UNIVERSIDADE FEDERAL DE SÃO CARLOS CENTRO DE CIÊNCIAS BIOLÓGICAS E DA SAÚDE PROGRAMA DE PÓS-GRADUAÇÃO EM ECOLOGIA E RECURSOS NATURAIS VINÍCIUS DA FONTOURA SPERANDEI ANUROFAUNA E CAVERNAS NEOTROPICAIS: FATORES BIÓTICOS E ABIÓTICOS QUE MODULAM A OCUPAÇÃO DO HABITAT SUBTERRÂNEO SÃO CARLOS - SP 2024 Vinícius da Fontoura Sperandei ANUROFAUNA E CAVERNAS NEOTROPICAIS: FATORES BIÓTICOS E ABIÓTICOS QUE MODULAM A OCUPAÇÃO DO HABITAT SUBTERRÂNEO Tese apresentada ao Programa de Pós-Graduação em Ecologia e Recursos Naturais, ao Centro de Ciências Biológicas e da Saúde da Universidade Federal de São Carlos, para obtenção do título de Doutor em Ciências, área de concentração em Ecologia e Recursos Naturais. Orientador: Prof. Dr. Francisco Leonardo Tejerina-Garro Coorientador: Prof. Dr. Wilian Vaz-Silva São Carlos - SP 2024 AGRADECIMENTOS Inicialmente agradeço aos meus pais, Suzi e Neri, por todo o apoio e torcida ao longo destes 12 anos de vida acadêmica na área das Ciências Biológicas, por entenderem as escolhas e comemorar a cada conquista. À minha esposa Isabela, minha parceira ao longo desse período que colocou o pé na estrada junto comigo e estamos trilhando um caminho árduo e de muitas vitórias. Ao meu orientador Dr. Francisco Leonardo por todos os ensinamentos, apoio, paciência nas explicações e incentivo com todos os desafios, com certeza estaremos juntos em novos projetos e ideias futuras que virão. Ao meu coorientador Wilian Vaz-Silva pelas colaborações ao longo do trabalho e aos novos desafios que virão, além de todos os revisores, professores das disciplinas e membros de banca que contribuíram ao longo destes anos para a construção desta tese. Ao Programa de Pós-Graduação em Ecologia e Recursos Naturais da Universidade Federal de São Carlos pelo apoio e recepção durante o doutoramento. Em especial a professora Dra. Ana Teresa Lombardi, na qual me fomentou um aprendizado único e talvez o mais valioso em toda minha vida acadêmica quando mais foi necessário, a verdadeira maneira de se ensinar olhando para além de um crescimento intelectual e prático, mas um crescimento pessoal, obrigado professora por ter me incentivado e orientado meus passos durante essa jornada. Agradeço o apoio da Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Código de Financiamento 001 (PROAP/CAPES PPGERN), imprescindível para a realização e conclusão desta. Á Universidade de Rio Verde e aos amigos presentes nesta instituição que incentivam e fazem parte de maneira direita e indireta desta materialização, sou muito agradecido pelas oportunidades e aprendizados no dia a dia desta valorosa universidade. Á todos os amigos e colegas de profissão que incentivaram, colaboraram com ideias, discussões e apoiaram chegar até aqui, saibam que acredito fortemente em uma ciência colaborativa, onde se trabalhar em conjunto e é muito mais satisfatório percorrer esta jornada com todos vocês. RESUMO Cavernas formam uma rede de espaços entre rochas que ocupam desde canalículos á grandes dimensões que podem ser acessados por seres humanos e outros seres vivos. Este ambiente possui características singulares quando comparado à superfície, como ausência de luz, abastecimento energético oriundo do ambiente epígeo, temperatura amena e umidade relativa tendendo a saturação. Dos animais vertebrados que vivem no interior de cavernas neotropicais brasileiras, os anuros ainda formam um grupo taxonômico com pequeno arcabouço científico e a ocupação do ambiente por eles ainda é tratada como acidental ou temporária. O objetivo desta tese é identificar o panorama do estudo da ocupação de cavernas por anuros, realizando o levantamento de registros científicos e por meio dele elaborar uma lista de espécies. A partir dos registros foram extraídas variáveis categóricas de natureza abiótica, discutindo as condições que a caverna possui e que influenciam a presença dos anuros e variáveis biológicas de área de vida, história natural e de conservação do perfil de espécies registradas em cavernas. Foram encontradas 18 publicações científicas no intervalo de 42 anos com registros de anuros em cavernas e totalizaram 247 registros independentes. A lista de espécies conta com 54 registros até o epíteto específico (83 morfótipos) com destaque para as famílias Leptodactylidae (13 espécies), Hylidae (12) e Bufonidae (10) como mais ricas, registradas predominantemente em Minas Gerais e São Paulo, em biomas de Floresta Atlântica e zonas de ecótono Floresta Atlântica/Cerrado brasileiro, em litologia carbonática e ferruginosa, em zona de entrada e com a presença de corpos d’água na cavidade. A análise de componentes multivariados para os fatores abióticos aponta para a formação de dois agrupamentos, o primeiro formado por cavernas em biomas da Floresta Atlântica e Ecótono Floresta Atlântica/Caatinga, em litologias carbonática e mármore, com variação altitudinal de 23 a 246m, inseridas em matriz com formações vegetais (florestal, campestre e savânica) e grau de endemismo. O segundo agrupamento aponta para fatores abióticos, e é formado por cavidades em litologias ferruginosa, quartzítica e granítica, com a presença de matriz de paisagem florestal, campestre e afloramentos rochosos com água perene. Este cenário é ocupado por Bokermannohyla martinsi (restrita à região do Quadrilátero Ferrífero) e Ischnocnema manezinho endêmica em Santa Catarina. As variáveis categóricas para os fatores biológicos demonstraram agrupamentos com características antagônicas, em um dos grupos caracterizado por animais de área de distribuição restrita, endêmicos e pouco frequente ou raro na comunidade herpetológica local, de período diurno, hábitos criptozóicos e local de reprodução grutas e tocas ou rio/riacho ou chão de mata, além de status de conservação “em perigo – EN” da União Internacional para a Conservação da Natureza (IUCN). O segundo grupo é formado por anuros com área de distribuição ampla e moderada, sem endemismo e frequente no ambiente, animais noturnos, hábitos arborícolas e em habitats abertos, com local de reprodução brejo/lago e remanso de rios, sem grau de ameaça a conservação. Apresenta-se também um registro inédito de predação de um anuro Crossodactylus caramaschii por uma aranha-pescadora (Trechaleoides biocellata) em uma caverna do interior paulista, reforçando o arcabouço científico da interação de anuros com a comunidade subterrânea. Palavras-chave: diversidade anurofaunística, Lacuna Linneana, Lacuna Wallaceana, ocupação de habitat. ABSTRACT Caves form a network of spaces between rocks, ranging from tiny spaces like crevices to large spaces accessible to humans. This environment has unique characteristics compared to the surface, such as the absence of light, energy supply originating from the epigean environment, mild temperature, and relative humidity tending towards saturation. Among vertebrate animals that live inside natural cavities in Brazilian neotropical regions, anurans still form a taxonomic group with limited scientific framework, and their occupation of the environment is still considered accidental or temporary. The objective is to identify the panorama of cave occupation by anurans, conducting a survey of scientific records and using it to compile a species list. From the records, categorical variables of abiotic nature were extracted, discussing the cave conditions that influence the presence of anurans, as well as biological variables related to home range, natural history, and conservation profile of species recorded in caves. Eighteen scientific publications were found over 42 years with records of anurans in caves, totaling 247 independent records. The species list includes 54 records up to the specific epithet (83 morphotypes), with emphasis on the families Leptodactylidae (13 species), Hylidae (12), and Bufonidae (10) as the richest, predominantly recorded in Minas Gerais and São Paulo, in the Atlantic Forest biome and ecotonal zones between the Atlantic Forest and Brazilian Cerrado, in carbonate and ferruginous lithology, at entrance zones with presence of water bodies in the cavity. Multivariate analysis for abiotic factors indicates the formation of two clusters: the first formed by caves in the Atlantic Forest biome and Atlantic Forest/Caatinga ecotone, in carbonate and marble lithologies, with altitudinal variation from 23m to 246m, inserted in a matrix with vegetal formations (forest, grassland, and savanna) and presence of endemic anurans. The second cluster for abiotic factors is formed by cavities in ferruginous, quartzitic, and granitic lithologies, with presence of forest landscape matrix, grassland, and rocky outcrops with perennial water; this scenario is inhabited by Bokermannohyla martinsi (restricted to the Quadrilátero Ferrífero region) and Ischnocnema manezinho endemic to Santa Catarina. Categorical variables for biological factors showed groupings with antagonistic characteristics: one group characterized by animals with restricted distribution area, endemics and rare or infrequent in the local herpetological community; diurnal period; cryptic habits; reproduction sites in caves, burrows, or river/stream or forest floor; and "endangered - EN" conservation status according to IUCN. The second group consists of anurans with wide and moderate distribution areas, without endemism and common in the environment; nocturnal animals; arboreal habits; open habitat; reproduction sites in marshes/lakes and river backwaters; without conservation threat status. We also present a previously unreported predation event of an anuran Crossodactylus caramaschii by a fishing spider (Trechaleoides biocellata) inside a cave in São Paulo State, supporting the scientific framework of anuran interaction with the subterranean community. Keywords: anuran diversity, habitat occupancy, Linnean Gap, Wallacean Gap.. LISTA DE FIGURAS E TABELAS Capítulo I Table 1 – Absolute and relative values of independent records of nominal and ordinal variables extracted from the 18 scientific works considered and published from January 1980 to December 2022. 25 Table 2 – Anuran species found in Brazilian natural cavities by state, lithology, and biome. 26 Figure 1 – Number of species per family of Brazilian neotropical anurans recorded from underground cavities. 30 Figure 2 – Number of anuran records in natural cavities in Brazilian states. 31 Figure 3 – Caves according to the 2022 Cadastro Nacional de Informações Espeleológicas CANIE/Cecav (green dots) and with recorded anurans (yellow triangle) and caves in the Brazilian neotropical region. 33 Figure 4 – Number of records by type of lithology Lithologies: Aren = sandstone; Carb = carbonate; Cong = conglomerate; Ferr = ferruginous; Gran = granitic; Quar = quartzite; Marm = marble; ND = Not available. 35 Figure 5 – Number of scientific publications with records of anurans in Brazilian neotropical cavities per year from 1980 to 2022. 37 Capítulo II Table 1 - Variables, categories and subcategories used in the multivariate Multiple Component Analysis for records of anurans in Neotropical underground cavities. 70 Table 2 – List of anuran morphospecies found inside Brazilian caves according to scientific literature and records of the federative units found, biome, lithology, and number of records per morphospecies. 74 Figure 1 – dispersion of categorical variables explaining 12.91% of data variability of records of occurrence of anurans in neotropical caves. 78 Capítulo III Tabela 1 - Variáveis categóricas utilizadas na Análise de Correspondência Multivariável (ACM) para os fatores bióticos de anuros neotropicais registrados em cavernas brasileiras. 110 Figura 1 – Dispersão dos dados significativos da Análise de Componentes Multivariada (ACM) com explicação de 11,59% da variabilidade de dados. 111 Capítulo IV Figure 1 – Observation of predation involving an anuran, Crossodactylus caramaschii (Anura: Hylodidae), by a fishing spider (Trechaleoides biocellata (Aranae: Trechaleidae) in Fenda das Almas, a carbonate cave in the State of São Paulo, Brazil. 137 SUMÁRIO 1 INTRODUÇÃO GERAL 14 2 CAPÍTULO I – Characterization of anuran amphibian fauna in caves in Brazil 18 2.1 - Abstract 19 2.2 - Resumo 20 2.3 - Introduction 21 2.4 - Material and Methods 23 2.5 - Results and Discussion. 25 2.6 - Conclusion 37 2.7 - References 38 2.8 - Appendix I 47 2.9 - Appendix II 62 3 - CAPÍTULO II – Abiotic characteristics that influence the occupation of Brazilian neotropical caves by anurans 66 3.1 - Abstract 67 3.2 - Introduction 68 3.3 - Material and Methods 70 3.4 - Results 74 3.5 - Discussion 79 3.6 - Conclusion 82 3.7 - Bibliographic References 82 3.8 - Appendix I 92 4 - CAPÍTULO III – Anuros e cavernas neotropicais: tópicos de história natural dos anuros habitantes do ambiente cavernícola 101 4.1 - Abstract/Resumo 103 4.2 - Introdução 104 4.3 - Materiais e Métodos 107 4.4 - Resultados 109 4.5 - Discussão 111 4.6 - Conclusão 113 4.7 - Referência Bibliográfica 114 4.8 - Anexo I 118 4.9 - Anexo II 119 5 - CAPÍTULO IV – The first observation of predation on Crossodactylus caramaschii Bastos and Pombal, 1995 (Anura: Hylodidae) by a fishing spider (Araneae: Trechaleidae) in a neotropical cave 135 5.1 - Short Communication 136 5.2 - References 139 6 - CONCLUSÃO FINAL 141 7 - REFERÊNCIAS BIBLIOGRÁFICAS 144 INTRODUÇÃO GERAL O ambiente subterrâneo natural compreende sistemas ecológicos complexos e dinâmicos com grande variação de características físicas e biológicas (Ferreira, 2005; Ortuño et al., 2013; Culver e Pipan, 2019). Compõe o ambiente cavernícola o conjunto de espaços inter-conectados no espaço subterrâneo, preenchidos por água ou ar e consistem desde variações milimétricas como canalículos e fissuras até cavidades acessíveis ao homem (Juberthie e Decu, 1994; Juberthie, 2000; Culver e Pipan, 2009). Podem ser formadas em rochas de qualquer litologia por processos de dissolução química ou mecânicos como erosão, resfriamento de lava, sobreposição de rochas abatidas (Juberthie, 2000). O Brasil apresenta um elevado número de cavidades naturais quando comparado a outros países no cenário mundial, com um patrimônio espeleológico de 23.278 cavidades cadastradas no Cadastro Nacional de Informações Espeleológicas (ICMBio, 2022). Porém com potencial para este número ser significativamente maior, uma vez que o país possui muitas áreas ainda não prospectadas e não catalogadas. Características singulares às cavernas são elevadas estabilidades nas condições ambientais, tais como temperatura constante e amena, úmidade relativa do ar próxima da saturação, ausência permanente de luz natural e baixa ou nenhuma produção de energia primária nas zonas mais profundas (Poulson e White, 1969; Tobin et al., 2013, Culver e Pipan, 2019). O aporte energético do sistema é dependente dos mecanismos alóctones principalmente pelos sistemas de drenagem que carream matéria orgânica, além de fezes e carcaças de animais que transitam neste ambiente ou eventualmente caem nas cavidades (Culver e Pipan, 2009; Culver e Pipan, 2019). A variação do tamanho e quantidade de entradas das cavidades (incluindo a captação de rios para o sistema) e o abastecimento energético do sistema influenciará a heterogeneidade do habitat do ambiente e por sua vez a abundância e riqueza da fauna cavernícola (Gibert e Deharveng, 2002; Silva e Ferreira, 2016; Sperandei, 2019). Hábitos dos anuros que facilitam o sucesso de colonização como ser noturno e/ou criptico, possuir orientação não-visual, tolerar altas umidades e dieta generalista (Cullingford, 1962).Somadas estas características nos indica que espécies para ocupar e permanecer neste ambiente possuem pré-adaptação de ordem morfológica ou fisiológica às condições do ambiente subterrâneo (Barr, 1968; Prous et al., 2004). O meio hipógeo apresenta três “zonas” com diferenças bióticas e abióticas baseadas na incidência de luz natural. 1 - a zona de entrada (zona fótica) com incidência direta de luz solar, crescimento de pequenos líquens e até vegetais, alta influência da superfície. 2- zona de penumbra (zona disfótica) onde não há incidência direta de luz porém ainda há luminosidade. 3- zona profunda (zona afótica) onde não há permanentemente luminosidade alguma e maior estabilidade por ter pouca influência do meio epígeo (Poulson e White, 1969; Howart, 1979). Animais com diferentes graus de afinidade e adaptação habitam as cavernas, com relações ecológica-evolutivas com o habitat e diversos graus de especialização (fisiológicas, morfológicas, reprodutivas e comportamental) (Culver e Wilkens, 2000; Sket, 2008; Romero, 2009). Os invertebrados são os organismos mais abundantes, com importante participação na ciclagem de nutrientes e transferência energética dentro da rede ecológica (Poulson, 2005; Silva et al., 2011). Entre os vertebrados, os grupos taxonômicos mais estudados em cavernas são os quirópteros e peixes, porém podemos registrar ainda aves que nidificam nas entradas das cavidades, pequenos mamíferos comuns em terrenos rochosos, além de animais da herpetofauna. Em geral, as comunidades no ambiente subterrâneo apresentam riqueza mais alta próximas ás entradas e diminuindo á medida que fica mais profundo (Prous et al., 2004; Culver e Pipan, 2009; 2019). Diante das características ecológicas e evolutivas do mundo animal, desde os primórdios do estudo da biologia animal procurou-se classificar a natureza, a fim de encontrar padrões ecossistêmicos globais e em recortes da paisagem (Gleason, 1926; Rosen, 1996; Ladle e Hortal, 2013). Os animais encontrados em cavernas podem ser classificados como troglóbios, troglófilos e trogloxenos, segundo Holsinger e Culver (1988) (modificado a partir do sistema Schinner-Racovitza, 1907). Os troglóbios são animais com ocorrência restrita ao meio subterrâneo e que podem apresentar especializações morfológicas, fisiológicas e comportamentais como resposta ás pressões seletivas do ambiente e/ou ausência de pressões ambientais e ecológicas da superfície, conhecidos como troglomorfismos (Holsinger e Culver, 1988; Langecker, 1989). Os troglófilos são facultativos, podendo completar seu ciclo de vida dentro ou fora das cavernas. Os trogloxenos são aqueles que ocorrem no ambiente subterrâneo, mas precisam retornar periodicamente à superfície para completar seu ciclo de vida. Há ainda a classificação dos animais acidentais (Trajano e Bichuette, 2006) que caracteriza a fauna do ambiente epígeo que entra eventualmente nas cavernas de forma aleatória ou por períodos não-sístemicos e em alguns casos não conseguem se alimentar ou orientar levando á morte do indíviduo. Em algumas regiões do planeta é demonstrado que alguns vertebrados ocupam o ambiente subterrâneo regularmente, passando a utiliza-lo em parte ou na totalidade do seu ciclo de vida, sendo assim caracterizados como animais trogloxenos ou troglófilos (Resetarits Jr,1986; Espino Del Castillo et al., 2009; Suwannapoom et al., 2018). Até o momento nenhuma espécie de anuro troglomórfica foi registrada, indicando que especializações extremas ao ambiente subterrâneo podem não ocorrer neste grupo (Biswas, 2014). Estudos em regiões temperadas discutem a variação sazonal na utilização das cavidades pelos anuros, a qual no inverno é quando se encontra maior riqueza e abundância indicando a ocupação como abrigo contra a condição externa adversa (Resetarist Jr, 1986; Fenolio et al., 2005; Lunghi et al., 2017). Em região tropical Lúria-Manzano e Ramírez-Bautista (2017), no México, compararam a variação da dieta de Craugastor alfredi (Boulenger, 1898) e concluíram que populações hipógeas possuem hábitos alimentares diferentes das populações da superfície. Já as cavernas neotropicais apresentam larga abundância e riqueza de invertebrados, presas em potencial para os anuros (Bernarde, 2012) e formam um ambiente propício contra predadores de orientação visual como as aves. As condições fisiológicas deste grupo devido ao tegumento mais permeável podem levar a uma tendência a ocupação do habitat favorável (Eterovick et al., 2010), com altos níveis de umidade relativa, normalmente ultrapassando os 90% mesmo nas estações mais secas (Matavelli et al., 2015). Alguns invertebrados como hemípteras, coleópteras, himenópteras, além de aracnídeos em geral podem predar anuros (Toledo, 2005; Bernarde, 2012), já registrados no ambiente hipógeo, ainda que em cavidades neotropicais ainda não há registro de eventos de predação de anuros por invertebrados ou vice-versa. Registros de interação ecológica (predação, simbiose) podem elucidar como os anuros compõe a rede-ecológica dos sistemas subterrâneos e se as presas e predadores deste grupo se mantem (ou não) similares aos do meio externo (Lúria-Manzano e Ramírez-Bautista, 2017). Os anfíbios em geral são apontados pela IUCN (2021) como grupo de vertebrados mais ameaçado do mundo, em declínio populacional acentuado desde a década de 90 (Wake, 1991; Becker e Zamudio, 2011) e causas diversas como fragmentação e destruição de habitat (Moreira, 2016), introdução de espécies exóticas (Collins e Crump, 2009; Boelter et al., 2012; Both e Grant, 2012), modificação e deterioração no uso da terra (Meegaskimbura et al., 2002; Moresco et al., 2014; Perez-Iglesisas et al., 2016), contaminação do solo e corpos d’água (Hayes et al., 2010), mudanças climáticas severas (McMenamin et al., 2008; de Oliveira et al., 2016) e a alta contaminação pelo fungo Batrachochytrium dendrobatidis (BD) (Vredenburg et al., 2010; Olson et al., 2013) acabam ameaçando a conservação de anuros neotropicais. Com tantos fatores que ameaçam a conservação dos anuros há uma crescente demanda de atenção a estes vertebrados, que exige maior compreendimento do uso do habitat e pode indicar novos caminhos para a conservação dos anuros neotropicais na região brasileira, entre elas o uso e interação com o habitat subterrâneo. Neste sentido, objetiva-se apresentar dados de anuros registrados em cavernas neotropicais brasileiras e identificar lacunas no conhecimento da ecologia de anuros. Especificamente objetiva-se (i) avaliar qual o atual panorama da ciência com registros de anuros em cavernas neotropicais brasileiras, (ii) identificar quais características bióticas dos animais influenciam na ocupação do habitat subterrâneo e (iii) apontar quais características abióticas das cavidades naturais influenciam na ocupação de cavernas por anuros e por fim (iv) apresentar uma interação inédita de predação de um anuro neotropical por uma aranha-pescadora no interior de uma cavidade discutindo o comportamento registrado e quais variáveis de história de vida (comportamental, ecológica e taxonômica) influenciam biologicamente na ocupação do meio hipógeo por anuros neotropicais. CAPÍTULO I Characterization of anuran amphibian fauna in caves in Brazil Submetido em Biota Neotropica (Qualis B1 – Quadriênio 2017-2020; Fator de impacto: 1.20 – 2023) na seção “Articles” Normas e diretrizes de submissão em https://www.biotaneotropica.org.br/BN/about/submissions Characterization of anuran amphibian fauna in caves in Brazil Vinícius da Fontoura Sperandei1*, Willian Vaz Silva², Francisco Leonardo Tejerina-Garro¹,²,³ 1 Programa de Pós-Graduação em Ecologia e Recursos Naturais, Universidade Federal de São Carlos, 13565-905, Km 235, Rodovia Washington Luís, São Carlos, SP, Brazil. 2 Programa de Pós-Graduação em Ciências ambientais e saúde, Pontifícia Universidade Católica de Goiás, Av. Universitária 1.440, Setor Universitário, 74605-010 - Goiânia, Goiás, Brazil. 3 Programa de Pós-Graduação em Sociedade, tecnologia e meio ambiente, Universidade Evangélica de Goiás, 75083-515, Av. Universitária km 3,5 Cidade Universitária – Anápolis, Goiás, Brazil. *Corresponding author: vinicius.sperandei@gmail.com ORCID: Vinícius da Fontoura Sperandei: 0000-0002-3093-706X Willian Vaz Silva: 0000-0001-6235-5331 Francisco Leonardo Tejerina-Garro: 0000-0002-5159-8108 Abstract: The natural underground environment has unique characteristics when compared to surface environments. In these environments, the food web is simplified but includes organisms such as anuran amphibians. This work aims to characterize the presence of anurans in caves in the Brazilian neotropical region considering taxonomic, geographic, geological, and environmental information collected in the scientific literature. A total of 247 records of anurans in caves were found and distributed in 18 scientific papers throughout 42 years. Of this total, 177 (71.6%) records presented identification up to the specific epithet (54 species) with emphasis on the families Leptodactylidae (13 species; 24.1%), Hylidae (12; 22.2%) and Bufonidae (10; 18.5%). The records of anurans are predominant in the states of Minas Gerais (81 records; 34.8%) and São Paulo (35; 15.0%). The biome with the highest number of records is the Atlantic Forest with 78 (33.5%) followed by ecotonal zones between the Atlantic Forest and Cerrado (43 records; 18.5%). Environmental information is scarce in the consulted records, 12 contained information about the light zone of the cave where the anuran was found (9 in the entrance zone) and 63 with the information about the presence/absence of water bodies in the cave. The carbonaceous (109; 46.8%) and ferruginous (76; 33.9%) lithology of the caves is predominant. The records of anurans in caves come from articles (Full Paper type), in the biological knowledge area of community studies. The number of sampled caves is low (55 caves; 0.24%) compared to the total number of caves registered in Brazil. The results suggest that there is a need to implement studies for the understanding of the use of the underground habitat by neotropical anurans. Keywords: Atlantic Forest, carbonate geology, Speleology, Leptodactylidae, Linnean and Wallacean gap, Neotropical region Resumo: O ambiente subterrâneo natural apresenta características singulares quando comparado aos ambientes superficiais. Nesses ambientes a rede trófica é simplificada, mas inclui organismos como os anfíbios anuros. Este trabalho objetiva caracterizar a presença de anuros em cavernas da região neotropical brasileira considerando informações taxonômicas, geográficas, geológicas e ambientais coletadas na literatura científica. Foram encontrados 247 registros de anuros em cavernas, distribuídos em 18 trabalhos científicos em um intervalo de 42 anos. Desse total, 177 (71,6%) registros apresentaram identificação até o epíteto específico (54 espécies) com destaque para as famílias Leptodactylidae (13 espécies; 24,1%), Hylidae (12; 22,2%) e Bufonidae (10; 18,5%). Os registros de anuros são predominantes nos estados de Minas Gerais (81 registros; 34,8%) e São Paulo (35; 15,0%). O bioma com maior quantidade de registros é a Floresta Atlântica com 78 (33,5%) seguido de zonas ecotonais entre Floresta Atlântica e Cerrado (43 registros; 18,5%). As informações ambientais são escassas nos registros consultados, 12 continham a informação da zona de luminosidade da caverna em que o anuro foi encontrado (9 em zona de entrada) e 63 com a informação da presença/ausência de corpos d’água na caverna. A litologia carbonática (109; 46,8%) e ferruginosa (76; 33,9%) das cavernas é predominante. Os registros de anuros em cavernas são provenientes de artigos (tipo Trabalho Completo), em área do conhecimento biológico do estudo de comunidades. O número de cavernas amostradas é baixo (55 cavernas; 0,24%) em relação ao número total de cavernas cadastradas no Brasil. Os resultados sugerem que há necessidade de implementar estudos para o entendimento do uso do habitat subterrâneo pelos anuros neotropicais. Palavras-chave: espeleologia, Floresta Atlântica, geologia carbonática, lacuna Linneana e Wallaceana, Leptodactylidae, região neotropical Introduction Caves are natural subterranean cavities that make up the karst systems, originating from the chemical and physical dissolution of the matrix rock (Gilbert et al. 1994). In Brazil, these cavities are accessible to humans (BRASIL 2008) and are popularly known as caves, grunas, burrows, lapa, caves, and others. 23,278 natural cavities (caves; CECAV 2022) have been recorded and registered, with an estimated 100 thousand caves (Auler et al. 2001, ICMBIO/CECAV 2017). Natural cavities have their internal characteristics, but they interact with the external environment, providing photic and aphotic zones inside the cave in the deeper parts and where a stability of average temperature and humidity close to saturation is observed (Culver 1982, Culver & Pipan 2009, Tobin et al. 2013). The lack of natural light causes the total or partial absence of photosynthetic organisms, thus primary production is limited, with the available energy in the form of organic matter coming from the epigeal environment favored by physical and biological agents (Simon et al. 2007). This energy availability changes according to the geomorphology of the cave, creating unique conditions for each of them according to specific variations in connection with epigean ecosystems (Poulson & White 1969, Souza-Silva et al. 2011, Pellegrini et al. 2016). The entrance opening of the cavities (entrances) modulates several characteristics, in addition to the entry of nutrients, as it also acts as a filter for animals that enter by force of water, floods and that start to inhabit the cave environment even if for a short period (Simões et al. 2015). The entrance opening and the adjacent area of the cave can also act as nesting sites for birds, pollinating insects, and small non-flying mammals and shelter for various animals as it is a shaded area with mild conditions (Boulton et al. 2008, Medellin et al. 2017). The cave community is classified considering the characteristics of the species (morphological, physiological, evolutionary, and behavioral) and affinity with the underground habitat (Sket 2008, Culver & Pipan 2009, Romero Díaz 2009). Thus, and according to Sket (2008) based on the Schinner-Racovitza proposal, organisms are classified into i) trogloxenes, animals that are regularly found in the underground environment, but that needs to go to the surface environment to complete their life cycle; ii ) troglophiles, animals with affinity to the cave environment, but with underground and epigeal populations, and can even complete their life cycle entirely in cavities; iii ) troglobites, are obligate subterranean species, with morphological, behavioral and physiological variations in response to the selective pressures of the hypogean environment and are normally restricted to the local extension of the cave, d) “accidental” are animals that do not present pre-adaptations, but they “accidentally” enter the underground environment and end up not being able to establish themselves (Barr 1968, Sket 2008, Juan et al. 2010). Among the animals that make up the “accidental” fauna of caves are anurans and reptiles, despite the recent understanding of the systemic use of this environment by herpetofauna (Martins de Andrade et al. 2021, Dos Santos 2022a, b). Concerning amphibians, there are species with affinity to the cave-dwelling environment, such as, for example, Litoria cavernicola Tyler and Davies, 1979, with a habitat in sandstone caves in Australia, Rana italica Dubois 1987, in Italy (Lunghi et al. 2018), Eleutherodactylus cavernicola Lynn, 1954, with endemic distribution in Jamaica (Stanely et al. 2021) and Craugastor pelorus Campbell and Savage, 2000, in southern Mexico (Couto et al. 2023). In Brazil, Trevelin et al. (2021) categorizes Pristimantis cf. fenestratus (Steindachner, 1864) as trogloxene in caves in the state of Pará, Martins de Andrade et al. (2021) indicates fidelity of cave use by the endemic species Bokermannohyla martinsi (Bokermann, 1964) in Minas Gerais and Vaz-Silva et al. (2020) describes the species Oreobates antrum to the northwest of Goiás discussing its presence in rocky environments and limestone outcrops in the region, including the popular name “rãzinha das cavernas” which is recorded by Motta et al. (2020) for caves in southwestern Tocantins. Knowledge of the distribution patterns and composition of the cave community are important tools for measuring the importance of caves as an element of the landscape (Gibert & Deharveng 1994, 2002). About tropical cave fauna, more specifically in Brazil, is still incipient when compared to that of temperate regions (Deharveng & Bedos 2012), although, in the last two decades, there has been a significant increase in the number of biospeleological studies addressing the fauna of invertebrates and cave-dwelling vertebrates (Delgado-Jaramillo et al. 2017, Parizotto et al. 2017, Gallão et al. 2018). Despite this, there are few studies of Brazilian anurofauna in caves with many knowledge gaps (sensu Hortal et al. 2015) to be filled (Guerra et al. 2018, Bichuette et al. 2022). Therefore, the objective of this article is to build a checklist of anurans recorded in Brazilian neotropical caves and characterize the records considering taxonomic, geographic, geological and environmental information. In this way we will be able to identify possible concentrations of articles and areas that are still little studied and reinforce the theoretical framework by reducing the Linnean and Wallacean gaps. Material and Methods We use records of anurans in natural cavities in the Brazilian neotropical region mentioned in the literature. The scientific works chosen for data compilation were full papers and scientific notes, theses or dissertations and book chapters published until December 2022. The search for scientific works was carried out on Google Scholar and in the scientific repositories SCOPUS, Web of Science, and AmphibiaWeb, which constitute suitable repositories of scientific literature (Falagas et al. 2008, Delgado & Repiso 2013, Martín-Martín et al. 2021). The selection of works in more than one database for the systematic search avoids bias in the collected data and consequently its interpretation (Archambault et al. 2009, Mongeon & Paul-Hus 2016). Keywords were listed to search for scientific works in the repositories indicated independently or in combination and without limiting search time (Silva & Bianchi, 2001, da Silva et al. 2011). The keywords were: “anura”, “Brazilian caves”, “anurans”, “amphibians”, “frogs”, “underground cavities”, “underground habitat”, “underground environment”, “grotto”, “gruna”, “Toca”, “subterranean cavities”, “subterranean habitat”, “subterranean environment” and “burrow” (all terms were researched with their variations in Portuguese). From the selected literature, each record of a different species or different location in each of the studies represented an independent record, so the total number of records was the sum of the independent records in all articles found until December 2022. From the records taxonomic information (order, family, genus, specific epithet), geographic (Federative Unit, municipality, name of the cavity and biome), geological (type of cave lithology ), environmental (description of the location of the anuran in the luminous zone - entrance) were extracted cave, dim or aphotic; presence of water and relating to the scientific literature consulted (type of publication - Full Article, Short Communication, Textbook or Dissertation; focus of the study; year of publication). In all cases, the absence of information was identified as “not available” (ND). Based on this information (called variables) a data matrix was created (Appendix I). The data matrix for the descriptive analysis considered all records with information on anurans in Neotropical caves in the Brazilian region, regardless of the level of taxonomic identification being up to the specific epithet or not. To prepare the list of species, only records that contained the identification of the anuran up to the specific epithet were considered and those with identification at the family, genus, or morphotype level were excluded. The nomenclature of each species was checked and, when necessary, updated using the website database Amphibian Species of the World 6.1 (Frost, 2023), in addition to identification guides by Haddad et al. (2013), Freitas (2015), and Vaz-Silva et al. (2020). The independent records, data on categorical variables were extracted, which involved the taxonomic identification data of the anuran, the record of the cave studied, the location of the cave, and the type of study itself (Table 1). These records point to individual observations of specimens in the cave environment, which indicates that the repetition of total specific epithets does not reflect species richness, since we will have more than one record for the same species in some cases. From the data matrix, a descriptive analysis was carried out considering the categories of nominal variables (order, family, genus, specific epithet, cave, federative unit, municipality, lithology, biome, presence or absence of water, type of publication, focus of the study) and ordinals (photic zone and year of publication) used (Reis & Reis 2002, Silvestre 2007). Results and Discussion The literature review indicated 247 records of anurans inside Brazilian caves distributed in 18 scientific works (Appendix II). From the first work in which anurans are mentioned inside cavities in Brazil by Desen et al. (1980) until the last publication in the time frame of this work represented by Dos Santos et al. (2022) 42 years have passed. The growth in the number of publications in the last three years 2020 stands out – 2022 with eight of the 18 publications in this period. Among the taxonomic variables, a smaller amount of information is observed in the “Specific epithet” category, since complete identification is often complex, with morphologically similar species belonging to the same genus, requiring a more accurate identification via comparison of the collected individuals (when available) with species from reference zoological collections (Haddad et al. 2015). Table 1 - Absolute and relative values of independent records of nominal and ordinal variables extracted from the 18 scientific works considered and published from January 1980 to December 2022. Variable Absolutes Relatives (%) Order 247 100 Family 240 97.2 Genus 233 94.3 Specific epithet 177 71.7 Cave 99 40.1 State 184 74.5 County 182 73.7 Lithology 246 99.6 Light zone 12 4.9 Biome 183 74.1 Presence/Absence of water 63 25.5 The literature review indicates 54 species of anurans from caves in the Brazilian Neotropical region distributed in 11 families (Table 2 and Figure 1). In this region, 1114 species of anurans are described, constituting one of the largest in the world (Segalla et al. 2021) of which the 54 species surveyed in this study correspond to 4.8%. The latter is low when considering that anurans have ecophysiological characteristics that favor the use of the underground environment (Eterovick et al. 2010, Matavelli et al. 2015). Table 2 - Anuran species found in Brazilian natural cavities by state, lithology, and biome. States: BA = Bahia; GO = Goiás; MT = Mato Grosso; MS = Mato Grosso do Sul; MG = Minas Gerais, PA = Pará; PR = Paraná; RN = Rio Grande do Norte; SC = Santa Catarina; SP = São Paulo; SE = Sergipe; TO = Tocantins. Lithologies: Aren = sandstone; Carb = carbonate; Cong = conglomerate; Ferr = ferruginous; Gran = granitic; Quar = quartzite; Marb = marble. Biomes: Ama = Amazon Rainforest; Caa = Caatinga; Cerr = Cerrado; Atl = Atlantic Forest; Ecot = Ecotone . Family Genus and species state Lithology Biome Records Aromobatidae (S=1) Allobates marchesianus (Melin, 1941) - Ferr Ama 1 Brachycephalidae (S=4) Ischnocnema guentheri (Steindachner , 1864) MG Ferr - 1 Ischnocnema izecksohni (Caramaschi & Kisteumacher , 1989) MG Carb - 1 Ischnocnema juipoca (Sazima & Cardoso, 1978) MG Carb - Ferr - Cong Ecot (Cerr - Atl) 6 Ischnocnema manezinho (Garcia, 1996) SC Gran Atl 7 Bufonidae (S=10) Rhaebo guttatus (Schneider, 1799) PA Aren - Ferr Ama 3 Rhinella crucifer (Wied- Neuwied , 1821) SP Carb - Gran Atl 3 Rhinella diptycha (Cope, 1862) MG-SE Aren - Carb Cerr - Atl 6 Rhinella granulosa (Spix, 1824) - Carb Caa 1 Rhinella icterica (Spix, 1824) SP Carb Atl 2 Rhinella magnussoni Lima, Menin & Araújo, 2007 - Ferr Ama 1 Rhinella margaritifera (Laurenti, 1768) MS Carb Cerr 1 Rhinella marina (Linnaeus, 1758) PA Ferr Cerr 2 Rhinella ornata (Spix, 1824) MG-SC Gran - Ferr - Quar Atl 7 Rhinella rubescens (Lutz, 1925) MG Ferr - Quar Atl - Ecot (Cerr - Atl) 7 Craugastoridae (S=4) Haddadus binotatus (Spix, 1824) MG Ferr - Quar - 2 Oreobates antrum Vaz-Silva, Maciel, Andrade & Amaro, 2018 GO-TO Carb Cerr 2 Oreobates heterodactylus (Miranda-Ribeiro, 1937) MT Carb Cerr 2 Pristimantis fenestratus (Steindachner , 1864) MT-PA Cong - Ferr Ama 5 Cycloramphidae (S=4) Cycloramphus eleutherodactylus (Miranda-Ribeiro, 1920) SP-PR Carbs Atl 6 Thoropa megatympanum Caramaschi & Sazima , 1984 MG Ferr - 2 Thoropa miliaris (Spix, 1824) MG Ferr - Gran - Quar - 6 Thoropa taophora (Miranda-Ribeiro, 1923) SP Marm - Quar - Gran Atl 3 Dendrobatidae (S=2) Adelphobates galactonotus (Steindachner, 1864) PA Ferr Ama 2 Ameerega flavopicta (Lutz, 1925) PA Aren - Ferr Ama 3 Hylidae (S=12) Boana albopunctata (Spix, 1824) MG Carb Ecot (Cerr - Atl) 1 Boana boans (Linnaeus, 1758) - Ferr Ama 1 Boana faber (Wied- Neuwied , 1821) MG Carb Ecot (Cerr - Atl) 1 Bokermannohyla hylax (Heyer, 1985) SP-SC Carb - Gran Atl 5 Bokermannohyla martinsi (Bokermann , 1964) MG Ferr Cerr - Atl - Ecot (Cerr - Atl) 15 Dendropsophus nanus ( Boulenger , 1889) - Aren Cerr 1 Phyllomedusa burmeisteri Boulenger , 1882 - Carb Atl 1 Scinax catharinae (Boulenger, 1888) SC Gran Atl 3 Scinax eurydice (Bokermann, 1968) MG Carb - 1 Scinax fuscovarius (Lutz, 1925) MG-MT Aren - Carb - Cong - Ferr Caa - Atl - Ecot (Cerr - Atl) 18 Scinax ruber (Laurenti, 1768) MS Carb Cerr 1 Scinax x- signatus (Spix, 1824) SE Carb Ecot (Caa - Atl) 6 Hylodidae (S=2) Hylodes cardosoi Lingnau , Canedo & Pombal, 2008 SP Carb Atl 1 Hylodes heyeri Haddad, Pombal & Bastos, 1996 SP Carb Atl 1 Leptodactylidae (S=13) Leptodactylus flavopictus Lutz, 1926 SP Carbs Atl 1 Leptodactylus labyrinthicus (Spix, 1824) MT-MG-RN Carb - Cong - Ferr - Marm Cerr - Caa 7 Leptodactylus latrans (Steffen, 1815) MG Carbs Ecot (Cerr - Atl) 1 Leptodactylus macrosternum Miranda-Ribeiro, 1926 - Carbs Cerr – Caa 2 Leptodactylus mystacinus (Burmeister, 1861) SP-MG Carb - Ferr Cerr - Atl 6 Leptodactylus paranaru Magalhães, Lyra, Carvalho, Baldo, Brusquetti, Burella, Colli, Gehara, Giaretta, Haddad, Langone, López, Napoli, Santana, de Sá, and Garda, 2020 SC Great Atl 1 Leptodactylus pentadactylus (Laurenti, 1768) PA Ferr Ama 2 Leptodactylus syphax Bokermann , 1969 - Carb Ama - Caa 2 Leptodactylus troglodytes Lutz, 1926 - Carb Caa 1 Leptodactylus vastus Lutz, 1930 SE Carb Atl - Ecot (Caa - Atl) 4 Physalaemus cuvieri Fitzinger, 1826 SE-BA-MG Carb - Ferr Caa - Cerr - Ecot (Caa - Atl) 7 Physalaemus ephippifer (Steindachner, 1864) - Ferr Ama 1 Physalaemus nattereri (Steindachner, 1863) MG Ferr - 1 Odontophrynidae (S=1) Proceratophrys boiei (Wied- Neuwied, 1824) - Gran Atl 1 Pipidae (S=1) Pipa carvalhoi (Miranda-Ribeiro, 1937) - Ferr Ama 1 The family with the greatest species richness was Leptodactylidae with 13 species (23.6% of the total sampled) followed by Hylidae with 12 species (21.8%) and Bufonidae with 10 species (18.2%). The families with the lowest richness were Aromobatidae, Odontophrynidae, and Pipidae with only one species each (Figure 1). The species distribution of anurans found in caves follows the general distribution of species within anuran families in Brazil. Hylidae is the family with the largest number of species (373 species) in the Brazilian region, a fact common to all Neotropical subregions. This family is characterized by mostly arboreal individuals with broad morphological, behavioral, and ecological niche plasticity (Duellman 1999, Cohen et al. 2020). Leptodactylidae also has a high richness in Brazil with 181 species (Segalla et al. 2021). The anurans of this family have terrestrial or semi-aquatic habits, ranging from species measuring 20 to 1500mm, which allows for a greater niche breadth (Cohen et al. 2020). Bufonidae in turn has 100 species in Brazil. It is made up of more robust species, with thicker integuments and more resistant to water loss (Cohen et al. 2020). The Bufonidade family has a higher proportion of species in caves (10%) concerning the richness found in Brazil than Leptodactylidae (7.2%) and Hylidae (3.2%). Species from the three families aforementioned present different behaviors and size ranges and are present in all vegetation types of the Neotropical region (Duellman 1999, Bernarde 2012, Cohen et al. 2020). Families with just one representative are smaller groupings when compared to families with more species. The Pipidae family has four species in the Brazilian neotropical region (1 recorded inside caves – Matavelli et al. 2015, Segalla et al., 2021). The members of this family have aquatic habits and a dorsoventrally flattened body. Odontophrynidae forms a group endemic to South America that encompasses 48 species (Segalla et al. 2021). In general, they are species with terrestrial habits and spend much of their time foraging in litter or underground, which can resemble a cave environment in abiotic conditions such as the absence of light or shading and relative humidity tending to saturation (Bernarde 2012). Aromobatidae is a family with 31 species present in the Amazon and Atlantic Forest biomes (Segalla et al. 2021). Individuals measure up to 35mm and have diurnal habits (Bernarde 2012), suggesting that caves may serve as a refuge from exposure to sunlight or favor foraging. Figure 1: Number of species per family of Brazilian neotropical anurans recorded from underground cavities. Ar = Aromobatidae; Br: Brachycephalidae; Bu: Bufonidae; Cr: Craugastoridae; Cy: Cycloramphidae; From: Dendrobatidae; Hy: Hylidae; Ho: Hylodidae; Le: Leptodactylidae; Od: Odontophrynidae and Pi: Pipidae. Records with identification up to the specific epithet totaled 177 independent records (71.7%). In 70 records (28.3%), individuals were classified by morphotype, in 233 records (94.3%) by genus, and in 240 (97.2%) by family. The difference between taxonomic identifications up to morphotype (including identification at the family or genus level) and those up to the specific epithet is one of the problems related to the Linnean gap in the study of cave anurofauna in Brazil. This is because many species have very similar phenotypic characteristics, which prevents accurate identification in the field, which is aggravated when there is no collection of specimens and subsequent comparison with animals deposited in scientific collections for taxonomic identification purposes. This is an obstacle to the knowledge of anuran taxonomy (Bernarde 2012; Haddad et al. 2013) the indication of photomarks or collection for deposit in scientific collections by legislative bodies may indicate better recognition of species and reinforce the theoretical framework of the Linnean Gap. Records by geographic variables The anuran record inside a Brazilian cavity was accompanied by data from the Federative Unit (state) in 184 records (74.5%; Figure 2) and from the municipality 182 times (73.7%). The other data referred to a geopolitical region of the country (for example: Brazilian Northeast) or lithology (for example: in a carbonate cave). Records of anurans in cave environments are most abundant, mainly in the states of Minas Gerais (81 records; 34.8%) and São Paulo (35; 15.0%). Comparing the percentage of records of anurans found in caves by the number of caves, São Paulo has 897 caves and Minas Gerais 2836, which demonstrates a higher rate of records in the state of São Paulo. These federative units have a high density of caves in some sub-regions, with tourist activities and research conducted for many years, such as the Alto Ribeira Tourist State Park in São Paulo, founded in 1958, and Minas Gerais parks such as the Ibitipoca State Park (of 1973), Cavernas do Peruaçu National Park (established in 1999) and the Peter Lund State Natural Monuments (2005) and Gruta do Rei do Mato (2009), with the Peter Lund monuments being historical because it is the region in which Lund lived and explored mining carbonate caves during the 19th century (Holten & Sterll 2017). Figure 2: Number of anuran records in natural cavities in Brazilian states. BA: Bahia; GO: Goiás; MG: Minas Gerais; MS: Mato Grosso do Sul; MT: Mato Grosso; PA: Pará; PR: Paraná; RN: Rio Grande do Norte; SC: Santa Catarina; SE: Sergipe; TO: Tocantins; ND: not described. In 99 records (40.1%) the natural cavity of the anuran found was identified, either by explicit information on the name of the cave in the text, by mentioning the geographic coordinate, or by consulting an official database (National Registry of Speleological Information – ICMBio / Cecav). The number of recorded cavities found in this study, when compared to the total number of recorded caves in Brazil in 2022, represents 0.24% of cavities with recorded anurans (Figure 3). The low number of caves with the presence of anurans does not reflect what was expected for a taxonomic group with ecophysiological affinities with this environment (Eterovick et al., 2010), since in work aimed at surveying anurans in cavities or monitoring the occupation of underground habitat, the encounter rate is high (Martins de Andrade et al. 2021, Bichuette et al. 2022). Figure 3: Caves according to the 2022 Cadastro Nacional de Informações Espeleológicas CANIE/Cecav (green dots) and with recorded anurans (yellow triangle) and caves in the Brazilian neotropical region. A – concentration caves in Rio Grande do Norte; B – concentration caves in Sergipe; C – concentration caves in Minas Gerais; D – concentration caves in São Paulo state and E – concentration caves in Santa Catarina. The predominant biomes of anuran records were the Atlantic Forest with 78 records (33.5%), the Atlantic Forest/Cerrado ecotonal zones with 43 (18.5%), and Caatinga with 16 occurrences (6.9%), in addition to 19 records in the Amazon (8.2%), 17 in the Caatinga (7.3%) and 24 in the Cerrado (10.3%). The predominance of anuran occurrence in cavities of the Atlantic Forest follows the distribution of species from this taxonomic group in Brazilian biomes. In other words, this biome has the greatest richness of amphibian species (543 species; Haddad et al. 2012) with high rates of endemism (88%; Haddad et al. 2012). The Atlantic Forest is characterized by closed forests, several bodies of water, and high rates of precipitation and air humidity, which make it a hotspot for anurofauna (Haddad et al. 2012, Bichuette et al. 2022), these biomes richest in cave anurans also form the landscape in the states with the most records (São Paulo and Minas Gerais) Records by environmental variables The presence/absence of deposited water or bodies of water inside the cavity was reported in 63 of the 233 total records (25.5%) with 47 classifieds as intermittent (20.2%), 15 as perennial (6.4%), and one record (0.4%) indicating the absence of water in the cavity. The photic zone of the cave in which the anuran was located was reported in 12 records (4.9% of the total), with nine of them pointing to the entrance zone (3.9%), one to the dysphotic zone (0.4%) and one in the aphotic zone (0.4%). The low recording rate of this type of variable partially demonstrates the gap in knowledge regarding the use of cavities by anurans as habitat. The presence of water influences the reproduction and feeding processes, the latter mainly in tadpoles, on the rate of loss of body water, that is, it is important information about the life history of anurans and a bias for their biospeleological classification (Eterovick et al. 2010, Matavelli et al. 2015, dos Santos et al. The luminosity categories considered constitute information that allows determining the use of cave space by anurans. For example, the occupation of the cave entrance area by anurans allows i) easy access to an escape exit or to forage in the external environment (mainly at night), ii ) obtaining food since the natural light incident along with the fact that the cave entrance region is an ecotone favors a high richness of organisms consumed by anurans, and iii ) a shaded environment that serves for the anurans to hide from predators and protect themselves against desiccation due to the heat of sunlight (Eterovick et al. 2010, Simões et al. The occupation of deeper areas with indirect incidence of light (dysphotic zone) or no light (aphotic zone) may indicate a refuge mainly against visually-oriented predators (Bernarde 2012) or a feeding site (dysphotic zone, personal communication Francisco L. Tejerina -Garro). Records by geological variable Information about the lithology of the caves was found in 246 records (99.6%; Figure 4), with a predominance in carbonate (109 records; 46.8%) and ferruginous (76; 33.9%) regions. On the other hand, conglomerates have a low number of registrations (6; 2.4%). Figure 4: Number of records by type of lithology Lithologies: Aren = sandstone; Carb = carbonate; Cong = conglomerate; Ferr = ferruginous; Gran = granitic; Quar = quartzite; Marm = marble; ND = Not available. The two most frequent lithologies in records are associated with the geology of the region (including the administrative area of one or more states) where the caves are located. In Brazil, areas with ferruginous characteristics are more frequent in the iron quadrangle in Minas Gerais, in regions with Cerrado-type vegetation cover, in ecotones between Cerrado/Atlantic Forest and the Carajás iron region in the state of Pará with Amazonian dominance. In turn, carbonate regions form large masses (groups) of much larger extensions that develop across several regions and different neotropical biomes, such as the Bambuí formation that passes through different biomes (Caatinga, Cerrado, Atlantic Forest, in addition to the ecotonal zones between them) and states (Minas Gerais, Goiás, Tocantins, Bahia; Rubiolli et al. 2019, Dos Santos 2022b). Furthermore, the predominance of records (109) in carbonate rocks found in this study is related to the higher rate of speleogenesis favored by this type of lithology (Rubioli et al. 2019). Records by type of scientific literature Records of anuran amphibians in Brazilian natural cavities are distributed in four publication formats: Complete (Full Article; 11 works; 61.1% of total works), Scientific notes (Short Communication; 4; 22.2%), book chapter (2; 11.1%), master's dissertation (1; 5.6%). Regarding the area of knowledge, most of the works (14 works; 77.8%) address community ecology, including in some cases other taxonomic groups in addition to anurans, that is, they cover endemic or troglobite species. In three publications (16.6%), population records are focused on one species: i) the permanence and occupation of ferruginous cavities of Bookermannolyna martinsi in Minas Gerais (Martins de Andrade et al. 2021), ii) in the vocalization activity of Cycloramphus eleutherodactylus in carbonate caves of São Paulo (Lima et al. 2012) and iii) in increasing the distribution and seasonal activity of the Oreobates frog antrum in carbonate caves in southeastern Tocantins (Motta et al. 2020). In the area of taxonomy, only one study (5.6%) recorded anurans inside cavities. This is a description of a species from Bolivia of the same genus as the Brazilian species Oreobates antrum; in the comparison made between the two species it is mentioned that O. antrum is found in natural underground cavities in the Brazilian Cerrado (Pansonato et al. 2020, Vaz-Silva et al. 2020). Since the first publication in 1980 that pointed out the presence of anurans in cavities by Dessen et al. (1980) until 2022, 42 years have passed. During this period, 18 scientific works were published (Figure 5). Between 1980 and 2008, four studies were published with records of anurans inside natural cavities in the Brazilian Neotropical region (22.2% of the total), one per year and spaced apart (5 to 10 years between publications). A specific and important milestone for the study of the underground environment was Normative 6640 of 2008, which specifies studies for the environmental licensing of natural cavities and studies of speleological relevance (BRASIL 2008). This situation and effect is notable when from 2009 to 2022, we have a total of 14 works (77.8%), including the first works in the area of populations and taxonomy. The last three years is the first period in which more than one work is published in this area, per year, demonstrating a recent understanding of the need for recording and scientific communication of the occupation and presence of anurans in Brazilian neotropical caves, added to the facts that globally we have had a significant increase in scientific publication and a greater number of researchers in Brazil. Figure 5: Number of scientific publications with records of anurans in Brazilian neotropical cavities per year from 1980 to 2022. Conclusion The study demonstrates a significant increase since 2008, the year in which legislation was implemented that provides for the environmental survey of Brazilian caves and their speleological classification for environmental licensing, as well as the increase in researchers and scientific articles worldwide. The records were concentrated in carbonate and ferruginous lithologies that have the largest number of caves in the region and have commercial appeal, mainly in the states of Minas Gerais and São Paulo and are in biomes of the Atlantic Forest and Cerrado (ecotonal zone). The families with more species recorded in caves follow those with greater richness in the surface environment, indicating that there is a maintenance of comparative richness and there is no apparent predilection. It is recommended to supplement studies with anurans in Brazilian neotropical caves in regions still without records to reinforce the list of occurrences and legal parameters for data collection, whether specimens for comparison and identification in reference collections or specialized photographs to assist in identifications to reduce the Linnean and Wallacean gaps. Acknowledgments We thank the Postgraduate Program in Ecology and Natural Resources at the Federal University of São Carlos. This work is partially financed by the Coordination of Improvement of Higher Education Personnel – Brazil (CAPES) – Code 001. Authors’ Contribution Vinícius F. Sperandei: contributed to data collection; data analysis and interpretation; and manuscript preparation. Willian Vaz-Silva: contribution to manuscript preparation; critical revision, adding intellectual content. Francisco L. 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ND PA Curionópolis Ferruginous ND ND ND 2022 Communities Full Article 18 5 Anura Brachycephalidae Ischnocnema guentheri ND MG Itabirito Ferruginous ND ND ND 2022 Communities Full Article 18 6 Anura Brachycephalidae Ischnocnema izecksohni ND MG Pains Carbonatic ND ND ND 2022 Communities Full Article 18 7 Anura Brachycephalidae Ischnocnema juipoca ND MG Moeda Ferruginous ND ND ND 2022 Communities Full Article 18 8 Anura Brachycephalidae Ischnocnema juipoca ND MG São Joaquim Ferruginous ND ND ND 2022 Communities Full Article 18 9 Anura Brachycephalidae Ischnocnema sp. ND MG Pains Carbonatic ND ND ND 2022 Communities Full Article 18 10 Anura Brachycephalidae Ischnocnema sp. ND MG São Joaquim Ferruginous ND ND ND 2022 Communities Full Article 18 11 Anura Brachycephalidae Ischnocnema sp.1 ND MG Pains Carbonatic ND Ecotone - Brazilian savanna / Atlantic Forest ND 2022 Communities Book chapter 17 12 Anura Brachycephalidae Ischnocnema sp.2 ND MG Pains Carbonatic ND ND ND 2022 Communities Full Article 18 13 Anura Brachycephalidae Ischnocnema sp.2 ND MG Pains Carbonatic ND Ecotone - Brazilian savanna / Atlantic Forest ND 2022 Communities Book chapter 17 14 Anura Brachycephalidae Ischnocnema sp.3 ND MG Pains Carbonatic ND Ecotone - Brazilian savanna / Atlantic Forest ND 2022 Communities Book chapter 17 15 Anura Brachycephalidae Ischnocnema juipoca ND ND ND Carbonatic ND Ecotone - Brazilian savanna / Atlantic Forest ND 2015 Communities Full Article 9 16 Anura Brachycephalidae Ischnocnema juipoca ND ND ND Conglomerate ND Ecotone - Brazilian savanna / Atlantic Forest ND 2015 Communities Full Article 9 17 Anura Brachycephalidae Ischnocnema juipoca ND ND ND Ferruginous ND Ecotone - Brazilian savanna / Atlantic Forest ND 2015 Communities Full Article 9 18 Anura Brachycephalidae Ischnocnema juipoca ND ND ND Quartizite ND Ecotone - Brazilian savanna / Atlantic Forest ND 2015 Communities Full Article 9 19 Anura Brachycephalidae Ischnocnema sp. ND ND ND Arenitic ND Brazilian savanna ND 2015 Communities Full Article 9 20 Anura Brachycephalidae Ischnocnema sp.1 ND ND ND Carbonatic ND Brazilian savanna ND 2015 Communities Full Article 9 21 Anura Brachycephalidae Ischnocnema sp.2 ND ND ND Granitic ND Atlantic Forest ND 2015 Communities Full Article 9 22 Anura Brachycephalidae Ischnocnema sp.3 ND ND ND Quartizite ND Ecotone - Brazilian savanna / Atlantic Forest ND 2015 Communities Full Article 9 23 Anura Brachycephalidae Ischnocnema sp.4 ND ND ND Quartizite ND Ecotone - Brazilian savanna / Atlantic Forest ND 2015 Communities Full Article 9 24 Anura Brachycephalidae Ischnocnema sp.5 ND ND ND Ferruginous ND Ecotone - Brazilian savanna / Atlantic Forest ND 2015 Communities Full Article 9 25 Anura Brachycephalidae Ischnocnema manezinho Caverna Lage SC Florianópolis Granitic ND Atlantic Forest Perennial 2022 Communities Short communication 16 26 Anura Brachycephalidae Ischnocnema manezinho Caverna Monte Verde SC Florianópolis Granitic ND Atlantic Forest Intermittent 2022 Communities Short communication 16 27 Anura Brachycephalidae Ischnocnema manezinho Caverna Monte Verde SC Florianópolis Granitic ND Atlantic Forest Intermittent 2022 Communities Short communication 16 28 Anura Brachycephalidae Ischnocnema manezinho Caverna Pedras Grandes SC Florianópolis Granitic ND Atlantic Forest Perennial 2022 Communities Short communication 16 29 Anura Brachycephalidae Ischnocnema manezinho Caverna Praia Brava SC Florianópolis Granitic ND Atlantic Forest Perennial 2022 Communities Short communication 16 30 Anura Brachycephalidae Ischnocnema manezinho Caverna Saco dos Limões SC Florianópolis Granitic ND Atlantic Forest Ausente 2022 Communities Short communication 16 31 Anura Brachycephalidae Ischnocnema manezinho Sistema Água Corrente SC Florianópolis Granitic ND Atlantic Forest Perennial 2022 Communities Short communication 16 32 Anura Bufonidae Rhinella ornata ND MG ND Ferruginous ND Atlantic Forest ND 2021 Communities Full Article 13 33 Anura Bufonidae Rhinella rubescens ND MG ND Ferruginous ND Atlantic Forest ND 2021 Communities Full Article 13 34 Anura Bufonidae Rhinella crucifer ND MG Pains Carbonatic ND Ecotone - Brazilian savanna / Atlantic Forest ND 2022 Communities Book chapter 17 35 Anura Bufonidae Rhinella diptycha ND MG Arcos Carbonatic ND ND ND 2022 Communities Full Article 18 36 Anura Bufonidae Rhinella diptycha ND MG Montes Claros Carbonatic ND ND ND 2022 Communities Full Article 18 37 Anura Bufonidae Rhinella diptycha ND MG Pains Carbonatic ND ND ND 2022 Communities Full Article 18 38 Anura Bufonidae Rhinella diptycha ND MG Pains Carbonatic ND Ecotone - Brazilian savanna / Atlantic Forest ND 2022 Communities Book chapter 17 39 Anura Bufonidae Rhinella ornata ND MG Conceição do Mato Dentro Ferruginous ND ND ND 2022 Communities Full Article 18 40 Anura Bufonidae Rhinella ornata ND MG Itabirito Ferruginous ND ND ND 2022 Communities Full Article 18 41 Anura Bufonidae Rhinella ornata ND MG Mariana Ferruginous ND ND ND 2022 Communities Full Article 18 42 Anura Bufonidae Rhinella ornata ND MG Dores de Campos Granitic ND ND ND 2022 Communities Full Article 18 43 Anura Bufonidae Rhinella ornata ND MG Congonhas Quartizite ND ND ND 2022 Communities Full Article 18 44 Anura Bufonidae Rhinella rubescens ND MG Bárbara Ferruginous ND ND ND 2022 Communities Full Article 18 45 Anura Bufonidae Rhinella rubescens ND MG Itabirito Ferruginous ND ND ND 2022 Communities Full Article 18 46 Anura Bufonidae Rhinella rubescens ND MG Mariana Ferruginous ND ND ND 2022 Communities Full Article 18 47 Anura Bufonidae Rhinella rubescens ND MG Moeda Ferruginous ND ND ND 2022 Communities Full Article 18 48 Anura Bufonidae Rhinella rubescens ND MG Morro do Pilar Ferruginous ND ND ND 2022 Communities Full Article 18 49 Anura Bufonidae Rhinella margaritifera Gruta de São Miguel MS Bonito Carbonatic ND Brazilian savanna ND 2001 Communities Book chapter 4 50 Anura Bufonidae Rhinella crucifer ND ND ND Granitic ND Atlantic Forest ND 2015 Communities Full Article 9 51 Anura Bufonidae Rhinella diptycha ND ND ND Arenitic ND Brazilian savanna ND 2015 Communities Full Article 9 52 Anura Bufonidae Rhinella granulosa ND ND ND Carbonatic ND Caatinga ND 2015 Communities Full Article 9 53 Anura Bufonidae Rhaebo guttatus ND ND ND Arenitic ND Amazon ND 2015 Communities Full Article 9 54 Anura Bufonidae Rhaebo guttatus ND ND ND Ferruginous ND Amazon ND 2015 Communities Full Article 9 55 Anura Bufonidae Rhinella magnussoni ND ND ND Ferruginous ND Amazon ND 2015 Communities Full Article 9 56 Anura Bufonidae Rhinella marina ND ND ND Ferruginous ND Brazilian savanna ND 2015 Communities Full Article 9 57 Anura Bufonidae Rhinella rubescens ND ND ND Quartizite ND Ecotone - Brazilian savanna / Atlantic Forest ND 2015 Communities Full Article 9 58 Anura Bufonidae Rhinella sp. ND ND ND Carbonatic ND Atlantic Forest ND 2015 Communities Full Article 9 59 Anura Bufonidae Rhinella sp.1 ND ND ND Carbonatic ND Atlantic Forest ND 2015 Communities Full Article 9 60 Anura Bufonidae Rhaebo guttatus ND PA Curionópolis Ferruginous ND ND ND 2022 Communities Full Article 18 61 Anura Bufonidae Rhinella marina ND PA Curionópolis Ferruginous ND ND ND 2022 Communities Full Article 18 62 Anura Bufonidae Rhinella sp.1 Caverna Apartamento de Pedra RN Martins Marble ND Caatinga ND 2017 Communities Full Article 10 63 Anura Bufonidae Rhinella sp.1 Caverna Dorminhoco RN Martins Marble ND Caatinga ND 2017 Communities Full Article 10 64 Anura Bufonidae Rhinella sp.1 Caverna Três Inchus RN Martins Marble ND Caatinga ND 2017 Communities Full Article 10 65 Anura Bufonidae Rhinella sp.2 Caverna Apartamento de Pedra RN Martins Marble ND Caatinga ND 2017 Communities Full Article 10 66 Anura Bufonidae Rhinella ornata cf. Caverna Lage SC Florianópolis Granitic ND Atlantic Forest Perennial 2022 Communities Short communication 16 67 Anura Bufonidae Rhinella diptycha Caverna Casa de Pedra SE Itabaiana Carbonatic ZE Atlantic Forest ND 2009 Communities Short communication 5 68 Anura Bufonidae Rhinella crucifer Abismo das Ossadas (SP-133) SP Iporanga Carbonatic ND Atlantic Forest ND 1980 Communities Full Article 1 69 Anura Bufonidae Rhinella sp. Caverna do Betari SP Iporanga Carbonatic ND Atlantic Forest ND 1980 Communities Full Article 1 70 Anura Bufonidae Rhinella sp. Gruta Itambé (SP-179) SP Altinópolis Arenitic ZE Atlantic Forest Intermittent 1986 Communities Full Article 2 71 Anura Bufonidae Rhinella sp. Gruta Olho de Cabra (SP-178) SP Altinópolis Arenitic ZE Atlantic Forest Intermittent 1986 Communities Full Article 2 72 Anura Bufonidae Rhinella sp. Gruta dos Paiva (SP-42) SP Iporanga Carbonatic ND Atlantic Forest Intermittent 1991 Communities Full Article 3 73 Anura Bufonidae Rhinella icterica ND SP Iporanga Carbonatic ND Atlantic Forest ND 2010 Communities Full Article 6 74 Anura Bufonidae Rhinella icterica ND SP Iporanga Carbonatic ND Atlantic Forest ND 2010 Communities Full Article 6 75 Anura Ceratophryidae Ceratophrys sp. Caverna Marreca (SP-50) SP Iporanga Carbonatic ND Atlantic Forest ND 1980 Communities Full Article 1 76 Anura Ceratophryidae Ceratophrys cf. sp. Gruta do Tufo (SP-248) SP Iporanga Carbonatic ND Atlantic Forest Perennial 1991 Communities Full Article 3 77 Anura Craugastoridae Oreobates antrum Gruta dos Revolucionários GO Posse Carbonatic ND Brazilian savanna ND 2020 Population Short communication 11 78 Anura Craugastoridae Haddadus binotatus ND MG São Joaquim Ferruginous ND ND ND 2022 Communities Full Article 18 79 Anura Craugastoridae Haddadus binotatus ND MG Congonhas Quartizite ND ND ND 2022 Communities Full Article 18 80 Anura Craugastoridae Oreobates heterodactylus Caverna do Cadeado MT Cáceres Carbonatic ND Brazilian savanna ND 2020 Taxonomy Full Article 12 81 Anura Craugastoridae Oreobates heterodactylus Gruta Fazendinha MT Cáceres Carbonatic ND Brazilian savanna ND 2020 Taxonomy Full Article 12 82 Anura Craugastoridae Oreobates heterodactylus Gruta Fazendinha MT Cáceres Carbonatic ND Brazilian savanna ND 2020 Taxonomy Full Article 12 83 Anura Craugastoridae Pristimantis fenestratus ND MT Primavera do Leste Conglomerate ND ND ND 2022 Communities Full Article 18 84 Anura Craugastoridae Pristimantis fenestratus ND ND ND Ferruginous ND Amazon ND 2015 Communities Full Article 9 85 Anura Craugastoridae Pristimantis fenestratus ND PA Parauapebas Ferruginous ND Amazon ND 2021 Communities Full Article 14 86 Anura Craugastoridae Pristimantis fenestratus ND PA Curionópolis Ferruginous ND ND ND 2022 Communities Full Article 18 87 Anura Craugastoridae Pristimantis fenestratus ND PA Curionópolis Ferruginous ND ND ND 2022 Communities Full Article 18 88 Anura Craugastoridae Oreobates antrum Gruta das Rãs TO Aurora do Tocantins Carbonatic ND Brazilian savanna ND 2020 Population Short communication 11 89 Anura Cycloramphidae Thoropa megatympanum ND MG Moeda Ferruginous ND ND ND 2022 Communities Full Article 18 90 Anura Cycloramphidae Thoropa megatympanum ND MG Morro do Pilar Ferruginous ND ND ND 2022 Communities Full Article 18 91 Anura Cycloramphidae Thoropa miliaris ND MG Conceição do Mato Dentro Ferruginous ND ND ND 2022 Communities Full Article 18 92 Anura Cycloramphidae Thoropa miliaris ND MG Itabirito Ferruginous ND ND ND 2022 Communities Full Article 18 93 Anura Cycloramphidae Thoropa miliaris ND MG Antônio Dias Granitic ND ND ND 2022 Communities Full Article 18 94 Anura Cycloramphidae Thoropa miliaris ND MG Dores de Guanhães Granitic ND ND ND 2022 Communities Full Article 18 95 Anura Cycloramphidae Thoropa miliaris ND MG Virginópolis Granitic ND ND ND 2022 Communities Full Article 18 96 Anura Cycloramphidae Thoropa miliaris ND MG Congonhas Quartizite ND ND ND 2022 Communities Full Article 18 97 Anura Cycloramphidae Thoropa taophora ND ND ND Granitic ND Atlantic Forest ND 2015 Communities Full Article 9 98 Anura Cycloramphidae Thoropa taophora ND ND ND Marble ND Atlantic Forest ND 2015 Communities Full Article 9 99 Anura Cycloramphidae Thoropa taophora ND ND ND Quartizite ND Atlantic Forest ND 2015 Communities Full Article 9 100 Anura Cycloramphidae Cycloramphus eleutherodactylus Caverna Fazenda Morungava PR Sengés ND ND Atlantic Forest ND 2012 Population Short communication 8 101 Anura Cycloramphidae Cycloramphus sp. Caverna Ouro Grosso (SP-54) SP Iporanga Carbonatic ND Atlantic Forest ND 1980 Communities Full Article 1 102 Anura Cycloramphidae Cycloramphus sp. Caverna Santana (SP-41) SP Iporanga Carbonatic ND Atlantic Forest ND 1980 Communities Full Article 1 103 Anura Cycloramphidae Cycloramphus sp. Caverna Ouro Grosso (SP-54) SP Iporanga Carbonatic ND Atlantic Forest Intermittent 1986 Communities Full Article 2 104 Anura Cycloramphidae Cycloramphus sp. Gruta do Chapéu (SP-13) SP Apiaí Carbonatic ND Atlantic Forest Intermittent 1991 Communities Full Article 3 105 Anura Cycloramphidae Cycloramphus eleutherodactylus Caboclo (Centro/Estação) SP Iporanga Carbonatic ND Atlantic Forest ND 2010 Communities Full Article 6 106 Anura Cycloramphidae Cycloramphus eleutherodactylus Caverna Cafezal SP Iporanga Carbonatic ND Atlantic Forest ND 2010 Communities Full Article 6 107 Anura Cycloramphidae Cycloramphus eleutherodactylus Ouro Grosso (Centro/Estação) SP Iporanga Carbonatic ND Atlantic Forest ND 2010 Communities Full Article 6 108 Anura Cycloramphidae Cycloramphus eleutherodactylus Caverna Cafezal SP Iporanga Carbonatic ND Atlantic Forest Intermittent 2012 Population Short communication 8 109 Anura Cycloramphidae Cycloramphus eleutherodactylus Caverna Morro Preto (SP-21) SP Iporanga Carbonatic ND Atlantic Forest Intermittent 2012 Population Short communication 8 110 Anura Dendrobatidae Ameerega flavopicta ND ND ND Arenitic ND Amazon ND 2015 Communities Full Article 9 111 Anura Dendrobatidae Ameerega flavopicta ND ND ND Ferruginous ND Amazon ND 2015 Communities Full Article 9 112 Anura Dendrobatidae Adelphobates galactonotus ND ND ND Ferruginous ND Amazon ND 2015 Communities Full Article 9 113 Anura Dendrobatidae Ameerega flavopicta ND PA Curionópolis Ferruginous ND ND ND 2022 Communities Full Article 18 114 Anura Dendrobatidae Adelphobates galactonotus ND PA Curionópolis Ferruginous ND ND ND 2022 Communities Full Article 18 115 Anura Hylidae ND sp. Conjunto São Mateus - Imbira (GO-11) GO São Domingos Carbonatic ND Brazilian savanna ND 1980 Communities Full Article 1 116 Anura Hylidae Scinax fuscovarius SG-030 MG Santa Bárbara Ferruginous ND Atlantic Forest ND 2021 Communities Full Article 13 117 Anura Hylidae Bokermannohyla martinsi RM 33 MG Belo Horizonte Ferruginous ND Ecotone - Brazilian savanna / Atlantic Forest ND 2021 Communities Full Article 13 118 Anura Hylidae Bokermannohyla martinsi Gand-0008 MG Rio Acima Ferruginous ND Atlantic Forest ND 2021 Communities Full Article 13 119 Anura Hylidae Bokermannohyla martinsi Gand-0096 MG Rio Acima Ferruginous ND Atlantic Forest ND 2021 Communities Full Article 13 120 Anura Hylidae Bokermannohyla martinsi SG-051 MG Rio Acima Ferruginous ND Atlantic Forest ND 2021 Communities Full Article 13 121 Anura Hylidae Bokermannohyla martinsi Gand_0008 MG Nova Lima Ferruginous ND Brazilian savanna Intermittent 2021 Population Full Article 15 122 Anura Hylidae Bokermannohyla martinsi Gand_0056 MG Nova Lima Ferruginous ND Brazilian savanna Intermittent 2021 Population Full Article 15 123 Anura Hylidae Bokermannohyla martinsi Gand_0096 MG Nova Lima Ferruginous ND Brazilian savanna Intermittent 2021 Population Full Article 15 124 Anura Hylidae Bokermannohyla martinsi Apol_25 MG Nova Lima Ferruginous ND Atlantic Forest Intermittent 2021 Population Full Article 15 125 Anura Hylidae Bokermannohyla martinsi SG_016 MG Nova Lima Ferruginous ND Atlantic Forest Intermittent 2021 Population Full Article 15 126 Anura Hylidae Bokermannohyla martinsi SG_017 MG Nova Lima Ferruginous ND Atlantic Forest Intermittent 2021 Population Full Article 15 127 Anura Hylidae Bokermannohyla martinsi SG_018 MG Nova Lima Ferruginous ND Atlantic Forest Intermittent 2021 Population Full Article 15 128 Anura Hylidae Bokermannohyla martinsi SG_019 MG Nova Lima Ferruginous ND Atlantic Forest Intermittent 2021 Population Full Article 15 129 Anura Hylidae Bokermannohyla martinsi SG_020 MG Nova Lima Ferruginous ND Atlantic Forest Intermittent 2021 Population Full Article 15 130 Anura Hylidae Bokermannohyla martinsi SG_051 MG Nova Lima Ferruginous ND Atlantic Forest Intermittent 2021 Population Full Article 15 131 Anura Hylidae Boana albopunctata ND MG Pains Carbonatic ND Ecotone - Brazilian savanna / Atlantic Forest ND 2022 Communities Book chapter 17 132 Anura Hylidae Scinax eurydice ND MG Montes Claros Carbonatic ND ND ND 2022 Communities Full Article 18 133 Anura Hylidae Boana faber ND MG Pains Carbonatic ND Ecotone - Brazilian savanna / Atlantic Forest ND 2022 Communities Book chapter 17 134 Anura Hylidae Scinax fuscovarius ND MG Arcos Carbonatic ND ND ND 2022 Communities Full Article 18 135 Anura Hylidae Scinax fuscovarius ND MG Iguatama Carbonatic ND ND ND 2022 Communities Full Article 18 136 Anura Hylidae Scinax fuscovarius ND MG Matozinhos Carbonatic ND ND ND 2022 Communities Full Article 18 137 Anura Hylidae Scinax fuscovarius ND MG Montes Claros Carbonatic ND ND ND 2022 Communities Full Article 18 138 Anura Hylidae Scinax fuscovarius ND MG Pains Carbonatic ND ND ND 2022 Communities Full Article 18 139 Anura Hylidae Scinax fuscovarius ND MG Pedro Leopoldo Carbonatic ND ND ND 2022 Communities Full Article 18 140 Anura Hylidae Scinax fuscovarius ND MG Conceição do Mato Dentro Ferruginous ND ND ND 2022 Communities Full Article 18 141 Anura Hylidae Scinax fuscovarius ND MG Igarapé Ferruginous ND ND ND 2022 Communities Full Article 18 142 Anura Hylidae Scinax fuscovarius ND MG Itabirito Ferruginous ND ND ND 2022 Communities Full Article 18 143 Anura Hylidae Scinax fuscovarius ND MG Mariana Ferruginous ND ND ND 2022 Communities Full Article 18 144 Anura Hylidae Scinax fuscovarius ND MG Pains Carbonatic ZE Ecotone - Brazilian savanna / Atlantic Forest ND 2022 Communities Book chapter 17 145 Anura Hylidae Scinax fuscovarius ND MG Pains Carbonatic ZD Ecotone - Brazilian savanna / Atlantic Forest ND 2022 Communities Book chapter 17 146 Anura Hylidae Scinax fuscovarius ND MG Pains Carbonatic ZA Ecotone - Brazilian savanna / Atlantic Forest ND 2022 Communities Book chapter 17 147 Anura Hylidae Scinax ruber Gruta de São Miguel MS Bonito Carbonatic ZE Brazilian savanna ND 2001 Communities Book chapter 4 148 Anura Hylidae Scinax fuscovarius ND MT Primavera do Leste Arenitic ND ND ND 2022 Communities Full Article 18 149 Anura Hylidae Scinax fuscovarius ND MT Primavera do Leste Conglomerate ND ND ND 2022 Communities Full Article 18 150 Anura Hylidae Boana boans ND ND ND Ferruginous ND Amazon ND 2015 Communities Full Article 9 151 Anura Hylidae Phyllomedusa burmeisteri ND ND ND Carbonatic ND Atlantic Forest ND 2015 Communities Full Article 9 152 Anura Hylidae Scinax fuscovarius ND ND ND Carbonatic ND Caatinga ND 2015 Communities Full Article 9 153 Anura Hylidae Scinax fuscovarius ND ND ND Ferruginous ND Ecotone - Brazilian savanna / Atlantic Forest ND 2015 Communities Full Article 9 154 Anura Hylidae Bokermannohyla martinsi ND ND ND Ferruginous ND Ecotone - Brazilian savanna / Atlantic Forest ND 2015 Communities Full Article 9 155 Anura Hylidae Dendropsophus nanus ND ND ND Arenitic ND Brazilian savanna ND 2015 Communities Full Article 9 156 Anura Hylidae Bokermannohyla sp. ND ND ND Ferruginous ND Ecotone - Brazilian savanna / Atlantic Forest ND 2015 Communities Full Article 9 157 Anura Hylidae Bokermannohyla sp.1 ND ND ND Ferruginous ND Ecotone - Brazilian savanna / Atlantic Forest ND 2015 Communities Full Article 9 158 Anura Hylidae Scinax catharinae Caverna Pedras Grandes SC Florianópolis Granitic ND Atlantic Forest Perennial 2022 Communities Short communication 16 159 Anura Hylidae Scinax catharinae Caverna Praia Brava SC Florianópolis Granitic ND Atlantic Forest Perennial 2022 Communities Short communication 16 160 Anura Hylidae Scinax catharinae Sistema Água Corrente SC Florianópolis Granitic ND Atlantic Forest Perennial 2022 Communities Short communication 16 161 Anura Hylidae Bokermannohyla hylax Caverna Saco Grande SC Florianópolis Granitic ND Atlantic Forest Perennial 2022 Communities Short communication 16 162 Anura Hylidae Bokermannohyla hylax Caverna Saco Grande SC Florianópolis Granitic ND Atlantic Forest Perennial 2022 Communities Short communication 16 163 Anura Hylidae Bokermannohyla hylax Sistema Água Corrente SC Florianópolis Granitic ND Atlantic Forest Perennial 2022 Communities Short communication 16 164 Anura Hylidae Bokermannohyla hylax Sistema Água Corrente SC Florianópolis Granitic ND Atlantic Forest Perennial 2022 Communities Short communication 16 165 Anura Hylidae Scinax x-signatus Gruta da Janela SE Laranjeiras Carbonatic ND Ecotone Caatinga / Atlantic Forest Intermittent 2011 Communities Master’s Thesis 7 166 Anura Hylidae Scinax x-signatus Gruta da Pedra Furada SE Laranjeiras Carbonatic ND Ecotone Caatinga / Atlantic Forest Intermittent 2011 Communities Master’s Thesis 7 167 Anura Hylidae Scinax x-signatus Gruta da Raposa SE Laranjeiras Carbonatic ND Ecotone Caatinga / Atlantic Forest Intermittent 2011 Communities Master’s Thesis 7 168 Anura Hylidae Scinax x-signatus Gruta da Raposinha SE Laranjeiras Carbonatic ND Ecotone Caatinga / Atlantic Forest Intermittent 2011 Communities Master’s Thesis 7 169 Anura Hylidae Scinax x-signatus Gruta do Tramandaí SE Laranjeiras Carbonatic ND Ecotone Caatinga / Atlantic Forest Intermittent 2011 Communities Master’s Thesis 7 170 Anura Hylidae Scinax x-signatus Gruta dos Aventureiros SE Laranjeiras Carbonatic ND Ecotone Caatinga / Atlantic Forest Perennial 2011 Communities Master’s Thesis 7 171 Anura Hylidae ND sp. Gruta Colorida (SP-129) SP Iporanga Carbonatic ZE Atlantic Forest Intermittent 1991 Communities Full Article 3 172 Anura Hylidae ND sp. Gruta do Bocão (SP-241) SP Iporanga Carbonatic ND Atlantic Forest Intermittent 1991 Communities Full Article 3 173 Anura Hylidae Bokermannohyla hylax Caverna Chapéu Mirim II (SP-15) SP Apiaí Carbonat