#ZooKeys ZooKeys 1217: 215-246 (2024) DOI: 10.3897/zookeys.1217.119844 Research Article Uncovering the rich amphibian fauna of two semideciduous forest fragments in southwestern Bahia, Brazil Carlos Augusto Souza-Costa’, Mirco Solé’®, Caio Vinicius de Mira-Mendes?4®, Anténio Jorge Suzart Argélo’, luri Ribeiro Dias’® ee wo NY — Graduate Program in Zoology, Universidade Estadual de Santa Cruz, Rodovia Jorge Amado, km 16, 45662-900, Ilhéus, Bahia, Brazil Museum Koenig Bonn (ZFMK), Leibniz Institute for the Analysis of Biodiversity Change, Adenauerallee 160, 53113 Bonn, Germany Graduate Program in Tropical Aquatic Systems, Universidade Estadual de Santa Cruz, Rodovia Jorge Amado, km 16, 45662-900 Ilhéus, Bahia, Brazil Department of Biology, Universidade Estadual do Maranhao, Sao Luis, Maranhao 65055-310, Brazil Corresponding author: luri Ribeiro Dias (iurirdias@hotmail.com) OPEN Qaccess Academic editor: Angelica Crottini Received: 19 March 2024 Accepted: 10 July 2024 Published: 4 November 2024 ZooBank: https://zoobank. org/8A9A02E3-392C-49B3-9131- BF228F99ED2A Citation: Souza-Costa CA, Sole M, Vinicius de Mira-Mendes C, Argdlo AJS, Ribeiro Dias | (2024) Uncovering the rich amphibian fauna of two semideciduous forest fragments in southwestern Bahia, Brazil. Zookeys 1217: 215-246. https://doi. org/10.3897/zookeys.1217.119844 Copyright: © Carlos Augusto Souza-Costa et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0). Abstract Fauna inventories reduce biodiversity knowledge gaps by providing comprehensive data on species distribution, richness, and abundance. Furthermore, they identify undocu- mented species and enhance understanding of ecosystem dynamics and conservation needs. The richness and abundance of amphibian species were studied in two Semide- ciduous Seasonal Forest areas in the municipalities of Potiragua (Serra Azul) and Itaran- tim (Serra do Mandim) in southwestern Bahia, Brazil. Active visual and acoustic surveys were conducted in 24 forest interior transects, two stream transects, and two permanent ponds investigated in the study area. Opportunistic encounters during team movements were also recorded. The richness was 46 amphibian species distributed in 14 families and 26 genera. Approximately half of the species were shared between the two areas, while 11 species were exclusive to Serra Azul and another nine were found only in Serra do Mandim. Cluster analysis for 42 locations in Atlantic Forest, Caatinga, and Cerrado, in a presence/absence matrix with 216 species, revealed that the composition of the am- phibians found in Serra do Mandim and Serra Azul is similar to other sampled locations in the northeastern region of Minas Gerais, close to the study site, which are considered transitional between the Atlantic Forest and the Caatinga. Our results demonstrate that the remaining forest fragments in the region, although small and isolated, still sustain a high richness of amphibians with species restricted to the Atlantic Forest and Bahia, such as Bahius bilineatus, Ololygon strigilata, Aplastodiscus weygoldti and Vitreorana eurygnatha, and others considered typical of the Caatinga, such as Leptodactylus trog- lodytes and Physalaemus cicada. Additionally, we sampled potential new species, filled occurrence gaps, and expanded the geographical range of Pseudis fusca. Key words: Anura, biodiversity, inventory, species distribution, species richness, transi- tional forest Introduction Amphibians are considered good environmental indicators due to their perme- able skin, exposed eggs and embryos, and generally biphasic life cycle, allow- ing these organisms to respond to disturbances in both terrestrial and aquatic PALES: Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil ecosystems (Wells 2007; Da-Silva et al. 2012; Fonte et al. 2019). Additionally, climate change can affect them, and studies demonstrate that some communi- ties in the Neotropical region are already close to their physiological temperature limits (Duarte et al. 2012; Gutiérrez-Pesquera et al. 2016; Carilo-Filho et al. 2021). The greatest threat to amphibians and fauna in general is habitat loss and fragmentation, which reduces shelter availability, food supply, isolates popula- tions, and affects their genetic variability (Young et al. 2004; Becker et al. 2007). Amphibians stand as the most endangered class of vertebrates, with 40.7% of their species at risk of extinction (Luedtke et al. 2023). Although the proportion of species classified as Data Deficient (DD) has decreased in the most recent Global Amphibian Assessment (from 22.5% to 11.3%), the high number of spe- cies still listed as DD poses a challenge for researchers and hinders effective conservation efforts (Hoffmann et al. 2010; Luedtke et al. 2023). The Atlantic Forest, originally spanning approximately 1.3 million km?, has undergone significant reduction, with estimates suggesting that only between 11.4% and 16% of its original coverage remains (Ribeiro et al. 2009). Neverthe- less, the forest remnants still house an exuberant biological diversity, including endemic and threatened species, as well as species with restricted distribution to specific ecosystems (Myers et al. 2000; Haddad et al. 2013; Zappi et al. 2015). In the southwest of Bahia, the Atlantic Forest is mainly composed of the Sea- sonal Forest (Deciduous and Semideciduous), which connects to interior for- ests, such as Caatinga and Cerrado (SOS MATA ATLANTICA and INPE 2018). This forest formation has physical and biological characteristics of adjacent regions, allowing faunal elements from other ecosystems to occur in these ar- eas (Willians 1996). Nevertheless, even with the damage caused by human ac- tivities, such as pasture creation, logging, and mining (Miles et al. 2006; Silva et al. 2006), few protected areas have been established in the region, such as the REBIO (Biological reserve) Mata Escura and Alto Cariri National Park (ICMBio 2003, 2010) and the RPPN (Private Reserve of Natural Heritage) Mata do Pas- sarinho (ICMBio 2016). Amphibian surveys in Bahia have revealed significant species richness (e.g., Rojas-Padilla et al. 2020; Protazio et al. 2021; Bastos and Zina 2022), with re- cords of new species (e.g., V6rds et al. 2017; Zucchetti et al. 2023; Santos et al. 2023) and expansions of the geographic distribution (e.g., Dias et al. 2010; Dias et al. 2011; Almeida et al. 2022). Interestingly, a family previously known only from the Amazon was recorded in the state of Bahia through the description of a new species (Caramaschiet al. 2013). Inventories contribute to the knowledge of species richness of a given region, as well as the understanding of the functional structure and population dynamics of amphibians (Droege et al. 1998; Hadd- ad 1998; Camardelli and Napoli 2012). These studies are essential for planning conservation decisions and policies aimed at mitigating anthropogenic effects on species and for the creation of strategic areas for environmental protection (Silvano and Segalla 2005; IUCN 2024). Moreover, they assist in gathering infor- mation that enables the reduction of gaps in the distribution and composition of the anurofauna in the country (Rodrigues 2003; Tabarelli and Silva 2003). Thus, the aim of this study was to conduct an inventory of the amphibians in two remaining Semideciduous Forests in the southwest of Bahia, comparing the amphibian community of these remnants with others from the Atlantic For- est, Caatinga, and Cerrado. ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 216 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil Data World Imagery and SRTM 4 - 2015 Datum WGS1984 Universidade de Santa Cruz Lab. Climatolgy 82°80'00 82°60'00 82°40°00 Materials and methods Study area The study was carried out in two fragments of Atlantic Forest in the Southwest region of Bahia: Fugiama farm (15°37'58"S, 39°59'01"W), with approximately 120 hectares of forest, located in the Serra do Mandim, municipality of Itaran- tim, and Serra Azul farm (15°52'01"S, 39°55'54"W), with about 160 hectares of forest, located in the Serra Azul, municipality of Potiragua, both located in the state of Bahia (Figs 1, 2). While the mountains themselves reach up to approxi- mately 1100 m in altitude, the areas accessed during the study were at around 800 m in altitude (Table 1). The region borders the state of Minas Gerais and is located between two neighboring basins: the Pardo River basin and the Jequitinhonha River basin (IBGE 1997). Both areas are inserted in the Phytogeographic Domain of the Semideciduous Forest, characterized by the presence of climate type Am, with one to three dry months (K6ppen 1936) and are ~ 28 km apart from each other. The climatic characteristics of the region encompass humid and sub-humid climates, defined by rainfall between 800 and 1100 mm and thermal averages from 23.5 to 25 °C (IBGE 1997). The vegetation of the study area is characterized as Semideciduous Forest (SOS MATA ATLANTICA and INPE 2018). It occurs latitudinally parallel to the formation zone of the Ombrophilous Forest (coastal), at a distance ranging from 20 km to 140 km from the coast (Mori and Silva 1980; Argdlo 2004). 37°00'00 | 39°00'00 42°23'22 Figure 1. Study areas in the southwest region of Bahia, Brazil. Serra do Mandim belongs to the municipality of Itarantim, Bahia and Serra Azul, one of its portions inserted in the municipality of Potiragua, Bahia. ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 217 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil Figure 2. Study areas in the southwest region of Bahia. Fugiama farm in Serra do Mandim (a, b, c) a semideciduous forest fragment b stream c permanent pond. Serra Azul farm in Serra Azul (d, e, f) d semideciduous forest fragment e stream f permanent pond. Table 1. Sampling points of the amphibian survey, coordinates, altitude, and sampling methods in Serra do Mandim, municipality of Itarantim and Serra Azul, municipality of Potiragua, state of Bahia. TF = Transects in the forest; TS = Tran- sects in the streams; P = permanent ponds. Locality | Sampling points Coordinates (Latitude; Longitude) | Altitude (m) Sampling method Serra do Mandim-BA 01 | 15°37'39.2'S, 39°58'41.6"W | 728m TF 02 = 15°37'38.9'"S, 39°58'39.2"W | 704m TF 03 | 15°37'40.8"S, 39°58'37.5"W 672m TF 4 8 | 15°37'47.9'S, 39°58'35.2"W 584m TF 05 15°37'50.4'S, 39°5902.2"W sds 758m | TF 06 | 15°37'49.5"S, 39°58'59.9"W 681m TF 07 15°37'47.6"S, 39°58'57.9"W 635m TF 08 15°37'47.8'S, 39°58'50.7"W 560 m le 09 | 15°37'47.6'S, 39°58'48.9"W 587m TF 10 15°37'47.5'S, 39°58'46.1"W 574m TF A 15°37'36.9'S, 39°58'43.5"W 755m | le 12 | 15°37'51.4"S, 39°58'35.9"W 513m TF 25 15°37'54.1'S, 39°58'34.0"W 485m TS 27 15°39'15.4"S, 39°59'00.8"W | 250m p Serra Azul-BA 13 | 15°52'21.8"S, 39°54'30.6"W | 731m TF | 14 15°52'19.7'S, 39°54'27.1"W goo m TF 15 15°52'22.0"S, 39°54'22.7"W 739m TF 16 | 15°52'24.7"S, 39°54'22.2"W | 690 m TF 17 15°52'27.7"S, 39°54'22.5"W | 672m Te 18 15°52'26.3"S, 39°54'18.6"W 780m TF 19 | 15°52'28.1"S, 39°54'10.9"W | 692m TF 20 15°52'25.9'S, 39°54'29.3"W 761m | TF 21 15°52'29.0"S, 39°54'27.7"W | 650m | TF DD | 15°52'34.5"S, 39°54'22.0"W | 668m TF 23 15°52'33.2"S, 39°54'25.0"W 652m Tr 24 | 15°52'29.7"S, 39°54'24.9"W 648 m TF 28 | 15°52'33.7'S, 39°54'24.2"W 658m TS 26 | 15°51'52.5"S, 39°53'26.1"W 256m p ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 218 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil Sample design A total of six field campaigns were carried out between December 2014 and March 2016 in Serra do Mandim and Serra Azul. Each expedition lasted approx- imately seven days, with approximately three or four days dedicated to active search in each mountain range. A total of 24 transects were used between altitudes of 500-800 meters with a length of 50 meters and a width of 5 meters, spaced 100 meters apart in a forested area, two transects of 120 meters in streams, and two ponds were selected in a lowland area (Table 1). Twelve transects were sampled in forest, one in a stream, and a permanent pond in each of the areas (Figs 1, 2). The transects in the forest were surveyed for 40 min, totaling 96 h. The streams in the forest were sampled for 90 min each, totaling 18 h. The perma- nent ponds were sampled for 30 min each, totaling six h. The sampling was car- ried out by two researchers. The total sampling effort was 44 days of sampling and 240 h/person. Amphibians were sampled using visual and acoustic active search methods (Heyer et al. 1994; Crump and Scott 1994) and by opportunis- tic encounters (i.e., along roads or trails outside the transects). All animals were collected with a license from the Brazilian Institute of Envi- ronment and Renewable Natural Resources-IBAMA and/or ICMBio (No. 13708), together with the permission of the administrators of the local farms. Some specimens were euthanized through the administration of a lethal dose of 5% xylocaine to the ventral region, followed by fixation in 10% formaldehyde and preservation in 70% alcohol. All specimens were deposited in the Zoology Mu- seum of the State University of Santa Cruz - MZUESC (Appendix 1). Data analysis For each species, the total number of individuals observed visually and acous- tically was recorded in each sampling unit and environment. To evaluate sam- ple sufficiency, rarefaction curves were constructed based on individuals. Four curves were made for each study area: a general curve considering all individuals sampled in the three standardized methodologies (transect in the forest, transect in the streams and ponds), and three others using individuals collected in each separate methodology. To extrapolate species richness, four non-parametric estimators were used: Chao 2, Jackknife 1, Jackknife 2, and Bootstrap (Magurran 1988; Colwell and Coddington 1994; Toti et al. 2000; Go- telli and Colwell 2001). The analysis was performed using presence/absence data for species during each sampling campaign, with 1000 randomizations. The species richness recorded in the study area was compared to 42 lo- cations, mostly in the northeast region of Brazil and a smaller number in the northeast region of Minas Gerais, sampled in the Atlantic Forest, Caatinga, and Cerrado (see Table 3). The information extracted from these locations was used to create a binary matrix of presence/absence with 216 amphibian spe- cies. The analysis of similarity, considering the specific composition of these areas, was performed using the Jaccard index to calculate dissimilarity and the UPMGA (Unweighted Pair Group Method with Arithmetic) linkage method. Species found in inventories with taxonomic doubts (sp., gr., and aff.) were ex- cluded from the analyses. Subsequently, an ANOSIM (Analysis of Similarities) ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 219 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil test was performed considering 9999 permutations, to determine whether the composition of samples recovered in the similarity analysis differs significantly among the groups (Atlantic Forest, Caatinga, and Cerrado). All analyses were conducted using the PAST 4.12 software (Hammer et al. 2001) The species were identified based on their original descriptions, redescriptions, or recent taxonomic revisions. In addition to the original descriptions, references consulted for species identification are provided in Table 2. Additionally, the col- lected material was compared with specimens identified at MZUESC. For nomen- clature, we followed Frost (2023), who also maintains an updated database con- taining all available synonyms for amphibians worldwide. Regarding Adelophryne spp. we follow Lourenco-De-Moraes et al. (2018). The conservation status of the species was Classified according to IUCN (2024). Furthermore, we verified which species are endemic of the Atlantic Forest based on Rossa-Feres et al. (2017). Results A total of 1785 individuals across 46 amphibian species were recorded, encom- passing one species of Gymnophiona (Siphonops annulatus) and 45 anuran species across 14 families (Table 2, Figs 3, 4). The majority of the identified species (n = 24; 53%) are endemic to the Atlantic Rainforest (Rossa-Feres et al. 2017). The Hylidae family was the most representative with 36.9% (n = 17), followed by the Leptodactylidae family, with 21.7% (n = 10) of the amphibians found. The richness identified stands among the highest ever recorded for the northeastern region of Brazil (Table 3). Furthermore, all recognized species identified are listed as Least Concern (LC) on the IUCN Red List (IUCN 2024). The amphibian richness recorded was similar in the two analyzed areas: Serra do Mandim (n = 34) and Serra Azul (n = 37), with 25 species shared between the areas and the rest divided, with nine exclusive to Serra do Mandim and 11 to Ser- ra Azul (Tables 2, 4). Among the total species recorded in each area, four species (Ololygon strigilata, Phyllodytes luteolus, Trachycephalus nigromaculatus, and Leptodactylus cf. mystaceus) were sampled exclusively through opportunistic encounters in Serra do Mandim, while another five (Rhinella crucifer, Gastrothe- ca pulchra, Scinax x-signatus, Trachycephalus nigromaculatus and Dermatonotus muelleri) also corresponded to opportunistic encounters in Serra Azul. In Serra do Mandim most species were found in the monitored pond in the region. Although this was the smallest sampled fragment, some species con- sidered rare and/or with restricted distribution were found only in this location, such as Dendrophryniscus proboscideus, and typical stream species such as Vitreorana eurygnatha and Ololygon strigilata. The anuran richness recorded in Serra Azul was 37 species, and the only spe- cies of Gymnophiona recorded in the study was found in this area (Table 2; Fig. 4). Two species were only detected by vocalization, Phyllodytes maculosus and Gastrotheca pulchra. The first vocalized in bromeliads during forest transect sampling, while the latter was registered vocalizing in the canopy of the forest. A higher number of individuals was recorded in Serra Azul (n = 868) com- pared to Serra do Mandim (n = 786). The most common species during the study were Pristimantis vinhai (n = 454), Haddadus binotatus (n = 296), and Den- dropsophus oliveirai (n = 167). A total of 131 specimens were collected during opportunistic encounters, 78 at Serra Azul and 53 at Serra do Mandim (Table 2). ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 220 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil Figure 3. Amphibians registered in Serra do Mandim and Serra Azul in southwestern Bahia, Brazil. a |schnocnema ver- rucosa (MZUESC 15874) b Ischnocnema sp. (gr. parva - MZUESC 15896) e¢ Dendrophryniscus proboscideus (MZUESC 14688) d Rhinella crucifer (MZUESC 15148) e R. diptycha (MZUESC 15503) f R. granulosa (MZUESC 15055) g Haddadus binotatus (MZUESC 15646) h Vitreorana eurygnatha (MZUESC 14691) i Thoropa miliaris (MZUESC 15782) j Aplastodiscus weygoldti (MZUESC 15787) k Boana crepitans (MZUESC 14675) IB. exastis (MZUESC 15108) m Dendropsophus branneri (MZUESC 14683) n D. elegans (MZUESC 14679) o Phyllodytes luteolus (MZUESC 17501) p Ololygon strigilata (MZUESC 15001) q Pithecopus nordestinus (MZUESC 14682) r Pseudis fusca (MZUESC 16528). ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 271 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil Table 2. Species of amphibians from Serra do Mandim and Serra Azul, southwest Bahia, Brazil. Legend: SM = Sampling Method (OE = Oportunistic encounters, TF = Transect in the forest, P = Ponds, TS = Transect in the streams). HAB = Habi- tat (LL = Leaf litter or understory, B = Bromeliads or epiphytes, S = Streams, F = Forest, P = Ponds or open area). N = Num- ber of registered specimens. Additional ID references = Additional references consulted to identify species. # = only acoustic record; t = species only found in the inner forests; * endemic to the Atlantic Rainforest. F ; Serra do Mandim Serra Azul § Order/Family/Species Additional ID references SM HAB_| N SM HAB | N ANURA Brachycephalidae Ischnocnema verrucosa TF LL | 02 Tr LL | 06 Lynch 1972; Canedo et al 2010; (Reinhardt & Liitken, 1862)" Araujo et al. 2023 Ischnocnema sp. (gr. parva) * : 5 - 5 Ae EL.) e207 Heyer et al. 1990; Silva-Soares et al. 2021 Bufonidae Dendrophryniscus proboscideus OE LF LL | 08 - - - Izecksohn 1976; Caramaschi 2012 (Boulenger, 1882)" Rhinella crucifer TFTS PLL | 07 OE PLL | 01 Baldissera et al. 2004; Oliveira et al. 2014 (Wied-Neuwied, 1821)* Rhinella diptycha (Cope, 1862) P PLL | 02 P PLL | 09 | Stevaux 2002; Lavilla and Brusquetti 2018 Rhinella granulosa (Spix, 1824) - - - | OE TEP | PLL | 61 Narvaes and Rodrigues 2009; Sao Pedro et al. 2011 Craugastoridae Haddadus binotatus (Spix, 1824)" TF LE) 51° | OETRIS:| Ek. | 145 Heyer et al. 1990; Dias et al. 2012 Centrolenidae Vitreorana eurygnatha (A. Lutz, 1925)” TS S| 15 - - - Heyer et al. 1990; Zucchetti et al. 2023 Cycloramphidae | | Thoropa miliaris (Spix, 1824)" OETETS |stt| 10] o&TS | SLL| 05 Feio et al. 2006a Eleutherodactylidae Adelophryne sp.8* TF LL | 08 | OE,TF a) ee Lourenco-de-Moraes et al. 2018 Adelophryne sp.2* TF,TS EL p35) “ORME LL | 29 Lourenco-de-Moraes et al. 2018 Hemiphractidae Gastrotheca pulchra - - - OE B 01 Duellman 2015 Caramaschi & Rodrigues, 2007#* Hylidae Aplastodiscus weygoldti TS S | 06 aps S 08 Orrico et al. 2006 (Cruz & Peixoto, 1987)™ Boana crepitans (Wied-Neuwied, 1824) OE,P P 1°18 | OE, TEP P 21 Orrico et al. 2017 Boana exastis Hs F | 01 : - - Loebmann et al. 2008 (Caramaschi & Rodriguez, 2003)* Boana faber (Wied-Neuwied, 1821)* OE TRE 1) BF) 4a OE,TF PF | 06 | Martins and Haddad 1988; Heyer et al. 1990 Dendropsophus branneri (Cochran, 1948) OEP P | 64 OEP P 83 | Bastos and Pombal 1996; Nunes et al. 2007; Orrico et al. 2021 Dendropsophus elegans OE,P P | 52 OE,P P 24 | Gomes and Peixoto 1991; Dias et al. 2017a; (Wied-Neuwied, 1824)* Pirani et al. 2022 Dendropsophus oliveirai OE,P P | 60 OEP P 107 Santana et al. 2011; Orrico et al. 2021 (Bokermann, 1963) Ololygon strigilata (Spix, 1824)" OE S | 01 - - - Pimenta et al. 2007 Phyllodytes maculosus Cruz, Feio & ; - x TF B 01 Dias et al. 2020 Cardoso, 2007#* Phyllodytes luteolus OE B | 02 3 = A Bokermann 1966a; Blotto et al. 2021 (Wied-Neuwied, 1821)* ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 299 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil Order/Family/Species Pithecopus nordestinus (Caramaschi, 2006) Pseudis fusca Garman, 1883* Scinax eurydice (Bokermann, 1968)* Scinax pachycrus (Miranda-Ribeiro, 1937a) Scinax x-signatus (Spix, 1824) Sphaenorhynchus prasinus Bokermann, 1973* Trachycephalus nigromaculatus von Tschudi, 1838* Leptodactylidae Leptodactylus fuscus (Schneider, 1799) Leptodactylus latrans (Steffen, 1815) Leptodactylus macrosternum Miranda-Ribeiro, 1926 Leptodactylus cf. mystaceus (Spix, 1824) Leptodactylus mystacinus (Burmeister, 1861) Leptodactylus troglodytes Lutz, 1926 Leptodactylus viridis Jim & Spirandeli-Cruz, 1973* Physalaemus cicada Bokermann, 1966c Physalaemus cf. erikae Cruz & Pimenta, 2004* Physalaemus kroyeri (Reinhardt & Liitken, 1862) Microhylidae Dermatonotus muelleri (Boettger, 1885) Odontophrynidae Proceratophrys schirchi (Miranda-Ribeiro, 1937b)* Pipidae Pipa carvalhoi (Miranda-Ribeiro, 1937a) Strabomantidae Bahius bilineatus (Bokermann, 1975) * Pristimantis vinhai (Bokermann 1975) * Pristimantis sp. (gr. ramagii)* GYMNOPHYONA Siphonopidae Siphonops annulatus (Mikan 1822)* Serra do Mandim SM —_ HAB OEP Pp OE,TF PF OEP Pp OEP Pp OE PF p p OE,TS,P PS TS PS OE S p p p p OEP Pp OE,TRTS SLL p p OETF LL OE,TRTS LL ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 Serra Azul N SM HAB 30,5°=«=~* Pp : P P 03) TF PF : P P 07 OE Pp 16 OE,P Pp 01 OE PF 58 OEP P 07 PS 01 RS 01 - - 03) P p - | OEP | P 07 : : - | OEP | P 18 OE,P Pp OE,P P : OE p 59 OETS) | “Stk 04; - 17 : : 7/76 | OETERTS: |) Tel - OE,TF ble - TE EL 06 02 09 178 01 Additional ID references Vilaga et al. 2006; Vaz-Silva et al. 2020 Caramaschi and Cruz 1998; Garda et al. 2010 Magrini et al. 2011; Novaes-e-Fagundes et al. 2016; Menezes et al. 2016 Carneiro et al. 2004; Novaes-e-Fagundes et al. 2016 Araujo-Vieira et al. 2020a; Novaes-e- Fagundes et al. 2021 Araujo-Vieira et al. 2020b Bokermann 1966b Heyer 1978; Heyer et al. 1990; De-Sa et al. 2014 Magalhaes et al. 2022 Magalhaes et al. 2022 Toledo et al. 2005; De-Sa et al. 2014; Cassini et al. 2013 Abrunhosa et al. 2001; De-Sa et al. 2014; Cassini et al. 2013 De-Sa et al. 2014 Magalhaes et al. 2022 Nascimento et al. 2005; Hepp and Pombal 2020 Nascimento et al. 2005; Hepp and Pombal 2020 Nascimento et al. 2005; Hepp and Pombal 2020; Braga et al. 2024 Vaz-Silva et al. 2020; Dubeux et al. 2021 Izecksohn and Peixoto 1980; Sichieri et al. 2021 Lima et al. 2020 Dias et al. 2017b Trevisan et al. 2020 Trevisan et al. 2020 Maciel and Hoogmoed 2011 223 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil Table 3. The number of amphibian species (S), study duration (SD in months), and region (R) type of different study sites in northeastern Brazil, including the northeastern portion of Minas Gerais. Localities listed as RPPN are Private Natural Heritage Reserves, those labelled as APA are Environmental Protection Areas, EE represents Ecological Stations, and PN denotes National Parks. Region abbreviations include Atlantic Forest (AF), Caatinga (CA), and Cerrado (CE). Localities, states of Brazil RPPN Serra Bonita, BA RE Michelin, BA APA Lagoa Encantada and River Almada, BA Serra da Jibdia, BA Serra do Timbo, BA PN Serra das Lontras, BA PN Grande Sertao Veredas, BA/GO/MG Serra Mandim and Serra Azul, BA Middle Jequitinhonha River, MG Chapada Diamantina, BA RPPN Frei Caneca, PE Complex Limoeiro, MG RPPN Estacao Veracel, BA Complex Nossa Senhora Fatima, MG Planalto de Ibiapaba, CE Tocantins River Basin, MA/TO Complex Cariri, BA/MG Morro do Mara, BA EE Serra Geral do Tocantins, TO Guaratinga, BA Macaiba, RN Conde, BA Sao Desiderio, BA Camamu, BA Serra do Brejo Novo, BA PN Chapada Diamantina, BA Cruz das Almas, BA Complex Bandeira, BA/MG Complex Santana, MG Complex Mumbuca, MG RPPN Sapiranga, BA PN Descobrimento, BA Itapebi, BA EE Raso da Catarina, BA PN Catimbau, PE Curimatau, PB Middle Jaguaribe River, CE RPPNs in Betania and Floresta, PE Jatoba, PE RPPN Serra das Almas, CE Paulo Afonso, BA Cariri Paraibano, PB ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 S) 80 69 59 995 995 49 47 46 46 44 42 39 39 38 38 38 36 36 36 34 34 33 oe 32 B32 31 31 30 28 2/ 29 25 24 21 21 21 19 19 18 18 17 16 SD 16 30 01 ~ 20 years 12 07 ~ 11 years 08 29 06 12 03 01 02 24 06 03 15 02 01 14 04 02 01 19 01 39 02 02 02 05 01 01 13 01 02 01 07 04 02 20 23 CA/CE AF/CA AF/CA AF/CA AF/CA Source Dias et al. 2014b Camurugi et al. 2010; Mira-Mendes et al. 2018 Dias et al. 2014a Junca, 2006; Freitas et al. 2018 Freitas et al. 2019 Rojas-Padilla et al. 2020 Brandao et al. 2020 This study Feio and Caramaschi 1995 Junca 2005 Santos and Santos 2011 Feio et al. 2006b Silvano and Pimenta 2003 Feio et al. 2006b Loebmann and Haddad 2010 Brasileiro et al. 2008 Feio et al. 2006b Bastos and Zina 2022 Valdujo et al. 2011 Silvano and Pimenta 2003 Magalhaes et al. 2013 Gondim-Silva et al. 2016 Valdujo et al. 2009 Silvano and Pimenta 2003 Lantyer-Silva et al. 2013 Magalhaes et al. 2015 Protazio et al. 2021 Feio et al. 2006b Feio et al. 2006b Feio et al. 2006b Junca 2006 Silvano and Pimenta 2003 Sivano and Pimenta 2003 Garda et al. 2013 Pedrosa et al. 2014 Arzabe et al. 2005 Santana et al. 2015 Borges-Nojosa and Santos 2005 Silva et al. 2011 Borges-Nojosa and Cascon 2005 Protazio et al. 2010 Vieira et al. 2007 224 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil ? Figure 4. Amphibians registered in Serra do Mandim and Serra Azul in southwestern Bahia, Brazil a Scinax pachycrus (MZUESC 16525) b S. x-signatus (MZUESC 17503) ¢ Sphaenorhynchus prasinus d Trachycephalus nigromaculatus (MZU- ESC 15064) e Leptodactylus mystacinus (MZUESC 16529) f L. troglodytes (MZUESC 15003) g L. viridis (MZUESC 15848) h Physalaemus cf. erikae (MZUESC 15878) i P. kroyeri (MZUESC 15784) j Dermatonotus muelleri (MZUESC 15070) k Pro- ceratophrys schirchi (MZUESC 146839) I Pipa carvalhoi m Pristimantis vinhai (MZUESC 15642) nP. sp (gr. ramagii — MZU- ESC 16523) o Siphonops annulatus (MZUESC 15900). Table 4. Number of species and abundance found in the study area through standardized methodologies and extrapolat- ed richness using richness estimators. TF: Transect in the forest; TS: Transect in the streams and P: Ponds. Serra Azul Serra do Mandim Estimators TF TS P Total TF TS P Total Richness fe} 5 18 32 13 9 15 30 Chao 2 5 sht2 7, 5.0+0.2 25.8+7.4 39.6+6.3 15:813:5 9.84+1.5 1562122 35.8+6.0 Jackknife1 17.24+2.0 5.8+0.8 24.7+4.9 41.2+4.4 VPEZEVS TALOET 7 i SNe] 35.8427 Jackknife2 18.9 5.9 28.9 45.8 19.4 12.4 17.9 39.4 Bootstrap 14.9 5.4 20.8 36.2 14.9 1OE2: 1a: 32.6 Abundance 433 43 392 868 350 106 330 786 ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 295 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil The overall rarefaction curve obtained for each area showed a tendency to- wards stabilization but did not reach the asymptote (Fig. 5A). The curves made for the different employed methodologies did not show a stabilization trend, except for the species recorded in ponds (Fig. 5D). The richness estimators suggested the occurrence of between 39-46 species in Serra Azul and be- tween 36-39 species in Serra do Mandim (Table 4). Thus, during field activities, between 78.2-94.4% of the estimated richness for each area was sampled. A summary of the richness estimates for each area and the employed method- ologies can be found in Table 4. Cluster analysis yielded three main groups, which are mainly based on dif- ferent regions in which the samplings were carried out (Fig. 6). The ANOSIM a) General Sampling b)Transect in the forest 5 15) SA 30 wm oO 25 as) oO Q. ” 20 Po te 5 15 = = Z 10 5 100 200 300 400 500 600 700 800 900 50 100 150 200 250 300 350 400 450 500 Specimens Specimens c)Transect in the streams d) Ponds 10 20) 9 M 18) SA 8 167 7 a 144 ES 6 © 12} a A 4 5 oo 10 D 4 4 8 3 ny Z 2 4 1 2 0 10 20 30 40 50 60 70 80 90 100 50 100 150 200 250 300 350 400 Specimens Snecimens Figure 5. Rarefaction curve based on amphibian individuals for the two fragments of Semi-deciduous Seasonal Forest in Serra do Mandim and Serra Azul in southwest Bahia, Brazil a general sampling: Serra do Mandim and Serra Azul (without considering opportunistic records) b transect in the forest c transect in the streams and d ponds. The central line corresponds to the average obtained with 1000 randomizations, and the lines above and below correspond to the 95% confidence interval. The blue line represents Serra Azul (SA), and the green line represents Serra do Mandim (SM). ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 226 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil 01 02 0.3 la 1 lb 2 3a 3b 3 ac test demonstrates significant differences in species composition between the locations sampled in the Caatinga, Cerrado and Atlantic Forest (R = 0.662, P = 0.0001). Group 1 is formed by anuran assemblages sampled mainly in the Caatinga, where two main subdivisions can be highlighted, the first (1a) formed by localities along the middle Jequitinhonha River - MG and two localities in the Chapada Diamantina-BA, the latter hosting several endemic species (e.g., Haddadus aramunha, Leptodactylus oreomantis, Rupirana cardosoi) contribut- ing to the unique anurofauna distinct from other sampled Caatinga locations (1b). Group 2 is formed by the anuran assemblages from the Cerrado. Group 3 is comprised of anuran assemblages within the Atlantic Forest, and three main subdivisions can be highlighted. The first subdivision (3a) Similarity 0.4 0.5 06 O07 O8 09 M. Jequitinhonha River - MG Chapada Diamantina - BA PN Chapada Diamantina - BA Macaiba - RN M. Jaguaribe River - CE Planalto da Ibiapaba - CE Cariri Paraibano - PB Paulo Afonso - BA Jatoba - PE PN Catimbau - PE EE Raso da Catarina - BA RPPN Betania and Floresta - PE Curimatau - PB RPPN Serra das Almas - CE EE Serra Geral do Tocantins - TO Sao Desidério - BA PN Grande Sertaéo Veredas - BA/GO/MG Tocantins River Basin - MA/TO PN Descobrimento - BA RPPN Veracel - BA Guaratinga - BA Itapebi - BA Camamu - BA Complex Nossa Senhora de Fatima - MG Complex Mumbuca - MG Complex Cariri - BA/MG Complex Santana - MG Complex Bandeira - BA/MG Complex Limoeiro - MG Serra Azul and Serra do Mandim - BA APA Lagoa Encantada and River Almada - BA RPPN Serra Bonita - BA RE Michelin - BA Serra da Jiboia - BA Serra do Timbo - BA PN Serra das Lontras - BA RPPN Frei Caneca - PE Morro do Mara - BA Serra do Brejo Novo - BA Conde - BA Cruz das Almas - BA RPPN Sapiranga - BA Figure 6. Similarity dendrogram for 42 locations, comparing amphibian composition primarily in the northeast region of Brazil. Jaccard index for dissimilarity calculation and UPGMA (Unweighted Pair Group Method with Arithmetic) cluster- ing method (cophenetic correlation coefficient = 0.798). Green: locations sampled in the Atlantic Forest; Red: locations sampled in the Cerrado; Yellow: locations sampled in the Caatinga. ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 227 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil encompasses studies conducted in Dense Ombrophilous Forest in the ex- treme south of Bahia (e.g., RPPN Veracel, National Park of Descobrimento). It also includes transition areas influenced by Seasonal Forests located fur- ther inland, such as localities in the northeast of Minas Gerais (e.g., Complex Santana, Bandeira, and Limoeiro), and Serra Azul and Serra do Mandim in Bahia. The second subdivision (3b) formed by areas located in the south of the state of Bahia, primarily sampled in the Dense Ombrophilous Forest, and encompasses locations with the highest species richness in our analysis. Finally, the third subdivision (3c) encompasses transition areas between the Atlantic Forest and Caatinga (Brejo Novo, Morro do Mara), as well as Restin- ga areas (Conde) and Semideciduous Forest regions (RPPN Frei Caneca and Cruz das Almas). Taxonomic remarks Some specimens encountered during field activities posed challenges for iden- tification. Recently, Magalhaes et al. (2020) revisited the Leptodactylus latrans species group, employing multiple lines of evidence (DNA, acoustic, and mor- phological data), providing morphological diagnoses for the lineages encoun- tered in their study. This facilitated the identification of two species from this group for the study area: L. macrosternum and L. latrans. Physalaemus erikae was described in 2004 based on morphological and acoustic data, with the type locality in Guaratinga, Bahia (~ 100 km from the study area). This species closely resembles P. kroyeri but differs in having a shorter snout-vent length, a shorter advertisement call duration, and the pres- ence of inguinal glands (Cruz and Pimenta 2004). Although our study area is close to the type locality, most specimens did not exhibit a visible inguinal gland, thus being considered as P. kroyeri. However, we encountered two spec- imens that exhibited an inguinal gland. For one of these, we recorded the ad- vertisement call, which had a duration varying 0.37-0.46 s (MZUESC 15878). Recently, Hepp and Pombal (2020) reported that the advertisement call dura- tion of P. erikae ranges from 0.478 to 0.566 s (n = 37 calls from four males), and for P. kroyeri it ranges from 0.673 to 0.759 s (n = 190 calls from eight males). Our data fall within the lower range of the reported variation for P. erikae. Bra- ga et al. (2024) analyzed the vocalization of P kroyeri from Cruz das Almas - Bahia (near the type locality) and found an advertisement call duration of 0.614 to 0.882 s (n = 83 calls from ten males), indicating a longer call duration in this species. However, we emphasize the need for a more comprehensive taxonomic revision of these two species, especially to increase the available data on the morphological and acoustic variation of P erikae, particularly its acoustic parameters, especially those from its type locality, as well as the ne- cessity of molecular sampling of topotypes of this species for inclusion in a phylogenetic approach. Four species were not identified at the species level (Adelophryne sp. 2, A. sp. 8, Ischnocnema sp. [gr. parva] and Pristimantis sp. [gr ramagii]) and are considered candidate species that require further taxonomic investigation (Lourencgo-De-Moraes et al. 2018; Trevisan et al. 2020; IR Dias, personal com- munication). Additionally, three other species (Phyllodytes luteolus, Ischnoc- nema verrucosa and Vitreorana eurygnatha) found in the area exhibit high ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 228 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil intraspecific molecular diversity and may represent a similar case to the afore- mentioned (Blotto et al. 2021; IR Dias, personal communication; Zucchetti and Castroviejo-Fisher 2024). Discussion During the present study, 46 amphibian species were recorded in Serra Azul and Serra do Mandim in southwestern Bahia. This diversity is likely to be even greater, as the species accumulation curves have not stabilized. Thus, in these forest remnants, isolated and immersed in a matrix dominated by pastures for livestock, approximately 25% of the species that occur in Bahia can be found (Dias et al. 2014b), showing the importance of maintaining and conserving the fragments of the region. The recorded richness ranks among the top ten highest ever found in the Northeast region of Brazil, with a high proportion of species endemic to the Atlantic Rainforest. However, it is important to note that the sampling effort among the different stud- ied areas is highly unequal, making comparisons difficult. Localities where low sampling effort was employed, such as APA (Environmental Protection Area) Lagoa Encantada and Almada River (1 month) and RPPN Veracel (1 month), have a similar richness to places where sampling effort was much higher, such as the Michelin Ecological Reserve and RPPN Frei Caneca. It is likely that the anuran fauna in these areas where low sampling effort was employed is underestimated. Among the species found, three are restricted to the Atlantic Forest of the state of Bahia: Bahius bilineatus, Physalaemus cf. erikae, Ololygon strigilata (Junca and Pimenta 2004; Cruz and Pimenta 2004; Pimenta et al. 2007), and five others have a geographical distribution between southern Bahia and Espiri- to Santo or northeastern Minas Gerais: Ischnocnema verrucosa, Aplastodiscus weygoldti, Phyllodytes maculosus, Dendrophryniscus proboscideus, and Lepto- dactylus viridis (Caramaschi and Canedo 2006; Cruz et al. 2007; Moura et al. 2009; Lantyer-Silva et al. 2011; Silva et al. 2012). Some of the species found occur mainly in open areas, such as Rhinella diptycha and Dermatonotus muelleri (Andrade and Carnaval 2004; Feio et al. 2006b), while others are considered typical of the Caatinga, such as Leptodac- tylus troglodytes, Scinax pachycrus, and Physalaemus cicada (Feio and Cara- maschi 1995; Rodrigues 2003; Peixoto and Arzabe 2004; Linares and Mello 2011). The encounter of species restricted to the Atlantic Forest, and others typical of the Caatinga, was expected since the study site is located in a tran- sition area between these two regions. According to the similarity analysis, the composition of the anuran fauna of Serra Azul and Serra do Mandim is similar to other sampled areas in the region in ecotonal areas, which also share fau- nistic elements of these two morphoclimatic domains (Feio and Caramaschi 1995; Feio et al. 2006b). Within the state of Bahia, some species found in the study area have re- cords limited to fewer than three occurrence points, such as Trachycephalus nigromaculatus, and Pseudis fusca (Garda et al. 2010; Dias et al. 2010). Pseu- dis fusca (Fig. 3R), an aquatic species, is found in the states of Bahia, Minas Gerais, and Espirito Santo, Brazil (Garda et al. 2010). In Bahia, the species is known from two municipalities: Guaratinga and Teixeira de Freitas (Garda et al. ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 229 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil 2010). The encounter of P fusca in Serra Azul represents the third record of the species for the state of Bahia, marking an approximately 250 km straight-line increase in its geographical distribution from its type locality (Aracuai — Minas Gerais). These new records contribute to a better understanding of the distri- bution patterns of these species in the region, especially for P fusca, with this new occurrence point now representing the northern limit of the distribution of this species. Amphibian inventories in southern Bahia have shown high species richness and endemism in the region (e.g., Silvano and Pimenta 2003; Camurugi et al. 2010; Rojas-Padilla et al. 2020; present study), which may be associated with the fact that the central region of the Atlantic Forest is estimated as a zone of climatic stability during the Quaternary glaciations, serving as a large refuge for amphibian species in the Atlantic Forest at the end of the Pleistocene, con- tributing to the maintenance of regional diversity (Carnaval and Moritz 2008; Carnaval et al. 2009). Serra do Mandim and Serra Azul still have conserved forest fragments that form a complex network of streams that compose the Jequitinhonha river ba- sins further south (IBGE 1997) and the Rio Pardo basin further north of the region (Pedreira 1999; Cetra et al. 2010). This great abundance of water bod- ies and the conservation of the study area contributed to the record of typical forest species (e.g., Adelophryne sp. 2, A. sp. 8, Ischnocnema verrucosa, Pristi- mantis vinhai, and Gastrotheca pulchra) and typical stream species (e.g., Aplas- todiscus weygoldti, Ololygon strigilata, and Vitreorana eurygnatha). In addition, a high abundance of direct development species using leaf litter (e.g., Pristiman- tis vinhai, P. sp. [gr. ramagii] and Haddadus binotatus), including species that are difficult to sample such as /schnocnema verrucosa and Dendrophyrniscus proboscideus, was found, demonstrating that the sampled forest fragments still have adequate conditions for the maintenance of these populations in the region. Despite this, no protected area has been established to ensure the con- servation of these elements of the region’s fauna. Actions are required to se- cure the preservation of these species. Our results demonstrate that the remaining forest fragments in the region, although small and isolated, still support a high richness of amphibians with species restricted to the Atlantic Forest and Bahia, such as Bahius bilineatus and Ololygon strigilata, and others considered typical of the Caatinga, such as Dermatonotus muelleri, Leptodactylus troglodytes, and Physalaemus cicada. Acknowledgments We thank Cintia de Melo Souto Brige (in memoriam) and family (Fugiama farm) and José Cordeiro de Almeida Filho (in memoriam) and family (Serra Azul farm) for allowing the execution of the study on their properties, and Camila Souza Batista for her support in the field. We thank Kaique Brito Silva for assistance in preparing the map. CASC thanks CAPES-Coordenagao de Aperfeigoamento de Pessoal de Nivel Superior and FAPESB-Fundagao de Amparo a Pesquisa do Estado da Bahia, for scholarships, as well as the State University of San- ta Cruz and the Zoology Graduate Program-PPGZOO for the opportunity and support granted. MS and IRD acknowledge the Brazilian National Council for Scientific and Technological Development (CNPq - PQ 309365/2019-8 and PQ: ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 230 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil 315362/2021-9, respectively) for research scholarships. CVM-M acknowledg- es funding from Universidade Estadual do Maranhao (UEMA) (senior productiv- ity grant 05/2023- PPG/UEMA). Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Funding This work was supported by Conselho Nacional de Desenvolvimento Cientifico e Tec- noldgico; Coordenacao de Aperfeigoamento de Pessoal de Nivel Superior; Fundacao de Amparo a Pesquisa do Estado da Bahia. Author contributions Conceptualization: CASC, AJSA, CVMM, IRD, MS. Data curation: CASC. Formal analy- sis: CASC, IRD. Funding acquisition: MS. Investigation: MS, CASC. Methodology: CASC. Project administration: MS, CASC. Supervision: MS, IRD. Validation: CASC. Visualization: CASC. Writing - original draft: MS, AJSA, CVMM, IRD, CASC. Writing - review and editing: AJSA, IRD, MS, CVMM, CASC. Author ORCIDs Mirco Solé ® https://orcid.org/0000-0001-7881-6227 Caio Vinicius de Mira-Mendes © https://orcid.org/0000-0002-7707-6439 luri Ribeiro Dias © https://orcid.org/0000-0002-2825-3494 Data availability All of the data that support the findings of this study are available in the main text. References Abrunhosa PA, Wogel H, Pombal Jr JP (2001) Vocalizacao de quatro espécies de anuros do Estado do Rio de Janeiro, Sudeste do Brasil (Amphibia, Hylidae, Leptodactylidae). Boletim do Museu Nacional do Rio de Janeiro 472: 1-12. Almeida AP. Bastos DFO, Junior PBA, Vila Nova MF, Dias IR, Zina J (2022) New re- cords and geographic distribution map of Proceratophrys sanctaritae Cruz & Na- poli, 2010 (Anura, Odontophrynidae). Check List 18(6): 1243-1247. https://doi. org/10.15560/18.6.1243 Andrade G, Carnaval AC (2004) Rhinella jimi. The IUCN Red List of Threatened Species 2004. [accessed on 29 July 2016] https://doi.org/10.2305/IUCN.UK.2004.RLTS. T54674A11184744.en Araujo AP, Ferreira RB, Canedo C, Zocca C, Lacerda JVA (2023) After 160 years of ‘si- lence’: The advertisement call of the frog Ischnocnema verrucosa. Herpetological Bul- letin 163(163): 31-34. https://doi.org/10.33256/hb163.3134 Araujo-Vieira K, Pombal Jr JP Caramaschi U, Novaes-e-Fagundes G, Orrico VGD, Faivovich J (2020a) A neotype for Hyla x-signata Spix, 1824 (Amphibia, Anura, Hylidae). Pa- ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 231 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil péis Avulsos de Zoologia 60[e20206056]: 1-30. https://doi.org/10.11606/1807- 0205/2020.60.56 Araujo-Vieira K, Luna MC, Caramaschi U, Haddad CFB (2020b) A new genus of Lime Treefrogs (Anura: Hylidae: Sphaenorhynchini). Zoologischer Anzeiger 286: 81-89. https://doi.org/10.1016/j.jcz.2020.04.002 Argdlo AJS (2004) As serpentes dos cacauais do sudeste da Bahia. Editus, llhéus, Ba- hia, Brazil, 259 pp. Arzabe C, Skuk G, Santana GG, Delfim FR, Lima YCC, Abrantes SHF (2005) Herpetofauna da area do Curimatau, Paraiba. In: Araujo FS, Rodal MJN, Barbosa MRV (Org) Andalise das Variacoes da Biodiversidade do Bioma Caatinga: suporte a estratégias regionais de conservacao. Ministério do Meio Ambiente, Brasilia, 259-273. Baldissera Jr FA, Caramaschi U, Haddad CFB (2004) Review of the Bufo crucifer spe- cies group, with descriptions of two new related species (Amphibia, Anura, Bu- fonidae). Rio de Janeiro. Arquivos do Museu Nacional. Museu Nacional (Brazil) 62(3): 255-282. Bastos RP Pombal Jr JP (1996) A new species of Hyla (Anura: Hylidae) from eastern Bra- zil. Amphibia-Reptilia 17(4): 325-331. https://doi.org/10.1163/156853896X00054 Bastos DFO, Zina J (2022) Amphibian fauna in an ecotonal and mountainous area in south-central Bahia State, northeastern Brazil. Herpetology Notes 15: 365-376. Becker CG, Fonseca CR, Haddad CFB, Batista RF, Prado PI (2007) Habitat Split and the Global Decline of Amphibians. Science 318(5857): 1775-1777. https://doi. org/10.1126/science.1149374 Blotto BL, Lyra ML, Cardoso MCS, Rodrigues MT, Dias IR, Marciano-Jr E, Vechio FD, Orri- co VGD, Brandao RA, Assis CL, Lantyer-Silva ASF, Rutherford MG, Gagliardi-Urrutia G, Solé M, Baldo D, Nunes |, Cajade R, Torres A, Grant T, Jungfer KH, Silva HR, Haddad CFB, 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 Boettger O (1885) Liste von Reptilien und Batrachiern aus Paraguay. Zeitschrift fiir Naturwissenschaften 58: 213-248. Bokermann WCA (1963) Nova espécie de Hyla da Bahia, Brasil. Atas da Sociedade de Biologia do Rio de Janeiro 7: 6-8. Bokermann WCA (1966c) Notas sdbre trés espécies de Physalaemus de Maracas, Bahia (Amphibia, Leptodactylidae). Revista Brasileira de Biologia 26(3): 253-259. Bokermann WCA (1966a) O género Phyllodytes Wagler, 1830 (Anura, Hylidae). Rio de Janeiro. Anais da Academia Brasileira de Ciéncias 38(2): 335-344. Bokermann WCA (1966b) Una nueva especie de Trachycephalus de Bahia, Brasil (Am- phibia, Hylidae). Neotrépica 12(39): 124-210. Bokermann WCA (1968) Three new Hyla from the Plateau of Maracas, central Bahia, Brazil. Journal of Herpetology 1(1/4): 25-31. https://doi.org/10.2307/1563259 Bokermann WCA (1973) Duas novas espécies de Sphaenorhynchus da Bahia (Anura, Hylidae). Revista Brasileira de Biologia 33: 589-594. Bokermann WCA (1975) Trés espécies novas de Eleutherodactylus do sudeste da Bahia, Brasil (Anura, Leptodactylidae). Revista Brasileira de Biologia 34(1): 11-18. Borges-Nojosa DM, Cascon P (2005) Herpetofauna da area Reserva da Serra das Al- mas, Ceara. In: Araujo FS, Rodal MUN, Barbosa MRV (Org) Analise das Variacées da Biodiversidade do Bioma Caatinga: suporte a estratégias regionais de conservagao. Brasilia, Ministério do Meio Ambiente, 243-258. Borges-Nojosa DM, Santos EM (2005) Herpetofauna da area de Betania e Floresta, Per- nambuco. In: Araujo FS, Rodal MJN, Barbosa MRV (Org) Andlise das Variagdes da ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 232 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil Biodiversidade do Bioma Caatinga: suporte a estratégias regionais de conservagao. Brasilia, Ministério do Meio Ambiente, 275-289. Boulenger GA (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the Collection of the British Museum. Second Edition. Taylor and Francis, London, 578 pp. Braga HSN, Vieira MVSA, Silva TAF, Protazio AS, Protazio AS (2024) Acoustic partition- ing explains the coexistence between two Physalaemus species (Anura, Leptodactyli- dae) in the Atlantic Forest in Eastern Bahia State, Brazil. Anais da Academia Brasileira de Ciéncias 95(1): e20211348. https://doi.org/10.1590/0001-3765202320211348 Brandao RA, Leao PF, Gedraite L (2020) Anfibios da Trijungao dos estados da Bahia, Goias e Minas Gerais e do Parque Nacional Grande Sertao Veredas. In: Brandao RA, Francoso RD, Machado TH, Santos NJ (Org) Historia Natural do Sertao da Trijuncao do Nordeste, Centro—Oeste e Sudeste do Brasil. Sao Paulo, PerSe 12 ed.: 88-127. Brasileiro CA, Lucas EM, Oyamaguchi HM, Thomé MTC, Dixo M (2008) Anurans, North- ern Tocantins River Basin, states of Tocantins and Maranhao, Brazil. Check List 4(2): 185-197. https://doi.org/10.15560/4.2.185 Burmeister H (1861) Reise durch die La Plata-Staaten mit besonderer Riicksicht auf die physische Beschaffenheit und den Culturzustand der Argentinischen Republik. Aus- gefiihrt in den Jahren 1857, 1858, 1859 und 1860. Vol. 2. H. W. Schmidt, Halle, 538 pp. Camardelli M, Napoli MF (2012) Amphibian Conservation in the Caatinga Biome and Semiarid Region of Brazil. Herpetologica 68(1): 31-47. https://doi.org/10.1655/HER- PETOLOGICA-D-10-00033.1 Camurugi F, Lima TM, Mercés EA, Junca FA (2010) Anuros da Reserva Ecoldgica da Michelin, Municipio de Igrapiuna, Estado da Bahia, Brasil. Biota Neotropica 10(2): 305-312. https://doi.org/10.1590/S1676-06032010000200032 Canedo CB, Pimenta VS, Leite FSF, Caramaschi U (2010) New species of Ischnocnema (Anura: Brachycephalidae) from the state of Minas Gerais, southeastern Brazil, with comments on the /. verrucosa species series. Copeia 2010(4): 629-634. https://doi. org/10.1643/CH-09-159 Caramaschi U (2006) Redefinicao do grupo de Phyllomedusa hypochondrialis, com re- descricao de P megacephala (Miranda-Ribeiro, 1926), revalidacdo de P azurea Cope, 1862 e descricdo de uma nova espécie (Amphibia, Anura, Hylidae). Arquivos do Mu- seu Nacional. Museu Nacional 64: 159-179. Caramaschi U (2012) A new species of beaked toad, Rhinella (Anura: Bufonidae), from the State of Bahia, Brazil. Zoologia. Sociedade Brasileira de Zoologia. Curitiba 29(4): 343-348. https://doi.org/10.1590/S1984-46 70201 2000400007 Caramaschi U, Canedo C (2006) Reassessment of the taxonomic status of the gen- era Ischnocnema Reinhardt and Lutken, 1862 and Oreobates Jiménez-de-la-Espada, 1872 with notes on the synonymy of Leiuperus verrucosus Reinhardt and Litken, 1862 (Anura: Leptodactylidae). Zootaxa 1116(1): 43-54. https://doi.org/10.11646/ zootaxa.1116.1.3 Caramaschi U, Cruz CAG (1998) Notas taxonémicas sobre Pseudis fusca Garman e P bolbodactyla A. Lutz, com a descrigao de uma nova espécie correlata (Anura, Pseud- idae). Revista Brasileira de Zoologia 15(4): 929-944. https://doi.org/10.1590/S0101- 81751998000400011 Caramaschi U, Rodrigues MT (2003) A new large treefrog species, genus Hyl/a Laurenti, 1768, from southern Bahia, Brazil (Amphibia, Anura, Hylidae). Rio de Janeiro. Arquiv- os do Museu Nacional. Museu Nacional 61(4): 255-260. Caramaschi U, Rodrigues MT (2007) Taxonomic status of the species of Gastrotheca Fitzinger, 1843 (Amphibia, Anura, Amphignathodontidae) of the Atlantic Rain Forest ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 233 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil of Eastern Brazil, with description of a new species. Boletim do Museu Nacional, Nova Serie. Zoologia 525: 1-19. Caramaschi U, Orrico VGD, Faivovich J, Dias IR, Solé M (2013) A new species of Allophryne (Anura: Allophrynidae) from the Atlantic Rain Forest Biome of eastern Brazil. Herpeto- logica 69(4): 480-491. https://doi.org/10.1655/HERPETOLOGICA-D-13-00029 Carilo-Filho LM, Carvalho BT, Azevedo BKA, Gutiérrez-Pesquera LM, Mira-Mendes CV, Solé M, Orrico VG (2021) Natural history predicts patterns of thermal vulnerability in amphibians from the Atlantic Rainforest of Brazil. Ecology and Evolution 11(23): 16462-16472. https://doi.org/10.1002/ece3.7961 Carnaval AC, Moritz C (2008) Historical climate modelling predicts patterns of current biodiversity in the Brazilian Atlantic forest. Journal of Biogeography 35(7): 1187- 1201. https://doi.org/10.1111/j.1365-2699.2007.01870.x Carnaval AC, Hickerson MJ, Haddad CFB, Rodrigues MT, Moritz C (2009) Stability Pre- dicts Genetic Diversity in the Brazilian Atlantic Forest Hotspot. Science 323(5915): 785-789. https://doi.org/10.1126/science.1166955 Carneiro MCL, Magalhdes PS, Junca FA (2004) Descricao do girino e vocalizacao de Scinax pachycrus (Miranda-Ribeiro, 1937) (Amphibia, Anura, Hylidae). Rio de Janeiro. Arquivos do Museu Nacional. Museu Nacional 62(3): 241-246. Cassini CS, Orrico VGD, Dias IR, Solé M, Haddad CFB (2013) Phenotypic variation of Lep- todactylus cupreus Caramaschi, Sao-Pedro and Feio, 2008 (Anura, Leptodactylidae). Zootaxa 3616(1): 073-084. https://doi.org/10.11646/zootaxa.3616.1.6 Cetra M, Sarmento-Soares LM, Martins-Pinheiro RF (2010) Peixes de riachos e novas Unidades de Conservagao no sul da Bahia. Pan-American Journal of Aquatic Scienc- es 5(1): 11-21. Cochran DM (1948) A new subspecies of tree frog from Pernambuco, Brazil. Journal of the Washington Academy of Sciences 38: 316-318. Colwell RK, Coddington JA (1994) Estimating terrestrial biodiversity through extrapola- tion. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 345(1311): 101-118. https://doi.org/10.1098/rstb.1994.0091 Cope ED (1862) Catalogues of the reptiles obtained during the explorations of the Pa- rana, Paraguay, Vermejo and Uruguay Rivers, by Capt. Thos. J. Page, U.S.N.; and of those procured by Lieut. N. Michler, U.S. Top. Eng., Commander of the expedition conducting the survey of the Atrato River. Proceedings. Academy of Natural Sciences of Philadelphia 14: 346-35. Crump ML, Scott Jr NJ (1994) Visual Encounter Surveys. In: Heyer WR, Donnelly MA, McDiar- mid RW, Hayek LC, Foster MS (Eds) Measuring and monitoring biological diversity: stan- dard methods for amphibians. Smithsonian Institution Press, Washington USA, 84-92. Cruz CAG, Peixoto OL (1987) Espécies verdes de Hyla: o complexo “albofrenata” (Amphib- ia, Anura, Hylidae). Arquivos de Universidade Federal Rural do Rio de Janeiro 8: 59-70. Cruz CAG, Pimenta BVS (2004) A new species of Physalaemus Fitzinger, 1826 from Southern Bahia state, Brazil (Anura, Leptodactylidae). Journal of Herpetology 38(4): 480-486. https://doi.org/10.1670/214-02A Cruz CAG, Feio RN, Cardoso MCS (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. Museu Nacional 64(4): 321-324. Da-Silva FR, Almeida-Neto M, Prado VHM, Haddad CFB, Rossa-Feres DC (2012) Humid- ity levels drive reproductive modes and phylogenetic diversity of amphibians in the Brazilian Atlantic Forest. Journal of Biogeography 39(9): 1720-1732. https://doi. org/10.1111/j.1365-2699.2012.02726.x ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 234 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil De-Sa RO, Grant T, Camargo A, Heyer WR, Ponssa ML, Stanley EL (2014) Systematics of the Neotropical genus Leptodactylus Fitzinger, 1826 (Anura: Leptodactylidae): Phylog- eny, the relevance of non-molecular evidence, and species accounts. South American Journal of Herpetology 9(s1): 1-128. https://doi.org/10.2994/SAJH-D-13-00022.1 Dias IR, Vilaga TRA, Silva JRS, Barbosa RS, Solé M (2010) Amphibia, Anura, Hylidae, Tra- chycephalus nigromaculatus Tschudi, 1838: Distribution extension. Check List 6(3): 412-413. https://doi.org/10.15560/6.3.412 Dias IR, Medeiros TT, Solé M, Pimenta BVS (2011) Amphibia, Anura, Hylidae, Bokerman- nohyla lucianae (Napoli and Pimenta 2003): Distribution extension and geographic distribution map. Check List 7(2): 108-109. https://doi.org/10.15560/7.2.108 Dias IR, Lourenco-de-Moraes R, Solé M (2012) Description of the advertisement call and morphometry of Haddadus binotatus (Spix, 1824) from a population from southern Bahia, Brazil. North-Western Journal of Zoology 8(1): 107-111. Dias IR, Mira-Mendes CV, Solé M (2014a) Rapid inventory of herpetofauna at the APA (En- vironmental Protection Area) of the Lagoa Encantada and Rio Almada, Southern Bahia, Brazil. Herpetology Notes 7: 627-637. http://biotaxa.org/hn/article/view/8557/10467 Dias IR, Medeiros TT, Vila Nova MF, Solé M (2014b) Amphibians of Serra Bonita, south- ern Bahia: A new hotpoint within Brazil's Atlantic Forest hotspot. ZooKeys 449: 105- 130. https://doi.org/10.3897/zookeys.449.7494 Dias IR, Haddad CFB, Argdélo AUS, Orrico VGD (2017a) The 100": An appealing new spe- cies of Dendropsophus (Amphibia: Anura: Hylidae) from northeastern Brazil. PLoS ONE 12(3): 1-20. https://doi.org/10.1371/journal.pone.0171678 Dias IR, Mira-Mendes CV, Souza-Costa CA, Junca FA, Solé M (2017b) The advertisement call and comments on the distribution of Eleutherodactylus bilineatus Bokermann, 1975, an endemic frog of Bahia State, Brazil (Amphibia, Anura). ZooKeys 677(1): 151-159. https://doi.org/10.3897/zookeys.677.12309 Dias IR, Novaes-e-Fagundes G, Neto AM, Zina J, Garcia C, Recoder RS, Dal Vechio F, Rodrigues MT, Solé M (2020) A new large canopy-dwelling species of Phyllodytes Wagler, 1930 (Anura, Hylidae) from the Atlantic Forest of the state of Bahia, north- eastern Brazil. PeerJ 8(e8642): 1-27. https://doi.org/10.7717/peerj.8642 Droege S, Cyr A, Larivée J (1998) Checklists: An under-used tool for the inventory and monitoring of plants and animals. Conservation Biology 12(5): 1134-1138. https:// doi.org/10.1046/j.1523-1739.1998.96402.x Duarte H, Tejedo M, Katzenberger M, Marangoni F, Baldo D, Beltran JF, Marti DA, Rich- ter-Boix A, Gonzalez-Voyer A (2012) Can amphibians take the heat? Vulnerability to climate warming in subtropical and temperate larval amphibian communities. Global Change Biology 18(2): 412-421. https://doi.org/10.1111/j.1365-2486.2011.02518.x Dubeux MJM, Nascimento FAC, Goncalves U, Mott T (2021) Identification key for anuran amphibians in a protected area in the northeastern Atlantic Forest. Papéis Avulsos de Zoologia 61[e20216176]: 1-10. https://doi.org/10.11606/1807-0205/2021.61.76 Duellman WE (2015) Marsupial Frogs. Gastrotheca & Allied Genera. Johns Hopkins Uni- versity Press, Baltimore, 432 pp. https://doi.org/10.1353/boo0k.40894 Feio RN, Caramaschi U (1995) Aspectos Zoogeograficos dos Anfibios do Médio Rio Jequitinhonha, Nordeste de Minas Gerais, Brasil. Revista Ceres 42(239): 53-61. http://www.ceres.ufv.br/ojs/index.php/ceres/article/view/2238/272 Feio RN, Napoli MF, Caramaschi U (2006a) Consideragdes taxonémicas sobre Thoro- pa miliaris (Spix, 1824), com revalidacao e redescricdo de Thoropa taophora (Miran- da-Ribeiro, 1923) (Amphibia, Anura, Leptodactylidae). Arquivos do Museu Nacional. Museu Nacional 64(1): 41-60. ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 235 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil Feio RN, Nascimento LB, Cruz CAG, Ferreira PL, Pantoja DL (2006b) Anfibios das areas prioritarias dos rios Jequitinhonha e Mucuri. In: Pinto LPS, Bedé L (Org) Biodiversi- dade e Conservacao nos Vales dos Rios Jequitinhonha e Mucuri. Brasilia, Ministerio do Meio Ambiente 1: 94-119. Fonte LFM, Mayer M, Létters S (2019) Long-distance dispersal in amphibians. Frontiers of Biogeography 11(4): 1-14. https://doi.org/10.21425/F5FBG44577 Freitas MA, Abegg AD, Dias IR, Moraes EPF (2018) Herpetofauna from Serra da Jiboia, an Atlantic Rainforest remnant in the state of Bahia, northeastern Brazil. Herpetology Notes 11(1): 59-72. https://www.biotaxa.org/hn/article/view/32254/39365 Freitas MA, Silva TFS, Fonseca PM, Hamdan B, Filadelfo T, Abegg AD (2019) Herpe- tofauna of Serra do Timbo, an Atlantic Forest remnant in Bahia State, northeast- ern Brazil. Herpetology Notes 12: 245-260. https://www.biotaxa.org/hn/article/ view/32254/39365 Frost DR (2023) Amphibian Species of the World: An Online Reference. Version 6.2. Electronic Database. https://amphibiansoftheworld.amnh.org/ Garda AA, Santana DJ, Sao-Pedro VA (2010) Taxonomic characterization of paradox- ical frogs (Anura, Hylidae, Pseudae): geographic distribution, external morpholo- gy, and morphometry. Zootaxa 2666(1): 1-28. https://doi.org/10.11646/zoot- axa.2666.1.1 Garda AA, Costa TB, Santos-Silva CR, Mesquita DO, Faria RG, Concei¢gao BM, Silva IRS, Ferreira AS, Rocha SM, Palmeira CNS, Rodrigues R, Ferrari SF, Torquato S (2013) Her- petofauna of protected areas in the Caatinga |: Raso da Catarina Ecological Station (Bahia, Brazil). Check List 9(2): 405-414. https://doi.org/10.15560/9.2.405 Garman S (1883) A species of Pseudis from the Rio Arassuahy, Brazil. Science Observer. Journal of Science 4: 47. Gomes MR, Peixoto OL (1991) Larvas de Hyla do grupo /eucophyllata com a descri¢ao da de H. elegans, 1824 e notas sobre a variagao do padrao de colorido do adulto nes- ta espécie (Anura, Hylidae). Revista Brasileira de Biologia 51(1): 257-262. Gondim-Silva FAT, Andrade ARS, Abreu RO, Nascimento JS, Corréa GR Menezes L, Trev- isan CC, Camargo SS, Napoli MF (2016) Composition and diversity of anurans in the Restinga of the Conde municipality, Northern coast of the state of Bahia, North- eastern Brazil. Biota Neotropica 16(3): 1-16. https://doi.org/10.1590/1676-0611- BN-2016-0157 Gotelli NJ, Colwell RK (2001) Quantifying biodiversity: Procedures and pitfalls in the measurement and comparison of species richness. Ecology Letters 4(4): 379-391. https://doi.org/10.1046/j.1461-0248.2001.00230.x Gutiérrez-Pesquera LM, Tejedo M, Olalla-Tarraga MA, Duarte H, Nicieza A, Solé M (2016) Testing the climate variability hypothesis in thermal tolerance limits of tropical and temperate tadpoles. Journal of Biogeography 43(6): 1166-1178. https://doi. org/10.1111/jbi.12700 Haddad CFB (1998) Biodiversidade dos Anfibios no Estado de Sado Paulo. In: Joly CA, Bicudo CEM (Org) Biodiversidade do Estado de Sao Paulo, Brasil: sintese do conhec- imento ao final do século XX, 6: vertebrados. Sao Paulo, Fapesp: 15-26. Haddad CFB, Toledo LF, Prado CPA, Loebmann D, Gasparini JL, Sazima | (2013) Guia de Anfibios da Mata Atlantica: diversidade e biologia. Anolisbooks, Sao Paulo, 544 pp. Hammer @, Harper DAT, Ryan PD (2001) Past: Paleontological Statistics Software Pack- age for Education and Data Analysis. Palaeontologia Electronica 4(1) [art. 4]: 1-9. [178kb]. http://palaeo-electronica.org/2001_1/past/issue1_01.htm ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 236 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil Hepp F, Pombal J (2020) Review of bioacoustical traits in the genus Physalaemus Fitzing- er, 1826 (Anura: Leptodactylidae: Leiuperinae). Zootaxa 4725(1): 1-106. https://doi. org/10.11646/zootaxa.4725.1.1 Heyer WR (1978) Systematics of the fuscus group of the frog genus Leptodactylus (Am- phibia, Leptodactylidae). Natural History Museum of Los Angeles County Science Bulletin 29: 1-85. https://doi.org/10.5479/si.00810282.301 Heyer WR, Rand AS, Cruz CAG, Peixoto OL, Nelson CE (1990) Frogs of Boracéia. Ar- quivos de Zoologia 31(4): 231-410. https://doi.org/10.11606/issn.2176-7793. v31i4p231-410 Heyer WR, Donnelly MA, McDiarmid RW, Hayek LC, Foster MS (1994) Measuring and monitoring biological diversity: standard methods for amphibians. Smithsonian Insti- tution Press Washington. Hoffmann M, Hilton-Taylor C, Angulo A, Bohm M, Brooks TM, Butchart SH, Carpenter KE, Chanson J, Collen B, Cox NA, Darwall WR, Dulvy NK, Harrison LR, Katariya V, Pollock CM, Quader S, Richman NI, Rodrigues ASL, Tognelli MF, Vié J-C, Aguiar JM, Allen DJ, Allen GR, Amori G, Ananjeva NB, Andreone F, Andrew P Ortiz ALA, Baillie JEM, Baldi R, Bell BD, Biju SD, Bird JP, Black-Decima P Blanc JJ, Bolanos F, Bolivar-G W, Burfield IJ, Burton JA, Capper DR, Castro F, Catullo G, Cavanagh RD, Channing A, Chao NL, Chenery AM, Chiozza F, Clausnitzer V, Collar NJ, Collett LC, Collette BB, Fernandez CFC, Craig MT, Crosby MJ, Cumberlidge N, Cuttelod A, Derocher AE, Diesmos AC, Donaldson JS, Duckworth JW, Dutson G, Dutta SK, Emslie RH, Farjon A, Fowler S, Freyhof J, Garshelis DL, Gerlach J, Gower DJ, Grant TD, Hammerson GA, Harris RB, Heaney LR, Hedges SB, Hero J-M, Hughes B, Hussain SA, Icochea M J, Inger RF, Ishii N, Iskandar DT, Jenkins RKB, Kaneko Y, Kottelat M, Kovacs KM, Kuzmin SL, La Marca E, Lamoreux JF, Lau MWN, Lavilla EO, Leus K, Lewison RL, Lichtenstein G, Livingstone SR, Lukoschek V, Mallon DR McGowan PJK, Mclvor A, Moehlman PD, Molur S, Alonso AM, Musick JA, Nowell K, Nussbaum RA, Olech W, Orlov NL, Papenfuss TJ, Parra-Olea G, Perrin WF, Polidoro BA, Pourkazemi M, Racey PA, Ragle JS, Ram M, Rathbun G, Reynolds RP Rhodin AGJ, Richards SJ, Rodriguez LO, Ron SR, Rondinini C, Rylands AB, Sadovy de Mitcheson Y, Sanciangco JC, Sand- ers KL, Santos-Barrera G, Schipper J, Self-Sullivan C, Shi Y, Shoemaker A, Short FT, Sillero-Zubiri C, Silvano DL, Smith KG, Smith AT, Snoeks J, Stattersfield AJ, Symes AJ, Taber AB, Talukdar BK, Temple HJ, Timmins R, Tobias JA, Tsytsulina K, Tweddle D, Ubeda C, Valenti SV, Paul van Dijk PR Veiga LM, Veloso A, Wege DC, Wilkinson M, Williamson EA, Xie F, Young BE, Akgcakaya HR, Bennun L, Blackburn TM, Boitani L, Dublin HT, da Fonseca GAB, Gascon C, Lacher Jr TE, Mace GM, Mainka SA, Mc- Neely JA, Mittermeier RA, Reid GMG, Rodriguez JP Rosenberg AA, Samways MJ, Smart J, Stein BA, Stuart SN (2010) The impact of conservation on the status of the world’s vertebrates. Science 330(6010): 1503-1509. https://doi.org/10.1126/ science.1194442 IBGE [Instituto Brasileiro de Geografia e Estatistica] (1997) Diagndstico Ambiental da Bacia do Rio Jequitinhonha: diretrizes gerais para a ordenagao territorial. Salvador, Diretoria de Geociéncias, 17 Divisado de Geociéncias do Nordeste-DIGEO 1/NE.1, 64 pp. ICMBio [Instituto Chico Mendes de Conservacao da Biodiversidade] (2003) Decretos MMA s/n° de 05 de Junho de 2003. http://www.icmbio.gov.br/portal/rebio-da-mata escura?highlight=WyJtY XRhliwiZXNjdXJhliwibWFOYSBIc2N1cmEixXQ [accessed on 25 March 2018] ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 237 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil ICMBio [Instituto Chico Mendes de Conservacao da Biodiversidade] (2010) Decreto MMA s/n° de 11 de Junho de 2010. http://www.icmbio.gov.br/portal/parna-do-al- to-cariri?highlight=WyJjYXJpcmkixXQ [accessed on 25 March 2018] ICMBio [Instituto Chico Mendes de Conservacao da Biodiversidade] (2016) Portaria MMA n° 110 de 22 de Dezembro de 2016. http://sistemas.icmbio.gov.br/site_media/ portarias/2017/01/19/Portaria_RPPN_Mata_do Passarinho_01.pdf [accessed on 25 March 2018] IUCN (2024) The IUCN Red List of Threatened Species. Version 2024-1. http://www. iucnredlist.org [accessed on 02 July 2024] Izecksohn E (1976) O status sistematico de Phryniscus proboscideus Boulenger (Am- phibia, Anura, Bufonidae). Revista Brasileira de Biologia 36(2): 341-34. Izecksohn E, Peixoto OL (1980) Sobre a utilizagdo do nome Stombus precrenulatus Mi- randa-Ribeiro, 1937 e a validez da espécie (Amphibia, Anura, Leptodactylidae). Revis- ta Brasileira de Biologia 40(3): 605-609. Jim J, Spirandeli-Cruz EF (1973) Uma nova espécie de Leptodactylus do Estado da Ba- hia, Brasil (Amphibia, Anura). Revista Brasileira de Biologia 39(3): 707-710. Junca FA (2005) Anfibios e Répteis. In: Junca FA, Funch L, Rocha W (Org) Biodiversidade e Conservacgao da Chapada Diamantina. Série Biodiversidade. Brasilia, Ministério do Meio Ambiente 13(1): 339-376. Junca FA (2006) Diversidade e uso de habitat por anfibios anuros em duas localidades de Mata Atlantica, no norte do Estado da Bahia. Biota Neotropica 6(2): 1-8. https:// doi.org/10.1590/S1676-06032006000200018 Junca FA, Pimenta B (2004) Ischnocnema bilineta. In: IUCN Red List of Threatened Spe- cies (International Union for Conservation of Nature IUCN). https://doi.org/10.2305/ IUCN.UK.2004.RLTS.T56462A11468784.en [accessed on 10 August 2022] K6ppen WP (1936) Das geographische System der Klimate. In: Ko6ppen WG, Geiger RM (Eds) Handbuch der Klimatologie Band |, Teil C. Gebriider Borntraeger, Berlin, 44 pp. Lantyer-Silva ASF, Lourengo-De-Moraes R, Siqueira-Jr S, Solé M (2011) Amphibia, Anu- ra, Bufonidae, Rhinella boulengeri Chaparro, Pramuk, Gluesenkamp and Frost, 2007: Distribution extension, state of Bahia, Brazil. Check List 7(6): 825-826. https://doi. org/10.15560/7.6.825 Lantyer-Silva ASF, Siqueira-Jr S, Zina J (2013) Checklist of amphibians in a transitional area between the Caatinga and the Atlantic Forest, central-Southern Bahia, Brazil. Check List 9(4): 725-732. https://doi.org/10.15560/9.4.725 Lavilla EO, Brusquetti F (2018) On the identity of Bufo diptychus Cope, 1862 (Anura: Bufonidae). Zootaxa 4442(1): 161-170. https://doi.org/10.11646/zootaxa.4442.1.9 Lima LR, Bruschi DP Nascimento FAC, Araujo PVS, Costa LP Thomé MITC, Garda AA, Zattera ML, Mott T (2020) Below the waterline: Cryptic diversity of aquatic pipid frogs (Pipa carvalhoi) unveiled through an integrative taxonomy approach. Sys- tematics and Biodiversity 18(8): 1-13. https://doi.org/10.1080/14772000.2020.1 795742 Linares AM, Mello HES (2011) Physalaemus cicada Bokermann, 1966 (Anura: Leiuper- idae): Distribution extension with new South limit and geographic distribution map. Check List 7(6): 859-861. https://doi.org/10.15560/7.6.859 Loebmann D, Haddad CFB (2010) Amphibians and reptiles from a highly diverse area of the Caatinga domain: Composition and conservation implications. Biota Neotropica 10(3): 227-256. https://doi.org/10.1590/S1676-06032010000300026 Loebmann D, Zina J, Araujo OGS, Toledo LF, Haddad CFB (2008) Acoustic Repertory of Hypsiboas exastis (Caramaschi and Rodrigues, 2003) (Amphibia, Hylidae). South ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 238 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil American Journal of Herpetology 3(2): 96-100. https://doi.org/10.2994/1808-9798( 2008)3[96:AROHEC]2.0.CO;2 Lourencgo-De-Moraes R, Dias IR, Mira-Mendes CV, de Oliveira RM, Barth A, Ruas DS, Vences M, Solé M, Bastos RP (2018) Diversity of miniaturized frogs of the genus Adelophryne (Anura: Eleutherodactylidae): A new species from the Atlantic Forest of northeast Brazil. PLoS ONE 13(9): e0201781. https://doi.org/10.1371/journal. pone.0201781 Luedtke JA, Chanson J, Neam K, Hobin L, Maciel AO, Catenazzi A, Borzée A, Hamidy A, Aowphol A, Jean A, Sosa-Bartuano A, Fong G A, de Silva A, Fouquet A, Angulo A, Kidov AA, Munoz Saravia A, Diesmos AC, Tominaga A, Shrestha B, Gratwicke B, Tjaturadi B, Martinez Rivera CC, Vasquez Almazan CR, Senaris C, Chandramouli SR, Striss- mann C, Cortez Fernandez CF, Azat C, Hoskin CJ, Hilton-Taylor C, Whyte DL, Gower DJ, Olson DH, Cisneros-Heredia DF, Santana DJ, Nagombi E, Najafi-Majd E, Quah ESH, Bolafios F, Xie F, Brusquetti F, Alvarez FS, Andreone F, Glaw F, Castafieda FE, Kraus F, Parra-Olea G, Chaves G, Medina-Rangel GF, Gonzalez-Duran G, Ortega-Andrade HM, Machado IF, Das I, Dias IR, Urbina-Cardona JN, Crnobrnja-lsailovi¢ J, Yang J-H, Ji- anping J, Wangyal JT, Rowley JJL, Measey J, Vasudevan K, Chan KO, Gururaja KV, Ovaska K, Warr LC, Canseco-Marquez L, Toledo LF, Diaz LM, Khan MMH, Meegaskum- bura M, Acevedo ME, Napoli MF, Ponce MA, Vaira M, Lampo M, Yanez-Munoz MH, Scherz MD, Rodel M-O, Matsui M, Fildor M, Kusrini MD, Anmed MF, Rais M, Kouamé NGG, Garcia N, Gonwouo NL, Burrowes PA, Imbun PY, Wagner P Kok PJR, Joglar RL, Auguste RJ, Brandao RA, Ibanez R, von May R, Hedges SB, Biju SD, Ganesh SR, Wren S, Das S, Flechas SV, Ashpole SL, Robleto-Hernandez SJ, Loader SP, Inchaustegui SJ, Garg S, Phimmachak S, Richards SJ, Slimani T, Osborne-Naikatini T, Abreu-Jar- dim TPF, Condez TH, De Carvalho TR, Cutajar TP Pierson TW, Nguyen TQ, Kaya U, Yuan Z, Long B, Langhammer P, Stuart SN (2023) Ongoing declines for the world’s amphibians in the face of emerging threats. Nature 622(7982): 308-314. https://doi. org/10.1038/s41586-023-06578-4 Lutz A (1925) Batraciens du Brésil. Comptes Rendus et Mémoires Hebdomadaires des Séances de la Société de Biologie et des ses Filiales. Paris 93(2): 137-139. Lutz A (1926) Observacdes sobre batrachios brasileiros/Observations on brazilian batrachians. Memorias do Instituto Oswaldo Cruz 19(2): 139-174. https://doi. org/10.1590/S0074-02761926000200001 Lynch JD (1972) Generic partitioning of the South American leptodactyloid frog genus Eupsophus Fitzinger, 1843 (sensu Jato). Bulletin of the Southern California Academy of Sciences 71: 2-11. Maciel AO, Hoogmoed MS (2011) Taxonomy and distribution of caecilian amphibians (Gymnophiona) of Brazilian Amazonia, with a key to their identification. Zootaxa 2984(1): 1-53. https://doi.org/10.11646/zootaxa.2984.1.1 Magalhaes FM, Dantas AKBP Brito MRM, Medeiros PHS, Oliveira AF, Pereira TCSO, Queiroz MHC, Santana DJ, Silva WP, Garda AA (2013) Anurans from an Atlantic For- est-Caatinga ecotone in Rio Grande do Norte State, Brazil. Herpetology Notes 6: 1-10. Magalhaes FM, Laranjeiras DO, Costa TB, Junca FA, Mesquita DO, Rohr DL, Silva WP Vie- ira GHC, Garda AA (2015) Herpetofauna of protected areas in the Caatinga IV: Cha- pada Diamantina National Park, Bahia, Brazil. Herpetology Notes 8: 243-261. http:// www.biotaxa.org/hn/article/view/9184/13441 Magalhaes FM, Lyra ML, Carvalho TR, Baldo D, Brusquetti F, Burella P Colli GR, Gehara MC, Giaretta AA, Haddad CFB, Langone JA, Lopez JA, Napoli MF, Santana DJ, De-Sa RO, Garda AA (2020) Taxonomic review of South American butter frogs: phylogeny, ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 239 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil biogeographic patterns, and species delimitation in the Leptodactylus latrans species group (Anura: Leptodactylidae). Herpetological Monograph 34(1): 131-177. https:// doi.org/10.1655/0733-1347-31.4.131 Magalhades FM, Camurugi F, Lyra ML, Baldo D, Gehara MC, Haddad CFB, Garda AA (2022) Ecological divergence and synchronous Pleistocene diversification in the widespread South American butter frog complex. Molecular Phylogenetics and Evolution 169(2): 107398. https://doi.org/10.1016/j.ympev.2022.107398 Magrini L, Carvalho-e-Silva SP, Béda AF, Giaretta AA (2011) Calls of five species of the Scinax ruber (Anura: Hylidae) clade from Brazil with comments on their taxonomy. Zootaxa 3066(1): 37-51. https://doi.org/10.11646/zootaxa.3066.1.3 Magurran AE (1988) Ecological Diversity and its Measurement. Springer, Dordrecht, 179 pp. https://doi.org/10.1007/978-94-015-7358-0 Martins MR, Haddad CFB (1988) Vocalizations and reproductive behaviour in the smith frog, Hyla faber Wied (Amphibia: Hylidae). Amphibia-Reptilia 9(1): 49-60. https://doi. org/10.1163/156853888X00206 Menezes L, Canedo C, Batalha-Filho H, Garda AA, Gehara MC, Napoli MF (2016) Mul- tilocus phylogeography of the treefrog Scinax Eurydice (Anura, Hylidae) reveals a Plio-Pleistocene diversification in the Atlantic Forest. PLoS ONE 11(6): e0154626. https://doi.org/10.1371/journal.pone.0154626 Mikan JC (1822) Delectus Florae et Faunae Brasiliensis. Fasicule 2. Antonii Strauss, Vindobonae, 80 pp. Miles L, Newton AC, DeFries RS, Ravilious C, May I, Blyth S, Kapos V, Gordon JE (2006) A global overview of the conservation status of tropical dry forests. Journal of Biogeog- raphy 33(3): 491-505. https://doi.org/10.1111/j.1365-2699.2005.01424.x Mira-Mendes CB, Ruas DS, Oliveira RM, Castro IM, Dias IR, Baumgarten JE, Junca FA, Solé M (2018) Amphibians of the Reserva Ecoldgica 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 Miranda-Ribeiro A (1926) Notas para servirem ao estudo dos Gymnobatrachios (Anura) brasileiros. Arquivos do Museu Nacional. Museu Nacional 27: 1-227. Miranda-Ribeiro A (1937b) Espécies novas do género “Stombus” da série de apéndices oculares reduzidos. O Campo 8(88): 24. Miranda-Ribeiro A (1937a) Sobre uma colleccao de vertebrados do nordeste brasileiro. Primeira parte: Peixes e batrachios. O Campo 1937: 54-56. Mori SA, Silva LAM (1980) O Herbario do Centro de Pesquisas do Cacau em Itabuna, Brasil. Comissao Executiva do Plano da Lavoura Cacaueira. Boletin Técnico 78: 1-31. https://doi.org/10.2307/2806174 Moura MRD, Santana DJ, Ferreira PL, Feio RN (2009) Amphibia, Anura, Leptodactyli- dae, Leptodactylus viridis Jim and Spirandeli-Cruz, 1979: Distribution extension, new state record, and geographic distribution map. Check List 5(4): 780-782. https://doi. org/10.15560/5.4.780 Myers N, Mittermeyer RA, Fonseca GAB, Kent J (2000) Biodiversity hotspots for con- servation priorities. Nature 403(6772): 853-858. https://doi.org/10.1038/35002501 Narvaes P Rodrigues MT (2009) Taxonomic revision of Rhinella granulosa species group (Amphibia, Anura, Bufonidae), with a description of a new species. Sao Paulo. Arqui- vos de Zoologia 40(1): 1-73. https://doi.org/10.11606/issn.2176-7793.v40i1p1-73 Nascimento LB, Caramaschi U, Cruz CAG (2005) Taxonomic review of the species group of the genus Physalameus Fitzinger, 1826 with revalidation of the genera Engysto- mops Jimenez-de-la-Espada, 1872 and Eupemphix Steindachner, 1836 (Amphib- ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 240 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil ia, Anura, Leptodactylidae). Arquivos do Museu Nacional. Museu Nacional 63(2): 297-320. Novaes-e-Fagundes G, Zina J (2016) Advertisement call of Scinax camposseabrai (Bo- kermann, 1968) (Anura: Hylidae), with comments on the call of three species of the Scinax ruber clade. Zootaxa 4084(2): 258-266. https://doi.org/10.11646/zoot- axa.4084.2.5 Novaes-e-Fagundes G, Araujo-Vieira K, Entiauspe Neto OM, Roberto IJ, Orrico VGD, Solé M, Haddad CFB, Loebmann D (2021) A new species of Scinax Wagler (Hylidae: Sci- naxini) from the tropical forests of northeastern Brazil. Zootaxa 4903(1): 001-041. https://doi.org/10.11646/zootaxa.4903.1.1 Nunes |, Santiago RS, Junca FA (2007) Advertisment calls of four hylid frogs from the state of Bahia, northeastern Brazil (Amphibia, Anura, Hylidae). South American Jour- nal of Herpetology 2(2): 89-96. https://doi.org/10.2994/1808-9798(2007)2[89:A COFHF]2.0.C0;2 Oliveira RM, Ruas DS, Mira-Mendes CV, Solé M (2014) Advertisement call of Rhinella cru- cifer (Wied-Neuwied, 1821) (Anura: Bufonidae) from southern Bahia, Brazil. Zootaxa 3784(1): 97-98. https://doi.org/10.11646/zootaxa.3784.1.9 Orrico VGD, Carvalho-e-Silva AMPT, Carvalho-e-Silva SP (2006) Redescription of the ad- vertisement call of Aplastodiscus arildae (Cruz & Peixoto) and description of the call of Aplastodiscus weygoldti (Cruz & Peixoto) with general notes about the genus in Southeastern Brazil (Anura, Hylidae). Revista Brasileira de Zoologia 23(4): 994-1001. https://doi.org/10.1590/S0101-81752006000400003 Orrico VGD, Nunes |, Mattedi C, Fouquet A, Lemos AW, Rivera-Correa M, Lyra ML, Loeb- mann D, Pimenta BVS, Caramaschi U, Rodrigues MT, Haddad CFB (2017) Integrative taxonomy supports the existence of two distinct species within Hypsiboas crepitans (Anura: Hylidae). Salamandra (Frankfurt) 53(1): 99-113. Orrico VGD, Grant T, Faivovich J, Rivera-Correa M, Rada M, Lyra ML, Cassini CS, Valdujo PH, Schargel WE, Machado DJ, Wheeler WC, Barrio-Amoros CL, Loebmann D, Mora- vec J, Zina J, Solé M, Sturaro MJ, Peloso PLV, Sudrez P Haddad CFB (2021) The phylogeny of Dendropsophini (Anura: Hylidae: Hylinae). Cladistics 37(1): 73-105. https://doi.org/10.1111/cla.12429 Pedreira AJ (1999) Evolucao sedimentar e tecténica da bacia metassedimentar do Rio Pardo: Uma sintese. Revista Brasileira de Geociencias 29(3): 339-344. https://doi. org/10.25249/0375-7536.199929339344 Pedrosa IMMC, Costa TB, Faria RG, Franga FGR, Laranjeiras DO, Oliveira TCSP, Palmeira CNS, Torquato S, Mott T, Vieira GHC, Garda AA (2014) Herpetofauna of protected areas in the Caatinga III: The Catimbau National Park, Pernambuco, Brazil. Biota Neo- tropica 14(4): 1-12. https://doi.org/10.1590/1676-06032014004614 Peixoto OL, Arzabe C (2004) Scinax pachycrus. The IUCN Red List of Threatened Spe- cies. https://doi.org/10.2305/IUCN.UK.2004.RLTS.T55984A11391526.en [accessed on 01 March 2016] Pimenta BVS, Faivovich J, Pombal Jr JP (2007) On the identity of Hyla strigilata Spix, 1824 (Anura: Hylidae): redescription and neotype designation for a “ghost” taxon. Zootaxa 1441(1): 35-49. https://doi.org/10.11646/zootaxa.1441.1.3 Pirani RM, Tonini JFR, Thomaz AT, Napoli MF, Encarnagao LC, Knowles LL, Werneck FP (2022) Deep genomic divergence and phenotypic admixture of the treefrog Dendrop- sophus elegans (Hylidae: Amphibia) coincide with riverine boundaries at the Brazil- ian Atlantic Forest. Frontiers in Ecology and Evolution 10(765977): 1-15. https://doi. org/10.3389/fevo.2022.765977 ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 241 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil Protazio AS, Santos-Protazio A, Silva CRS, Ribeiro ES, Nogueira EMS, Moura GJB (2010) Anurofauna do Municipio de Paulo Afonso-BA, Bioma Caatinga, Nordeste do Brasil. Revista Nordestina de Zoologia 4(2): 31-38. Protazio AS, Protazio AS, Silva LS, Conceigao LC, Braga HSN, Santos UG, Ribeiro AC, Almeida AC, Gama V, Vieira MVSA, Silva TAF (2021) Amphibians and reptiles of the Atlantic Forest in Rec6ncavo Baiano, east Brazil: Cruz das Almas municipality. ZooK- eys 1060: 125-153. https://doi.org/10.3897/zookeys.1060.62982 Reinhardt JT, Liitken CF (1862) Bidrag til Kundskab om Brasiliens Padder og Kryb- dyr. Forste Afdeling: Padderne og Oglerne. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kj@benhavn, Serie 2 3: 143-242. Ribeiro MC, Metzger JP, Martensen AC, Ponzoni FJ, Hirota MM (2009) The Brazilian At- lantic Forest: How much is left, and how is the remaining forest distributed? Impli- cations for conservation. Biological Conservation 142(6): 1141-1153. https://doi. org/10.1016/j.biocon.2009.02.021 Rodrigues MT (2003) Herpetofauna da Caatinga. In: Leal IR, Tabarelli M, Silva JMC (Eds) Ecologia e conservagao da Caatinga. Editora Universitaria da UFPE, Recife, 181-236. Rojas-Padilla O, Menezes VQ, Dias IR, Argdlo AJS, Solé M, Orrico VGD (2020) Amphibi- ans and reptiles of Parque Nacional da Serra das Lontras: An importante center of en- demism within the Atlantic Forest in southern Bahia, Brazil. ZooKeys 1002: 159-185. https://doi.org/10.3897/zookeys.1002.53988 Rossa-Feres DC, Garey MV, Caramaschi U, Napoli MF, Nomura F, Bispo AA, Brasileiro CA, Thome MTC, Sawaya RJ, Conte CE, Cruz CAG, Nascimento LB, Gasparini JL, Almeida AP, Haddad CFB (2017) Anfibios da Mata Atlantica: Lista de espécies, historico dos estudos, biologia e conservacao. In: Monteiro-Filho ELA, Conte CE (Eds) Revis6es em Zoologia: Mata Atlantica. Editora UFPR, Curitiba, 237-314. Santana DJ, Mesquita DO, Garda AA (2011) Advertisement call of Dendropsophus ol- iveirai (Anura, Hylidae). Zootaxa 2997(1): 67-68. https://doi.org/10.11646/zoot- axa.2997.1.5 Santana DJ, Mangia S, Silveira-Filho RR, Barros LCS, Andrade |, Napoli MF, Junca FA, Garda AA (2015) Anurans from the Middle Jaguaribe River Region, Ceara State, Northeastern Brazil. Biota Neotropica 15(3): 1-8. https://doi.org/10.1590/1676- 06032015001715 Santos SPL, Santos EM (2011) Anurofauna da Reserva Particular do Patrimdénio Natural Frei Caneca, Municipio de Jaqueira, Estado de Pernambuco, Brasil. In: Moura GJB, Santos EM, Oliveira MAB, Cabral MCC (Org) Herpetofauna de Pernambuco. Brasilia, Ministério do Meio Ambiente: 187-198. Santos L, Roseno RS, Solé M, Dias IR (2023) Another new species (and it’s not over yet) of Phyllodytes Wagler, 1930 (Anura, Hylidae) from the Atlantic Forest of south- ern Bahia, northeastern Brazil. Zootaxa 5374(4): 519-532. https://doi.org/10.11646/ zootaxa.5374.4.4 Sao Pedro VA, Medeiros PH, Garda AA (2011) The advertisement call of Rhinella granulosa (Anura, Bufonidae). Zootaxa 3092(1): 60-62. https://doi.org/10.11646/zootaxa.3092.1.4 Schneider JG (1799) Historia Amphibiorum Naturalis et Literarariae. Fasciculus Primus. Continens Ranas, Calamitas, Bufones, Salamandras et Hydros in Genera et Species Descriptos Notisque suis Distinctos. Friederici Frommann, Jenae, 264 pp. https://doi. org/10.5962/bhI.title. 78757 Sichieri GRF, Cruz CAG, Pimenta BVS, Nunes | (2021) Advertisement call description of two Proceratophrys species (Anura: Odontophrynidae). Zootaxa 4975(2): 397-400. https://doi.org/10.11646/zootaxa.4975.2.10 ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 242 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil Silva JF, Farifias MR, Felfili JM, Klink CA (2006) Spatial heterogeneity, land use and con- servation in the cerrado region of Brazil. Journal of Biogeography 33(3): 536-548. https://doi.org/10.1111/j.1365-2699.2005.01422.x Silva CRS, Moura GJB, Andrade EVE, Nogueira EMS (2011) Aspectos Ecoldégicos da Anurofauna dos municipios de Jatoba-PE e Paulo Afonso-BA, Nordeste do Brasil. In: Moura GJB, Santos EM, Oliveira MAB, Cabral MCC (Org) Herpetofauna de Pernambu- co. Brasilia, Ministério do Meio Ambiente 1(1): 177-185. Silva GR, Luna-Dias C, Hepp FSFS, Carvalho-e-Silva AMPT, Carvalho-e-Silva SP (2012) New record of Aplastodiscus weygoldti (Cruz & Peixoto, 1987) in the municipality of Mimoso do Sul, Espirito Santo State, southeastern Brazil (Anura, Hylidae). Herpetol- ogy Notes 5: 371-373. Silva-Soares T, Ferreira RB, Ornellas IS, Zocca C, Caramaschi U, Cruz CAG (2021) A new species of Ischnocnema (Anura: Brachycephalidae) from the mountainous re- gion of Atlantic Forest, southeastern Brazil, with a new phylogeny and diagnose for Ischnocnema parva series. Zootaxa 5082(3): 201-222. https://doi.org/10.11646/ zootaxa.5082.3.1 Silvano DL, Pimenta BVS (2003) Diversidade de anfibios na Mata Atlantica do Sul da Bahia. In: Prado Pl, Landau EC, Moura RT, Pinto LPS, Fonseca GAB, Alger K (Eds) Corredor de Biodiversidade na Mata Atlantica do Sul da Bahia CD-ROM. IESB/CI/ CABS/ UFMG/UNICAMP7 IIhéus, Bahia, Brazil, 1-22. Silvano DL, Segalla MV (2005) Conservacao de anfibios no Brasil. Megadiversidade 1(1): 79-86. Sos Mata Atlantica [Fundacado Sos Mata Atlantica], INPE [Instituto Nacional de Pesqui- sas Espaciais] (2018) Atlas dos remanescentes florestais da Mata Atlantica: periodo 2016-2017. Relatorio Técnico, ArcPlan, Sao Paulo, 63 pp. Spix JBv (1824) Animalia nova sive Species novae Testudinum et Ranarum quas in itine- re per Brasiliam annis MDCCCXVII-MDCCCXxX jussu et auspiciis Maximiliani Josephi |. Bavariae Regis. Miinchen, F. S. Hibschmann, ii, 53, 39 plates (numbered 1-17 and 1-22) pp. Steffen GA (1815) De Ranis nonnullis Observationes Anatomicae quas Consensu Gra- tiosae Facultatis Medicae. Joannis Friderici Starckii, Berlin, 24 pp. Stevaux MN (2002) A new species of Bufo Laurenti (Anura, Bufonidae) from northeastern Brazil. Revista Brasileira de Zoologia 19(suppl 1): 235-242. https://doi.org/10.1590/ $0101-81752002000500018 Tabarelli M, Silva JMC (2003) Areas e acées prioritarias para a conservacdo da Caat- inga. In: Leal |, Tabarelli M, Silva JMC (Eds) Ecologia e Conservacao da Caatinga. Editora Universitaria da UFPE, Recife, 777-796. Toledo LF, Castanho LM, Haddad CFB (2005) Recognition and distribution of Lepto- dactylus mystaceus (Anura; Leptodactylidae) in the State of Sao Paulo, South- eastern Brazil. Biota Neotropica 5(1): 57-62. https://doi.org/10.1590/S1676- 06032005000100006 Toti DS, Coyle FA, Miller JA (2000) A structured inventory of Appalachian grass bald and heath bald spider assemblages and a test of species richness esti- mator performance. The Journal of Arachnology 28(3): 329-345. https://doi. org/10.1636/0161-8202(2000)028[0329:ASIOAG]2.0.C0;2 Trevisan CC, Batalha-Filho H, Garda AA, Menezes L, Dias IR, Sole M, Canedo C, Junca FA, Napoli MF (2020) Cryptic diversity and ancient diversification in the northern Atlantic Forest Pristimantis (Amphibia, Anura, Craugastoridae). Molecular Phylogenetics and Evolution 148(4): 106811. https://doi.org/10.1016/j.ympev.2020.106811 ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 243 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil Valdujo PH, Recoder RS, Vasconcellos MM, Portella AS (2009) Amphibia, Anura, Sao Desidério, western Bahia uplands, northeastern Brazil. Check List 5(4): 903-911. https://doi.org/10.15560/5.4.903 Valdujo PH, Camacho A, Recoder RS, Teixeira M Junior, Ghellere JMB, Mott T, Nunes PMS, Nogueira CC, Rodrigues MT (2011) Anfibios da Estacao Ecoldgica Serra Geral do Tocantins, regiao do Jalapao, Estados do Tocantins e Bahia. Biota Neotropica 11(1): 251-261. https://doi.org/10.1590/S1676-06032011000100025 Vaz-Silva W, Maciel NM, Nomura F, Morais AR, Guerra Batista V, Santos DL, Andrade SP, Oliveira AAB, Brandaéo RA, Bastos RP (2020) Guia de identificacaéo das es- pécies de anfibios (Anura e Gymnophiona) do estado de Goias e do Distrito Fed- eral, Brasil Central. Sociedade Brasileira de Zoologia, Curitiba, 229 pp. https://doi. org/10.7476/9786587590011 Vieira WLS, Arzabe C, Santana GG (2007) Composicao e distribuicao espaco-tempo- ral de anuros no Cariri Paraibano, Nordeste do Brasil. Oecologia Brasiliensis 11(3): 383-396. https://doi.org/10.4257/oeco.2007.1103.08 Vilaga TRA, Silva JES, Solé M (2006) Vocalization and territorial behaviour of Phyllo- medusa nordestina Caramaschi, 2006 (Anura: Hylidae) from southern Bahia, Brazil. Journal of Natural History 45(29-30): 1823-1834. https://doi.org/10.1080/0022293 3.2011.561018 von Tschudi JJ (1838) Classification der Batrachier mit Beriicksichtigung der fossi- len Thiere dieser Abtheilung der Reptilien. Petitpierre, Neuchatel, 99 pp. https://doi. org/10.5962/bhi.title.4883 Vords J, Dias JR, Solé M (2017) A new species of Phyllodytes (Anura: Hylidae) from the Atlantic Rainforest of Southern Bahia, Brazil. Zootaxa 4337(4): 584-594. https://doi. org/10.11646/zootaxa.4337.4.9 Wells KD (2007) The ecology and behavior of amphibians. The University of Chicago Press Books, Chicago, 1400 pp. Wied-Neuwied MAP (1821) Reise nach Brasilien in den Jahren 1815 bis 1817. Vol. 2. Henrich Ludwig Bronner, Frankfurt, 345 pp. Wied-Neuwied MAP (1824) Abbildungen zur Naturgeschichte Brasiliens. Heft 8. Lan- des-Industrie-Comptoir, Weimar, 614 pp. Willians PH (1996) Mapping variations in strength and breadth of biogeographic tran- sition zones using species turnover. Proceedings. Biological Sciences 263(1370): 579-588. https://doi.org/10.1098/rspb.1996.0087 Young BE, Stuart SN, Chanson JS, Cox NA, Boucher TM (2004) Joyas que estan desa- pareciendo: El estado de los anfibios en el Nuevo Mundo. NatureServe, Arlington, Virginia, USA, 53 pp. Zappi DC, Filardi FLR, Leitman P, Souza VC, Walter BMT, Pirani JR, Morim MP, Queiroz LP Cavalcanti TB, Mansano VF, Forzza RC (2015) Growing knowledge: An over- view of Seed Plant diversity in Brazil. Rodriguésia 66(4): 1085-1113. https://doi. org/10.1590/2175-7860201 56641 1 Zucchetti VM, Castroviejo-Fisher S (2024) Lost in time: unraveling the identity of Vitreo- rana parvula (Boulenger, 1895) (Anura: Centrolenidae). Zootaxa 5415(3): 351-391. https://doi.org/10.11646/zootaxa.5415.3.1 Zucchetti VM, Rojas-Padilla O, Dias IR, Solé M, Orrico VGD, Castroviejo-Fisher S (2023) An elusive giant: A new species of Vitreorana Guayasamin et al., 2009 (Anura: Cen- trolenidae) from the northern Atlantic Forest with an osteological description and comments on integumentary spicules. Zootaxa 5249(3): 301-334. https://doi. org/10.11646/zootaxa.5249.3.1 ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 244 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil Appendix 1 List of vouchers deposited in the Museu de Zoologia da Universidade Estadual de Santa Cruz-MZUESC. AMPHIBIANS BRACHYCEPHALIDAE-/schnocnema verrucosa: MZUESC 15874, 15875, 15885, 15886, 15895, 15898, 15899. Ischnocnema sp. (gr. parva): MZUESC 15896. BUFONIDAE-Dendrophryniscus proboscideus: MZUESC 14688, 15126, 15127, 15128, 15798, 15803, 15859, 16532. Rhinella crucifer: MZUESC 15031, 15148. Rhinella granulosa: MZUESC 15030, 15055, 15504, 15505, 15506, 15850. Rhi- nella diptycha: MZUESC 15503. CRAUGASTORIDAE-Bahius bilineatus: MZUE- SC 15095, 15096, 15097, 15794, 15809, 15838, 15842, 15855, 15856, 15860, 15882, 15883. Haddadus binotatus: MZUESC 15014, 15015, 15032, 15034, 15044, 15047, 15051, 15053, 15054, 15057, 15058, 15104, 15105, 15106, 15107, 15131, 15142, 15465, 15471, 15472, 15473, 15475, 15476, 15478, 15479, 15482, 15526, 15622, 15623, 15634, 15635, 15643, 15644, 15646, 15650, 15651, 15656, 15659, 15660, 15778, 15802. Pristimantis vinhai: MZUE- SC 15000, 15098-15103, 15109-15112, 15114, 15115, 15120, 15121, 15132, 15133, 15140, 15141, 15143, 15144, 15464, 15481, 15502, 15640, 15642, 15657, 15663, 15664, 15667, 15780, 15785, 15786, 15797, 15799-15801, 15806-15808, 15813-15815, 15837, 15840, 15841, 15843, 15853, 15854, 15857, 15858, 15861, 15879, 15897, 15889. Pristimantis sp. (gr. ramagii): MZU- ESC 15033, 15474, 15641, 15658, 15662, 15665, 15783, 15849, 15851, 15852, 16523. CENTROLENIDAE-Vitreorana eurygnatha: MZUESC 14691, 14698, 14699, 14700, 14701, 15810-15812. CYCLORAMPHIDAE-Thoropa miliaris: MZUESC 14674, 15130, 15477, 15782, 15789, 15845, 15846, 15872, 15873, 15880, 15881. ELEUTHERODACTYLIDAE-—Adelophryne sp.8: MZUESC 15035, 15036, 15039, 15041, 15043, 15048, 15050, 15052, 15116, 15462, 15466, 15480, 15528-15531, 15625-15627, 15631, 15633, 15637-15639, 1564/7, 15652, 15661, 15796, 15830, 15834, 15835, 15839, 15862, 16522, 16530. Ade- lophryne sp.2: MZUESC 15024, 15037, 15038, 15040, 15042, 15045, 15046, 15049, 15090, 15091, 15094, 15113, 15118, 15119, 15123-15125, 15457- 15460, 15467-15470, 15524, 15525, 15527, 15533, 15624, 15628, 15629, 15632, 15636, 15645, 15648, 15649, 15655, 15795, 15831, 15832, 15836. HYL- IDAE-Aplastodiscus weygoldti: MZUESC 14690, 15787, 15788, 15844, 15887. Boana crepitans: MZUESC 14673, 14675, 14998, 14999, 15002, 15145, 15486. Boana exastis: MZUESC 15108. Boana faber: MZUESC 14676, 15066, 15068, 15147, 15666, 15884. Dendropsophus branneri: MZUESC 14683, 14684, 15022, 15023, 15509, 15510, 15512, 15818, 15824. Dendropsophus elegans: MZUE- SC 14678, 14679, 15018, 15019, 15020, 15021, 15138, 15507, 15508, 15820. Dendropsophus oliveirai: MZUESC 14686, 15010, 15011, 15487-15501, 15511, 15791, 15792, 15823, 15825. Ololygon strigilata: MZUESC 15001. Phyllodytes luteolus: MZUESC 17501, 17502. Pithecopus nordestinus: MZUESC 14680- 14682, 14685, 14695, 14997, 15136, 15137, 15826, 15827, 16527. Pseudis fus- ca: MZUESC 16528. Scinax eurydice: MZUESC 14672, 14693, 15876, 16531. Scinax pachycrus: MZUESC 15790, 16525. Scinax x-signatus: MZUESC 14692, 14694, 15027, 15139, 15892, 15894, 17503. Sphaenorhynchus prasinus: MZU- ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 245 Carlos Augusto Souza-Costa et al.: Amphibians of Serra Azul and Serra do Mandim in Southwestern Bahia, Brazil ESC 14677, 15004-15009, 15016, 15017, 15135, 15484, 15485, 15793, 15821, 15822. Trachycephalus nigromaculatus: MZUESC 15063, 15064, 15065, 15483, 15888. LEPTODACTYLIDAE-Leptodactylus cf. mystaceus: MZUESC 14696. Leptodactylus fuscus: MZUESC 15012, 15067, 15513-15516. Leptodactylus latrans: MZUESC 15871, 15893. Leptodactylus macrosternum: MZUESC 15146, 16524. Leptodactylus mystacinus: MZUESC 16529, 16533. Leptodactylus trog- lodytes: MZUESC 15003, 15028, 15029. Leptodactylus viridis: MZUESC 15848, 15869, 15870. Physalaemus cicada: MZUESC 15059-15062. Physalaemus cf. erikae: MZUESC 15013, 15878. Physalaemus kroyeri: MZUESC 15517, 15518, 15521, 15523, 15784, 15819, 15877. MICROHYLIDAE — Dermatonotus muelleri: MZUESC 15069, 15070. ODONTOPHRYNIDAE-Proceratophrys schirchi: MZUE- SC 14689, 14702-14705, 15071, 15072, 15122, 15134, 15779, 15781, 15804, 15816, 15817, 15833, 15847, 15863-15868. PIPIDAE-Pipa carvalhoi: MZUESC 15129. SIPHONOPIDAE-Siphonops annulatus: MZUESC 15900. ZooKeys 1217: 215-246 (2024), DOI: 10.3897/zookeys.1217.119844 246