Herpetofauna of a Cerrado-Caatinga transition area in the municipality of Pedro II, Piauí state, Northeastern Brazil

Herpetofauna de uma área de transição Cerrado-Caatinga no município de Pedro II, estado do Piauí, Nordeste do Brasil

Abstract

Anthropogenic disturbances represent a direct threat to populations of amphibians and reptiles. Therefore, herpetofaunal checklists are important to understand the species distribution, to fill sampling gaps, and aiding protection initiatives for these animals. Herein, we present the first herpetofaunal checklist of the municipality of Pedro II, Piauí state, Northeastern Brazil. It is inserted in the Complexo da Serra Grande bioregion within a transitional area between the Caatinga and Cerrado biomes. We searched for amphibians and reptiles using a non-systematic methodology over five years, from January 2019 and April 2023. We found 82 species, 27 amphibians, and 55 reptiles, representing the third region with the highest species richness in Piauí state. Except for the species with deficient data or not evaluated, all amphibians and reptiles are classified as least concern regarding the risk extinction. Most amphibians and reptiles reported in Pedro II are widely distributed in Brazilian biomes, but three snakes are endemic to the Caatinga biome and transitional zones: Boiruna sertaneja, Erythrolamprus viridis, and Epictia borapeliotes. Additionally, it is the only region in Piauí state where the species Kinosternon scorpioides, Epictia borapeliotes, and Micrurus lemniscatus occur. Almost half of the species of amphibians and reptiles known in Piauí state occur in the municipality of Pedro II, which might be associated with different phytophysiognomies within the Caatinga and Cerrado biomes influencing this region. Overall, our work contributes to unveiling the rich herpetofauna of this understudied area from the Complexo da Serra Grande bioregion, raising awareness of the need for effective conservation plans to maintain and preserve this real herpetofaunal “hotspot” in Piauí state.

Keywords
Amphibians; conservation; ecotonal areas; Ibiapaba complex; Serra dos Matões; reptiles

Resumo

Perturbações antropogênicas representam uma ameaça direta às populações de anfíbios e répteis. Portanto, inventários da herpetofauna são importantes para entender a distribuição das espécies, preencher lacunas amostrais e auxiliar iniciativas de proteção desses animais. Aqui, apresentamos a primeira lista de espécie da herpetofauna do município de Pedro II, Piauí, Nordeste do Brasil. Pedro II está inserido na biorregião do Complexo da Serra Grande, em uma área de transição entre os biomas Caatinga e Cerrado. Buscamos anfíbios e répteis usando uma metodologia não sistematizada durante cinco anos, entre janeiro de 2019 e abril de 2023. Encontramos 82 espécies de herpetofauna, 27 anfíbios e 55 répteis, representando a terceira região com maior riqueza de espécies no Piauí. Com exceção das espécies com dados deficientes ou não avaliadas, todos os anfíbios e répteis são classificados como de menor preocupação quanto ao risco de extinção. A maioria dos anfíbios e répteis relatados em Pedro II estão amplamente distribuídos nos biomas brasileiros, mas três serpentes são endêmicas do bioma Caatinga e zonas de transição: Boiruna sertaneja, Erythrolamprus viridis e Epictia borapeliotes. Além disso, é a única região onde as espécies Kinosternon scorpioides, Epictia borapeliotes e Micrurus lemniscatus ocorrem no estado do Piauí. Quase metade das espécies de anfíbios e répteis conhecidas no estado do Piauí ocorrem no município de Pedro II, o que pode estar associado a diferentes fisionomias vegetais dentro dos biomas Caatinga e Cerrado influenciando esta região. No geral, nosso trabalho contribui para revelar a rica herpetofauna desta área pouco estudada da biorregião do Complexo da Serra Grande, aumentando a conscientização sobre a necessidade de planos de conservação eficazes para manter e preservar este verdadeiro “hotspot” da herpetofauna no estado do Piauí.

Palavras-chave
Anfíbios; conservação; áreas ecotonais; complexo da Ibiapaba; Serra dos Matões; répteis

Introduction

The Caatinga and Cerrado biomes comprise part of the dry diagonal of the Neotropical region, and they are responsible for harboring a large part of the biodiversity known in South America (Werneck 2011). However, although they are considered biomes with high biological diversity, their knowledge is still quite underestimated, as evidenced by the countless species described each year (Mângia et al. 2018, Barbo et al. 2022, Dubeux et al. 2022). The transition between biomes and ecosystems does not have well-defined limits, and it is consensus ecotonal areas can harbor a unique and peculiar biodiversity, including endemism (Smith et al. 2001, Yarrow and Marín 2007, Nogueira et al. 2010, Gutiérrez and Marinho-Filho 2017). However, these biomes have been impacted by global climatic changes, forest fires, and anthropogenic disturbances, which cause the loss of natural habitats and, consequently, fauna and flora (Araujo et al. 2023, Fiorillo et al., 2023).

Amphibians and reptiles are two of the most diverse vertebrate groups, with 8877 (Frost, 2024) and 12,386 (Uetz et al. 2025) species registered, respectively. These animals have important ecosystem functions, participating in diverse food networks and providing ecosystem services such as biological pest control, seed dispersal, and water quality (Cortéz-Gómez et al. 2015), thus being important environmental bioindicators (Saber et al. 2017, Jacob et al. 2024). The Caatinga and Cerrado biomes host a high diversity of species, including many endemics (Guedes, Nogueira, and Marques 2014; Azevedo et al. 2016; Garda et al. 2018); however, human activities have impacted these herpetofaunal populations (Hill et al. 2019; Fiorillo et al. 2023; With et al. 2024). Amongst the main threats in the Caatinga and Cerrado biomes are habitat fragmentation, substitution of natural areas for agriculture and pasture, pollution, and exotic species (Araujo et al. 2023, Fiorillo et al., 2023). Therefore, carrying out herpetofauna inventories is important for understanding the health and dynamics of the local population and for studies on the distribution and degree of threat to species, enabling the filling of geographic sampling gaps and subsidizing species conservation and preservation plans (Oitaven et al. 2021).

With extensive areas covered by the Cerrado and Caatinga biomes, the Piauí state harbors a rich and diverse vegetation (Castro et al. 2007), reflecting on its herpetofauna. Most studies in Piauí state dealing with amphibians and reptiles were carried out in protected areas (Pantoja et al. 2022; Rocha and Prudente 2010, Dal Vechio et al. 2013, Cavalcanti et al. 2014, Dal Vechio et al. 2016, Araújo et al. 2020a, b, Marques et al. 2023), with few works in unprotected areas (e.g., Rodrigues and Prudente 2011, Benício, Silva and Fonseca 2014, Benício, Silva and Fonseca 2015, Benício, Lima and Fonseca 2015). These studies highlight the rich diversity of amphibians and reptiles in Piauí state. However, there are some regions understudied, such as the Complexo da Serra Grande bioregion, where most of herpetofaunal studies were performed just in the Ibiapaba plateau, Ceará state (e.g., Loebmann and Haddad 2010, Castro et al. 2019). As part of this complex, the municipality of Pedro II is under the influence of different phytophysiognomies within a transitional area between the Caatinga and Cerrado biomes (IBGE 2019). However, herpetological studies are incipient, typically limited to research on range extensions and natural history (e.g., Araújo et al. 2020a, c, Araújo et al. 2022a, b, Uchôa et al. 2023, Oliveira et al. 2023). Thus, we present the first herpetofaunal checklist of the municipality of Pedro II, Piauí state, Northeastern Brazil, with data about the conservation status and distribution of amphibians and reptiles.

Material and Methods

1. Study area

This study was conducted in the municipality of Pedro II (PedII), located in the Center-North region of Piauí state, Northeastern Brazil (04°25’23’’ S, 41°27’34’’ W, WGS84 datum, Figure 1). It is inserted within the Environmental Protection Area (EPA) Serra da Ibiapaba and belongs to the Complexo da Serra Grande bioregion, which comprises a group of mountain ranges inserted in the Ibiapaba plateau with altitudes ranging from 200 to 820 m (Lima and Guerra 2020), with the highest altitude is restricted to Serra dos Matões (Gomes 2004). Pedro II is located in the Parnaíba Sedimentary Basin, in a transitional area between the Caatinga and Cerrado biomes, characterized by the presence of typical Cerrado vegetation at mid and high altitudes, whereas the lowlands are characterized by the presence of Caatinga on the rocky outcrops (Gomes 2004, IBGE 2019). Concentrated between December and May, the annual mean precipitation is 1043 mm (Aguiar and Gomes, 2004) and the annual mean temperatures range from 18 to 30 °C (Freitas, Gomes and Aquino 2016).

Figure 1
Geographic location map of amphibian and reptile collection points in the municipality of Pedro II, state of Piauí, Northeastern Brazil, highlighting the Serra da Ibiapaba Environmental Protection Area (EPA). States of the Brazilian Northeast: Maranhão - MA; Piauí - PI; Ceará - CE, Rio Grande do Norte - RN; Paraíba - PB; Pernambuco - PE; Alagoas - AL; Sergipe - SE.

2. Sampling

We searched for amphibians and reptiles in a non-systematic method over five years, from January 2019 to April 2023, using complementary sampling methods in different natural areas of Pedro II, Piauí state. For amphibians, we carried out species searches mainly from January 2019 to January 2021, conducting searches biweekly during the rainy season (January–May) and monthly during the dry season (June–December), with an average duration of two days using visual search and auditory search methods (Crump and Scott Jr. 1994). We conducted complementary random searches during the rainy seasons of 2022 and 2023. Samplings started around 18h00 and ended at 23h00 depending on the vocalization activity of anurans and the number of environments visited per night (see Table 1 to consult the geographical coordinates of the visited sampling points). Due to the qualitative nature of this study, the collection time was not standardized, being carried out by three to six people.

Table 1
Collection points for amphibians and reptiles in the municipality of Pedro II, state of Piauí, Northeastern Brazil. The species collected are represented by number according to tables 2 and 3.

Collection point	Taxon	Species collected	Latitude	Longitude
Açude Joana	Anura/Reptilia	6, 9, 12, 17, 26, 59, 69, 70	4°25′36.98″S	41°27′55.22″W
Açude Joana	Anura	7, 14, 22, 17, 24	4°25′41.07″S	41°28′5.81″W
Açude Joana	Reptilia	32	4°25′39.60″S	41°28′01.70″W
Bairro Areia Branca	Anura/Reptilia	1, 13, 14, 16, 41, 49	4°26′23.00″S	41°27′31.70″W
Bairro Centro	Anura	25	4°25′32.70″S	41°27′34.27″W
Bairro Centro	Anura	5	4°25′42.24″S	41°27’4.84″W
Bairro Centro	Reptilia	34, 39, 50	4°25′45.60″S	41°27′36.90″W
Bairro Engenho Novo	Reptilia	67, 81	4°26′59.44″S	41°27′22.14″W
Bairro Santa Fé	Reptilia	36	4°25′05.01″S	41°26′57.83″W
Bairro São Gonçalo	Reptilia	4, 60, 70, 76	4°26′36.04″S	41°28′20.43″W
Bairro São Gonçalo	Reptilia	75	4°26′34.46″S	41°28′20.43″W
Cachoeira do Buriti	Reptilia	78	4°19′41.83″S	41°27′48.49″W
Cachoeira Urubu Rei	Anura	13, 24	4°19′12.08″S	41°27′49.45″W
Cachoeira Urubu Rei	Reptilia	32, 43, 51	4°19′44.50″S	41°27′33.90″W
Comunidade Aroeira	Anura	27	4°26′23.16″S	41°24′28.47″W
Comunidade Buritizinho	Reptilia	32	4°24′49.40″S	41°26′28.30″W
Comunidade Buritizinho	Anura	22	4°24′52.22″S	41°26′31.96″W
Comunidade Caranguejo	Reptilia	54, 56	4°19′39.64″S	41°26′50.62″W
Comunidade Chã do Lambedor	Reptilia	29	4°27′53.27″S	41°28′29.83″W
Comunidade Corrente	Reptilia	30, 41, 64	4°28′51.05″S	41°24′42.71″W
Comunidade Corrente	Reptilia	40	4°29′15.50″S	41°24′38.00″W
Comunidade Corrente	Anura/Reptilia	2, 5, 6, 8, 9, 11, 21, 25, 27, 52, 57, 66, 71, 72, 74, 77	4°29′6.34″S	41°24′38.84″W
Comunidade Grota	Reptilia	54	4°29′56.00″S	41°28′59.00″W
Comunidade Pequis	Anura/Reptilia	5, 16, 18, 19, 30, 31, 33, 34, 40, 42, 43, 44, 45, 47, 49, 54, 65, 68, 70, 73, 79, 80	4°30′34.00″S	41°29′20.00″W
Comunidade Pimenta*	Reptilia	37, 48, 58, 61	4°25′13.04″S	41°27′14.10″W
Comunidade Riacho Fundo*	Anura/Reptilia	9, 22, 19, 35, 36, 39, 47, 48, 50	4°25′24.34″S	41°28′46.67″W
Comunidade Santana	Reptilia	53, 57, 74	4°27′21.57″S	41°26′8.60″W
Comunidade Santana	Reptilia	73	4°27′39.11″S	41°25′45.84″W
Comunidade Santana	Reptilia	60	4°27′56.51″S	41°26′7.34″W
Comunidade São Luís	Reptilia	67	4°32′31.35″S	41°20′43.00″W
Comunidade Serra dos Matões	Anura	6, 9, 19	4°23′40.82″S	41°26′48.78″W
Comunidade Serra dos Matões	Anura/Reptilia	8, 9, 15, 21, 22, 57	4°24′4.94″S	41°26′45.48″W
Comunidade Serra dos Matões	Anura	23	4º21′58.00″S	41º26′36.00W
Comunidade Serra dos Matões	Anura	8, 30	4º23′55.32″S	41º26′48.30″W
Comunidade Tamboril	Reptilia	73	4°29′15.58″S	41°34′13.99″W
Comunidade Torres	Reptilia	55	4°27′27.60″S	41°25′37.09″W
Comunidade Torres	Reptilia	53,	4°28′3.13″S	41°25′13.20″W
Comunidade Vale do São Francisco	Anura	5	4°39′46.09″S	41°12′51.67″W
Bairro Cristo Reis	Anura	3, 23, 10, 21, 26	4°25′44.13″S	41°27′8.29″W
Localidade Samambaia	Anura	13, 24	4°19′46.36″S	41°27′43.50″W
Parque Municipal do Pirapora	Anura/Reptilia	20, 46, 76	4°25′54.00″S	41°27′24.00″W
Bairro Vila Operária	Reptilia	76	4°26′2.14″S	41°27′13.68″W
Trilha da Samambaia	Anura/Reptilia	14, 20, 33, 63	4°19′45.70″S	41°27′20.70″W

We searched for reptiles from September 2020 to November 2021 using four sampling methods: passive collections using pitfall trap, visual active search, occasional encounters, and collection by third parties (Foster 2012). We installed one pitfall traps in two sampling points located in the rural zone of Pedro II (“Pimenta” and “Riacho Fundo” communities; Table 1). Each pitfall trap consisted of three lines 0.6 m high plastic canvas guide fence arranged in a radial “Y” shape, with each line 10 m long, buckets of 60 L buried at ground level at each extremity, and one at the interception point of each line (see Corn 1994, Cechin and Martins 2000). We kept the pitfall traps open for five consecutive days and covered them on the weekend from September to October 2020 in the “Pimenta” community and from October 2020 to January 2021 in the “Riacho Fundo”, totaling 32 and 59 sampling days, respectively, totaling 8,736 hours/trap performed. Additionally, we searched for reptiles in different microhabitats such as rock outcrops, fallen logs, trees, exposed soil, and leaf litter in natural areas of Pedro II from February to March 2021 and October to November 2021 during the morning and night. We considered as occasional encounters the recording of live or dead reptiles by one of the researchers during other activities than those linked to the previous sampling methods (Foster 2012). We also considered dead specimens collected by third parties (Fitzgerald 2012) within the Pedro II limits. In this case, only complete and uninjured specimens that could be correctly identified were included in the list.

We manually collected the amphibians and captured the reptiles using herpetological hooks (collection license #61838-4/21), then transported them to the lab, where they were euthanized through intraperitoneal injection of 2% lidocaine for reptiles and topical application for amphibians, following the method described by Jared et al. (2023). We fixed the collected individuals in formal 10% and conserved them in alcohol 70%. Voucher specimens were housed in the Coleção Biológica of the Instituto Federal de Educação, Ciência e Tecnologia do Piauí, Campus Pedro II (CBPII). Nomenclature follows Frost (2024) and Uetz et al. (2025). Data on the conservation status and distribution of species were extracted from the Biodiversity Extinction Risk Assessment System - SALVE (ICMBio 2024).

Statistical Analysis

We used a sample-size-based rarefaction and extrapolation sampling curve, with a 95% confidence interval from 500 replicates, to estimate the rate of growth of richness from increasing the number of samples (Chao et al. 2014). For this analysis, we considered each day of species records, regardless of sampling method, as continuous samples. We used the R package iNEXT 2.0.2 (Chao et al. 2014, Hsieh, Ma and Chao 2016) to produce the rarefaction curves of amphibians and reptiles. We used the non-parametric estimators Chao 1 and Jackknife 1 (Magurran and McGill 2011) with 10,000 replications (Colwell et al. 2012) to estimate the species richness of amphibians and reptiles of Pedro II using the R package Vegan (Oksanen et al. 2016).

We calculated the Local Contribution to Beta Diversity (LCBD) (sensuLegendre and De-Cáceres 2013) of amphibians and reptiles to compare the species composition dissimilarity from Pedro II with other localities in Piauí state whose herpetofauna are known (see Araújo et al. 2020a). We used each separated group because most localities have different herpetofaunal checklists. The LCBD values were obtained by the Jaccard dissimilarity index of the Baselga family for presence/absence data (Legendre 2014). Thereafter, we performed the Jaccard Similarity coefficient (Magurran and McGill 2011) followed by a cluster analysis using the Unweighted Pair Group Average Method (UPGAM). The LCBD and Jaccard index were calculated using R packages Adespatial (Dray et al. 2019), BiodiversityR (Kindt and Coe 2005), and Vegan (Oksanen et al. 2016). For this analysis, we used data about 144 species distributed in three matrixes containing 52 amphibians, 39 lizards, and 53 snakes (see Data Availability section; Suppl. 1-3). Other reptiles were not included because few species are known in Piauí state. Species composition and richness data were obtained from the following localities: municipalities of José de Freitas - JFrei (Rocha and Santos 2004), Castelo do Piauí - CasPi (Rodrigues and Prudente 2011), Caxingó - Caxin, Ribeiro Gonçalves - RiGon, and São Raimundo Nonato - SRNon (Roberto, Ribeiro and Loebmann 2013), Barras - Barra (Benício, Silva and Fonseca 2014; Benício et al. 2015), Picos - Picos (Benício, Silva and Fonseca 2015, Benício, Lima and Fonseca 2015), Batalha - Batal (Silva, Carvalho and Rodrigues 2015), and Floriano - Flori (Lima et al. 2019), Uruçuí-Una Ecological Station - UUnES (Dal Vechio et al. 2013), National Parks of Serra da Capivara - SCaNP (Cavalcanti et al. 2014), Serra das Confusões - SCoNP (Dal Vechio et al. 2016, Marques et al. 2023), and Sete Cidades - SCiNP (Rocha and Prudente 2010, Araújo et al. 2020a), Delta do Parnaíba Environmental Protection Area - DPEPA (Araújo et al. 2020b), and Middle Gurguéia River Basin - MeGur (Madella-Auricchio, Auricchio and Soares 2017).

We reviewed each species distribution, and we excluded species without specific identification (“gr.”, “aff.” and “sp.”) to avoid biases regarding nomenclature. The nomenclature of Leptophis ahaetulla (Linnaeus, 1758) and L. dibernadoiAlbuquerque, Santos, Borges-Nojosa & Ávila, 2022 followed the distribution provided by Albuquerque et al. (2022); Ameivula pyrrhogularis (Silva & Ávila-Pires, 2013) and A. ocellifera (Spix, 1825) followed Silva and Ávila-Pires (2013). Thamnodynastes sp. was recognized as Dryophilax phoenix (Franco, Trevine, Montingelli & Zaher, 2017) according to Franco et al. (2017), and Micrurus ibiboboca (Merrem, 1820) was recognized as M. bonitaNascimento, Graboski, Silva Jr & Prudente, 2024 according to Nascimento et al. (2024). On the other hand, we included Dermatonotus aff. muelleri and Leptodactylus aff. mystaceus in the analysis because their populations in Piauí state are considered unique lineages (Oliveira et al. 2018, Silva et al. 2020), but we excluded Elachistocleis spp. because of their taxonomic uncertainties (Novaes-e-Fagundes et al. 2023). We excluded data on anuran species composition from Serra da Capivara National Park to avoid bias due to its low species richness.

Results

1. Amphibian species composition

We found 27 species of amphibian anurans belong to five families: Bufonidae (N = 3 spp.), Hylidae (N = 9 spp.), Leptodactylidae (N = 12 spp.), Microhylidae (N = 2 spp.), and Odontophrynidae (N = 1 sp.) (Table 2, Figure 2). The rarefaction curve shows a slight tendency to stabilize (Figure 3a), and the richness estimators Chao1 (27.47 ± 1.85) and Jacknife1 (31.89 ± 2.29) suggest including at least four anuran species in Pedro II; thus, we sampled around 87% of the anuran species that likely occur in the region. All species are classified as least concern (LC) regarding the imminent risk of extinction (ICMBio 2024). Most of the species are widely distributed in the Brazilian biomes, and although some are considered typical of the Caatinga and Cerrado environments (Uchôa et al. 2022; Santana et al. 2024), they occur in adjacent biomes or transitional areas. No endemism occurs in Pedro II (Table 2).

Table 2
Amphibian species recorded in the municipality of Pedro II, Piauí State, Northeastern Brazil, with their respective authors; vouchers; conservation status (IUCN and ICMbio): LC (Least Concern), NE (Not Evaluated) and DD (Data Deficient); and distribution in Brazilian biomes: CA (Caatinga), CE (Cerrado), AT (Atlantic Forest), AM (Amazon Forest), and widely distribution (WD). Species without information (?).

Taxa		Voucher	Conservation status (ICMBio)	Biome
	BUFONIDAE
1	Rhinella diptycha (Cope, 1862)	CBPII 098	LC	WD
2	Rhinella granulosa (Spix, 1824)	CBPII 020	LC	AT, CA, CE
3	Rhinella mirandaribeiroi Gallardo, 1965	CBPII 009	LC	WD
	HYLIDAE
4	Boana raniceps Cope, 1862	CBPII 022	LC	WD
5	Corythomantis greeningi Boulenger, 1896	CBPII 074	LC	AT, CA, CE
6	Dendropsophus minutus Peters, 1872	CBPII 096	LC	WD
7	Dendropsophus nanus (Boulenger, 1889)	CBPII 030	LC	WD
8	Dendropsophus soaresi Carmaschi and Jim, 1983	CBPII 019	LC	AT, CA, CE
9	Pithecopus gonzagai Andrade, Haga, Ferreira, Recco-Pimentel, Toledo, and Bruschi, 2020	CBPII 028	NE	AT, CA, CE
10	Scinax x-signatus Spix, 1824	CBPII 016	LC	WD
11	Scinax aff. similis (Cochran, 1952)	CBPII 097	?	?
12	Trachycephalus typhonius Linnaeus, 1758	CBPII 218	LC	WD
	LEPTODACTYLIDAE
13	Adenomera hylaedactyla (Cope, 1868)	CBPII 163	LC	WD
14	Adenomera juikitam Carvalho and Giaretta, 2013	CBPII 033	LC	CE
15	Leptodactylus fuscus (Schneider, 1799)	CBPII 005	LC	WD
16	Leptodactylus macrosternum (Miranda-Ribeiro, 1926)	CBPII 027	LC	WD
17	Leptodactylus aff. mystaceus Spix, 1824	CBPII 162	?	?
18	Leptodactylus syphax Bokermann, 1969	CBPII 024	LC	WD
19	Leptodactylus troglodytes Lutz, 1926	CBPII 025	LC	WD
20	Leptodactylus vastus Lutz, 1930	CBPII 256	LC	WD
21	Physalaemus albifrons (Spix, 1824)	CBPII 017	LC	AT, CA, CE
22	Physalaemus cuvieri Fitzinger, 1826	CBPII 023	LC	WD
23	Pleurodema diplolister Peters, 1870	CBPII 011	LC	AT, CA, CE
24	Pseudopaludicola mystacalis Cope, 1887	CBPII 166	LC	WD
	MICROHYLIDAE
25	Dermatonotus aff. muelleri Boettger, 1885	CBPII 026	?	?
26	Elachistocleis aff. piauiensis Caramaschi and Jim, 1983	CBPII 014	?	?
	ODONTOPHRYNIDAE
27	Proceratophrys cristiceps (Müller, 1883)	CBPII 001	LC	AT, CA, CE

Figure 2
Anuran amphibians recorded in the municipality of Pedro II, Piauí State, Northeastern Brazil. A. Rhinella diptycha; B. R. mirandaribeiroi; C. Boana raniceps; D. Corythomantis greeningi; E. Dendropsophus minutus; F. D. nanus; G. D. soaresi; H. Scinax x-signatus; I. S. aff. similis; J. Adenomera hylaedactyla; K. A. juikitam; L. Leptodactylus fuscus; M. L. macrosternum; N. L. aff. mystaceus; O. L. troglodytes; P. L. vastus; Q. Physalaemus albifrons; R. P. cuvieri; S. Pleurodema diplolister; T. Pseudopaludicola mystacalis; U. Dermatonotus aff. muelleri; V. Elachistocleis aff. piauiensis; W. Proceratophrys cristiceps; X. Pithecopus gonzagai.

Figure 3
Sample-size-based rarefaction and extrapolation sampling curves, with their respective confidence intervals (95%), performed with incidence data of anurans (A) and squamate reptiles (B) collected in the municipality of Pedro II, Piauí State, Northeastern Brazil.

Regarding the amphibian species composition similarity, we observed a high species turnover in the localities sampled in Piauí state (75.6% of total beta diversity). The loss of species contributes just 24.4% of the total beta diversity (Figure 4a). Differences in species composition were mainly due to Flori (LCBD = 0.1436) and SRNon (LCBD = 0.1274), which have the lowest species richness, and by RiGon (LCBD = 0.1412) and UUnES (LCBD = 0.1297), which presented the highest numbers of exclusive or shared species (p.LCBD < 0.05). Just Leptodactylus fuscus (Schneider, 1799), Leptodactylus troglodytes Lutz, 1926, and Physalaemus cuvieri Fitzinger, 1826 were common to all sampled areas. Other 13 species were exclusive to one locality (see Data Availability section). Additionally, we observed that anuran composition of Pedro II was more similar to those areas located in Caatinga than Cerrado environments (Figure 4a). Amongst the study areas, we observed it as more similar to SCoNP (j = 0.8571), Picos (j = 7407), Caxin (j = 0.7241), SCiNP (j = 0.6774), and Barra (j = 0.6333).

Figure 4
Ternary diagram of beta diversity and similarity (Jaccard index and “UPGMA” clustering method) between different inventories carried out in Piauí, northeastern Brazil. A - anurans, B - lizards, C - snakes. Each point represents a pairwise comparison of the sites and the dotted lines represent the links between the mean values (black square).

2. Reptile species composition

We found 55 reptile species belong to three orders: Crocodylia with one only species belonging to the Alligatoridae family; Testudines with five species belonging to the families Chelidae (N = 3 spp.), Kinosternidae (N = 1 sp.), and Testudinidae (N = 1 sp.); and Squamata with 46 species recorded from 16 families (Table 3, Figures 5–6). Among the amphisbaenas, we found only three species belonging to the Amphisbaenidae family. Among lizards, we recorded 20 species from the following families: Gekkonidae (N = 3 spp.), Gymnophthalmidae (N = 4 spp.), Iguanidae (N = 1 sp.), Phyllodactylidae (N = 1 sp.), Polychrotidae (N = 1 sp.), Sphaerodactylidae (N = 2 spp.), Scincidae (N = 3 spp.), Teiidae (N = 3 spp.), and Tropiduridae (N = 2 spp.). Snakes correspond to 56.5% of the Squamata recorded with 26 species from the following families: Boidae (N = 2 spp.), Colubridae (N = 6 spp.), Dipsadidae (N = 14 spp.), Elapidae (N = 2 spp.), Leptotyphlopidae (N = 1 sp.), and Viperidae (N = 1 sp.). The rarefaction curve also has a slight tendency to stabilize (Figure 3b), and the richness estimators Chao1 (53.48 ± 4.93) and Jacknife1 (62.79 ± 3.64) suggest including of seven and 16 reptile species in Pedro II, respectively. We sampled around 79% of the reptile species that likely occur in the region. Except for Micrurus bonita, which do not have sufficient data to evaluate their conservational status, all species are classified as having the least concern regarding the imminent risk of extinction (ICMBio 2024). We found three snakes typical to the Caatinga biome: Boiruna sertaneja Zaher, 1996, Erythrolamprus viridis (Günther, 1862), and Epictia borapeliotes (Vanzolini, 1996), whereas other reptiles have a wide distribution in the Brazilian biomes or occur in more than two biomes and transitional areas (Table 3).

Tabela 3
Reptile species recorded in the municipality of Pedro II, Piauí State, Northeastern Brazil, with their respective authors; vouchers; conservation status (IUCN and ICMbio): LC (Least Concern), NE (Not Evaluated) and DD (Data Deficient); and distribution in Brazilian biomes: CA (Caatinga), CE (Cerrado), AT (Atlantic Forest), AM (Amazon Forest), and widely distribution (WD). Species viewed and not collected (*). Invasive species (-).

Taxa		Voucher	Conservation status (ICMBio)	Biome
	Order Crocodylia
	ALLIGATORIDAE
1	Caiman crocodilus (Linnaeus, 1758)	*	LC	AM, CE, CA
	Order Squamata
	Amphisbaenia
	AMPHISBAENIDAE
2	Amphisbaena alba Linnaeus, 1758	CBPII 230	LC	WD
3	Amphisbaena vermicularis Wagler, 1824	CBPII 214	LC	WD
4	Leposternon polystegum (Duméril, 1851)	CBPII 216	LC	WD
	“Lizard”
	GEKKONIDAE
5	Hemidactylus agrius Vanzolini, 1978	CBPII 136	LC	CA, CE, AT
6	Hemidactylus brasilianus (Amaral, 1935)	CBPII 182	LC	CA, CE, AT
7	Hemidactylus mabouia (Moreau de Jonnès, 1818)	CBPII 148	-	WD
	GYMNOPHTHALMIDAE
8	Colobosaura modesta (Reinhardt and Lütken, 1862)	CBPII 145	LC	WD
9	Colobosauroides cearensis Cunha, Lima-Verde and Lima, 1991	CBPII 146	LC	CA, CE
10	Micrablepharus maximiliani (Reinhardt and Lütken, 1862)	CBPII 129	LC	WD
11	Vanzosaura multiscutata (Amaral, 1933)	*	LC	CA, CE, AT
	IGUANIDAE
12	Iguana iguana (Linnaeus, 1758)	CBPII 170	LC	WD
	PHYLLODACTYLIDAE
13	Phyllopezus pollicaris (Spix, 1825)	CBPII 183	LC	WD
	POLYCHROTIDAE
14	Polychrus acutirostris Spix, 1825	CBPII 093	LC	WD
	SCINCIDAE
15	Brasiliscincus heathi (Schmidt and Inger, 1951)	CBPII 151	LC	CA, CE, AT
16	Copeoglossum arajara (Reboucas-Spieker, 1981)	CBPII 142	LC	CA, CE
17	Copeoglossum nigropunctatum (Spix, 1825)	CBPII 177	LC	WD
	SPHAERODACTYLIDAE
18	Coleodactylus meridionalis (Boulenger, 1888)	CBPII 186	LC	WD
19	Gonatodes humeralis (Guichenot, 1855)	CBPII 733	LC	WD
	TEIIDAE
20	Ameiva ameiva (Linnaeus, 1758)	CBPII 127	LC	WD
21	Ameivula pyrrhogularis (Basto da Silva and Ávila-Pires, 2013)	CBPII 134	LC	CA, CE
22	Salvator merianae Duméril and Bibron, 1839	CBPII 171	LC	WD
	TROPIDURIDAE
23	Tropidurus hispidus (Spix, 1825)	CBPII 131	LC	WD
24	Tropidurus semitaeniatus (Spix, 1825)	CBPII 179	LC	CA, CE, AT
	Serpentes
	BOIDAE
25	Boa constrictor Linnaeus, 1758	CBPII 213	LC	WD
26	Epicrates assisi Machado, 1945	CBPII 236	LC	CA, CE, AT
	COLUBRIDAE
27	Drymoluber brazili (Gomes, 1918)	CBPII 187	LC	CA, CE
28	Drymarchon corais (Boie, 1827)	CBPII 306	LC	WD
29	Leptophis dibernardoi Albuquerque, Santos, Borges-Nojosa and Ávila, 2022	CBPII 732	LC	CA, CE, AT
30	Oxybelis aeneus (Wagler, 1824)	CBPII 243	LC	WD
31	Spilotes pullatus (Linnaeus, 1758)	CBPII 249	LC	WD
32	Tantilla melanocephala (Linnaeus, 1758)	CBPII 311	LC	WD
	DIPSADIDAE
33	Apostolepis cearensis Gomes, 1915	CBPII 172	LC	CA, CE, AT
34	Adelphostigma occipitalis (Jan, 1863)	CBPII 189	LC	WD
35	Boiruna sertaneja Zaher, 1996	*	LC	CA, CE, AT
36	Dryophylax phoenix Franco, Trevine, Montingelli and Zaher, 2017	CBPII 330	LC	CA, CE, AT
37	Erythrolamprus poecilogyrus (Wied-Neuwied, 1824)	CBPII 225	LC	WD
38	Erythrolamprus viridis (Günther, 1862)	CBPII 209	LC	WD
39	Leptodeira tarairiu Costa, Graboski, Grazziotin, Zaher, Rodrigues and Prudente, 2022	CBPII 228	LC	AT, CA, CE
40	Oxyrhopus trigeminus Duméril, Bibron and Duméril, 1854	CBPII 196	LC	WD
41	Philodryas nattereri (Steindachner, 1870)	CBPII 215	LC	WD
42	Philodryas olfersii (Lichtenstein, 1823)	CBPII 246	LC	WD
43	Pseudoboa nigra (Duméril, Bibron and Duméril, 1854)	CBPII 190	LC	WD
44	Psomophis joberti (Sauvage, 1884)	CBPII 223	LC	WD
45	Rodriguesophis iglesiasi (Gomes, 1915)	CBPII 220	LC	CA, CE
46	Xenodon merremii (Wagler, 1824)	CBPII 211	LC	WD
	ELAPIDAE
47	Micrurus bonita (Merrem, 1820)	CBPII 188	DD	CA, CE
48	Micrurus lemniscatus (Linnaeus, 1758)	CBPII 242	LC	WD
	LEPTOTYPHLOPIDAE
49	Epictia borapeliotes (Vanzolini, 1996)	CBPII 125	LC	CA, CE, AT
	VIPERIDAE
50	Crotalus durissus Linnaeus, 1758	CBPII 212	LC	WD
	Order Testudines
	CHELIDAE
51	Mesoclemmys perplexa Bour and Zaher, 2005	CBPII 307	LC	CA, CE
52	Mesoclemmys tuberculata (Luederwaldt, 1926)	CBPII 217	LC	CA, CE, AT
53	Phrynops geoffroanus (Schweigger, 1812)	CBPII 226	LC	WD
	KINOSTERNIDAE
54	Kinosternon scorpioides (Linnaeus, 1766)	CBPII 319	LC	WD
	TESTUDINIDAE
55	Chelonoidis carbonarius (Spix, 1824)	*	LC	WD

Figure 5
Reptiles recorded in the municipality of Pedro II, Piauí State, Northeastern Brazil. A. Mesoclemmys perplexa; B. M. tuberculata; C. Kinosternon scorpioides; D. Amphisbaena alba; E. A. vermicularis; F. Leposternon polystegum; G. Hemidactylus agrius; H. H. brasilianus; I. Colobosauroides cearensis; J. Micrablepharus maximiliani; K. Vanzosaura multiscutata; L. Iguana iguana; M. Phyllopezus pollicaris; N. Polychrus acutirostris; O. Brasiliscincus heathi; P. Copeoglossum arajara; Q. Coleodactylus meridionalis; R. Gonatodes humeralis; S. Ameiva ameiva; T. Ameivula pyrrhogularis; U. Tropidurus hispidus; V. Tropidurus semitaeniatus; W. Boa constrictor; X. Epicrates assisi.

Figure 6
Reptiles recorded in the municipality of Pedro II, Piauí State, Northeastern Brazil. A. Drymoluber brazili; B. Drymarchon corais; C. Leptophis dibernardoi; D. Oxybelis aeneus; E. Spilotes pullatus; F. Apostolepis cearensis; G. Dryophylax phoenix; H. Erythrolamprus poecilogyrus; I. E. viridis; J. Leptodeira tarairiu; K. Oxyrhopus trigeminus; L. Philodryas nattereri; M. P. olfersii; N. Pseudoboa nigra; O. Psomophis joberti; P. Xenodon merremii; Q. Micrurus bonita; R. M. lemniscatus; S. Epictia borapeliotes; T. Crotalus durissus.

Regarding reptile species composition, we observed a high lizard species turnover (76.82% of the total beta diversity). The loss of species contributes 23.18% of the total beta diversity (Figure 4b). Differences in species composition were mainly due to UUnES (LCBD = 0.1387) and SCapNP (LCBD = 0.1371), which had the highest species richness (p.LCBD < 0.05). Just the lizards Iguana iguana (Linnaeus, 1758), Ameiva ameiva (Linnaeus, 1758), and Salvator merianae Duméril & Bibron, 1839 were common to all sampled areas, whereas nine species were exclusive to one unique sampled area (see Data Availability section). The lizard fauna of Pedro II was more similar to Batal (j = 0.7000; Figure 4b). Regarding snakes, differences in species composition were mainly due to SCapNP (LCBD = 0.1054) and Picos (LCBD = 0.1050), which had the highest species richness. The species turnover contributes 65.36% of the total beta diversity, whereas the loss of species was 34.64% (Figure 4c). Just Oxyrhopus trigeminus Duméril, Bibron & Duméril, 1854 was common to all sampled areas, whereas 14 species were exclusive to one unique sampled area (see Data Availability section). The snake fauna of Pedro II was more similar to SCiNP (j = 0.7; Figure 4c).

Discussion

Given the increasing impacts on natural landscapes caused by human activity and the imminent climate crisis, the need to obtain basic knowledge about Neotropical biodiversity is becoming increasingly urgent. In this context, the state of Piauí still has large unknown areas (Pantoja et al. 2022), including areas with high species richness and/or priority areas for conservation (MMA 2023). In general, we observed a high species richness in Pedro II, representing about 45% of the amphibian and reptile species recorded in Piauí state (Roberto, Ribeiro and Loebmann 2013, Guedes, Entiauspe-Neto and Costa 2023). Ecotone areas are usually diverse because species from different biomes coexist in these environments, increasing species richness (Kark and Rensburg 2006, Kark 2013, Marques et al. 2023). Therefore, Pedro II is an ecotone area of Caatinga and Cerrado influenced by different phytophysiognomies, which might explain the high species richness in this region. Additionally, Pedro II is located in the Complexo da Serra Grande bioregion (Lima and Guerra 2020), in which some areas are considered real herpetofaunal “hotspots” (Loebmann and Haddad 2010). This region was formed from the removal of different surface layers of the Ibiapaba plateau (Andrade and Lins 1977, Lima 1987) that previously constituted a single geological unit, facilitating the sharing of species between previously connected areas.

The total of species recorded is expressive (82 species) compared with other herpetological studies in Piauí state. Araújo et al. (2020b) recorded 96 herpetofaunal species in the Delta do Parnaíba Environmental Protection Area (DPEPA), which comprises a total area of 3138 km² between the states of Ceará, Piauí, and Maranhão. At the opposite extreme of Piauí state, Marques et al. (2023) reported 94 species of amphibians and reptiles in the region of the Serra das Confusões National Park (SCoNP), a protected area of around 5500 km². Although Pedro II comprises a total area of about 1550 km², we found the third region with the highest herpetofauna in Piauí state. Therefore, we reinforce the biological importance of ecotone areas to the maintenance and conservation of biodiversity, besides the ecosystem services provided by them.

Comparing the species composition separately (between groups: “amphibians”, “snakes”, and “lizards”) with other checklists, we observed Pedro II shared with Caxingó (Roberto, Ribeiro and Loebmann 2013) the fourth highest amphibian richness in Piauí state (25 species; see Statistical analysis topic). In this case, DPEPA (Araújo et al. 2020b; 30 species), SCiNP and ScoNP (Dal Vechio et al. 2016, Araújo et al. 2020a, Marques et al. 2023, 27 species), and RiGon (Roberto, Ribeiro and Loebmann 2013, 26 species) have the highest amphibian richness. Pedro II shared with DPEPA (Araújo et al., 2020b) the second-highest snake richness in Piauí state (27 species), behind only UUnEE (Dal Vechio et al. 2013, 34 species). Similarly, we found the second-highest lizard richness in Piauí state (20 species), being behind only ScoNP (Dal Vechio et al. 2016, Marques et al. 2023, 24 species). Other reptiles were not included in these comparisons separately because they represent incidental or random sporadic reports. However, Pedro II had five (83%) of the six continental Testudines and three (60%) of the five Amphisbaenidae reported in Piauí state (Guedes, Entiauspe-Neto and Costa 2023).

Pedro II stands out by the presence of unique species in the state, for instance, Kinosternon scorpioides (Linnaeus, 1766) (Andrade 2019), Epictia borapeliotes (Araújo et al. 2020c), and Micrurus lemniscatus (Linnaeus, 1758) (Araújo et al. 2022a) have wide geographical distribution, however, their occurrences were recently recorded and correspond to the only known records for Piauí state (Guedes, Entiauspe-Neto and Costa 2023). In addition, Drymoluber brazili (Gomes, 1918) is a snake poorly known in the Northeast region, having its distribution updated until the Pedro II limits, being the northmost report of this species (Araújo et al. 2022b). However, we believe the species richness in Pedro II is still underestimated because species with the potential to occur in the region have not yet been found. For instance, Eunectes murinus (Linnaeus, 1758) and Bothrops lutzi (Miranda-Ribeiro, 1915) were recorded around 50 km in the Sete Cidades National Park (Rocha and Prudente 2010) and around 100 km in the municipality of Castelo do Piauí (Rodrigues and Prudente 2011). Additionally, other herpetofaunal species recorded in the Ibiapaba plateau (see Loebmann and Haddad 2010) might occur in Pedro II due to its geographical proximity and similarity in phytophysiognomies in the highlands of the municipality.

Most species recorded in Pedro II have wide geographic distribution (45 species) or occur only in the Caatinga and Cerrado biomes (12 species). We observed the species composition similarity varied according to the studied group, for example, the amphibian fauna was more similar to the Serra das Confusões National Park (Marques et al. 2023), Sete Cidades National Park (Araújo et al. 2020a), and to the municipalities of Picos (Benício, Silva and Fonseca 2015), Barras (Benício, Silva and Fonseca 2014), and Caxingó (Roberto, Ribeiro and Loebmann 2013). The lizard community was more similar to the municipality of Batalha (Silva, Carvalho and Rodrigues 2015), whereas the snake fauna was more similar to the Sete Cidades National Park (Rocha and Prudente 2010) and the municipality of José de Freitas (Rocha and Santos 2004). In any case, the herpetofauna in Piauí state consists of typical species of Caatinga, Cerrado, and transitional zones, besides those widely distributed in Brazilian biomes. However, comparisons between these sampled areas are difficult because they were conducted in different sampling periods, having distinct sample sizes and methodologies. Furthermore, there is still a large sampling gap in the state, in which most studies were carried out within conservation units, especially those with restricted use (Pantoja et al. 2022).

Pedro II is located in a region of great natural and economic importance for the Piauí, sometimes generating a conflict between economic development and conservation in the municipality. The municipality is considered the only producer of Precious Opal gemstone in all of Brazil (Milanez and Oliveira 2009, Sousa et al. 2020) extracted from several mining areas in rural areas of the municipality (Santos et al. 2024). Some negative environmental impacts have been identified in these mining areas, such as the strong alteration of the natural landscape due to changes in topography, rainfall erosion processes, soil pollution, and displacement of local fauna (Santos et al. 2024). Additionally, the municipality also has extensive natural areas that have been impacted in recent years due to the increase in real estate speculation in natural preservation areas, such as near hillsides, springs, and/or river courses, and irregular tourism practices developed in an unsustainable manner (Lima and Gerra 2020).

In this context, the collection of data on local biodiversity, including herpetofauna (Santos et al. 2019), is the initial step towards developing adequate conservation and land use control measures. Although it belongs to the Complexo da Serra Grande bioregion and is included within a federal conservation unit, this is the first long-term study to characterize the amphibian and reptile fauna of the municipality of Pedro II, highlighting the importance of faunal surveys for regions that are still little explored. As discussed in recent years, the decline in reptile and amphibian populations in the world has been growing alarmingly, and many of these losses are due to anthropic actions and climate change (Gibbons et al. 2000, Luedtke et al. 2023). Therefore, measures to conserve areas that have great species richness should be taken especially in ecotonal areas, since these areas usually harbor a high species diversity (Smith et al. 2001, Walker et al. 2003, Yarrow and Marín 2007), including amphibians and reptiles (Matavelli et al. 2019, Madani et al. 2024).

Supplementary Material

The following online material is available for this article:

Supplementary material 1 – Table of presence/absence of amphibian species at all localities in the state of Piauí used in the analyses comparing species richness and composition.

Supplementary material 2 – Table of presence/absence of serpentes species at all localities in the state of Piauí used in the analyses comparing species richness and composition.

Supplementary material 3 – Table of presence/absence of lizard species at all localities in the state of Piauí used in the analyses comparing species richness and composition

Acknowledgments

We thank the Instituto Federal de Educação, Ciência e Tecnologia do Piauí-IFPI for providing a grant through the Programa de Apoio à Pesquisa, Estruturação e Reestruturação Laboratorial-PROAGRUPAR-INFRA (edital nº 29/2021) and to Instituto Chico Mendes de Conservação da Biodiversidade by collection license (#61838-4/21). MAP thanks the Instituto Federal de Educação, Ciência e Tecnologia do Piauí-IFPI for providing a scholarship through the Institutional Scientific Initiation Program (Edital nº 7843 - PIBIC IFPI 2020). CAC (Process: 88887.765685/2022-00), SCMA (Process: 88887.706181/2022-00), and NLAR (Process: 8888.7827305/2023-00) thanks the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for providing research fellowship. FPS (Process: 073.6819.2024.0005705-43), LRU (Process: BO0450/2022), and JLPF (Process: 073.6819.2024.0005706-24) thanks the Fundação de Amparo à Pesquisa do Estado da Bahia (FAPESB) for providing a research fellowship. KCA thanks Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) and Fundação de Amparo à Pesquisa do Estado do Piauí (FAPEPI) for providing research fellowship (Process: 150013/2023-0). We thank all those who contributed directly and indirectly to the completion of this work, including all former Biotecpi members who helped with data collection activities.

Data availability

Supporting data are available at https://doi.org/10.48331/scielodata.5VSHR0

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Edited by

• Associate Editor
  Pedro Nunes

Publication Dates

• Publication in this collection
  16 May 2025
• Date of issue
  2025

History

• Received
  26 Aug 2024
• Accepted
  16 Apr 2025