Another piece in the puzzle: A new species of Phyllodytes Wagler, 1830 (Anura, Hylidae) from the Atlantic Forest of southern Bahia, northeastern Brazil
Abstract
We describe Phyllodytes gravata sp. nov., a new species of bromeligenous treefrog from the Atlantic Forest of southern Bahia, Brazil. The new species is characterised by its small body size (males 19.5–22.0 mm snout-vent length; female 23.5 mm), a rounded snout with an apical tubercle, an inconspicuous row of tubercles along the tarsus, with a single prominent tubercle at the tibiotarsal articulation, a dorsolateral brown stripe, and an advertisement call composed of long series of pulsed notes (22–34 notes/call; call duration 5.2–7.3 s; dominant frequency 2.75–3.83 kHz). The species is currently known from only two nearby localities within the Central Corridor of the Atlantic Forest (CCAF), in the municipality of Porto Seguro, district of Trancoso, Bahia, a major tourist destination, where it inhabits bromeliads in sandy-soil ecosystems known as mussunungas. Its apparently microendemic distribution and strict association with bromeliads render it vulnerable to several anthropogenic threats, such as bromeliad harvesting, increasing tourism, urbanisation, and agriculture. This discovery underscores the persistent hidden diversity of the CCAF and reinforces the urgent need to protect its highly special microhabitats.
References
Alarcón-Elbal P. M., Sosa M.A.R., Tiburcio J.C.D., Sevilla J.R. & Ee B.W. 2024. Diversity of Aquatic Macroinvertebrates Associated with Bromeliaceae Plants in Jarabacoa, Dominican Republic. Caribbean Journal of Science 54 (2): 369–406. https://doi.org/10.18475/cjos.v54i2.a17
Albertim K.M., Andrade E.V.E., Melo I.V.C. & Moura G.J.B. 2010. Anurans and lizards associated to bromeliads in Atlantic Rainforest fragment in Pernambuco State, Northeastern Brazil. Sitientibus Série Ciências Biológicas 10 (2): 289–298. https://doi.org/10.13102/scb7976
Alves-Silva R. & Silva H.R. 2009. Life in bromeliads: reproductive behaviour and the monophyly of the Scinax perpusillus species group (Anura: Hylidae). Journal of Natural History 43 (3–4): 205–217. https://doi.org/10.1080/00222930802568808
Arango A.A., Guzmán G.I. & González A. 2011. Diversidad y abundancia de la Artropofauna en bromelias de Bosques de Manglar de la Bahía de Buenaventura (Valle, Colombia). Boletín del Museo de Entomología de la Universidad del Valle 12 (1): 1–11.
Bastos D.F.O. & Zina J. 2022. Amphibian fauna in an ecotonal and mountainous area in south-central Bahia State, northeastern Brazil. Herpetology Notes 15: 365–376.
Blotto B.L., Lyra M.L., Cardoso M.C.S., Rodrigues M.T., Dias I.R., Marciano-Jr E., Vechio F.D., Orrico V.G.D., Brandão R.A., Assis C.L., Lantyer-Silva A.S.F., Rutherford M.G., Gagliardi-Urrutia G., Solé M., Baldo D., Nunes I., Cajade R., Torres A., Grant T., Jungfer K.H., Silva H.R., Haddad C.F.B. & Faivovich J. 2021. The phylogeny of the Casque headed Treefrogs (Hylidae: Hylinae: Lophyohylini). Cladistics 37 (1): 36–72. https://doi.org/10.1111/cla.12409
Bokermann W.C.A. 1966. O gênero Phyllodytes Wagler, 1830 (Anura, Hylidae). Anais da Academia Brasileira de Ciências 38: 335–344.
Borges R.L., Gaem P.H. & Roque N. 2020. A new species of Tocoyena (Rubiaceae, Gardenia) from the Brazilian Atlantic Forest. Phytotaxa 470 (3): 243–248. https://doi.org/10.11646/phytotaxa.470.3.4
Campos T.F., Lima M.G., Nascimento A.C. & Santos E.M. 2014. Larval morphology and advertisement call of Phyllodytes acuminatus Bokermann, 1966 (Anura: Hylidae) from northeastern Brazil. Zootaxa 3779: 93–100. https://doi.org/10.11646/zootaxa.3779.1.10
Caramaschi U. & Peixoto O.L. 2004. A new species of Phyllodytes (Anura: Hylidae) from the state of Sergipe, northeastern Brazil. Amphibia-Reptilia 25: 1–8.
Caramaschi U., Silva H.R. & Britto-Pereira M.C. 1992. A new species of Phyllodytes (Anura: Hylidae) from southern Bahia, Brazil. Copeia 1992 (1): 187–191.
Carilo-Filho L.M., de Carvalho B.T., Azevedo B.K.A., Gutiérrez-Pesquera L.M., Mira-Mendes C.V., Solé M. & Orrico V.G.D. 2021. Natural history predicts patterns of thermal vulnerability in amphibians from the Atlantic Rainforest of Brazil. Ecology and Evolution 11: 16462–16472. https://doi.org/10.1002/ece3.7961
Cruz C.A.G., Feio R.F. & Cardoso M.C.S. 2007. Description of a new species of Phyllodytes Wagler, 1830 (Anura, Hylidae) from the Atlantic Rain Forest of the states of Minas Gerais and Bahia, Brazil. Arquivos do Museu Nacional, Rio de Janeiro 64: 321–324.
Cruz D., Marciano J.R.E. & Napoli M.F. 2014. Advertisement and courtship calls of Phyllodytes wuchereri (Peters, 1873) (Anura: Hylidae). Zootaxa 3774: 97–100. https://doi.org/10.11646/zootaxa.3774.1.8
Dias I.R., Novaes-e-Fagundes G., Neto A.M., Zina J., Garcia C., Recoder R.S., Vechio F.D., Rodrigues M.T. & Solé M. 2020. A new large canopy-dwelling species of Phyllodytes Wagler, 1930 (Anura, Hylidae) from the Atlantic Forest of the state of Bahia, Northeastern Brazil. PeerJ 8: e8642. https://doi.org/10.7717/peerj.8642
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. American Museum of Natural History 294: 1–240. http://dx.doi.org/10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2
Faivovich J., Gasparini J.L. & Haddad C.F.B. 2010. A new species of the Scinax perpusillus group (Anura: Hylidae) from Espírito Santo, Brazil. Copeia 2010 (1): 97–102.
Ferreira R.B., Schneider J.A.P. & Teixeira R.L. 2012. Diet, fecundity and use of bromeliads by Phyllodytes luteolus (Anura: Hylidae) in southeastern Brazil. Journal of Herpetology 46: 19–24. https://doi.org/10.1670/09-040
Ferreira R.B., Faivovich J., Beard K.H. & Pombal Jr. J.P. 2015. The first bromeligenous species of Dendropsophus (Anura: Hylidae) from Brazil’s Atlantic Forest. PLoS ONE 10 (12): e0142893. https://doi.org/10.1371/journal.pone.0142893
Forti L.R., Foratto R.M., Marquez R., Pereira V.R. & Toledo L.F. 2018. Current knowledge on bioacoustics of the subfamily Lophyohylinae (Hylidae, Anura) and description of Ocellated treefrog Itapotihyla langsdorffii vocalizations. PeerJ 6:e4813. https://doi.org/10.7717/peerj.4813
Fundação SOS Mata Atlântica, INPE. 2021. Atlas dos remanescentes florestais da Mata Atlântica: período 2019/2020, relatório técnico. São Paulo: Fundação SOS Mata Atlântica, São Paulo.
Godoy F.I. & Piratelli A.J. 2020. Avifauna associada a muçunungas: Composição e uso do habitat. Oecologia Australis 25 (1): 7–21. https://doi.org/10.4257/oeco.2021.2501.02
Gondim-Silva F.A.T., Andrade A.R.S., Abreu R.O., Nascimento J.S., Corrêa G.P., Menezes L., Trevisan C.C., Camargo S.S. & Napoli M.F. 2016. Composition and diversity of anurans in the Restinga of the Conde municipality, northern coast of the state of Bahia, northeastern Brazil. Biota Neotropica 16 (3): e20160157. http://dx.doi.org/10.1590/1676-0611-BN-2016-0157
Hammer Ø., Harper D.A.T. & Ryan P.D. 2001. PAST: paleontological statistics software package for education and data analysis. Palaeontologia Electronica 4 (1): 4
Heyer W.R., Rand A.S., Cruz C.A.G., Peixoto O.L. & Nelson C.E. 1990. Frogs of Boracéia. Arquivos de Zoologia 31: 231–410.
Jorge J.S., Sales R.F.D., Silva V.T.C. & Freire E.M.X. 2021. Lizards and bromeliads in the Neotropics: literature review and relevance of this association to conservation. Symbiosis 84 (2): 119–130. https://doi.org/10.1007/s13199-021-00779-z
Juncá F.A., Napoli M.F., Cedraz J. & Nunes I. 2012. Acoustic characteristics of the advertisement and territorial calls of Phyllodytes tuberculosus Bokermann, 1966 (Amphibia: Anura: Hylidae). Zootaxa 3506: 87–88. https://doi.org/10.11646/zootaxa.3506.1.7
Koh L.P., Dunn R.R., Sodhi N.S., Colwell R.K., Proctor H.C. & Smith V.S. 2004. Species coextinctions and the biodiversity crisis. Science 305: 1632–1634. https://www.doi.org/10.1126/science.1101101
Köhler J., Jansen M., Rodríguez A., Kok P.J.R., Toledo L.F., Emmrich M., Glaw F., Haddad C.F.B., Rödel M.O. & Vences M. 2017. The use of bioacoustics in anuran taxonomy: theory, terminology, methods and recommendations for best practice. Zootaxa 4251: 1–124. https://doi.org/10.11646/zootaxa.4251.1.1
Kok P.J.R. & Kalamandeen M. 2008. Introduction to the taxonomy of the amphibians of Kaieteur National Park, Guyana. Abc Taxa 5: 1–278.
Laviski B.F.S., Monteiro E.M., Pinho L.C., Baptista R.L.C., Mayhé-Nunes A.J., Racca-Filho F. & Nunes-Freitas A.F. 2021. Bromeliad habitat regulates the richness of associated terrestrial and aquatic fauna. Austral Ecology 46 (5): 860–870. https://doi.org/10.1111/aec.13033
Lima M.G., Lingnau R. & Skuk G.O. 2008. The Advertisement call of Phyllodytes edelmoi (Anura, Hylidae). South American Journal of Herpetology 3:118-121
Lima R.G.V.N., Lima L.F. & Zickel C.S. 2021. Chrysophyllum lancisepalum (Sapotaceae, Crysophylloideae): a new endemic species from the Brazilian Atlantic forest. Phytotaxa 478 (1): 131–140. https://doi.org/10.11646/phytotaxa.478.1.9
Lopez L.C.S., Alves R.R.D.N. & Rios R.I. 2009. Micro-environmental factors and the endemism of bromeliad aquatic Fauna. Hydrobiologia 625: 151–156. https://doi.org/10.1007/s10750-009-9704-1
Magalhães F.D.M., Juncá F.A. & Garda A.A. 2015. Tadpole and vocalisations of Phyllodytes wuchereri (Anura: Hylidae) from Bahia, Brazil. Salamandra 51: 83–90.
Mageski M.M., Ferreira R.B., Costa L.C., Jesus P.R. & Ferreira P.D. 2017. Frog assemblage associated with bromeliads in a sandy coastal plain in the state of Espírito Santo, southeastern Brazil. Papéis Avulsos de Zoologia 57 (34): 445–449. https://doi.org/10.11606/0031-1049.2017.57.34
Marciano-Jr E., Lantyer-Silva A.S. & Solé M. 2017. A new species of Phyllodytes Wagler, 1830 (Anura, Hylidae) from the Atlantic Forest of southern Bahia, Brazil. Zootaxa 4238: 135–142. https://doi.org/10.11646/zootaxa.4238.1.11
Marques de Abreu L., Guerra V. & Solé M. 2024. Influence of conspecifics and road noise on the properties of the advertisement call of Phyllodytes luteolus (Anura, Hylidae). Ecology and Evolution 14: e11261. https://doi.org/10.1002/ece3.11261
Martini A.M.Z., Fiaschi P., Amorim A. & Paixão J.F. 2007. A hot-point within a hot-spot: a high diversity site in Brazil’s Atlantic Forest. Biodiversity and Conservation 16: 3111–3128. https://doi.org/10.1007/s10531-007-9166-6
Martins A., Pontes R., Mattedi C., Murta-Fonseca R.A., Fratani J., Ramos L.O., Brandão A.L.R., Maciel D.B. & Pinto R.R. 2019. Herpetofauna community from coastal restinga remnants in Northeast Rio de Janeiro state, Brazil. Journal of Coastal Conservation 23: 1019–1037. https://doi.org/10.1007/s11852-019-00708-9
Mira-Mendes C.B., Ruas D.S., Oliveira R.M., Castro I.M., Dias I.R., Baumgarten J.E., Juncá F.A. & Solé M. 2018. Amphibians of the Reserva Ecológica Michelin: a high diversity site in the lowland Atlantic Forest of southern Bahia, Brazil. ZooKeys 753:1–21. https://doi.org/10.3897/zookeys.753.21438
Nunes I., Santiago R.S. & Juncá F.A. 2007. Advertisement calls of four hylid frogs from the state of Bahia, northeastern Brazil (Amphibia, Anura, Hylidae). South American Journal of Herpetology 2: 89–96. https://doi.org/10.2994/1808-9798(2007)2[89:ACOFHF]2.0.CO;2
Oliveira F.B. & Navas C.A. 2004. Plant selection and seasonal patterns of vocal activity in two populations of the bromeligen treefrog Scinax perpusillus (Anura, Hylidae). Journal of Herpetology 38: 331–339. https://doi.org/10.1670/205-03A
Oliveira J.C.F., Winck G.R., Pereira-Ribeiro J. & Rocha C.F.D. 2017. Local Environmental Factors Influence the Structure of Frog Communities on the Sandy Coastal Plains of Southeastern Brazil. Herpetologica 73 (4): 307–312. https://doi.org/10.1655/Herpetologica-D-16-00075.1
Opolski-Neto T. & Melo Jr. J.C.F. 2022. Influência das condições edáficas na composição florística e estrutural de uma comunidade de restinga sobre costão rochoso no sul do Brasil. Revista Brasileira de Geografia Física 15 (02): 1108–1127. https://doi.org/10.26848/rbgf.v15.2.p1108-1127
Orrico V.G.D., Dias I.R. & Marciano-Jr E. 2018. Another new species of Phyllodytes (Anura: Hylidae) from the Atlantic Forest of northeastern Brazil. Zootaxa 4407: 101–110. https://doi.org/10.11646/zootaxa.4407.1.6
Paz A., Brown J.L., Cordeiro C.L.O., Aguirre-Santoro J., Assis C., Amaro R.C., Raposo do Amaral F., Bochorny T., Bacci L.F., Caddah M.K., d’Horta F., Kaehler M., Lyra M., Grohmann C.H., Reginato M., Silva-Brandão K.L., Freitas A.V.L., Goldenberg R., Lohmann L.G., Michelangeli F.A., Miyaki C., Rodrigues M.T., Silva T.S. & Carnaval A.C. 2021. Environmental correlates of taxonomic and phylogenetic diversity in the Atlantic Forest. Journal of Biogeography 48: 1377–1391. https://doi.org/10.1111/jbi.14083
Peixoto A.L. & Simonelli M. 2007. Florestas de tabuleiro. In: Simonelli M. & Fraga C. N. Espécies da Flora Ameaçadas de Extinção no Estado do Espírito Santo. Instituto de Pesquisa da Mata Atlântica. Vitória.
Peixoto O.L. & Cruz C.A.G. 1988. Descrição de duas espécies novas do gênero Phyllodytes Wagler (Amphibia, Anura, Hylidae). Revista Brasileira de Biologia 48: 265–272.
Peixoto O.L., Caramaschi U. & Freire E.M.X. 2003. Two new species of Phyllodytes (Anura: Hylidae) from the state of Alagoas, northeastern Brazil. Herpetologica 59: 235–246. https://doi.org/10.1655/0018-0831(2003)059[0235:TNSOPA]2.0.CO;2
Peres E.A., Pinto-da-Rocha R., Lohmann L.G., Michelangeli F.A., Miyaki C.Y. & Carnaval A.C. 2020. Patterns of species and lineage diversity in the Atlantic Rainforest of Brazil. In: Rull V. & Carnaval A. (eds) Neotropical Diversification: Patterns and Processes. Fascinating Life Sciences: 415–447. Springer, Cham. https://doi.org/10.1007/978-3-030-31167-4_16
Pimenta B.V.S., Napoli M.F. & Haddad C.F.B. 2009. A new species of casque-headed tree frog, genus Aparasphenodon Miranda-Ribeiro (Amphibia: Anura: Hylidae), from the Atlantic Rainforest of southern Bahia, Brazil. Zootaxa 2123: 46–54. https://doi.org/10.11646/zootaxa.2123.1.2
Roberto I.J. & Ávila R.W. 2013. The advertisement call of Phyllodytes gyrinaethes Peixoto, Caramaschi & Freire, 2003 (Anura, Hylidae). Zootaxa 3669: 193–196. https://doi.org/10.11646/zootaxa.3669.2.13
Rocha C.F.D., Cogliatti-Carvalho L., Nunes-Freitas A.F., Rocha-Pessôa T.C., Dias A.S., Ariani C.V. & Morgado L.N. 2004. Conservando uma larga porção da diversidade biológica através da conservação de Bromeliaceae. Vidalia 2 (1): 52–72.
Rocha C.F.D., Militão C.M., Vrcibradic D., Van Sluys M., Pereira-Ribeiro J., Dias E.J.R., Marra R.V., Bergallo H.G., Winck G.R., Galdino C.A.B., Cunha-Barros M., Kiefer M.C., Telles F.B.S., Almeida-Santos P., Hatano F.H., Menezes V.A., Siqueira C.C., Miranda J.P., Maia-Carneiro T. & Oliveira J.C.F. 2021. A summary of reptile and anuran amphibian species from Brazilian sandy coastal plains: 31 years of sampling efforts of the “Laboratório de Ecologia de Vertebrados, Universidade do Estado do Rio de Janeiro”. Brazilian Journal of Biology 81 (4): 1144–1165. https://doi.org/10.1590/1519-6984.229617
Rojas-Padilla O., Menezes V.Q., Dias I.R., Argôlo A.J.S., Solé M. & Orrico V.G.D. 2020. Amphibians and reptiles of Parque Nacional da Serra das Lontras: an important center of endemism within the Atlantic Forest in southern Bahia, Brazil. ZooKeys 1002: 159–185. https://doi.org/10.3897/zookeys.1002.53988
Sabagh L.T., Ferreira R.B. & Rocha C.F.D. 2017. Host bromeliads and their associated frog species: further considerations on the importance of species interactions for conservation. Symbiosis 73: 201–211. https://doi.org/10.1007/s13199-017-0500-9
Santos L.S., Roseno R.S., Solé M. & Dias I.R. 2023. Another new species (and it’s not over yet) of Phyllodytes Wagler, 1930 (Anura: Hylidae) from the Atlantic Forest of southern Bahia, northeastern Brazil. Zootaxa 5374 (4): 519–532. https://doi.org/10.11646/zootaxa.5374.4.4
Saporetti-Junior A.W., Schaefer C.E.G.R., Souza A.L., Soares M.P., Araújo D.S.D. & Meira-Neto J.A.A. 2012. Influence of soil physical properties on plants of the Mussununga ecosystem, Brazil. Folia Geobotanica 47 (1): 29–39. https://doi.org/10.1007/s12224-011-9106-9
Sato J., Talora D., Anjos A., Martins A.S., Abreu L.M. & Solé M. 2025. Where to call tonight? Selection of breeding sites by Phyllodytes luteolus (Anura: Hylidae), a bromeligenous tree frog from Brazil. Journal of Natural History 95: 819–832. https://doi.org/10.1080/00222933.2025.2471949
Savage J.M. & Heyer W.R. 1967. Variation and distribution in the tree frog genus Phyllomedusa in Costa Rica, Central America. Beiträge zur neotropischen Fauna 5: 111–131. https://doi.org/10.1080/01650526709360400
Savage J.M. & Heyer W.R. 1997. Digital webbing formulae for anurans: a refinement. Herpetological Review 28: 131.
Simon J.E. & Gasparini J.L. 2003. Descrição da vocalização de Phyllodytes kautskyi Peixoto e Cruz, 1988 (Amphibia, Anura, Hylidae). Boletim do Museu de Biologia Mello Leitão 16: 47–54.
Simonelli M. & Peixoto A.L. 2008. Floristic composition and structure of the tree component of a Muçununga forest in the Linhares Forest Reserve, Espírito Santo, Brazil. In: Thomas W.W. (ed.) The Atlantic Coastal Forest of Northeastern Brazil: 345–364. New York Botanical Garden Press, New York.
Steindachner F. 1862. Über zwei noch unbeschriebene Batrachier aus der Sammlung des K. K. zoologischen Museum zu Wien. Archivio per la Zoologia, l’Anatomia e la Fisiologia 2: 77–82.
Tonini J.F.R., Provete D.B., Maciel N.M., Morais A.R., Goutte S., Toledo L.F. & Pyron R.A. 2020. Allometric escape from acoustic constraints is rare for frog calls. Ecology and Evolution 10 (8): 3686–3695. https://doi.org/10.1002/ece3.6155
Tschudi J.J. von. 1838. Classification der Batrachier mit Berücksichtigung der fossilen Thiere dieser Abtheilung der Reptilien. Neuchâtel: Petitpierre.
Veloso H.P., Rangel-Filho A.L.R. & Lima J.C.A. 1991. Classificação da Vegetação Brasileira, Adaptada a um Sistema Universal. IBGE, Departamento de Recursos Naturais e Estudos Ambientais, Rio de Janeiro.
Vörös J., Dias I.R. & Solé M. 2017. A new species of Phyllodytes (Anura: Hylidae) from the Atlantic Rainforest of southern Bahia, Brazil. Zootaxa 4337: 584–594. https://doi.org/10.11646/zootaxa.4337.4.9
Weygoldt P. 1981. Beobachtungen zur Fortpflanzungsbiologie von Phyllodytes luteolus (Wied, 1824) im Terrarium (Amphibia: Salientia: Hylidae). Salamandra 17: 1–11.
Copyright (c) 2026 Laisa Souza dos Santos, Miguel Trefaut Rodrigues, Iuri Ribeiro Dias

This work is licensed under a Creative Commons Attribution 4.0 International License.
Creative Commons Copyright Notices
Authors who publish with this journal agree to the following terms:
- Authors retain copyright and grant the journal right of first publication with the work simultaneously licensed under a Creative Commons Attribution License (CC BY 4.0) that allows others to share the work with an acknowledgement of the work's authorship and initial publication in this journal.
- Authors are able to enter into separate, additional contractual arrangements for the non-exclusive distribution of the journal's published version of the work (e.g., post it to an institutional repository or publish it in a book), with an acknowledgement of its initial publication in this journal.
- Authors are NOT PERMITTED to post their submitted work online (e.g., in institutional repositories or on personal websites) prior to publication, as it may lead to nomenclatural issues.
