���21338) and five adult females (UFMG 21333��� 21336 and 21339) collected from the type locality by H. Thomassen from November 2017 to February 2018. Additional material. One genetic voucher (UFMG 16995) from Santa B��rbara do Leste, Minas Gerais State (Fig. 1). Etymology. The word watu, a noun in apposition, is derived from the dialect of the Bor��n indigenous people and is a reference to the Doce River (Costa-Reis & Genovez 2013). In 2015, the Doce River experienced the worst environmental disaster that ever took place in Brazil, the collapse of a mining tailings dam owned by Samarco (and co-owned by the Brazilian Vale and Australian BHP Billiton). The catastrophic dam failure released around 60 million cubic meters of iron ore tailings (toxic slurry) directly into the Doce River watershed, killing 20 people and affecting biodiversity across hundreds of kilometers around the river drainage, riparian lands, and the Atlantic coast, in addition to a severe contamination of the soil and water table within the affected region. The epithet is a tribute to the resistance of the Bor��n people and to the Watu (Doce River) in southeastern Brazil. Diagnosis. Leptodactylus watu is characterized by the following combination of character states: (1) small to moderate size, male SVL = 38.5���42.2 mm, female SVL = 40.5���47.9 mm; (2) sole of foot with few, non-obvious tubercles; (3) advertisement call composed of single, pulsed notes; (4) note duration varying from 27���70 ms; (5) note dominant frequency varying from 495���1098 Hz; (6) frequency upsweep varying from 0���560 Hz. Comparisons with members of the L. mystaceus complex. Leptodactylus watu is distinguished from other species belonging to the L. mystaceus complex, except L. barrioi, L. kilombo and L. notoaktites, by having the sole of the foot covered with few, non-obvious tubercles, whereas L. didymus, L. elenae, L. mystaceus, and L. spixi have prominent tubercles (Heyer 1978, 1983; Heyer et al. 1996). The new species can be further distinguished from some species by a smaller size (Table 2): L. didymus (male SVL = 45.9���52.2 mm, mean = 49.3; female SVL = 45.8���53.5 mm, mean = 49.8; Heyer et al. 1996), L. mystaceus (male SVL = 42.4���52.2 mm, mean = 47.4; female SVL = 44.8��� 56.1 mm, mean = 50.0; Heyer et al. 1996), and L. notoaktites (male SVL = 42.5���54.2 mm, mean = 47.5; female SVL = 43.4���55.8 mm, mean = 48.0; de S�� et al. 2014). Members of the L. mystaceus complex are mainly distinguished from each other by their distinct advertisement calls (Fig. 6). Acoustic comparisons in the L. fuscus group. The advertisement call of Leptodactylus watu is composed of single, pulsed notes (Table 3; Fig. 6h), differing from species with trill calls (L. cunicularius, L. cupreus, L. oreomantis, and L. plaumanni; Fig. 7; Carvalho et al. 2013) and species with nonpulsed calls (L. albilabris, L. bufonius, L. camaquara, L. didymus, L. elenae, L. furnarius, L. fuscus, L. gracilis, L. jolyi, L. kilombo sp. nov., L. laticeps, L. latinasus, L. longirostris, L. marambaiae, L. mystacinus, L. notoaktites, L. poecilochilus, L. sertanejo, L. spixi, L. syphax, L. tapiti, and L. troglodytes). From other species with pulsed calls, L. watu is distinguished by having its call formed by partly fused pulses, whereas the calls of L. barrioi, L. mystaceus and L. cf. mystaceus are formed by complete pulses (Fig. 6). From species also having calls with partly fused pulses (L. caatingae, L. fragilis, and L. labrosus), L. watu is distinguished by a shorter note (27���70 ms) from L. fragilis (> 100 ms; Heyer et al. 2006). The calls of L. watu, L. caatingae, and L. labrosus are the most similar to each other in the L. fuscus group, but these species can be distinguished, although value ranges partially overlap, by differences in the dominant frequency (L. watu: 495���1098 Hz, mean = 763; L. caatingae: 1148���1430 Hz, mean = 1315, in our sample; 943���1620 Hz, mean = 1423 Hz, in the original description; Heyer & Junc�� 2003) or duration (L. watu: 27���70 ms, mean = 40; L. labrosus: 64���133 ms, mean = 100; Carvalho & Ron 2011) of their calls. Still, these species differ markedly in morphology and size (see previous paragraph; Heyer 1978; Heyer & Junc�� 2003). Description of holotype. Body robust. Snout sub-elliptical in dorsal and ventral views (Fig. 10 a���b), acuminate in lateral view (Fig. 10e). Canthus rostralis rounded; loreal region flat; tympanic annulus well-defined, circular (TD = 62.4% ED); supratympanic fold from the posterior corner of the eye, passing over the dorsal edge of the tympanic annulus, and ending at the base of the arm; vocal sac subgular; vocal slits present; vomerine teeth in two nearly straight rows, almost connected, medial and posterior to choanae and almost parallel to sagittal plane. Tongue rounded, free at its posterior third. Relative finger lengths II = IV Colors. In life, dorsal surfaces (body and limbs) light brown with dark gray blotches poorly delimited (Figs. 4a, c, 5h). Upper region of the loreal region black, lower region light cream, lips dark brown. Black stripe on the supratympanic fold, delimited dorsally by a white line. Light cream stripe on the dorsolateral fold, more conspicuous posteriorly. Flank light gray. Tympanic membrane dark brown on the center and upper rim, light brown laterally and on ventral rim. Throat immaculate white with nonpigmented portions near the jaw. Ventral surface of arms, legs (except tarsus), and belly nonpigmented; tarsus with scattered dark brown stains. Weakly defined dark brown transversal bars on the upper surface of hindlimbs and forearm. Posterior surface of thigh with a yellow longitudinal stripe bordered by two black stains; upper portion of with black spots and stains. Ventral surface of hand, foot, and tarsus dark brown interspersed with nonpigmented areas. In preservative, dorsal surfaces (body and hindlimbs) light gray with dark gray blotches poorly delimited; forearms light cream with scattered dark gray dots and spots (Fig. 10). Upper portion of the loreal region black, lower half cream-colored, lips blackish gray. Black stripe on the supratympanic fold. White stripe on the dorsolateral fold, more conspicuous posteriorly. Flank light gray. Tympanic membrane light brown with the center and upper edge dark gray. Ventral surfaces cream-colored. Throat with few gray dots and stains near the jaw laterally. Ventral surface of hindlimbs with few dark gray spots. Dark brown transversal bars on the dorsal surface of hindlimbs and forearm. Posterior surface of thigh with a white longitudinal stripe bordered by two black stains. Ventral surface of hand, foot, and tarsus dark brown interspersed with nonpigmented areas. Intraspecific variation. In life, dorsal coloration varies from brown, sometimes conspicuous reddish shades, or dark gray. The dorsal light brown coloration in life becomes light gray in preservative. Dark brown blotches on the dorsum can be present, more conspicuous in preserved specimens. Dark brown transversal bars on the upper surface of thigh and tibia are more evident in preserved species. Lateral line of tubercles is poorly evident in most specimens. Inner metacarpal tubercle is rounded in UFMG 21329. Advertisement call. Description based on 129 calls recorded from two males (Table 3). The call (Fig. 6h) consists of single, pulsed notes given at a rate of 124���275 per minute. Note duration varies from 27���70 ms. Rise time varies from 37���80% of note duration. Notes are formed by 2���7 partly fused pulses emitted at a rate of 57���153 per second. The dominant frequency coincides with the fundamental harmonic at 495���1098 Hz. Notes have no frequency modulation or a modest frequency upsweep throughout their duration, varying from 0���560 Hz. Habitat and natural history. At the type locality, Leptodactylus watu was found calling during the rainy season in the region from late October to late February. Calling is strongly associated with rainy days or few days after heavy pours. Males were found calling around flooded areas or close to moist ground in both open areas (transitional marshlands between the forest and lakes) and inside the forest. All calling males were hidden either under dense vegetation, ground leaf litter or inside underground chambers. They were never found calling in pastures. The congeneric species found calling in syntopy with L. watu were L. fuscus, L. latrans, and L. labyrinthicus. Distribution. Leptodactylus watu is only known from two localities in a region characterized by lowland interior Atlantic forests associated with the upper-middle Doce River drainage, in the Brazilian state of Minas Gerais (Fig. 1): (1) the type locality, at Parque Estadual do Rio Doce, and (2) Santa B��rbara do Leste, approximately 70 km southeast from the type locality. Remarks. The identity of populations assigned to Leptodactylus aff. mystaceus (Santana et al. 2010), L. aff. spixi (Ferreira et al. 2019), and L. spixi (R��dder et al. 2007; Silva-Soares & Scherrer 2013; Pereira-Ribeiro et al. 2019) from different localities in southeastern and northeastern Brazil should be addressed using calls and/or DNA sequences, as these records could correspond to L. barrioi or L. watu, or additional candidate new species., Published as part of Da Silva, Leandro A., Magalh��es, Felipe M., Thomassen, Hans, Leite, Felipe S. F., Garda, Adrian A., Brand��o, Reuber A., Haddad, C��lio F. B., Giaretta, Ariovaldo A. & De Carvalho, Thiago R., 2020, Unraveling the species diversity and relationships in the Leptodactylus mystaceus complex (Anura: Leptodactylidae), with the description of three new Brazilian species, pp. 151-189 in Zootaxa 4779 (2) on pages 167-170, DOI: 10.11646/zootaxa.4779.2.1, http://zenodo.org/record/3833431, {"references":["Guimaraes, C. S., Assis, C. L., Thomassen, H., Leite, F. S. F. & Feio, R. N. (2019) Anfibios do Parque Estadual do Rio Doce, Minas Gerais. Universidade Federal de Vicosa, Departamento de Biologia Animal, Museu de Zoologia Joao Moojen, Vicosa, Minas Gerais, 57 pp.","Costa-Reis, R. & Genovez, P. F. (2013) Territorio sagrado: exilio, diaspora e reconquista Krenak no vale do Rio Doce, Resplendor, MG. Boletim Goiano de Geografia, 33, 1 - 15. https: // doi. org / 10.5216 / bgg. v 33 i 1.23628","Heyer, W. R. (1978) Systematics of the fuscus group of the frog genus Leptodactylus (Amphibia, Leptodactylidae). Science Bulletin, Natural History Museum of Los Angeles County, 29, 1 - 85.","Heyer, W. R. (1983) Clarification of the names Rana mystacea Spix, 1824, Leptodactylus amazonicus Heyer, 1978, and a description of a new species, Leptodactylus spixi (Amphibia: Leptodactylidae). Proceedings of the Biological Society of Washington, 96, 270 - 272.","Heyer, W. R., Garcia-Lopez, J. M. & Cardoso, A. J. (1996) Advertisement call variation in the Leptodactylus mystaceus species complex (Amphibia: Leptodactylidae) with a description of a new sibling species. Amphibia-Reptilia, 17, 7 - 31. https: // doi. org / 10.1163 / 156853896 X 00252","de Sa, R. O., Grant, T., Camargo, A., Heyer, W. R., Ponssa, M. L. & Stanley, E. (2014) Systematics of the Neotropical genus Leptodactylus Fitzinger, 1826 (Anura: Leptodactylidae): Phylogeny, the relevance of non-molecular evidence, and species accounts. South American Journal of Herpetology, 9, S 1 - S 128. https: // doi. org / 10.2994 / SAJH-D- 13 - 00022.1","Carvalho, T. R., Leite, F. S. F. & Pezzuti, T. L. (2013) A new species of Leptodactylus Fitzinger (Anura, Leptodactylidae, Leptodactylinae) from montane rock fields of the Chapada Diamantina, northeastern Brazil. Zootaxa, 3701 (3), 349 - 364. https: // doi. org / 10.11646 / zootaxa. 3701.3.5","Heyer, W. R., Heyer, M. M. & de Sa, R. O. (2006) Leptodactylus fragilis. Catalogue of American Amphibians and Reptiles, 830, 1 - 26.","Heyer, W. R. & Junca, F. A. (2003) Leptodactylus caatingae, a new species of frog from eastern Brazil (Amphibia: Anura: Leptodactylidae). Proceedings of the Biological Society of Washington, 116, 317 - 329.","Carvalho, T. R. & Ron, S. R. (2011) Advertisement call of Leptodactylus labrosus Jimenez de la Espada, 1875 (Anura, Leptodactylidae): an unusual advertisement call within the L. fuscus group. Herpetology Notes, 4, 325 - 326.","Santana, D. J., Sao Pedro, V. A., Hote, P. S., Roberti, H. M., Sant'Anna, A. C., Figueiredo-de-Andrade, C. A. & Feio, R. N. (2010) Anurans in the region of the High Muriae River, state of Minas Gerais, Brazil. Herpetology Notes, 3, 1 - 10.","Ferreira, R. B., Monico, A. T., Silva, E. T., Lirio, F. C. F., Zocca, C., Mageski, M. M., Tonini, J. F. R., Beard, K. H., Duca, C. & Silva- Soares, T. (2019) Amphibians of Santa Teresa, Brazil: the hotspot further evaluated. ZooKeys, 857, 139 - 162. https: // doi. org / 10.3897 / zookeys. 857.30302","Rodder, D., Teixeira, R. L., Ferreira, R. B., Dantas, R. B., Pertel, W. & Guarniere, G. J. (2007) Anuran hotspots: the municipality of Santa Teresa, Espirito Santo, southeastern Brazil. Salamandra, 43, 91 - 110.","Silva-Soares, T. & Scherrer, P. V. (2013) Amphibians of Parque Estadual do Forno Grande, State of Espirito Santo, Southeastern Brazil: Species composition and conservation. North-Western Journal of Zoology, 9, 113 - 120.","Pereira-Ribeiro, J., Ferreguetti, A. C., Linause, T. M., Cozer, J. S., Bergallo, H. G. & Rocha, C. F. D. (2019) Diversity and distribution of anurans from Mata das Flores State Park, Espirito Santo, southeastern Brazil. Oecologia Australis, 23, 292 - 300. https: // doi. org / 10.4257 / oeco. 2019.2302.08"]}

Leptodactylus barrioi sp. nov. (Figs. 2, 4e, 5a, 6g, 9) [http://zoobank.org/ urn:lsid:zoobank.org:act: 4948F138-66DC-41DD-8426-0DE12B0FED67] Holotype. AAG-UFU 502, an adult male from Duas Barras (22.062862°S, 2.520780°W, approximately 625 m a.s.l.), Rio de Janeiro State, southeastern Brazil, collected by T. R. de Carvalho and L. B. Martins on 10 September 2011. Paratypes. Three adult males (AAG-UFU 503, 506, and 694) and two adult females (AAG-UFU 504–505) collected at the type locality from 10 September to 7 December 2011. One adult male (AAG-UFU 1102) from Chiador (21.921698°S, 42.926676°W, nearly 495 m a.s.l.), Minas Gerais State, southeastern Brazil, collected by T. R. de Carvalho on 8 March 2012. One adult male (AAG-UFU 6183) from the District of Povoação (19.551215°S, 39.846788°W, nearly 20 m a.s.l.), in Linhares, Espírito Santo State, southeastern Brazil, collected by T. R. de Carvalho, B. F. V. Teixeira, and L. B. Martins on 6 December 2017. Additional material. Genetic vouchers from Ibitirama, Espírito Santo State (UFMG 17268), and Cataguases, Minas Gerais State (AAGARDA 3300–3301); see Fig. 1. Etymology. The epithet is a homage to Avelino Barrio, who was a pioneer in characterizing calls of Leptodactylus and Physalaemus species in two comparative studies (Barrio 1965a, b). This researcher was very attentive to the relevance of acoustic data for species discrimination in both leptodactylid genera. Diagnosis. Leptodactylus barrioi is characterized by the following combination of character states: (1) sole of foot with few, non-obvious tubercles; (2) advertisement call composed of single, pulsed notes; (3) note duration varying from 62–126 ms; (4) note dominant frequency varying from 586–1383 Hz; (5) frequency upsweep varying from 141–750 Hz. Comparisons with members of the L. mystaceus complex. Leptodactylus barrioi can be distinguished by having the sole of the foot covered with few, non-obvious tubercles, whereas the other species have prominent tubercles (Heyer 1978, 1983; Heyer et al. 1996), except L. kilombo and L. notoaktites. The new species cannot be distinguished from other members of the L. mystaceus complex by morphometric traits (Table 2), but differs from the other species in acoustic traits (Fig. 6). Acoustic comparisons in the L. fuscus group. The advertisement call of Leptodactylus barrioi is composed of single, pulsed notes (Table 3; Fig. 6g), differing from the trill calls of L. cunicularius, L. cupreus, L. oreomantis, and L. plaumanni (Fig. 7; Carvalho et al. 2013), and from the nonpulsed calls of L. albilabris, L. bufonius, L. camaquara, L. didymus, L. elenae, L. furnarius, L. fuscus, L. gracilis, L. jolyi, L. kilombo, L. laticeps, L. latinasus, L. longirostris, L. marambaiae, L. mystacinus, L. notoaktites, L. poecilochilus, L. sertanejo, L. spixi, L. syphax, L. tapiti, and L. troglodytes. From other species with pulsed calls, L. barrioi is distinguished by having its call formed by complete pulses (i.e., with silent gaps in between pulses), whereas those of L. caatingae, L. fragilis, and L. labrosus are formed by partly fused pulses (Heyer 1978; Heyer & Juncá 2003; Carvalho & Ron 2011). The only other congeners of the L. fuscus group with the call formed by complete pulses are L. mystaceus and L. cf. mystaceus, from which L. barrioi is distinguished by a shorter note duration (L. barrioi: 62–126 ms; L. mystaceus: 182–218 ms; L. cf. mystaceus: 136–260 ms; Table 3). Description of holotype. Body robust. Snout sub-elliptical in dorsal and ventral views (Fig. 9 a–b), acuminate in lateral view (Fig. 9e). Canthus rostralis rounded; loreal region flat; tympanic annulus well-defined, circular (TD = 60.8% ED); supratympanic fold from the posterior corner of the eye, passing over the dorsal edge of the tympanic annulus, and ending at the base of the arm; vocal sac subgular; vocal slits present; vomerine teeth in two nearly convex rows medial and posterior to choanae and almost parallel to sagittal plane. Tongue ovoid, free at its posterior third. Relative finger lengths IV Colors. In preservative (Fig. 9), dorsal surfaces of body light grayish brown with light brown blotches and black spots. Upper portion of the loreal region black, lower portion cream-colored, lips blackish brown. Black stripe on the supratympanic fold delimited dorsally by a white line. Flank light gray. Tympanic membrane dark brown in the middle and upper rim, light brown laterally and lower rim. Throat, ventral surface of forelimbs, hindlimbs, and belly cream-colored. Brown transversal bars on the upper surfaces of forearms and hindlimbs. Posterior surface of thigh with a white longitudinal stripe bordered by black stains; upper portion of the posterior surface of thigh with gray spots and irregular stripe. Ventral surface of hand, foot, and tarsus dark gray interspersed with nonpigmented areas. Intraspecific variation. In life, dorsal coloration varies from light gray to reddish, dark brown. Reddish brown shades become light gray or light brown in preservative. Lateral line of tubercles most times restricted to posterior flanks (AAG-UFU 503) or poorly evident (AAG-UFU 505). Females (AAG-UFU 504–505) and most males (AAG- UFU 503, 506, 694, and 1102) have tiny white tubercles on the sole of foot, whereas the male AAG-UFU 6183 has the sole of foot nearly smooth. In most specimens, throat predominantly light-colored without dark gray spots and dots concentrated near the jaw (dark gray spots and dots present in AAG-UFU 6183). Advertisement call. Description based on 522 calls recorded from seven males (Table 3). The call consists of single, pulsed notes given at a rate of 145–196 per minute (Fig. 6g). Note duration varies from 62–126 ms. Rise time varies from 26–94% of note duration. Notes are formed by 6–12 complete pulses emitted at a rate of 75–125 per second. The dominant frequency coincides with the fundamental harmonic at 586–1383 Hz. Notes usually have a modest frequency upsweep throughout their duration, varying from 141–750 Hz. Habitat and natural history. At the type locality, individuals were found calling from inside underground chambers in open areas always adjacent to the forest edge, where water was accumulated as small puddles. The surroundings can be described as grass tufts in poorly drained marshy soil. Although all three study sites are characterized as moderately to highly modified by anthropogenic activities, small fragments of Atlantic forest were present in these areas. In the District of Povoação (Linhares, Espírito Santo), calling males were heard from underneath dense leaf litter in cacao plantations. Distribution. Leptodactylus barrioi is known to occur within the interior Atlantic forests of southeastern Minas Gerais, west of the Serra do Mar mountain range in Rio de Janeiro State, in the northern sector of the Mantiqueira mountain range in extreme south of Espírito Santo, and also in the coastal Atlantic forests alongside the lower Doce River in the same Brazilian state (all regions within southeastern Brazil; Fig. 1).

Adenomera phonotriccus, n. sp. Figures 1‒3 REFERRED MATERIAL: Voucher specimens with DNA sequenced by Fouquet et al. (2014) and assigned to Adenomera sp. F (in part). SUGGESTED VERNACULAR NAME: Tody-tyrant-voiced nest-building frog. HOLOTYPE: MPEG 41155 (field number TRC 135), adult male, from the municipality of Palestina do Pará (5.70228° S, 48.24949° W; 166 m), on the western margin of the lower Araguaia River, state of Pará, northern Brazil, collected by P.L. V. Peloso, M.J. Sturaro, P. V. Cerqueira, G. Gonçalves, A.A. Giaretta, P. Marinho, and T. R. de Carvalho on January 9, 2018 (Zoobank LSID registration: urn:lsid:zoobank.org:act: 330AE4AB-1ABB-4B90-9245-1DBC2A1F661B). SOUND RECORDING: Adenomera _phonotriccusPalestinaPA2bAAGm661 MK 2. PARATOPOTYPES: MPEG 41156 and CFBH 43130‒43131, all adult males, collected (5.70300° S, 48.24833° W; 176 m) on January 9‒10, 2018, by the same collectors. ETYMOLOGY: The epithet phonotriccus is the combination of Greek phono - (from phoné, “sound, voice”) and triccus (a small bird whose species is not identifiable, though in modern times the name is applied to tyrant flycatchers; Jobling, 2010). This name is to be treated as a noun in apposition and is an allusion to the similarity between the vocalization of the new species and those of tody-tyrants. Tody-tyrants of the Neotropical genus Hemitriccus have peculiar vocalizations, which are reminiscent of the vocalization of Adenomera phonotriccus, especially the trilled song of H. cohnhafli (see Zimmer et al., 2013). Interestingly, several Hemitriccus species are morphologically very similar, and consequently best differentiated by their vocalizations, a pattern also observed in Adenomera. DIAGNOSIS: Adenomera phonotriccus is differentiated from its congeners by the following combination of characters: (1) small size (adult male SVL 19.8–21.6 mm; table 1); (2) robust body shape; (3) toe tips unexpanded to slightly swollen (character states B, C; Heyer, 1973); (4) distal antebrachial tubercle on underside of forearm; (5) throat and belly cream colored, mottled white and gray in some parts; (6) two color morphotypes (presence/absence of dorsolateral stripes); (7) advertisement call consisting of a single type of pulsed note, emitted regularly, not in calling bouts; (8) advertisement notes composed of complete pulses (pulses with periods of silence in between); (9) long-lasting call duration (213‒433 ms). COMPARISONS: Adenomera phonotriccus has adult males (SVL 19.8–21.6 mm; table 1) smaller than those of A. coca (23.6–25.6 mm; Angulo and Reichle, 2008), A. lutzi (25.7–33.5 mm; Kok et al., 2007), and A. simonstuarti (SVL 25.9–26.2 mm; Angulo and Icochea, 2010). Adenomera phonotriccus has a robust body shape (figs. 1–2), whereas A. diptyx, A. martinezi, and A. saci have a slender body (Carvalho and Giaretta, 2013a). Adenomera phonotriccus has toe tips unexpanded or slightly swollen (character states B, C), but not expanded into flattened discs (character state D) as in A. ajurauna, A. andreae, A. marmorata, A. lutzi, A. nana, and A. simonstuarti (Angulo et al., 2003; Kok et al., 2007; Kwet, 2007; Berneck et al., 2008; Angulo and Icochea, 2010; appendix 1). Adenomera phonotriccus is distinguished from congeners (except A. cotuba and A. lutzi; Kok et al., 2007; Carvalho and Giaretta, 2013b) by having antebrachial tubercles on underside of forearm (fig. 3), presumably absent in all remaining species. Adenomera phonotriccus differs from A. heyeri and A. lutzi by having ventral surfaces cream colored (mottled white and gray in some parts), yellow in the latter two species (Boistel et al., 2006; Kok et al., 2007). From A. engelsi, which does not have distinctive dorsolateral stripes and pigmentation on throat (Kwet et al., 2009), Adenomera phonotriccus differs by the occurrence of both color features (fig. 1). Adenomera cotuba does not have dorsolateral stripes (Carvalho and Giaretta, 2013b), whereas A. phonotriccus has two color morphotypes (presence/ absence of dorsolateral stripes). These species are better distinguished by their distinct vocalizations (see below; fig. 4). The advertisement call of Adenomera phonotriccu s (fig. 4; table 2) distinguishes it from all congeners by having pulsed notes composed of complete pulses (periods of silence in between), whereas the other species’ notes are formed by incomplete pulses, i.e., not separated one from the next by periods of silence (fig. 4). From the remaining species of Adenomera, the new spe- cies differs by having pulsed advertisement notes, whereas those species’ calls consist of nonpulsed notes (table 3). The advertisement call of A. phonotriccus is distinguished from that of A. cotuba by being composed of single notes emitted regularly (table 2), whereas A. cotuba’ s call is made up of many notes emitted as calling bouts, i.e., multinote advertisement call (Carvalho and Giaretta, 2013b: fig. 4A). In addition, to date, Adenomera phonotriccus has the longest note duration in the genus, whose range does not overlap with note-duration values for any congeners (table 3). DESCRIPTION OF HOLOTYPE: MPEG 41155 (figs. 1A, 2, 3A). Adult male. Body robust (figs. 1A, 2A). Snout subovoid in dorsal view (fig. 2A), acuminate in lateral view (fig. 2D). Nostrils closer to the snout tip than to the eyes; fleshy ridge on snout tip; canthus rostralis not marked; loreal region slightly concave; supratympanic fold from the posterior corner of the eye to the base of the arm; oval postcommissural gland; vocal sac subgular with a fold from jaw extending to forearm on each side, vocal slits present; vomerine teeth in two straight rows medial and posterior to choanae and oblique to sagittal plane. Tongue elongated, free behind. Relative finger lengths IV a Sinusoidal oscillation (Boistel et al., 2006) corresponds to incomplete pulses in Adenomera, based on (quasi-) periodic sinusoidal FMs (sometimes there is strong modulation unaccompanied by distinct pulses though). b Based on the call of Adenomera sp. 2 from Joinville. c See Materials and Methods for a detailed explanation. d Frequencies lower than 1 kH do not pertain to this species’ call (appendix 2). COLOR OF HOLOTYPE IN PRESERVATIVE (fig. 2): Snout tip with a faded white coloration (coincident with the fleshy ridge). Dorsum marble gray with a few darker specks and spots. Sacral stripe on the posterior third of body length, cream-colored. Black lumbar glands. Dorsal surface of limbs with dark brown stripes/blotches on a slightly lighter brown background. Dorsolateral, broad stripe from the posterior corner of the eye, passing over the flanks, predominantly cream-colored to a somewhat orange hue (faded coloration; orange in life). Upper and lower jaws covered with white-colored spots/blotches, whitish postcommissural gland, outlined at the bottom by a dark coloration. Tympanum reddish brown. Throat, belly, and ventral surface of limbs cream, with melanophores, throat with a black-dotted pattern, solid dark-colored laterally, coinciding with the expanded vocal sac. Posterior surface of thighs with a few dark brown spots on a light brown background, and paracloacal glands cream colored. COLOR OF HOLOTYPE IN LIFE (fig. 1A): Dorsum covered with black specks and spots irregularly distributed on medium brown background. Arms and legs reddish brown. Sacral and dorsolateral stripe, and postcommissural gland orange. Iris copper. Tympanum brown. Ventral surfaces cream colored and with mottling on throat and belly (laterally). Groin yellow. VARIATION IN THE TYPE SERIES: Despite minor variation in dorsal coloration and distribution of spots, specks, and warts on dorsal surfaces, characteristics of the three paratypes largely agree with the description of the holotype. Dorsolateral stripe is absent in two paratypes (MPEG 41156 and CFBH 43130). Shape and size of antebrachial tubercle varies among the type series; some of them are very discrete (fig. 2). ADVERTISEMENT CALL: The description is based on recordings of three males (appendix 3). Sample sizes for each acoustic trait and descriptive statistics, i.e., means and standard deviations, are provided in table 2. The call (fig. 4) consists of a single type of pulsed note, which is emitted at a rate of 26–35 calls per minute. Note duration varies from 213–433 msec. Notes have a smooth attack and decay, reaching the amplitude peak at 43%–85% of their duration. Pulse number varies from 14–26; pulse duration from 4–26 msec, emitted at a rate of 51–67 pulses per second; silence between pulses varies from nearly zero to 13 msec. The fundamental frequency of the note occupies a narrow bandwidth harmonic, with a peak frequency of 1857– 1997 Hz; the dominant frequency varies from 3639–4113 Hz and is contained within the second harmonic. Notes usually have a slight positive frequency modulation along their duration, but a few notes do not have any modulation, and a single note has negative modulation. Linear frequency modulation varies from -43–861 Hz. HABITAT AND NATURAL HISTORY: Adenomera phonotriccus is associated with forest habitats in Brazil’s eastern Amazonia in the state of Pará. The type locality is located on the west margin of the lower Araguaia River. This region is also occupied by savanna landscapes or ecotones between the Cerrado and Amazonian rainforest. The species was heard along an open area transect (by recent logging activity), especially at the forest edge, and inside the forest remnant. Males called while exposed or under leaf litter, and increased their calling activity during and shortly after rainfalls. The vocalization of Adenomera phonotriccus is very peculiar when heard in the field. Our first impression was that the calls might not be produced by a leptodactylid frog, or even an anuran species. Indeed, the vocalization is similar to that of some Neotropical suboscine birds (e.g., Rhynchocyclidae, Tyrannidae). There were two partially syntopic species of Adenomera also in calling activity at the study site: Adenomera sp. (A. heyeri clade) and A. aff. hylaedactyla. We heard the three species calling at the same period (late afternoon), even though A. phonotriccus and Adenomera sp. ceased most calling activity at dusk, whereas A. aff. hylaedactyla continued calling through the first hours of the night. The last species occupied open areas, e.g. pasture. In contrast, A. phonotriccus and Adenomera sp., are associated with forest habitats. DISTRIBUTION (fig. 5): Adenomera phonotriccus is known with certainty only from the type locality and Marabá. However, specimens from other regions in the state of Pará were also assigned to this lineage, referred as Adenomera sp. F, by Fouquet et al. (2014). Given the high levels of cryptic species diversity in Adenomera and complex genetic structure within the lineages identified by Fouquet et al. (2014), we conservatively restrict the species’ distribution to the type locality region until additional data, especially vocalizations, are obtained for the other populations assigned to Adenomera sp. F, a lineage widely distributed in central-east Pará, northern Brazil. CONSERVATION STATUS: Because the species is apparently restricted to forested habitats, or its immediate margins (forest edge), we infer that the current population trend for this species is “declining.” Major threats are deforestation (destruction of habitats) due to conversion of large areas where the species occur into pasture and agricultural land. The species appears to be locally abundant (although difficult to observe), which means that as long as its habitat is preserved the species should have its future secured. The known range of the species is hard to determine with current evidence. It may be much wider, if all the clade F specimens refer to A. phonotriccus, in which case the species would be found from the western margin of the Tocantins River to the western margins of the Tapajós River (Fouquet et al., 2014: fig. 2). Alternatively, it could have a much narrower distribution (southern Pará) if only specimens in subclade F3 correspond to A. phonotriccus —in the latter case, the species would be probably restricted to small forest fragments in one of the most highly threatened portions of the Amazonia (the so-called Arc of Deforestation). This is a typical example of the issues raised by Peloso (2010) and Angulo and Icochea (2010), where lack of taxonomic resolution hinders a proper assessment of threats to cryptic species. For now, we must unfortunately suggest this new species is yet another case of data-deficient taxa—a lot more studies and data are needed to define the conservation status of A. phonotriccus. REMARKS: Calls of the voucher PV 2597 (specimen no. DT 2123; recordings PLPDR 100; see appendix 3) were unambiguously assigned to A. phonotriccus (“F3” in fig. 6). This voucher was cited as collected in Araguatins (state of Tocantins), on the east margin of Araguaia River. However, this specimen was actually collected in Palestina do Pará, state of Pará—the animal was collected by P.L.V.P. in 2010 (coordinates given by Fouquet et al.’s appendix 1 are correct). The basecamp for this collection site was in Araguatins, and this may have been the source of confusion with the sample provenance (P.L.V.P., personal obs.). Another voucher specimen used in the phylogeny (PV 2412, specimen no. DT 2016), also assigned here to A. phonotriccus (“F3” in fig. 6) was collected in Marabá—this municipality is located in the state of Pará, not Amazonas (AM), as mentioned in appendix 1 of Fouquet et al. (2014). Of special interest is the acquisition of additional information for the subclade F1, which occurs in partial sympatry with A. phonotriccus (fig. 6). Meanwhile, we conservatively assign only subclade F3 (sensu Fouquet et al., 2014) to A. phonotriccus, and this species is an endemic of the central-east part of the state of Pará in association with the west margin of the lower Araguaia River (fig. 6). With respect to subclade F2, we suspect that it may not be conspecific with A. phonotriccus, given its distribution range associated with another region in the state of Pará (Xingu-Tapajós interfluve; fig. 6). Therefore, we maintain the original status of unconfirmed candidate species of both subclades F1 and F2 for the moment.

De Carvalho, Thiago R., Giaretta, Ariovaldo A., Angulo, Ariadne, Haddad, Célio F.B., Peloso, Pedro L.V. (2019): A new Amazonian species of Adenomera (Anura: Leptodactylidae) from the Brazilian state of Pará: a tody-tyrant voice in a frog. American Museum Novitates 2019 (3919): 1-24, DOI: https://doi.org/10.1206/3919.1