Abstract:

Among amphibian species from high elevation areas in the Brazilian Atlantic Forest there is a high percentage of threatened and endemic species, but there is still a relative scarcity of local inventories for these organisms. Here, we present data on anuran composition, relative abundance and estimated densities for leaf-litter frogs from an Atlantic Forest area within the APA Serra da Mantiqueira, in Rio de Janeiro state, Brazil, based on results of a short-term survey carried out at altitudes of 1,350-1,750 m, in November 2005 (with additional records from surveys made in 2010 and 2011). Three sampling methods were used during the 2005 survey: plot sampling, visual encounter surveys (VES; performed during the day, at the dusk, and at night), and pitfall traps with drift fences; only non-standardized visual searches were employed during the 2010 and 2011 surveys. We recorded 24 species, with the direct-developer Ischnocnema sp. (gr. lactea) being the most abundant. Most anurans (90% of all individuals) sampled by VES were captured during the crepuscular and nocturnal periods. The estimated density of the local leaf-litter frog assemblage based on plot sampling was 18.4 ind/100 m², which is one of the highest values currently reported for Atlantic Rainforest areas. This is the first study analyzing the anuran fauna composition of an Atlantic Forest area within the APA Serra da Mantiqueira and adds to the body of knowledge on the fauna of the southern region of Rio de Janeiro state.

Keywords:
amphibians; endemism; inventories; montane forest; southeastern Brazil

Resumo:

Entre as espécies de anfíbios de áreas de altas altitudes da Mata Atlântica brasileira há uma elevada porcentagem de espécies ameaçadas e endêmicas, mas ainda há relativa escassez de inventários locais desses organismos. Aqui, apresentamos dados sobre a composição de espécies, abundância relativa e densidade estimada para anfíbios anuros de serapilheira de área de Mata Atlântica na APA Serra da Mantiqueira, estado do Rio de Janeiro, Brasil, com base em resultados de estudo de curto prazo realizado a altitudes de 1.350-1.750 m, em novembro de 2005 (incluindo registros adicionais de coletas feitas em 2010 e 2011). Três métodos de amostragem foram utilizados na campanha de 2005: amostragem em parcelas, encontros visuais (realizados durante o dia, no crepúsculo e à noite) e armadilhas de queda; apenas procuras visuais não padronizadas foram usadas nas coletas de 2010 e 2011. Foram registradas 24 espécies, sendo a mais abundante Ischnocnema sp. (gr. lactea), uma forma com desenvolvimento direto. A maioria dos anuros (90% de todos os indivíduos) amostrados pelo método de encontros visuais foi capturada durante os períodos crepuscular e noturno. A densidade estimada para anuros de serapilheira com base nas amostragens em parcelas foi de 18,4 ind/100 m2, um dos valores mais altos registrados até o momento para áreas de Mata Atlântica. Este é o primeiro estudo que analisa a composição da fauna de anuros de área de Mata Atlântica da APA Serra da Mantiqueira e contribui para o conhecimento da fauna da região sul do estado do Rio de Janeiro.

Palavras-chave:
anfíbios; endemismo; floresta serrana; inventários; sudeste do Brasil

Introduction

Like many other animals (e.g., Kattan & Franco 2001, McCain 2004MCCAIN, C.M. 2004. The mid-domain effect applied to elevational gradients: Species diversity of small mammals in Costa Rica. J. Biogeogr. 31(1):19-31., Liew et al. 2010LIEW, T.-S., SCHILTHUIZEN, M. & BIN LAKIM, M. 2010. The determinants of land snail diversity along a tropical elevational gradient: insularity, geometry and niches. J. Biogeogr. 37(6):1071-1078.), anuran amphibians typically present comparatively lower species richness at higher altitudes (Duellman 1988DUELLMAN, W.E. 1988. Patterns of species diversity in anuran amphibians in the American Tropics. Ann. Missouri Bot. Gard. 75:79-104.; Zancolli et al. 2014ZANCOLLI, G., STEFFAN-DEWENTER, I. & RÖDEL, M.O. 2014. Amphibian diversity on the roof of Africa: Unveiling the effects of habitat degradation, altitude and biogeography. Divers. Distrib. 20(3):297-308., Siqueira et al. 2021SIQUEIRA, C.C., VRCIBRADIC, D., ALMEIDA-GOMES, M. & ROCHA, C.F.D. 2021. Assessing the importance of reproductive modes for the evaluation of altitudinal distribution patterns in tropical frogs. Biotropica 53(3):786-797.). On the other hand, highland habitats are often important centers of endemism for these organisms, especially in the tropics, with many species being restricted to these areas (e.g., Fu et al. 2006FU, C., HUA, X., LI, J., CHANG, Z., PU, Z. & CHEN, J. 2006. Elevational patterns of frog species richness and endemic richness in the Hengduan Mountains, China: Geometric constraints, area and climate effects. Ecography 29(6):919-927., Cruz & Feio 2007CRUZ, C.A.G. & FEIO, R.N. 2007. Endemismos em anfíbios em áreas de altitude na Mata Atlântica no sudeste do Brasil. In: Herpetologia no Brasil II (L.B. Nascimento & M.E. Oliveira, eds). Sociedade Brasileira de Herpetologia, Belo Horizonte, p.117-126., Bernal & Lynch 2008BERNAL, M.H. & LYNCH, J.D. 2008. Review and analysis of altitudinal distribution of the Andean anurans in Colombia. Zootaxa 1826:1-25.).

Recent surveys have provided lists of amphibian species, mostly preliminary, for some high montane Atlantic Forest areas (i.e., above 1,000 m) in southeast Brazil (Cruz et al. 2009CRUZ, C.A.G., FEIO, R.N. & CARAMASCHI U. 2009. Anfíbios do Ibitipoca. Bicho do Mato Editora, Belo Horizonte., Siqueira et al. 2011aSIQUEIRA, C.C., VRCIBRADIC, D., ALMEIDA-GOMES, M., MENEZES, V.A., BORGES-JUNIOR, V.N.T., HATANO, F.H., PONTES, J.A.L., GOYANNES-ARAÚJO, P., GUEDES, D.M., VAN SLUYS, M. & ROCHA, C.F.D. 2011a. Species composition and density estimates of the anurofauna of a site within the northernmost large Atlantic Forest remnant (Parque Estadual do Desengano) in the state of Rio de Janeiro, Brazil. Biota Neotrop. 11(4):131-137.,b, Garey et al. 2014GAREY, M.V., PROVETE, D.B., MARTINS, I.A., HADDAD, C.F.B. & ROSSA-FERES, D.C. 2014. Anurans from the Serra da Bocaina national park and surrounding buffer area, southeastern Brazil. Check List 10(2):308-216., 2016GAREY, M.V. & PROVETE, D.B. 2016. Species composition, conservation status, and sources of threat of anurans in mosaics of highland grasslands of southern and southeastern Brazil. Oecol. Aust. 20(2):94-108., Folly et al. 2016FOLLY, M, BEZERRA, A.M., RUGGERI, J., HEPP, F., CARVALHO-E-SILVA, A.M.P.T., GOMES, M.R. & CARVALHO-E-SILVA, S.P. 2016. Anuran fauna of the high-elevation areas of the Parque Nacional da Serra dos Órgãos (PARNASO), Southeastern Brazil. Oecol. Aust. 20(2):247-258., Lima et al. 2021LIMA, L.M.C., YVES, A., ALMEIDA, V.G., NEVES, M.O. & SOUSA, B.M. 2021. Anurans of a protected area from Mantiqueira Complex, Atlantic Forest of Minas Gerais state, Brazil. Biota Neotrop. 21(3):e20200993. https://doi.org/10.1590/1676-0611-BN-2020-0993 (last access on 15/07/2021)
https://doi.org/10.1590/1676-0611-BN-202... ). These habitats are known for their high rates of endemism of anurans, particularly in portions of the Serra do Mar (Siqueira et al. 2011bSIQUEIRA, C.C., VRCIBRADIC, D., DORIGO, T.A. & ROCHA, C.F.D. 2011b. Anurans from two high-elevation areas of Atlantic Forest in the state of Rio de Janeiro, Brazil. Zoologia 28(4):457-464., Carvalho-e-Silva et al. 2020CARVALHO-E-SILVA, S.P., CARVALHO-E-SILVA, A.M.P.T., FOLLY, M., LUNA-DIAS, C., BEZERRA, A.M., GOMES M.R., CARAM, J., PEIXOTO, O.L. & IZECKSOHN, E. 2020. Parque Nacional da Serra dos Órgãos: the highest Amphibian diversity within an Atlantic Forest protected area. Biota Neotrop. 20(3):e20201033. https://doi.org/10.1590/1676-0611-BN-2020-1033 (last access on 12/07/2021)
https://doi.org/10.1590/1676-0611-BN-202... ) and Serra da Mantiqueira regions (Neves et al. 2018NEVES, M.O., PEREIRA, E.A., SUGAI, J.L.M.M., ROCHA, S.B., FEIO, R.N. & SANTANA, D.J. 2018. Distribution pattern of anurans from three mountain complexes in southeastern Brazil and their conservation implications. An. Acad. Bras. Ciênc. 90(2):1611-1623., Silva et al. 2018SILVA, E.T., PEIXOTO, M.A.A., LEITE, F.S.F., FEIO, R.N. & GARCIA, P.C. 2018. Anuran distribution in a highly diverse region of the Atlantic Forest: The Mantiqueira Mountain Range in Southeastern Brazil. Herpetologica 74(4):294-305.).

The mountain complex of Serra da Mantiqueira contains the highest areas of the Atlantic Forest domain (reaching nearly 2,900 m of altitude), harboring at least 234 amphibian species, 88 of them endemic to the region (Silva et al. 2018SILVA, E.T., PEIXOTO, M.A.A., LEITE, F.S.F., FEIO, R.N. & GARCIA, P.C. 2018. Anuran distribution in a highly diverse region of the Atlantic Forest: The Mantiqueira Mountain Range in Southeastern Brazil. Herpetologica 74(4):294-305.). In the Serra da Mantiqueira complex, the Serra do Itatiaia region is reported as the area with the highest amphibian richness, with 61 species (nine endemic), many of them originally described from the Parque Nacional de Itatiaia (Neves et al. 2018NEVES, M.O., PEREIRA, E.A., SUGAI, J.L.M.M., ROCHA, S.B., FEIO, R.N. & SANTANA, D.J. 2018. Distribution pattern of anurans from three mountain complexes in southeastern Brazil and their conservation implications. An. Acad. Bras. Ciênc. 90(2):1611-1623.). However, there is still no published list of amphibian species for the region of Itatiaia since Neves et al. (2018)NEVES, M.O., PEREIRA, E.A., SUGAI, J.L.M.M., ROCHA, S.B., FEIO, R.N. & SANTANA, D.J. 2018. Distribution pattern of anurans from three mountain complexes in southeastern Brazil and their conservation implications. An. Acad. Bras. Ciênc. 90(2):1611-1623. presented the species richness value but listed only the endemic species. Adjacent to the Parque Nacional de Itatiaia, there is another Conservation Unit, the Área de Proteção Ambiental Serra da Mantiqueira (APA Serra da Mantiqueira). In spite of covering more than 400,000 ha and encompassing portions of three Brazilian states (MMA 2018MMA - Ministério do Meio Ambiente. 2018. Plano de Manejo da Área de Proteção Ambiental da Serra da Mantiqueira. ICMBIO, Brasília.), published data on the amphibian fauna from this Conservation Unit is, to our knowledge, available for only one locality in the state of Rio de Janeiro. Information on the anuran species occurring in that locality is presently restricted to studies reporting an extension of the geographical distribution of Paratelmatobius mantiqueira (Vrcibradic et al. 2010VRCIBRADIC, D., ARIANI, C.V., VAN SLUYS, M. & ROCHA, C.F.D. 2010. Amphibia, Leptodactylidae, Paratelmatobius mantiqueira Pombal and Haddad, 1999: Distribution extension. Check List 6(1):1-2.), some ecological aspects of Proceratophrys mantiqueira (Almeida-Gomes et al. 2007ALMEIDA-GOMES, M., VRCIBRADIC, D., BORGES JR, V.N.T., VAN SLUYS, M. & ROCHA, C.F.D. 2007. Proceratophrys melanopogon. Reproduction. Herpetol. Rev. 38(2):191-192., Almeida-Santos et al. 2017ALMEIDA-SANTOS, M., SIQUEIRA, C.C., ANJOS, L.A., VAN SLUYS, M. & ROCHA, C.F.D. 2017. Ecological aspects of the horned leaf-frog Proceratophrys mantiqueira (Odontophrynidae) in an Atlantic Rainforest area of southeastern Brazil. Salamandra 53(3):413-422.), and a record of snake predation on Scinax cardosoi (Dorigo et al. 2014DORIGO, T.A., VRCIBRADIC, D., BORGES-JR., V.N.T. & ROCHA, C.F.D. 2014. New records of anuran predation by snakes of the genus Thamnodynastes Wagler, 1830 (Colubridae: Dipsadinae) in the Atlantic Rainforest of southeastern Brazil. Herpetol. Notes 7:261-264.). Another study presented data on sampling efficiency during different periods of the day and the number of species sampled at the area but did not provide information on the local species composition (Rocha et al. 2015ROCHA, C.F.D., SIQUEIRA, C.C., ARIANI, C.V., VRCIBRADIC, D., GUEDES, D.D.M., KIEFER, M.C., ALMEIDA-GOMES, M., GOYANNES-ARAÚJO, P., BORGES-JR, V.N.T. & VAN SLUYS, M. 2015. Differential success in sampling of Atlantic Forest amphibians among different periods of the day. Braz. J. Biol. 75(2):261-267.).

The state of Rio de Janeiro, whose territory is entirely included within the Atlantic Forest domain, was recently reported to harbor 201 amphibian species (Dorigo et al. 2018DORIGO, T.A., VRCIBRADIC, D. & ROCHA, C.F.D. 2018. The amphibians of the state of Rio de Janeiro, Brazil: An updated and commented list. Pap. Avulsos Zool. 58:e20185805. https://doi.org/10.11606/1807-0205/2018.58.05 (last access on 02/11/2020)
https://doi.org/10.11606/1807-0205/2018.... ), and new species with occurrence in the state are still being steadily described (e.g., Folly et al. 2018FOLLY, M., HEPP, F. & CARVALHO-E-SILVA S.P. 2018. A new bromeligenous species of Fritziana Mello-Leitão, 1937 (Amphibia: Anura: Hemiphractidae) from high elevations in the Serra dos Órgãos, Rio de Janeiro, Brazil. Herpetologica 74(1):58-72., Pereira Silva et al. 2018PEREIRA SILVA, E.A., ROCHA, L.C.L., FOLLY, H., SILVA, H.R. & SANTANA, D.J. 2018. A new species of spotted leaf frog, genus Phasmahyla (Amphibia, Phyllomedusidae) from Southeast Brazil. PeerJ 6:e4900. https://doi.org/10.7717/peerj.4900 (last access on 12/07/2021)
https://doi.org/10.7717/peerj.4900... , Taucce et al. 2018TAUCCE, P.P.G., CANEDO, C., PARREIRAS, J.S., DRUMMOND, L.O., NOGUEIRA-COSTA, P. & HADDAD, C.F.B. 2018. Molecular phylogeny of Ischnocnema (Anura: Brachycephalidae) with the redefinition of its series and the description of two new species. Mol. Phylogenet. Evol. 128:123-146., Cruz et al. 2019CRUZ, C.A.G., CARAMASCHI, U., FUSINATTO, L.A. & BRASILEIRO, C.A. 2019. Taxonomic review of Dendrophryniscus brevipollicatus Jiménez de la Espada, 1870, with revalidation of D. imitator (Miranda-Ribeiro, 1920) and D. lauroi Miranda-Ribeiro, 1926, and description of four new related species (Anura, Bufonidae). Zootaxa 4648:27-62., Silva et al. 2020SILVA, L.A., MAGALHÃES, F.M., THOMASSEN, H., LEITE, F.S.F., GARDA, A.A., BRANDÃO, R.A., HADDAD, C.F.B., GIARETTA, A.A. & CARVALHO, T.R. 2020. Unraveling the species diversity and relationships in the Leptodactylus mystaceus complex (Anura: Leptodactylidae), with the description of three new Brazilian species. Zootaxa 4779(2):151-189., Nunes et al. 2021NUNES, I., GUIMARÃES, C.S., MOURA, P.H.A.G., PEDROZO, M., MOROTI, M.T., CASTRO, L.M., STUGINSKI, D.R. & MUSCAT, E. 2021. Hidden by the name: A new fluorescent pumpkin toadlet from the Brachycephalus ephippium group (Anura: Brachycephalidae). PLoS ONE 16(4):e0244812.). In recent years, lists of species of amphibians have been compiled or updated for several forested areas in the state of Rio de Janeiro (e.g., Almeida-Gomes et al. 2014ALMEIDA-GOMES, M., SIQUEIRA, C.C., BORGES-JR, V.N.T., VRCIBRADIC, D., FUSINATTO, L.A. & ROCHA, C.F.D. 2014. Herpetofauna of the Reserva Ecológica de Guapiaçu (REGUA) and its surrounding areas, in the state of Rio de Janeiro, Brazil. Biota Neotrop. 14(3):e20130078. https://doi.org/10.1590/1676-0603007813 (last access on 02/11/2020)
https://doi.org/10.1590/1676-0603007813... , Martins et al. 2014MARTINS, A., PONTES, R., MATTEDI, R., FRATANI, J., MURTA-FONSECA, R.A., RAMOS, L. & BRANDÃO, A.L.R. 2014. Anuran community of a coastal Atlantic Forest fragment in the state of Rio de Janeiro, southeastern Brazil. Salamandra 50(1):27-39., Caram et al. 2016CARAM, J., GOMES, M.R., LUNA-DIAS, C. & CARVALHO-E-SILVA, S.P. 2016. Updated list of anurans from Floresta Nacional Mário Xavier, Seropédica, Rio de Janeiro, Brazil: Changes from 1990 to 2012. Check List 12(6):1-7., Rocha et al. 2018ROCHA, C.F.D., TELLES, F.B.S., VRCIBRADIC, D. & COSTA, P.N. 2018. The herpetofauna from Ilha Grande (Angra dos Reis, Rio de Janeiro, Brazil): Updating species composition, richness, distribution and endemisms. Pap. Avulsos Zool. 58:e20185825. https://doi.org/10.11606/1807-0205/2018.58.25 (last access on 02/11/2020)
https://doi.org/10.11606/1807-0205/2018.... , Carvalho-e-Silva et al. 2020CARVALHO-E-SILVA, S.P., CARVALHO-E-SILVA, A.M.P.T., FOLLY, M., LUNA-DIAS, C., BEZERRA, A.M., GOMES M.R., CARAM, J., PEIXOTO, O.L. & IZECKSOHN, E. 2020. Parque Nacional da Serra dos Órgãos: the highest Amphibian diversity within an Atlantic Forest protected area. Biota Neotrop. 20(3):e20201033. https://doi.org/10.1590/1676-0611-BN-2020-1033 (last access on 12/07/2021)
https://doi.org/10.1590/1676-0611-BN-202... , Dorigo et al. 2021DORIGO, T.A., SIQUEIRA, C.C., OLIVEIRA, J.C.F., FUSINATTO, L.A., SANTOS-PEREIRA, M., ALMEIDA-SANTOS, M., MAIA-CARNEIRO, M., REIS, C.N.C. & ROCHA, C.F.D. 2021. Amphibians and reptiles from the Parque Nacional da Tijuca, Brazil, one of the world’s largest urban forests. Biota Neotrop. 21(2):e20200978. https://doi.org/10.1590/1676-0611-BN-2020-0978 (last access on 15/07/2021)
https://doi.org/10.1590/1676-0611-BN-202... ), contributing considerably to the knowledge of its amphibian fauna. However, gaps of knowledge for areas at high altitudes still remain, not only for the state of Rio de Janeiro, but for the Atlantic Forest biome as a whole (Lima et al. 2021LIMA, L.M.C., YVES, A., ALMEIDA, V.G., NEVES, M.O. & SOUSA, B.M. 2021. Anurans of a protected area from Mantiqueira Complex, Atlantic Forest of Minas Gerais state, Brazil. Biota Neotrop. 21(3):e20200993. https://doi.org/10.1590/1676-0611-BN-2020-0993 (last access on 15/07/2021)
https://doi.org/10.1590/1676-0611-BN-202... ).

In this study, we address some parameters of the amphibian assemblage from a locality within the APA Serra da Mantiqueira for which preliminary data on the anuran fauna has been previously published, as mentioned above (Almeida-Gomes et al. 2007ALMEIDA-GOMES, M., VRCIBRADIC, D., BORGES JR, V.N.T., VAN SLUYS, M. & ROCHA, C.F.D. 2007. Proceratophrys melanopogon. Reproduction. Herpetol. Rev. 38(2):191-192., Vrcibradic et al. 2010VRCIBRADIC, D., ARIANI, C.V., VAN SLUYS, M. & ROCHA, C.F.D. 2010. Amphibia, Leptodactylidae, Paratelmatobius mantiqueira Pombal and Haddad, 1999: Distribution extension. Check List 6(1):1-2., Dorigo et al. 2014DORIGO, T.A., VRCIBRADIC, D., BORGES-JR., V.N.T. & ROCHA, C.F.D. 2014. New records of anuran predation by snakes of the genus Thamnodynastes Wagler, 1830 (Colubridae: Dipsadinae) in the Atlantic Rainforest of southeastern Brazil. Herpetol. Notes 7:261-264., Rocha et al. 2015ROCHA, C.F.D., SIQUEIRA, C.C., ARIANI, C.V., VRCIBRADIC, D., GUEDES, D.D.M., KIEFER, M.C., ALMEIDA-GOMES, M., GOYANNES-ARAÚJO, P., BORGES-JR, V.N.T. & VAN SLUYS, M. 2015. Differential success in sampling of Atlantic Forest amphibians among different periods of the day. Braz. J. Biol. 75(2):261-267., Almeida-Santos et al. 2017ALMEIDA-SANTOS, M., SIQUEIRA, C.C., ANJOS, L.A., VAN SLUYS, M. & ROCHA, C.F.D. 2017. Ecological aspects of the horned leaf-frog Proceratophrys mantiqueira (Odontophrynidae) in an Atlantic Rainforest area of southeastern Brazil. Salamandra 53(3):413-422.). We present data on species composition, richness and relative abundances of anuran amphibians in that locality, as well as density estimates (individuals per 100 m²) for frogs inhabiting the forest floor leaf litter.

Material and Methods

1. Study area

The study was carried out in a locality within the APA Serra da Mantiqueira, a Conservation Unit encompassing parts of the states of Minas Gerais, Rio de Janeiro and São Paulo, southeastern Brazil. Created in 1985, the APA Serra da Mantiqueira (437,525 ha) comprises an area varying from 458 m up to 2,798 m of elevation, and covers, totally or partially, approximately 50 Conservation Units (MMA 2018MMA - Ministério do Meio Ambiente. 2018. Plano de Manejo da Área de Proteção Ambiental da Serra da Mantiqueira. ICMBIO, Brasília.). This study’s surveys were carried out within a private property, the Marimbondo Farm (22°21’50” S, 44°35’25” W; 34,475 ha), whose area is contained within both Resende and Itatiaia municipalities, state of Rio de Janeiro (Figure 1). Vegetation in the study area is characterized by Upper Montane Rain Forest (sensu Oliveira-Filho & Fontes 2000OLIVEIRA-FILHO, A.T. & FONTES, M.A.L. 2000. Patterns of floristic differentiation among Atlantic Forests in Southeastern Brazil and the influence of climate. Biotropica 32(4b):793-810.; Figure 2). Mean annual temperature in the region is 14.9°C, and mean annual precipitation is 1,813 mm (Attias et al. 2009ATTIAS, N., RAÍCES, D.S.L., PESSOA, F.S., ALBUQUERQUE, H., JORDÃO-NOGUEIRA, T., MODESTO, T.C. & BERGALLO, H.G. 2009. Potential distribution and new records of Trinomys species (Rodentia: Echimyidae) in the state of Rio de Janeiro. Zoologia 26(2):305-315.).

2. Research design, field methods and data analysis

Fieldwork was conducted at sites between 1,350 m and 1,750 m of elevation, in three different years: 2005, 2010, and 2011. In November 2005, to obtain a representative dataset of the anuran fauna for the study area (including both qualitative and quantitative data) we used three sampling methods. Two of them, visual encounter surveys (Crump & Scott 1994CRUMP, M.L. & SCOTT JR., N.J. 1994. Visual encounter surveys. In: Measuring and monitoring biological diversity: standard methods for amphibians (W.R. Heyer, M.A. Donnelly, R.W. McDiarmid, L.A.C. Hayek & M.S. Foster, eds.). Smithsonian Institution Press, Washington DC, p.84-92.) and plot sampling (Jaeger & Inger 1994JAEGER. R. & INGER, R.F. 1994. Standard techniques for inventory and monitoring: Quadrat Sampling In: Measuring and monitoring biological diversity: standard methods for amphibians (W.R. Heyer, M.A. Donnelly, R.W. McDiarmid, L.A.C. Hayek & M.S. Foster, eds). Smithsonian Institution Press, Washington DC, p.97-102.), were performed from 14-18 November 2005. The third method, pitfall traps with drift fences (Corn 1994CORN, P.S. 1994. Straight-line drift fences and pitfall traps. In Measuring and monitoring biological diversity: standard methods for amphibians (W.R. Heyer, M.A. Donnelly, R.W. McDiarmid, L.A.C. Hayek & M.S. Foster, eds). Smithsonian Institution Press, Washington DC, p.109-117.), was employed from 05-11 and from 14-19 November 2005. Additional sampling was carried out from 18-20 December 2010 and from 13-15 November 2011 using non-standardized methodology.

For visual encounter surveys (VES), 150 time-constrained searching bouts of 30 minutes each were carried out daily by ten people wearing headlamps, totaling 75 hours of sampling effort. Equal numbers of transects (50) were surveyed during the diurnal (11:00-16:30h), crepuscular (17:30-18:30h) and nocturnal (19:30-22:00h) periods. During transect sampling, each observer moved at a slow walking pace, carefully searching all types of potential microhabitats for anurans (e.g., leaf litter, water bodies, bromeliads, fallen logs, tree trunks, shrubs, rocks).

For the plot method, 30 quadrats of 5 x 5 m were established on the forest floor during the afternoon, totaling 750 m² of area sampled. The corners of each plot were marked with wooden stakes and the area inside was enclosed with a 50 cm high soft plastic fence, whose base was buried or attached to the ground. Soon after sunset, each plot was carefully searched for about half an hour by a crew of five persons using headlamps, moving on hands and knees, side-by-side. During the searches the leaf-litter was stirred with hand rakes, and leaves, stones and fallen branches were overturned; rock crevices and fissures among tree roots were also checked for anurans. Plot sampling was performed during five consecutive days, with six quadrats surveyed per night.

Three pitfall trap systems were established within the forest and remained open for a total of 11 days. Each system consisted of ten 30-liter buckets buried on the ground up to their rims, each bucket set ca. 5 m apart from the nearest one, with soft plastic drift fences about 50 cm high extended between them. Six buckets were set in line and the other four were placed at opposite ends of the fence, perpendicular to the main axis. Pitfalls were checked once per day for captured animals, always in the morning.

All anurans found by the three sampling methods were collected and identified (whenever possible). Estimates of leaf litter frog density (ind/100 m²) were produced based on the data obtained through plot method. An evaluation of sampling effectiveness was undertaken by cumulative and individual-based rarefaction curves performing 1,000 randomizations without replacement, using the program EstimateS 9.1.0 (Colwell 2013COLWELL, R.K. 2013. EstimateS: Statistical estimation of species richness and shared species from samples. Version 9.1.0. http://viceroy.colorado.edu/estimates/ (accessed on 05/07/2021)
http://viceroy.colorado.edu/estimates/... ). We carried out this procedure for VES and plot methods separately, using abundance data.

Further excursions for sampling the same area occurred in December 2010 and in November 2011 for collection of additional specimens. In these excursions, a three-person team performed haphazard visual searches (not standardized by time, unlike those of 2005) between 17:00 and 22:00h, at altitudes of 1,450-1,550 m, during two consecutive nights. Additionally, in 2011, funnel traps (made with five-liter plastic bottles) baited with ham were set at the bottom of a stream to capture tadpoles. Records obtained by these collections were incorporated into the local species list but were not used in quantitative analyses.

3. Taxonomic issues

Voucher specimens of anuran species collected during the study were deposited at the amphibian collection of the Museu Nacional, Universidade Federal do Rio de Janeiro (MNRJ), in the municipality of Rio de Janeiro (see Appendix).

Bokermannohyla circumdata can be potentially confused with B. luctuosa (Pombal & Haddad, 1993) (which also occurs in the region of Itatiaia; see Silva et al. 2018SILVA, E.T., PEIXOTO, M.A.A., LEITE, F.S.F., FEIO, R.N. & GARCIA, P.C. 2018. Anuran distribution in a highly diverse region of the Atlantic Forest: The Mantiqueira Mountain Range in Southeastern Brazil. Herpetologica 74(4):294-305.) since the taxonomic limits between these two species may be considered problematic (M. F. Napoli, pers. comm.). The larger Bokermannohyla individuals from the Marimbondo Farm were assigned to B. circumdata due to the presence (albeit not in all specimens) of bifid distal subarticular tubercles on fingers III and IV (see Napoli 2000NAPOLI, M. F. 2000. Taxonomia, variação morfológica e distribuição geográfica das espécies do grupo de Hyla circumdata (Cope, 1870) (Amphibia, Anura, Hylidae). PhD Thesis, Universidade Federal do Rio de Janeiro, Rio de Janeiro.). Also, Silva et al. (2018)SILVA, E.T., PEIXOTO, M.A.A., LEITE, F.S.F., FEIO, R.N. & GARCIA, P.C. 2018. Anuran distribution in a highly diverse region of the Atlantic Forest: The Mantiqueira Mountain Range in Southeastern Brazil. Herpetologica 74(4):294-305. cited our specimens as B. circumdata in their list of material examined, which indicates that those authors have seen that material and confirmed its identification. A second, smaller form of Bokermannohyla recorded during our surveys could not be positively identified to any currently described species in the genus and is treated herein as Bokermannohyla sp.

Phantasmarana (formerly Megaelosia) lutzae remained unrecorded for decades since the types and other associated specimens were collected in the Parque Nacional de Itatiaia (Izecksohn & Gouvêa 1987IZECKSOHN, E. & GOUVÊA, E. 1987 “1985”. Nova espécie de Megaelosia de Itatiaia, Estado do Rio de Janeiro. Arq. Univ. Fed. Rur. Rio de Janeiro 8(1-2):17-22.). Even though only tadpoles were recorded in our surveys, we felt confident in assigning them to P. lutzae as this is the only species of large-bodied hylodid currently known from the region of Itatiaia and its surroundings (Vittorazzi et al. 2021VITTORAZZI, S.E., AUGUSTO-ALVES, G., NEVES-DA-SILVA, D., CARVALHO-E-SILVA, A.M.P.T., RECCO-PIMENTEL, S.M., TOLEDO, L.F., LOURENÇO, L.B. & BRUSCHI, D.P. 2021. Paraphyly in the giant torrent-frogs (Anura: Hylodidae: Megaelosia) and the description of a new genus. Salamandra 57(2):274-284.). Moreover, some traits of the tadpoles, such as the presence of distinct dark blotches on the tail and the small marginal papillae (compared to those of Megaelosia goeldii tadpoles) match the brief description given by Izecksohn & Gouvêa (1987)IZECKSOHN, E. & GOUVÊA, E. 1987 “1985”. Nova espécie de Megaelosia de Itatiaia, Estado do Rio de Janeiro. Arq. Univ. Fed. Rur. Rio de Janeiro 8(1-2):17-22..

Results

A total of 24 anuran species belonging to eight families were recorded, of which six were registered only during samplings carried out in 2010 and 2011 (Table 1, Figures 3 and 4). For two species, Phantasmarana lutzae and Phasmahyla cochranae, only tadpoles were recorded. The anuran assemblage was dominated by species of the family Hylidae (ten species).

The VES method yielded records of 132 individuals from 17 anuran species, with Proceratophrys mantiqueira (N = 28, or 21.1% of all individuals found), Ischnocnema sp. (gr. lactea) (N = 21, or 15.9%), and Bokermannohyla circumdata (N = 18, or 13.6%) being the ones most frequently sampled (Table 1). Most anuran species recorded during transects were found during the nocturnal (N = 14 species) and crepuscular periods (N = 9), with eight species being recorded only at night and three species found only during crepuscular samplings (Table 1). Four species were recorded both during crepuscular and nocturnal periods, and only the two most abundant species, P. mantiqueira and Ischnocnema sp. (gr. lactea), were recorded on all three sampling periods (Table 1). Moreover, most individuals (regardless of the species) were found during the nocturnal (N = 78, or 59.1%) and crepuscular (N = 41, or 31.1%) transect searches.

A total of 138 individuals belonging to four anuran species were found during plot sampling (Table 1). The number of anurans per plot ranged from zero (7/30 or 23.3% of all plots) to 25 (1/30 or 3.3% of all plots). The estimated overall density of the local leaf litter frog assemblage was 18.4 ind/100 m². The highest density was that of Ischnocnema sp. (gr. lactea) (16.1 ind/100 m²), followed by Proceratophrys mantiqueira (2.0 ind/100 m²), and those two species together comprised about 98.5% of individuals found in plots. Both Ischnocnema aff. guentheri and Rhinella icterica had an estimated density of 0.1 ind/100 m².

Nine individuals of three species were captured in pitfall traps, with Proceratophrys mantiqueira (N = 7) being the most frequent (Table 1). Overall (i.e., including all methodologies pooled) the most abundant species recorded during the study were Ischnocnema sp. (gr. lactea) and P. mantiqueira (comprising 51.3% and 17.9% of all individuals collected, respectively).

Cumulative and individual-based species-rarefaction curves showed an asymptotic shape for VES, but it was non-asymptotic for plot-sampling data (Figure 5).

Discussion

This is the first study to provide an approach of species composition, richness and estimated density (on the forest floor) of amphibians for any area within the APA Serra da Mantiqueira, reporting a total of 24 anuran species for the local assemblage. The anuran assemblage in the study area was dominated by species of the family Hylidae, which is the richest anuran family in the Atlantic Forest domain (Rossa-Feres et al. 2017ROSSA-FERES, D.D.C., GAREY, M.V., CARAMASCHI, U., NAPOLI, M.F., NOMURA, F., BISPO, A.A., BRASILEIRO, C.A., THOMÉ, M.T.C., SAWAYA, R.J., CONTE, C.E., CRUZ, C.A.G., NASCIMENTO, L.B., GASPARINI, J.L., ALMEIDA, A.D.P. & HADDAD, C.F.B. 2017. Anfíbios da Mata Atlântica: Lista de espécies, histórico dos estudos, biologia e conservação. In Revisões em Zoologia: Mata Atlântica (E.L.A. Monteiro Filho & C.E. Conte, orgs). UFPR, Curitiba, p.237-314.). Most of the hylids here recorded were found in disturbed areas, and many individuals in five species (Boana polytaenia, Bokermannohyla circumdata, Bokermannohyla sp., Scinax cardosoi, and S. dolloi) were collected in a single marsh. The higher species richness of hylids found in tropical regions of the Americas, compared to that of temperate regions, may reflect the fact that these anurans originated in tropical South America and there has been a long time for species to accumulate through in-situ speciation (Wiens et al. 2006WIENS, J.J., GRAHAM, C.H., MOEN, D.S., SMITH, S.A. & REEDER, T.W. 2006. Evolutionary and ecological causes of the latitudinal diversity gradient in hylid frogs: Treefrog trees unearth the roots of high tropical diversity. Am. Nat. 168(5):579-596.).

Some of the anurans recorded in this study represent new or undetermined species. The anuran most frequently found during our fieldwork was an undetermined species of Ischnocnema of the lactea group (currently in the process of being formally described) that superficially resembles other members of that group such as I. melanopygia Targino, Costa & Carvalho-e-Silva, 2009 and I. concolor Targino, Costa & Carvalho-e-Silva, 2009 (which also occur in the Itatiaia region, but at higher altitudes; Targino et al. 2009TARGINO, M., NOGUEIRA-COSTA, P. & CARVALHO-E-SILVA, S.P. 2009. Two new species of the Ischnocnema lactea species series from Itatiaia highlands, southeastern Brazil (Amphibia, Anura, Brachycephalidae). South Am J Herpetol 4(2):139-150.). We also recorded members of the Ischnocnema guentheri species complex (see Gehara et al. 2013GEHARA, M., CANEDO, C., HADDAD, C.F.B. & VENCES, M. 2013. From widespread to microendemic: Molecular and acoustic analyses show that Ischnocnema guentheri (Amphibia: Brachycephalidae) is endemic to Rio de Janeiro, Brazil. Conserv. Genet. 14(5):973-982.), which also likely represents an undescribed taxonomic entity. Gehara et al. (2013)GEHARA, M., CANEDO, C., HADDAD, C.F.B. & VENCES, M. 2013. From widespread to microendemic: Molecular and acoustic analyses show that Ischnocnema guentheri (Amphibia: Brachycephalidae) is endemic to Rio de Janeiro, Brazil. Conserv. Genet. 14(5):973-982. restricted I. guentheri sensu stricto to the Parque Nacional da Tijuca, municipality of Rio de Janeiro, based on molecular and bioacoustic data. The specimens reported herein may belong to the Ischnocnema aff. guentheri lineage CS3 of Gehara et al. (2013)GEHARA, M., CANEDO, C., HADDAD, C.F.B. & VENCES, M. 2013. From widespread to microendemic: Molecular and acoustic analyses show that Ischnocnema guentheri (Amphibia: Brachycephalidae) is endemic to Rio de Janeiro, Brazil. Conserv. Genet. 14(5):973-982., since the samples from Itatiaia used in their study fell within that lineage in their phylogenetic tree. Bokermannohyla sp. (gr. circumdata) may also represent an undescribed taxon (A.C.C. Lourenço, pers. comm.).

Two other species recorded during our surveys have only been described within the last ten years, being previously confused with other taxa: Proceratophrys mantiqueira (split off from P. melanopogon by Mângia et al. 2014MÂNGIA, S., SANTANA, D.J., CRUZ, C.A.G. & FEIO, R.N. 2014. Taxonomic review of Proceratophrys melanopogon (Miranda-Ribeiro, 1926) with description of four new species (Amphibia, Anura, Odontophrynidae). Bol. Mus. Nac., N.S., Zool. 531:1-33.) and Brachycephalus rotenbergae (split off from B. ephippium by Nunes et al. 2021NUNES, I., GUIMARÃES, C.S., MOURA, P.H.A.G., PEDROZO, M., MOROTI, M.T., CASTRO, L.M., STUGINSKI, D.R. & MUSCAT, E. 2021. Hidden by the name: A new fluorescent pumpkin toadlet from the Brachycephalus ephippium group (Anura: Brachycephalidae). PLoS ONE 16(4):e0244812.). Paratelmatobius mantiqueira was also described relatively recently, based on specimens collected in 1953 (Pombal & Haddad 1999POMBAL, J.P. & HADDAD, C.F.B. 1999. Frogs of the genus Paratelmatobius (Anura: Leptodactylidae) with descriptions of two new species. Copeia 1999(4):1014-1026.), and remained unrecorded in the wild for more than 50 years until one specimen was collected during our 2005 surveys at the studied site in the APA Serra da Mantiqueira (Vrcibradic et al. 2010VRCIBRADIC, D., ARIANI, C.V., VAN SLUYS, M. & ROCHA, C.F.D. 2010. Amphibia, Leptodactylidae, Paratelmatobius mantiqueira Pombal and Haddad, 1999: Distribution extension. Check List 6(1):1-2.). Finally, Scinax dolloi is a taxon that remained in obscurity for more than a century, with no natural populations being associated with this name since its description by Werner (1903)WERNER F. 1903. Neue Reptilien und Batrachier aus dem naturhistorischen Museum in Brussel. Nebst bemerkungen uber einige andere Arten. Zool. Anz. 26:246-253., until its “rediscovery” in 2016 (Santos et al. 2021SANTOS, L.R., SILVA, P.H. & MARTINS, I.A. 2021. Vocalizations of Scinax dolloi (Anura, Hylidae) at its type locality Vila de Maringá, Itatiaia, Rio de Janeiro, Brazil, including bioacoustic comparisons with morphologically similar species. S. Am. J. Herpetol. 19:95-101.). The latter authors reported a population of the species from its type locality in Vila de Maringá, municipality of Itatiaia (see Caramaschi et al. 2013CARAMASCHI U., NUNES I., NASCIMENTO L.B. & CARVALHO JR., R.R. 2013. The type locality of Scinax dolloi (Werner, 1903) (Anura, Hylidae). Zootaxa 3691:299-300.), some 2.6 km in straight line from the locality where the present study was carried out. These cases, together with the aforementioned undescribed/undetermined taxa, highlight how incomplete the knowledge of the Atlantic Forest anuran fauna still is, particularly in the Serra da Mantiqueira region.

The genus Megaelosia has recently been rendered monotypic (for M. goeldii) due to non-monophyly, and a new genus (Phantasmarana) was erected to accommodate the other species, including M. lutzae (Vittorazzi et al. 2021VITTORAZZI, S.E., AUGUSTO-ALVES, G., NEVES-DA-SILVA, D., CARVALHO-E-SILVA, A.M.P.T., RECCO-PIMENTEL, S.M., TOLEDO, L.F., LOURENÇO, L.B. & BRUSCHI, D.P. 2021. Paraphyly in the giant torrent-frogs (Anura: Hylodidae: Megaelosia) and the description of a new genus. Salamandra 57(2):274-284.). Phantasmarana lutzae was described (as Megaelosia lutzae) by Izecksohn & Gouvêa (1987)IZECKSOHN, E. & GOUVÊA, E. 1987 “1985”. Nova espécie de Megaelosia de Itatiaia, Estado do Rio de Janeiro. Arq. Univ. Fed. Rur. Rio de Janeiro 8(1-2):17-22. based on a few adult and juvenile specimens collected between 1949 and 1964 and some tadpoles (not included in the type series) without collection date. This species remained unrecorded in the wild for more than five decades until Vittorazzi et al. (2021)VITTORAZZI, S.E., AUGUSTO-ALVES, G., NEVES-DA-SILVA, D., CARVALHO-E-SILVA, A.M.P.T., RECCO-PIMENTEL, S.M., TOLEDO, L.F., LOURENÇO, L.B. & BRUSCHI, D.P. 2021. Paraphyly in the giant torrent-frogs (Anura: Hylodidae: Megaelosia) and the description of a new genus. Salamandra 57(2):274-284. reported five tadpoles collected at the type locality (Parque Nacional de Itatiaia), which they used in a molecular analysis. Rosa et al. (2003)ROSA, C., AGUIAR-JR., O., GIARETTA, A.A. & RECCO-PIMENTEL, S.M. 2003. Karyotypic variation in the genus Megaelosia (Anura, Leptodactylidae) with the first description of a B chromosome in leptodactylid frogs. Copeia 2003(1):166-174. referred some tadpoles from the municipality of Pindamonhangaba, São Paulo state (ca. 100 km westward from Parque Nacional de Itatiaia), to Megaelosia lutzae, but Vittorazzi et al. (2021)VITTORAZZI, S.E., AUGUSTO-ALVES, G., NEVES-DA-SILVA, D., CARVALHO-E-SILVA, A.M.P.T., RECCO-PIMENTEL, S.M., TOLEDO, L.F., LOURENÇO, L.B. & BRUSCHI, D.P. 2021. Paraphyly in the giant torrent-frogs (Anura: Hylodidae: Megaelosia) and the description of a new genus. Salamandra 57(2):274-284. re-identified those specimens as Phantasmarana jordanensis (Heyer 1983) based on analyses of DNA samples. Thus, P. lutzae is currently known only from the type locality and its surroundings. Our record of P. lutzae tadpoles for the APA Serra da Mantiqueira represents the second confirmed report of the species since its description (but it does not constitute an actual range extension, as our study site is located adjacent to and just outside the limits of the Parque Nacional de Itatiaia). Although no adults were found, the fact that several tadpoles were collected in 2011 in spite of relatively little sampling effort suggests that P. lutzae may not be an uncommon species, at least in our study site. Further sampling is required to gather information on this poorly known species.

The treefrog Scinax cardosoi is another species that remains poorly known (Kirchmeyer et al. 2019KIRCHMEYER, J., RUGGERI, J., GOMES, M.R. & CARVALHO-E-SILVA, S.P. 2019. The tadpole of Scinax cardosoi (Carvalho e Silva and Peixoto, 1991), with description of internal oral morphology and taxonomic considerations (Anura: Hylidae). S. Am. J. Herpetol. 14(3):188-195.), being currently reported from a few isolated localities in the states of Espírito Santo, Minas Gerais, Rio de Janeiro and São Paulo, in southeastern Brazil (Frost 2021FROST, D.R. 2021. Amphibian Species of the World: an Online Reference. Version 6.1. Electronic database acessible at: http://research.amnh.org/herpetology/amphibia/index.html (accessed on 07/02/2021). American Museum of Natural History, New York.
http://research.amnh.org/herpetology/amp... ). Studies updating the geographic distribution of this species (Pinto et al. 2009PINTO, F.C.S., CAMPOS, R.A.R., BRAGA, F.S., QUEIRÓS, F.N.S. & NASCIMENTO, L.B. 2009. Amphibia, Anura, Hylidae, Scinax cardosoi Carvalho-e-Silva and Peixoto, 1991: distribution extension and new state records. Check List 5(4):866-868., Linares et al. 2011LINARES, A.M., BATEMARQUE, M.H.O., MELLO, H.E.S. & NASCIMENTO, L.B. 2011. Amphibia, Anura, Hylidae, Scinax cardosoi (Carvalho-e-Silva and Peixoto, 1991): Distribution extension. Check List 7(4):407-408., Moroti et al. 2017MOROTI, M.T., FERREIRA NETO, J.A., SANTOS, A.C., SOARES, C.P. & MACHADO, I.F. 2017. Amphibia, Anura, Hylidae, Scinax cardosoi (Carvalho-e-Silva & Peixoto, 1991): Distribution extension and new state record for São Paulo, Brazil. Check List 13(1):2042.) have omitted its presence in Visconde de Mauá, in the municipality of Resende, state of Rio de Janeiro, apparently overlooking the fact that one of its paratypes came from that locality (Carvalho-e-Silva & Peixoto 1991CARVALHO-E-SILVA, S.P. & PEIXOTO, O.L. 1991. Duas novas espécies de Ololygon para os Estados do Rio de Janeiro e Espírito Santo (Amphibia, Anura, Hylidae). Rev. Bras. Biol. 51(1):263-270.). The village of Visconde de Mauá is about 6 km distant from our study site. More recently, Kirchmeyer et al. (2019)KIRCHMEYER, J., RUGGERI, J., GOMES, M.R. & CARVALHO-E-SILVA, S.P. 2019. The tadpole of Scinax cardosoi (Carvalho e Silva and Peixoto, 1991), with description of internal oral morphology and taxonomic considerations (Anura: Hylidae). S. Am. J. Herpetol. 14(3):188-195. reported five newly metamorphosed individuals from the Parque Nacional de Itatiaia. Those records, together with those of our study, confirm the presence of S. cardosoi in the region of Itatiaia and its occurrence in other areas within the state of Rio de Janeiro besides its type locality in Teresópolis municipality (Carvalho-e-Silva & Peixoto 1991CARVALHO-E-SILVA, S.P. & PEIXOTO, O.L. 1991. Duas novas espécies de Ololygon para os Estados do Rio de Janeiro e Espírito Santo (Amphibia, Anura, Hylidae). Rev. Bras. Biol. 51(1):263-270.).

The estimated overall density of the local leaf litter frog assemblage was high (18.1 ind/100 m²) when compared to those found for most other sites sampled (also with large quadrats) within the Atlantic Forest in southeastern and southern Brazil (1.4-13.2 ind/100 m²; Giaretta et al. 1997GIARETTA, A.A., SAWAYA, R.J., MACHADO, G., ARAÚJO, M.S., FACURE, K.G., MEDEIROS, H.F. & NUNES, R. 1997. Diversity and abundance of litter frogs at altitudinal sites at Serra do Japi, Southeastern Brazil. Rev. Bras. Zool. 14(2):341-346., 1999GIARETTA, A.A., FACURE, K.G., SAWAYA, R.J., MEYER, J.H.D. & CHEMIN, N. 1999. Diversity and abundance of litter frogs in a montane forest of southeastern Brazil: seasonal and altitudinal changes. Biotropica 31(4):669-674., Rocha et al. 2001ROCHA, C.F.D., VAN SLUYS, M., ALVES, M.A.S., BERGALLO, H.G. & VRCIBRADIC, D. 2001. Estimates of forest floor litter frog communities: A comparison of two methods. Austral Ecol. 26(1):14-21., 2007ROCHA, C.F.D., VRCIBRADIC, D., KIEFER, M.C., ALMEIDA-GOMES, M., BORGES-JR, V.N.T., CARNEIRO, P.C.F., MARRA, R.V., SANTOS, P.A., SIQUEIRA, C.C., GOYANNES-ARAÚJO, P., FERNANDES, C.G.A., RUBIÃO, E.C.N. & VAN SLUYS, M. 2007. A survey of the leaf-litter frog community from an Atlantic forest area (Reserva Ecológica de Guapiaçu) in Rio de Janeiro State, Brazil, with an estimate of frog densities. Trop. Zool. 20(1):99-108., 2011ROCHA C.F.D., VRCIBRADIC D., KIEFER M.C., SIQUEIRA C.C., ALMEIDA-GOMES M., BORGES JÚNIOR V.N.T., HATANO F.H., FONTES A.F., PONTES J.A.L., KLAION T., GIL, L.O. & VAN SLUYS, M. 2011. Parameters from the community of leaf-litter frogs from Estação Ecológica Estadual Paraíso, Guapimirim, Rio de Janeiro State, southeastern Brazil. An Acad Bras Ciênc. 83(4):1259-1268., 2013ROCHA, C.F.D., VRCIBRADIC, D., KIEFER, M.C., ALMEIDA-GOMES, M., BORGES-JR, V.N.T., MENEZES, V.A., ARIANI, C.V., PONTES, J.A.L., GOYANNES-ARAÚJO, P., MARRA, R.V., GUEDES, D.M., SIQUEIRA, C.C. & VAN SLUYS, M. 2013. The leaf-litter frog community from Reserva Rio das Pedras, Mangaratiba, Rio de Janeiro State, Southeastern Brazil: species richness, composition and densities. North-West. J. Zool. 9(1):151-156., Almeida-Gomes et al. 2008ALMEIDA-GOMES, M., VRCIBRADIC, D., SIQUEIRA, C.C., KIEFER, M.C., KLAION, T., ALMEIDA-SANTOS, P., NASCIMENTO, D., ARIANI, C.V., BORGES-JUNIOR, V.N.T., FREITAS-FILHO, R.F., VAN SLUYS, M. & ROCHA, C.F.D. 2008. Herpetofauna of an Atlantic Rainforest area (Morro São João) in Rio de Janeiro State, Brazil. An. Acad. Bras. Ciênc. 80(2):291-300., 2010ALMEIDA-GOMES, M., ALMEIDA-SANTOS, M., GOYANNES-ARAÚJO, P., BORGES-JR, V.N.T., VRCIBRADIC, D., SIQUEIRA, C.C., ARIANI, C.V., DIAS, A.S., SOUZA, V.V., PINTO, R.R., VAN SLUYS, M. & ROCHA, C.F.D. 2010. Anurofauna of an Atlantic Rainforest fragment and its surroundings in northern Rio de Janeiro State, Brazil. Braz. J. Biol. 70(3):871-877., Santos-Pereira et al. 2011SANTOS-PEREIRA, M., CANDATEN, A., MILANI, D., OLIVEIRA, F.B., GARDELIN, J. & ROCHA, C.F.D. 2011. Seasonal variation in the leaf-litter frog community (Amphibia: Anura) from an Atlantic Forest area in the Salto Morato Natural Reserve, southern Brazil. Zoologia 28(6):755-761., Siqueira et al. 2011aSIQUEIRA, C.C., VRCIBRADIC, D., ALMEIDA-GOMES, M., MENEZES, V.A., BORGES-JUNIOR, V.N.T., HATANO, F.H., PONTES, J.A.L., GOYANNES-ARAÚJO, P., GUEDES, D.M., VAN SLUYS, M. & ROCHA, C.F.D. 2011a. Species composition and density estimates of the anurofauna of a site within the northernmost large Atlantic Forest remnant (Parque Estadual do Desengano) in the state of Rio de Janeiro, Brazil. Biota Neotrop. 11(4):131-137., Oliveira et al. 2013OLIVEIRA, J.C.F., PRALON, E., COCO, L., PAGOTTO, R.V. & ROCHA, C.F.D. 2013. Environmental humidity and leaf-litter depth affecting ecological parameters of a leaf-litter frog community in an Atlantic Rainforest area. J. Nat. Hist. 47(31-32):2115-2124.). Exceptions are two localities in the municipality of Cachoeiras de Macacu, state of Rio de Janeiro, that yielded comparable (17.1 ind/100 m²; Siqueira et al. 2009SIQUEIRA, C.C., VRCIBRADIC, D., ALMEIDA-GOMES, M., BORGES-JR, V.N.T., ALMEIDA-SANTOS, P., ALMEIDA-SANTOS, M., ARIANI, C.V., GUEDES, D.M., GOYANNES-ARAÚJO, P., DORIGO, T.A., VAN SLUYS, M. & ROCHA, C.F.D. 2009. Density and richness of leaf litter frogs (Amphibia: Anura) of an Atlantic Rainforest area in the Serra dos Órgãos, Rio de Janeiro State, Brazil. Zoologia 26(1):97-102.) or higher estimates (24.6 ind/100 m²; Siqueira et al. 2014SIQUEIRA, C.C., VRCIBRADIC, D., COSTA, P.N., MARTINS, A.R., DANTAS, L.F., GOMES, V.L.R., BERGALLO, H. G. & ROCHA, C.F.D. 2014. Environmental parameters affecting the structure of leaf-litter frog (Amphibia: Anura) communities in tropical forests: A case study from an Atlantic Rainforest area in southeastern Brazil. Zoologia 31(2):147-152.). However, the high frog density observed in this study reflects the high local abundance of Ischnocnema sp. (gr. lactea), since 88% of the frogs found in plots represented that species. Indeed, this small direct-developing anuran numerically dominated the anuran assemblage at the studied site in the APA Serra da Mantiqueira (as also evidenced by the VES methodology). Several other plot-sampling studies in Atlantic Forest areas have also evidenced a clear dominance of species with direct development of terrestrial eggs in the local leaf-litter frog assemblages (e.g., Giaretta et al. 1997GIARETTA, A.A., SAWAYA, R.J., MACHADO, G., ARAÚJO, M.S., FACURE, K.G., MEDEIROS, H.F. & NUNES, R. 1997. Diversity and abundance of litter frogs at altitudinal sites at Serra do Japi, Southeastern Brazil. Rev. Bras. Zool. 14(2):341-346., 1999GIARETTA, A.A., FACURE, K.G., SAWAYA, R.J., MEYER, J.H.D. & CHEMIN, N. 1999. Diversity and abundance of litter frogs in a montane forest of southeastern Brazil: seasonal and altitudinal changes. Biotropica 31(4):669-674., Rocha et al. 2001ROCHA, C.F.D., VAN SLUYS, M., ALVES, M.A.S., BERGALLO, H.G. & VRCIBRADIC, D. 2001. Estimates of forest floor litter frog communities: A comparison of two methods. Austral Ecol. 26(1):14-21., 2007ROCHA, C.F.D., VRCIBRADIC, D., KIEFER, M.C., ALMEIDA-GOMES, M., BORGES-JR, V.N.T., CARNEIRO, P.C.F., MARRA, R.V., SANTOS, P.A., SIQUEIRA, C.C., GOYANNES-ARAÚJO, P., FERNANDES, C.G.A., RUBIÃO, E.C.N. & VAN SLUYS, M. 2007. A survey of the leaf-litter frog community from an Atlantic forest area (Reserva Ecológica de Guapiaçu) in Rio de Janeiro State, Brazil, with an estimate of frog densities. Trop. Zool. 20(1):99-108., 2013ROCHA, C.F.D., VRCIBRADIC, D., KIEFER, M.C., ALMEIDA-GOMES, M., BORGES-JR, V.N.T., MENEZES, V.A., ARIANI, C.V., PONTES, J.A.L., GOYANNES-ARAÚJO, P., MARRA, R.V., GUEDES, D.M., SIQUEIRA, C.C. & VAN SLUYS, M. 2013. The leaf-litter frog community from Reserva Rio das Pedras, Mangaratiba, Rio de Janeiro State, Southeastern Brazil: species richness, composition and densities. North-West. J. Zool. 9(1):151-156., Almeida-Gomes et al. 2008ALMEIDA-GOMES, M., VRCIBRADIC, D., SIQUEIRA, C.C., KIEFER, M.C., KLAION, T., ALMEIDA-SANTOS, P., NASCIMENTO, D., ARIANI, C.V., BORGES-JUNIOR, V.N.T., FREITAS-FILHO, R.F., VAN SLUYS, M. & ROCHA, C.F.D. 2008. Herpetofauna of an Atlantic Rainforest area (Morro São João) in Rio de Janeiro State, Brazil. An. Acad. Bras. Ciênc. 80(2):291-300., 2010ALMEIDA-GOMES, M., ALMEIDA-SANTOS, M., GOYANNES-ARAÚJO, P., BORGES-JR, V.N.T., VRCIBRADIC, D., SIQUEIRA, C.C., ARIANI, C.V., DIAS, A.S., SOUZA, V.V., PINTO, R.R., VAN SLUYS, M. & ROCHA, C.F.D. 2010. Anurofauna of an Atlantic Rainforest fragment and its surroundings in northern Rio de Janeiro State, Brazil. Braz. J. Biol. 70(3):871-877., Siqueira et al. 2009SIQUEIRA, C.C., VRCIBRADIC, D., ALMEIDA-GOMES, M., BORGES-JR, V.N.T., ALMEIDA-SANTOS, P., ALMEIDA-SANTOS, M., ARIANI, C.V., GUEDES, D.M., GOYANNES-ARAÚJO, P., DORIGO, T.A., VAN SLUYS, M. & ROCHA, C.F.D. 2009. Density and richness of leaf litter frogs (Amphibia: Anura) of an Atlantic Rainforest area in the Serra dos Órgãos, Rio de Janeiro State, Brazil. Zoologia 26(1):97-102., 2011aSIQUEIRA, C.C., VRCIBRADIC, D., ALMEIDA-GOMES, M., MENEZES, V.A., BORGES-JUNIOR, V.N.T., HATANO, F.H., PONTES, J.A.L., GOYANNES-ARAÚJO, P., GUEDES, D.M., VAN SLUYS, M. & ROCHA, C.F.D. 2011a. Species composition and density estimates of the anurofauna of a site within the northernmost large Atlantic Forest remnant (Parque Estadual do Desengano) in the state of Rio de Janeiro, Brazil. Biota Neotrop. 11(4):131-137., 2014SIQUEIRA, C.C., VRCIBRADIC, D., COSTA, P.N., MARTINS, A.R., DANTAS, L.F., GOMES, V.L.R., BERGALLO, H. G. & ROCHA, C.F.D. 2014. Environmental parameters affecting the structure of leaf-litter frog (Amphibia: Anura) communities in tropical forests: A case study from an Atlantic Rainforest area in southeastern Brazil. Zoologia 31(2):147-152.), and this has also been observed in Amazonian (e.g., Rodriguez 1992RODRIGUEZ, L. 1992. Structure et organisation du peuplement d’anoures de Cocha Cashu, Parc National Manu, Amazonie péruvienne. Rev. Ecol. Terre-Vie 47(2):151-197., Gascon 1996GASCON, C. 1996. Amphibian litter fauna and river barriers in flooded and non-flooded Amazonian rainforests. Biotropica 28(1):136-140.) and Central American forest areas (e.g., Scott 1976SCOTT JR., N.J. 1976. The abundance and diversity of the herpetofauna of tropical forest litter. Biotropica 8(1):41-58., Toft 1981TOFT, C.A. 1981. Feeding ecology of Panamanian litter anurans: patterns in diet and foraging mode. J. Herpetol 15(2):139-144., Liebermann 1986LIEBERMAN, S.S. 1986. Ecology of the leaf litter herpetofauna of a Neotropical rainforest: La Selva, Costa Rica. Acta Zool. Mex. 15(1):1-71., Fauth et al. 1989FAUTH, J.E., CROTHER, B.I. & SLOWINSKI, J.B. 1989. Elevational patterns of species richness, evenness and abundance of the Costa Rican leaf-litter herpetofauna. Biotropica 21(2):178-185., Hofer & Bersier 2001HOFER, U. & BERSIER, L.F. 2001. Herpetofaunal diversity and abundance in tropical upland forests of Cameroon and Panama. Biotropica 33(1):142-152.). Direct-developing anurans may be more widely distributed within the forest due to their independence of water bodies for reproduction (Haddad & Prado 2005HADDAD, C.F.B. & PRADO, C.P.A. 2005. Reproductive modes in frogs and their unexpected diversity in the Atlantic Forest of Brazil. BioScience 55(3):207-217.), though other factors such as rates of dehydration/rehydration (Dabés et al. 2012) and a generalist diet (Alves-dos-Santos et al. 2021ALVES-DOS-SANTOS, T.F.S.S.A., FORTI, L.R. & NAPOLI, M.F. 2021. Feeding habits of the Robber Frog Pristimantis paulodutrai (Bokermann, 1975) in northeastern Brazil. Biota Neotrop. 21(2):e20201098. https://doi.org/10.1590/1676-0611-BN2020-1098 (last access on 22/12/2021)
https://doi.org/10.1590/1676-0611-BN2020... ) may also play a role.

Cumulative and rarefaction curves using data from VES method indicated that sampling effort was almost enough to obtain a representative sample of the local anuran fauna, with a tendency toward stabilization, despite plot methodology falling short of reaching the asymptote. Plot sampling is useful for sampling anurans typical of forest floor leaf litter in Atlantic Forest areas (Giaretta et al. 1997GIARETTA, A.A., SAWAYA, R.J., MACHADO, G., ARAÚJO, M.S., FACURE, K.G., MEDEIROS, H.F. & NUNES, R. 1997. Diversity and abundance of litter frogs at altitudinal sites at Serra do Japi, Southeastern Brazil. Rev. Bras. Zool. 14(2):341-346., 1999GIARETTA, A.A., FACURE, K.G., SAWAYA, R.J., MEYER, J.H.D. & CHEMIN, N. 1999. Diversity and abundance of litter frogs in a montane forest of southeastern Brazil: seasonal and altitudinal changes. Biotropica 31(4):669-674., Rocha et al. 2001ROCHA, C.F.D., VAN SLUYS, M., ALVES, M.A.S., BERGALLO, H.G. & VRCIBRADIC, D. 2001. Estimates of forest floor litter frog communities: A comparison of two methods. Austral Ecol. 26(1):14-21., 2007ROCHA, C.F.D., VRCIBRADIC, D., KIEFER, M.C., ALMEIDA-GOMES, M., BORGES-JR, V.N.T., CARNEIRO, P.C.F., MARRA, R.V., SANTOS, P.A., SIQUEIRA, C.C., GOYANNES-ARAÚJO, P., FERNANDES, C.G.A., RUBIÃO, E.C.N. & VAN SLUYS, M. 2007. A survey of the leaf-litter frog community from an Atlantic forest area (Reserva Ecológica de Guapiaçu) in Rio de Janeiro State, Brazil, with an estimate of frog densities. Trop. Zool. 20(1):99-108., 2013ROCHA, C.F.D., VRCIBRADIC, D., KIEFER, M.C., ALMEIDA-GOMES, M., BORGES-JR, V.N.T., MENEZES, V.A., ARIANI, C.V., PONTES, J.A.L., GOYANNES-ARAÚJO, P., MARRA, R.V., GUEDES, D.M., SIQUEIRA, C.C. & VAN SLUYS, M. 2013. The leaf-litter frog community from Reserva Rio das Pedras, Mangaratiba, Rio de Janeiro State, Southeastern Brazil: species richness, composition and densities. North-West. J. Zool. 9(1):151-156., Siqueira et al. 2009SIQUEIRA, C.C., VRCIBRADIC, D., ALMEIDA-GOMES, M., BORGES-JR, V.N.T., ALMEIDA-SANTOS, P., ALMEIDA-SANTOS, M., ARIANI, C.V., GUEDES, D.M., GOYANNES-ARAÚJO, P., DORIGO, T.A., VAN SLUYS, M. & ROCHA, C.F.D. 2009. Density and richness of leaf litter frogs (Amphibia: Anura) of an Atlantic Rainforest area in the Serra dos Órgãos, Rio de Janeiro State, Brazil. Zoologia 26(1):97-102., 2011aSIQUEIRA, C.C., VRCIBRADIC, D., ALMEIDA-GOMES, M., MENEZES, V.A., BORGES-JUNIOR, V.N.T., HATANO, F.H., PONTES, J.A.L., GOYANNES-ARAÚJO, P., GUEDES, D.M., VAN SLUYS, M. & ROCHA, C.F.D. 2011a. Species composition and density estimates of the anurofauna of a site within the northernmost large Atlantic Forest remnant (Parque Estadual do Desengano) in the state of Rio de Janeiro, Brazil. Biota Neotrop. 11(4):131-137.) and, despite our effort being not enough using this method, members of that guild of anurans were also recorded by VES, adding four leaf litter species to our list that were not recorded in plots (e.g., Ischnocnema juipoca, Paratelmatobius mantiqueira, Odontophrynus americanus and Proceratophrys boiei). Moreover, our additional effort with non-standardized samplings in 2010 and 2011, added two additional species that live in the forest floor (Brachycephalus rotenbergae and Physalaemus cuvieri), besides five more anuran taxa with arboreal or semi-aquatic habits (Boana pardalis, Dendropsophus minutus, Leptodactylus latrans, Phantasmarana lutzae and Phasmahyla cochranae). The exception is P. lutzae, which was sampled only as tadpoles, through the use of funnel traps (the only other species recorded only in larval stage, Phasmahyla cochranae, was sampled as a tadpole found during haphazard visual searching). Thus, our data showed that visual searches were sufficient to produce a representative sample of the anuran fauna of our study site, with pitfall traps and plot sampling providing no additional species records. However, in some other short-term inventories employing multiple sampling methods in tropical areas, plot and/or pitfall trap sampling added anuran species that had not been recorded by standardized visual searches or occasional encounters (Almeida-Gomes et al. 2008ALMEIDA-GOMES, M., VRCIBRADIC, D., SIQUEIRA, C.C., KIEFER, M.C., KLAION, T., ALMEIDA-SANTOS, P., NASCIMENTO, D., ARIANI, C.V., BORGES-JUNIOR, V.N.T., FREITAS-FILHO, R.F., VAN SLUYS, M. & ROCHA, C.F.D. 2008. Herpetofauna of an Atlantic Rainforest area (Morro São João) in Rio de Janeiro State, Brazil. An. Acad. Bras. Ciênc. 80(2):291-300., Ribeiro-Júnior et al. 2008RIBEIRO-JÚNIOR, M.A., GARDNER, T.A. & ÁVILA-PIRES. T.C.S. 2008. Evaluating the effectiveness of herpetofaunal sampling techniques across a gradient of habitat change in a tropical forest landscape. J. Herpetol. 42(4):733-749., Siqueira et al. 2009SIQUEIRA, C.C., VRCIBRADIC, D., ALMEIDA-GOMES, M., BORGES-JR, V.N.T., ALMEIDA-SANTOS, P., ALMEIDA-SANTOS, M., ARIANI, C.V., GUEDES, D.M., GOYANNES-ARAÚJO, P., DORIGO, T.A., VAN SLUYS, M. & ROCHA, C.F.D. 2009. Density and richness of leaf litter frogs (Amphibia: Anura) of an Atlantic Rainforest area in the Serra dos Órgãos, Rio de Janeiro State, Brazil. Zoologia 26(1):97-102., Rocha et al. 2011ROCHA C.F.D., VRCIBRADIC D., KIEFER M.C., SIQUEIRA C.C., ALMEIDA-GOMES M., BORGES JÚNIOR V.N.T., HATANO F.H., FONTES A.F., PONTES J.A.L., KLAION T., GIL, L.O. & VAN SLUYS, M. 2011. Parameters from the community of leaf-litter frogs from Estação Ecológica Estadual Paraíso, Guapimirim, Rio de Janeiro State, southeastern Brazil. An Acad Bras Ciênc. 83(4):1259-1268.). Nevertheless, pitfall traps are generally considered to have low cost-benefit for simple short-term surveys of tropical amphibians (Cechin & Martins 2000CECHIN, S.Z. & MARTINS, M. 2000. Eficiência de armadilhas de queda (pitfall traps) em amostragens de anfíbios e répteis no Brasil. Rev. Bras. Zool. 17(3):729-740., Rödel & Ernst 2004RÖDEL, M-O. & ERNST, R. 2004. Measuring and monitoring amphibian diversity in tropical forests. I. An evaluation of methods with recommendations for standarization. Ecotropica 10(1):1-14.), which is supported by the results of our study.

Phantasmarana lutzae and Scinax dolloi are the only species recorded in our study that are currently known only from the state of Rio de Janeiro and are possibly endemic to the Itatiaia region (Frost 2021FROST, D.R. 2021. Amphibian Species of the World: an Online Reference. Version 6.1. Electronic database acessible at: http://research.amnh.org/herpetology/amphibia/index.html (accessed on 07/02/2021). American Museum of Natural History, New York.
http://research.amnh.org/herpetology/amp... , Vittorazzi et al. 2021VITTORAZZI, S.E., AUGUSTO-ALVES, G., NEVES-DA-SILVA, D., CARVALHO-E-SILVA, A.M.P.T., RECCO-PIMENTEL, S.M., TOLEDO, L.F., LOURENÇO, L.B. & BRUSCHI, D.P. 2021. Paraphyly in the giant torrent-frogs (Anura: Hylodidae: Megaelosia) and the description of a new genus. Salamandra 57(2):274-284.), though Santos et al. (2021)SANTOS, L.R., SILVA, P.H. & MARTINS, I.A. 2021. Vocalizations of Scinax dolloi (Anura, Hylidae) at its type locality Vila de Maringá, Itatiaia, Rio de Janeiro, Brazil, including bioacoustic comparisons with morphologically similar species. S. Am. J. Herpetol. 19:95-101. suggested that S. dolloi may occur more widely. Neves et al. (2018)NEVES, M.O., PEREIRA, E.A., SUGAI, J.L.M.M., ROCHA, S.B., FEIO, R.N. & SANTANA, D.J. 2018. Distribution pattern of anurans from three mountain complexes in southeastern Brazil and their conservation implications. An. Acad. Bras. Ciênc. 90(2):1611-1623. considered the Serra do Itatiaia as the area of highest amphibian endemicity rate within the Mantiqueira/Espinhaço/Canastra mountain complex in southeastern Brazil, with nine endemic anuran species (Scinax dolloi was not mentioned, possibly because it was a species of problematic identity at the time). Among the undescribed species recorded in the APA Serra da Mantiqueira survey, at least one, Ischnocnema sp. (gr. lactea), may be endemic to the Itatiaia region, and it is possible that the degree of anuran endemicity in the area is still underestimated.

Regarding their conservation status, most species from the APA Serra da Mantiqueira are listed under “Least Concern” (LC) by the IUCN (2021)IUCN 2021. The IUCN red list of threatened species. Version 2021-3. Electronic database accessible at: https://www.iucnredlist.org/ (last access on 17/06/2021)
https://www.iucnredlist.org/... , though the status of Brachycephalus rotenbergae, Proceratophrys mantiqueira, Scinax dolloi, and the undescribed species have not been assessed. Phantasmarana lutzae (Carvalho-e-Silva & Carvalho-e-Silva 2004CARVALHO-E-SILVA, S.P. & CARVALHO-E-SILVA, A.M.T. 2004. Megaelosia lutzae. The IUCN Red List of Threatened Species 2004:e.T57183A11582381. https://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T57183A11582381.en (last access on 17/06/2021)
https://dx.doi.org/10.2305/IUCN.UK.2004.... ) and Paratelmatobius mantiqueira (Nascimento & Garcia 2004NASCIMENTO, L.B. & GARCIA, P. 2004. Paratelmatobius mantiqueira. The IUCN Red List of Threatened Species 2004: e.T57196A11586987. https://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T57196A11586987.en (last access on 17/06/2021)
https://dx.doi.org/10.2305/IUCN.UK.2004.... ) are both classified as “Data Deficient” (DD) with population trends classified as “unknown” and “decreasing”, respectively (IUCN 2021IUCN 2021. The IUCN red list of threatened species. Version 2021-3. Electronic database accessible at: https://www.iucnredlist.org/ (last access on 17/06/2021)
https://www.iucnredlist.org/... ). None of the species found in the present survey is listed as threatened in the Red Book of Threatened Brazilian Fauna (ICMBio/MMA, 2018ICMBio/MMA. 2018. Livro Vermelho de Fauna Brasileira Ameaçada de Extinção. v. 5. Anfíbios. ICMBio, Brasília.) either. Threatened species are usually the focus of conservation efforts and funding programs, but little attention is given to species considered “Data-Deficient”, contradicting the IUCN’s recommendation to give them the same degree of attention as threatened taxa (Pimenta et al. 2005PIMENTA, B.V.S., HADDAD, C.F.B., NASCIMENTO, L.B., CRUZ, C.A.G. & POMBAL JR., J.P. 2005. Comment on “Status and trends of amphibian declines and extinctions worldwide”. Science 309:1999., Siqueira et al. 2011bSIQUEIRA, C.C., VRCIBRADIC, D., DORIGO, T.A. & ROCHA, C.F.D. 2011b. Anurans from two high-elevation areas of Atlantic Forest in the state of Rio de Janeiro, Brazil. Zoologia 28(4):457-464.).

Lists of species present in a given area constitute a fundamental tool for the development and management of effective conservation interventions. Our inventory at an area within the APA Serra da Mantiqueira adds to the body of knowledge on the amphibian diversity of high-elevation forests in an Atlantic Forest area, which is still insufficiently known.

Supplementary Material

The following online material is available for this article:

Appendix I - Voucher specimens of species collected in the present study deposited at the Museu Nacional, Universidade Federal do Rio de Janeiro (MNRJ), Rio de Janeiro municipality, Rio de Janeiro state, Brazil.

Acknowledgments

This study was financed by Critical Ecosystem Partnership Fund - CEPF, Conservação Internacional and Aliança para a Conservação da Mata Atlântica. CFDR (Processes 302974/2015-6, 424473/2016 and 304375/2020-9) received financial support from the Conselho Nacional de Desenvolvimento Científico e Tecnológico - CNPq, and from Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) through the “Programa Cientistas do Nosso Estado” (Process E-26/202.803/2018). CCS received a post-doctoral fellowship from FAPERJ (Processes E-26/202.477/2019 and E-26/202.377/2021) and now receives a scholarship from the Programa de Apoio à Docência (PAPD-UERJ). We are especially thankful to T. Nakano and G. Huber (Marimbondo farm) for many facilities, logistical support, and for permit to work in the area. Our thanks to many colleagues for field assistance in different moments of the study, especially H.G. Bergallo and M.A.S. Alves. We also thank J.P. Pombal Jr., C.A.G. Cruz, U. Caramaschi, and A.C.C. Lourenço from Museu Nacional (MNRJ) and I.A. Martins of Universidade de Taubaté for their help with the identification of frogs.

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