2urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7Eurn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727AZooKeysZK1313-29891313-2970Pensoft Publishers10.3897/zookeys.1002.5099750997Research ArticleSerpentesBiodiversity & ConservationBrazilSnakes of the Pernambuco Endemism Center, Brazil: diversity, natural history and conservationFrançaRafaela C.https://orcid.org/0000-0001-5034-679612MoraisMayara1FrançaFrederico G. R.https://orcid.org/0000-0001-6989-24553RödderDennis2SoléMircomsole@uesc.brhttps://orcid.org/0000-0001-7881-622721Programa de Pós-graduação em Ecologia e Conservação da Biodiversidade, Universidade Estadual de Santa Cruz, Rodovia Jorge Amado, Km 16, CEP 45662-900 Ilhéus, Bahia, BrazilHerpetology Section, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, GermanyPrograma de Pós-graduação em Zoologia, Universidade Estadual de Santa Cruz, 11 Rodovia Jorge Amado, Km 16, CEP 45662-900 Ilhéus, Bahia, BrazilDepartamento de Engenharia e Meio Ambiente, Centro de Ciências Aplicadas e Educação, Universidade Federal da Paraíba – UFPB, Av. Santa Elizabete, s/n – Centro. CEP 58297-000, Rio Tinto, PB, BrazilDepartamento de Ciências Biológicas, Universidade Estadual de Santa Cruz, Rodovia Jorge Amado, km,16, 45662-900 Ilhéus, Bahia, Brazil
Corresponding author: Mirco Solé (msole@uesc.br)
Academic editor: R. Jadin
20201012202010021151587883AC7C-75BA-538F-BC82-DB985ABB93F156565B89-F2AA-4AD4-8012-0CBC274C136543469291002202014042020Rafaela C. França, Mayara Morais, Frederico G. R. França, Dennis Rödder, Mirco SoléThis is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.http://zoobank.org/56565B89-F2AA-4AD4-8012-0CBC274C1365
The Atlantic Forest is one of the largest and richest tropical rainforests on the planet, being one of the 25 world priorities for conservation. The Atlantic Forest portion located north of the São Francisco River corresponds to the Pernambuco Endemism Center (PEC). We describe the snake composition of the PEC, providing information about the diversity, natural history and geographical distribution of the species, based on records from five scientific collections and additional information from the literature. A total of 78 species of snakes distributed in eight families was registered in the Pernambuco Endemism Center. The Caatinga is the Brazilian biome that most shares species with the PEC, followed by Cerrado. On the other hand, seven species are considered endemic of this region. Most of the snake species in the PEC have been registered in forest (94.8%), followed by “Brejos Nordestinos” (46.1%), Tabuleiros (43.5%), Restingas (14.1%) and Mangroves (5.1%). The PEC snake fauna includes mainly terrestrial species (60.2%) and cryptozoic and/or fossorial species (21.7%), but also presents a high richness of semi-arboreal and arboreal species (29.5%). Vertebrates are the main food item consumed by the species (78% of species), among the main prey are mammals, lizards, and amphibians. Most species show a strictly nocturnal activity period (50%), followed by strictly diurnal (38%). The PEC is the most degraded and least known region of the Atlantic Forest, yet it has revealed a high richness of snake species, including seven endemic species. It is emphasized that regional conservation efforts need to be intensified, because few forests in the region are formally protected, and the majority consist of small and poorly protected fragments, which means that many species in the region may be in risk of extinction.
biodiversityinventorygeographic distributionnatural historySerpentesrichnessFundação de Amparo à Pesquisa do Estado da Bahia501100006181http://doi.org/10.13039/501100006181Coordenação de Aperfeiçoamento de Pessoal de Nível Superior501100002322http://doi.org/10.13039/501100002322Alexander von Humboldt-Stiftung100005156http://doi.org/10.13039/100005156Conselho Nacional de Desenvolvimento Científico e Tecnológico501100003593http://doi.org/10.13039/501100003593Citation
França RC, Morais M, França FGR, Rödder D, Solé M (2020) Snakes of the Pernambuco Endemism Center, Brazil: diversity, natural history and conservation. ZooKeys 1002: 115–158. https://doi.org/10.3897/zookeys.1002.50997
Introduction
The Atlantic Forest is considered one of the 25 priority areas for conservation worldwide (Myers et al. 2000). This biome was one of the largest tropical forests in the Americas, originally covering 150 million hectares along the Brazilian coast and parts of Paraguay and Argentina (Silva and Casteleti 2003). Today, the Atlantic Forest has been reduced to less than 12% of its original coverage (Ribeiro et al. 2009). Even having suffered an extensive fragmentation since long time ago, the Atlantic Forest still presents a great biodiversity, housing one of the highest percentages of endemic species in the world (Morellato and Haddad 2000).
Although practically the entire Brazilian coast was occupied by European colonization, it was in the northeast that the Atlantic Forest was more rapidly degraded, due to the economic cycle of brazilwood and sugar cane (Coimbra-Filho and Câmara 1996). This degradation is even more evident in the portion of the Atlantic Forest located north of the São Francisco River, where an important center of endemism is located in South America – The Pernambuco Endemism Center (hereafter PEC) (Prance 1982, Silva and Casteleti 2003). In this region, sugar cane is the main agricultural crop and other anthropic actions, such as animal and plant extractivism, have contributed to the reduction of biodiversity in the PEC (Coimbra-Filho and Câmara 1996, Tabarelli et al. 2002, 2006a). In the midst of this scenario, the PEC is considered the most devastated, least known and least protected sector of the Atlantic Forest, being one of the regions on the planet where conservation efforts are most urgent (Coimbra-Filho and Câmara 1996, Tabarelli et al. 2002, 2005).
Among reptiles, snakes are the group that currently presents the most underestimated risks of extinction, due to the scarcity of information on the natural history of most species, mainly because they have long periods of inactivity, are difficult to observe and live in low population densities (Seigel 1993). Although some studies carried out on Atlantic forest remnants of the PEC have provided important information about snakes in this region (e.g. Moura et al. 2011, Pereira Filho and Montingelli 2011, França et al. 2012, Roberto et al. 2012, 2015, Rodrigues et al. 2015, Pereira Filho et al. 2017, Mesquita et al. 2018, Sampaio et al. 2018, Freitas et al. 2019a), the knowledge about the diversity, distribution and natural history of PEC snake species remains scarce and fragmented. In this direction, scientific collections perform a fundamental role in obtaining information that is the basis for the description of new species, biodiversity inventories and identification of endemism areas (Rocha et al. 2014).
Herein, we describe the snake composition at the Pernambuco Endemism Center, providing information about the diversity, natural history and geographical distribution of the species, based on records from scientific collections and additional information from the literature.
Materials and methodsStudy area
The study area comprises the Atlantic Forest located north of the São Francisco River, which corresponds to the Pernambuco Endemism Center (PEC) (Fig. 1) (Prance 1982, Silva and Casteleti 2003), located between the states of Alagoas and Rio Grande do Norte. This region has a humid tropical climate (Köppen’s As’), with autumn-winter rains and rainfall ranging from 750 to 1500 mm per year (Tabarelli et al. 2006a).
Map of the location of the Pernambuco Endemism Center, with the original coverage of Atlantic Forest (gray), and the actual remnants (green).
https://binary.pensoft.net/fig/485060
The PEC region is composed by different native forest formations and ecosystems associated with the Atlantic Forest domain. A mosaic of ombrophilous and semi-deciduous forests is present in this region (Tabarelli et al. 2006a). Also, PEC comprises the “Brejos de Altitude” or “Brejos Nordestinos”, which are “islands” of humid forests established in the semi-arid region, surrounded by Caatinga vegetation (Andrade-Lima 1982). Although the vegetation of the PEC is composed mainly of humid tropical forests, we can also find open physiognomies along the coast, which are called “Restingas”, and in the interior, which are called “Tabuleiros”. The restingas are formed by strips of beaches and dunes covered by herbaceous and shrubby vegetation (Araujo 1992). The Tabuleiros are considered natural enclaves of savannah, characterized by herbaceous vegetation, with scattered trees and shrubs or grouped in patches that are structurally similar to the coastal restingas, but without the marine influence (Andrade-Lima 1982). On the coast along the PEC, we can also find areas of mangroves, with a diversified aggregation of trees and shrubs that form the dominant plant communities in saline solution of the tides (Tabarelli et al. 2006b).
According to Uchoa Neto and Tabarelli (2002), the PEC presents the largest amount of remaining area of Atlantic Forest in the state of Pernambuco (1,363.23 km²), followed by the states of Alagoas (807.95 km²), Rio Grande do Norte (567.67 km²) and Paraíba (566.09 km²).
Data collection
The data presented here is the result of verification of 3,118 snake specimens deposited in five scientific collections (Coleção Herpetológica da Universidade Federal da Paraíba – UFPB; Coleção do Laboratório de Anfíbios e Répteis da Universidade Federal do Rio Grande do Norte - CLAR; Coleção Herpetológica do Museu de História Natural da Universidade Federal de Alagoas – MUFAL; Coleção Herpetológica da Universidade Federal Rural de Pernambuco – CHUFRPE; Coleção Herpetológica da Universidade Federal de Pernambuco – CHUFPE) and literature data.
The information on the distribution and occurrence of species in each environment were obtained through the records of the scientific collections and literature data, and was subsequently georeferenced. We include records of occurrence of species in the literature only when we were able to confirm the record by direct observation, photo or through museum records or documented vouchers. Information on diet, habitat use, and litter size of the species was obtained from personal data, records of scientific collections and literature data. We categorized the snake size considering the mean body size of each species based on published data as small (< 500mm), moderate (501–1000mm) and large (> 1001mm).
In this work, we have differentiated the habitats of the species into five vegetation physiognomies found in this region: Forests (when the species were found in areas with a typical forest physiognomy, with a large vegetation cover, reaching 35 meters high in the canopy, presenting epiphytes, lianas and bromeliads); Coastal Restingas; Mangroves; Tabuleiros; Brejos Nordestinos (remnants of humid forests scattered in the Caatinga) (Fig. 2); and urban areas. In addition, we compared the snake fauna found in the PEC with these of five other natural ecoregions in Brazil (Amazon, Caatinga, Cerrado, Pampas, and Pantanal). These regions are divided on the basis of geomorphology, climate, and vegetation (IBGE 2004).
Vegetation physiognomies found in the Pernambuco Endemism Center. A forest B forest interior C Coastal Restingas D mangroves, E tabuleiros F Brejos Nordestinos. Photograph credits: Ivan L. Sampaio, in the Barra de Gramame (A), Frederico França, in the APA da Barra do Rio Mamanguape (B, C), Marcelo Melo, in the APA da Barra do Rio Mamanguape (D), Frederico França, in the Reserva Biológica Guaribas (E) and Adonias Teixeira, in the Parque Estadual Mata do Pau-Ferro (F).
The species Caaeteboia sp. found in the PEC, differs from Caaeteboiaamarali (at present the only representative of the genus) mainly because it presents 15 rows of dorsal scales without reduction, while C.amarali presents 17 rows of dorsal scales without reduction. In addition, there is a strong variation between the number of ventral and subcaudal scales between the two species (Pereira Filho et al. 2017).
We decided to use the name Micrurusibiboboca according to Silva Jr (2016). Although Silva Jr (2016) affirms that M.ibiboboca may be a species complex throughout the distribution of the species, the author still maintains the proper name. Thus, the species designated here as M.ibiboboca is the same mentioned in previous works as Micrurusaff.ibiboboca (e.g. França et al. 2012, Rodrigues et al. 2015, França and França 2019).
Results
We registered a total of 78 species of snakes of eight families, distributed in the PEC (Table 1, Figs 3–7). The most species rich family was Dipsadidae (47 species, 60% of total), followed by Colubridae (12 species, 15.4%), Viperidae (6 species, 7.7%), Boidae and Typhlopidae, both with four species (5.1%), Elapidae (3 species, 3.8 %) and Anomalepididae and Leptotyphlopidae, both with a single species (1.3%).
Snake species from the Pernambuco Endemism Center. ABoaconstrictorBCorallushortulanusCEpicratesassisiDEpicratescenchriaEChironiuscarinatusFChironiusexoletusGChironiusflavolineatus, HDendrophidionatlanticaIDrymarchoncoraisJDrymoluberdichrousKLeptophisahaetullaLPalusophisbifossatusMOxybelisaeneusNSpilotespullatusOSpilotessulphureus. Photograph credits: Frederico França (A, B, E, F, G, J, L, M, N, O), Vanessa Nascimento (D), Davi Pantoja (C, H, I), Rafaela França (K).
https://binary.pensoft.net/fig/485062
Many species of snakes that are found in PEC are also found in other Brazilian biomes. The Caatinga (58 species, 74.3% found in PEC) is the Brazilian biome that shares most species with the PEC, followed by Cerrado (44 species, 56.4%), Amazon Forest (35 species, 44.9%), Pantanal (35 species, 44.9%) and Pampas (13 species, 16.6%). On the other hand, some species (Atractuscaete, A.maculatus, Bothropsmuriciensis, Caaeteboia sp., Dendrophidionatlantica, Echinantheracephalomaculata and Micruruspotyguara) are found only in the PEC and are considered endemic of this region.
Snake species from the Pernambuco Endemism Center. ATantillamelanocephalaBApostolepiscearensisCApostolepislongicaudataDAtractusmaculatusEAtractuspotschiFBoirunasertanejaGCaaeteboia sp. HDipsasmikaniiIDipsasneuwiediJDipsassazimaiKDipsasvariegataLErythrolamprusaesculapiiMErythrolamprusalmadensisNErythrolampruspoecilogyrusOErythrolamprusreginae. Photograph credits: Frederico França (A, B, G, H, I), Anderson A. Santos (C, N), Rafaela França (D, E, J, K, L, M, O), Paulo R. S. Freitas (F).
https://binary.pensoft.net/fig/485063
Most of the snake species in the PEC have been registered in Forest areas (74 species, 94.8%), followed by Brejos Nordestinos (36 species, 46.1%), Tabuleiros (34 species, 43.5%), Restingas (11 species, 14.1%) and Mangroves (4 species, 5.1%). Six species were found in four different habitats and 31 species were found only in one habitat type (Table 1). Of these, 26 species were collected only in forested areas, three species only in the Brejos Nordestinos and one species was found only in restingas (Table 1).
Summary of the Information of Natural History of the Snakes in the Pernambuco Endemism Center. Abbreviations are: Habitats (BN = Brejos Nordestinos, F = forest, Tb = Tabuleiro, Rt = Restinga, Mg = Mangrove); Diet (abn = amphisbaenians, amp = amphibians, ann = annelids, art = arthropods, bi = birds, fi = fishes, mo = mollusks, li = lizards, mam = mammals, sn=snakes; Activity period (D = Diurnal, N = Nocturnal); Habits (AB = arboreal, SAB = semi-arboreal, AQ = aquatic, SAQ = semi-aquatic, CR = cryptozoic, FS = Fossorial, TE = terrestrial). * Endemic species of the Pernambuco Endemism Center (PEC).
Family/Species
Habitats
Diet
Habits
Diel activity
Anomalepididae
Liotyphlopstrefauti
F
art
FS
N
Boidae
Boaconstrictor
BN, F, Tb, Rt
mam, li, bi
SAB, TE
D, N
Corallushortulanus
F
mam, bi, li, amp
AB
N
Epicratesassisi
BN, F, Tb
mam, li, bi
TE
N
Epicratescenchria
F
mam, bi, li, amp
TE, SAB
N
Colubridae
Chironiuscarinatus
F
amp, bi, li, mam
TE, AB
D
Chironiusexoletus
BN, F, Tb
amp, li
AR, TE
D
Chironiusflavolineatus
BN, F, Tb
amp
SAB
D
Dendrophidionatlantica*
F
–
TE
D
Drymarchoncorais
F, Tb
amp, abnli, sn, bi, mam
TE
D
Drymoluberdichrous
BN, F, Tb
li, amp
TE
D
Leptophisahaetulla
BN, F
amp, li
AB,TE
D
Oxybelisaeneus
BN, F, Tb
li, amp, fi
AB
D
Palusophisbifossatus
F, BN
amp, mam, li
TE
D
Spilotespullatus
BN, F, Tb
mam, bi
SAB
D
Spilotessulphureus
F
mam, bi
SAB
D
Tantillamelanocephala
BN, F, Tb, Rt
art
FS
D, N
Dipsadidae
Apostolepiscearensis
F, Tb
sn, abn
FS
D
Apostolepislongicaudata
F
sn
FS
D
Atractuscaete*
F
ann
FS
N
Atractusmaculatus
F
ann
FS
N
Atractuspotschi
F
ann
FS
N
Boirunasertaneja
Tb, F
sn, li, mam
TE
N
Caaeteboia sp.*
F
–
TE
D
Dipsasmikanii
BN, F, Tb
mo
TE
N
Dipsasneuwiedi
F, BN
mo
TE
N
Dipsassazimai
F
mo
AB, TE
N
Dipsasvariegata
F
mo
AB, TE
N
Echinantheracephalomaculata*
F
amp
TE
D
Echinantheracephalostriata
F
amp
TE
D
Erythrolamprusaesculapii
F
sn, li
TE
D
Erythrolamprusalmadensis
F
amp
SAQ
D
Erythrolamprusmiliaris
F, BN
amp, fi
SAQ
D, N
Erythrolampruspoecilogyrus
BN, F, Tb, Mg
amp, li
TE
D, N
Erythrolamprusreginae
F
amp, li, fi
SAQ
D
Erythrolamprustaeniogaster
F, Tb, Rt
amp, fi
SAQ
D
Erythrolamprusviridis
BN, F
amp, li
TE
D
Helicopsangulatus
F, Mg, Rt
fi, amp
AQ
N
Helicopsleopardinus
Rt, F
fi, amp
AQ
N
Hydrodynastesgigas
F, Rt
amp, fi, sn, mam
AQ, TE
D
Imantodescenchoa
F, Tb
li, amp
AB
N
Leptodeiraannulata
F, Rt, BN
amp, li
AB, TE
N
Lygophisdilepis
BN, F
amp
TE
D
Oxyrhopusguibei
BN, F, Tb
mam, li
TE
D, N
Oxyrhopuspetolarius
BN, F, Tb
li, mam, bi, amp
TE
N
Oxyrhopustrigeminus
BN, F, Tb, Rt,
li, mam, bi
TE
D, N
Philodryasnattereri
BN, F, Tb
li, mam, amp, sn, bi
TE, SAB
D
Philodryasolfersii
BN, F, Tb, Mg
amp, li, bi, mam
TE, SAB
D
Philodryaspatagoniensis
F, Tb, Rt
amp, li, mam, bi, sn
TE
D
Phimophisguerini
F, Tb
li, mam
TE
N
Pseudoboanigra
BN, F, Tb
li, mam, sn
TE
N
Psomophisjoberti
F
amp, li
TE
D
Sibonnebulatus
F, Tb
mo
AB
N
Siphlophiscompressus
F, Tb
li, sn
AB, TE
N
Taeniophallusaffinis
BN, F, Tb
li, amp, abn, mam
CR
N
Taeniophallusoccipitalis
BN, F, Tb
li, amp, abn
CR
N
Thamnodynastesalmae
BN
amp, li
AB, TE
N
Thamnodynasteshypoconia
BN
amp, li
TE, AB
N
Thamnodynastespallidus
F, Tb
amp
TE, AB
N
Thamnodynastesphoenix
BN
amp
TE, AB
N
Xenodonmerremii
BN, F, Tb
amp
TE
D
Xenodonrabdocephalus
F
amp
TE
D
Xenopholisscalaris
F
amp
TE
N
Xenopholisundulatus
BN, F
amp
TE
N
Elapidae
Micruruscorallinus
F
abn, li, sn, amp
CR
D
Micrurusibiboboca
BN, F, Tb
abn, sn, li
CR
D, N
Micruruspotyguara*
F, Tb
sn
CR
D, N
Leptotyphlopidae
Epictiaborapeliotes
F, BN, Rt
art
FS
D, N
Typhlopidae
Amerotyphlopsamoipira
Rt
art
FS
N
Amerotyphlopsarenensis
BN, F
art
FS
N
Amerotyphlopsbrongersmianus
F, Tb
art
FS
N
Amerotyphlopspaucisquamus
F, Tb
art
FS
N
Viperidae
Bothropsbilineatus
F
mam, amp, bi, sn, li
AB
N
Bothropserythromelas
F
li, mam
TE
N
Bothropsleucurus
F, BN, Tb, Mg
amp, li, sn, bi, mam
TE
N
Bothropsmuriciensis*
F
amp, mam
TE
N
Crotalusdurissus
BN, F, Rt
mam
TE
N
Lachesismuta
F
mam
TE
N
The majority of snake species found in the PEC use the soil as substrate, of which 47 species (60.2%) are terrestrial and 17 (21.7%) are cryptozoic and/or fossorial. In addition, 23 species are arboreal or semi-arboreal (29.5%) and 16 (20.5%) are aquatic or semi-aquatic. The diet of PEC snakes consists mainly of vertebrates (61 species, 78.2%), of which 23 species are considered generalists, feeding on three or more types of prey, 21 species feed on two types of prey, 23 species are specialists in amphibians, two species are specialists in snakes and two species are specialists in mammals. Only 14 species feed on invertebrates, of which six species feed on arthropods, three species feed on annelids and five species feed on mollusks (Table 1). As for the period of activity, 39 (50 %) species are nocturnal, 30 (38.4%) species are diurnal and nine (11.5%) species are diurnal and nocturnal (Table 1).
Snake species from the Pernambuco Endemism Center. AErythrolamprustaeniogasterBErythrolamprusviridisCHelicopsangulatusDHelicopsleopardinusEHydrodynastesgigasFImantodescenchoaGLeptodeiraannulataHLygophisdilepisIOxyrhopusguibeiJOxyrhopuspetolariusKOxyrhopustrigeminusLPhilodryasnattereriMPhilodryasolfersiiNPhilodryaspatagoniensisOPhimophisguerini. Photograph credits: Frederico França (A, C, F, H, I, K, M, N, O), Vanessa Nascimento (B, D), Ivan L. Sampaio (E), Willianilson Pessoa (G), Rafaela França (J, L).
https://binary.pensoft.net/fig/485064
We present below a commented list of species of snakes that occur in PEC, with notes on natural history and distribution. The “N” corresponds to the number of individuals analyzed in the scientific collections. The species L.trefauti, A.caete, A.potschi, E.cephalomaculata, E.cephalostriata, T.almae, T.hypoconia, and T.phoenix were recorded only by literature data.
Snake species from the Pernambuco Endemism Center. APseudoboanigraBPsomophisjoberti, CSibonnebulatusDSiphlophiscompressusETaeniophallusaffinisFTaeniophallusoccipitalisGThamnodynastesalmaeHThamnodynasteshypoconiaIThamnodynastespallidusJThamnodynastesphoenixKXenodonmerremiiLXenodonrabdocephalusMXenopholisscalarisNXenopholisundulatusOMicruruscorallinus. Photograph credits: Frederico França (A, B, C, D, F, H, K, N), Vanessa Nascimento (L), Samuel Cardoso (G), Davi Pantoja (M), Rafaela França (I), Anderson A. Santos (E), Paulo R. S. Freitas (J), Adrian Garda (O).
Snake species from the Pernambuco Endemism Center. AMicrurusibibobocaBMicruruspotyguaraCEpictiaborapeliotesDAmerotyphlopsarenensisEAmerotyphlopsbrongersmianusFAmerotyphlopspaucisquamusGBothropsbilineatusHBothropserythromelasIBothropsleucurusJBothropsmuriciensisKCrotalusdurissusLLachesismuta. Photograph credits: Frederico França (A, B, E, F, H, I, K, L), Ivan L. Sampaio (C), Gentil A. Pereira Filho (D), Willianilson Pessoa (J), Rafaela França (G).
Liotyphlopstrefauti Freire, Caramaschi, Suzart & Argolo, 2007 - A small-sized fossorial species (total length = 366–389 mm; N = 3), with nocturnal activity (Freire et al. 2007). It has a restricted distribution, occurring in the Atlantic Forest and Caatinga (Abegg et al. 2017b). In the PEC it occurs in the states of Alagoas and Pernambuco (Fig. 8A), being found in Forest areas (Freire et al. 2007, Abegg et al. 2017b). Liotyphlopstrefauti, as observed in other congeneric species, feeds on eggs and arthropod larvae (Marques et al. 2019).
Geographic distribution records for snakes of the Pernambuco Endemism Center (PEC). ALiotyphlopstrefautiBBoaconstrictor and CorallushortulanusCEpicratescenchria and E.assisiDChironiuscarinatus and C.exoletusEChironiusflavolineatus and DendrophidionatlanticaFDrymarchoncorais and DrymoluberdichrousGLeptophisahaetulla and PalusophisbifossatusHOxybelisaeneus and TantillamelanocephalaISpilotessulphureus and S.pullatus.
Boaconstrictor Linnaeus, 1758 - A large semiarboreal species (average SVL = 1023 mm; N = 42), with nocturnal activity (Marques et al. 2001). It has a wide distribution, occurring in the Atlantic Forest, Amazon Forest, Caatinga, Cerrado and Pantanal (Cunha and Nascimento 1993, Marques et al. 2005, 2015, 2019, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 8B), being found in Forest, Brejos Nordestinos, Tabuleiros and Restinga Areas (Pereira Filho and Montingelli 2011, Rodrigues et al. 2015, Pereira Filho et al. 2017, Sampaio et al. 2018). This species can also occur in urban areas (França and França 2019). Boaconstrictor feeds on mammals, birds and lizards (Pizzatto et al. 2010). Its litter can range from 18 to 60 hatchlings (Vitt and Vangilder 1983, Pizzatto and Marques 2007, Fraga et al. 2013).
Corallushortulanus (Linnaeus, 1758) - A moderate-sized arboreal snake (SVL = 745 mm; N = 11), with nocturnal activity (Marques et al. 2019). It has a wide distribution, occurring in the Atlantic Forest, Amazon Forest, Caatinga, Cerrado and Pantanal (Marques et al. 2005, 2015, 2019, Fraga et al. 2013, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 8B), being found in Forest. Corallushortulanus feeds on mammals, birds, lizards and amphibians (Pizzatto et al. 2010). Its litter can range from 3 to 24 hatchlings (Pizzatto and Marques 2007, Fraga et al. 2013).
Epicratesassisi Machado, 1945 – A moderate-sized terrestrial species (average SVL = 691 mm; N = 135), with nocturnal activity (Marques et al. 2019). This species occurs in the Cerrado, Caatinga and Atlantic Forest (Guedes et al. 2014, Marques et al. 2015, 2019). In the PEC it occurs in all states (Fig. 8C), being found in Forest, Brejos Nordestinos, Tabuleiros, Restingas and urban areas (França et al. 2012, Rodrigues et al. 2015, Pereira Filho et al. 2017, Sampaio et al. 2018). Epicratesassisi feeds on mammals, birds, and lizards. Its litter can range from 7 to 14 hatchlings (Pizzatto and Marques 2007).
Epicratescenchria (Linnaeus, 1758) – A large semi-arboreal or terrestrial species (average SVL = 1105 mm; N = 6), with nocturnal activity (Marques et al. 2019). It has a wide distribution, occurring in the Atlantic Forest, Amazon Forest, Cerrado and Pantanal (Marques et al. 2005, 2015, 2019, Passos and Fernandes 2008). In the PEC it occurs in the states of Alagoas and Pernambuco (Fig. 8C), being found in Forest areas, but also in urban areas. Epicratescenchria feeds on mammals, birds, lizards and amphibians (Martins and Oliveira 1998, Pizzatto et al. 2010). Its litter can range from 8 to 25 hatchlings (Pizzatto and Marques 2007).
Family Colubridae Oppel, 1811
Chironiuscarinatus (Linnaeus, 1758) – A large terrestrial and arboreal species (average SVL = 1001 mm; N = 15), with diurnal activity (Marques et al. 2019). It has a disjunct distribution, occurring in the Amazon Forest and Atlantic Forest (Araújo et al. 2019). In the PEC it occurs in the states of Alagoas, Pernambuco and Paraíba (Fig. 8D), being found in Forest and urban areas when these are close to forests (Araújo et al. 2019). Chironiuscarinatus feeds on amphibians, birds, lizards and mammals (Dixon et al. 1993, Silva et al. 2010, Rodrigues et al. 2016). Its litter can have 5 to 12 eggs (Dixon et al. 1993, Goldberg 2007).
Chironiusexoletus (Linnaeus, 1758) – A moderate-sized arboreal and terrestrial species (average SVL = 614 mm; N = 16), with diurnal activity (Marques et al. 2019). It has a wide distribution, occurring in the Atlantic Forest, Caatinga, Cerrado, Pantanal and Amazon Forest (Cunha and Nascimento 1993, Marques et al. 2005, 2015, 2019, Guedes et al. 2014). In the PEC it occurs in the states of Alagoas, Pernambuco and Paraíba (Fig. 8D), being found in Forest, Brejos Nordestinos and Tabuleiro (Pereira Filho and Montingelli 2011, Rodrigues et al. 2015). Chironiusexoletus feeds mainly on amphibians, but occasionally on lizards (Marques and Sazima 2004, Rodrigues et al. 2016). Its litter can range from 4 to 12 eggs (Dixon et al. 1993, Goldberg 2007).
Chironiusflavolineatus (Linnaeus, 1758) – A moderate-sized semi-arboreal species (average SVL = 592 mm; N = 60), with diurnal activity (Marques et al. 2019). It presents a wide distribution, occurring in the Atlantic Forest, Cerrado, Caatinga, Pantanal and Amazon Forest (Cunha and Nascimento 1993, Dixon et al. 1993, Marques et al. 2005, 2015, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 8E), being found in Forest, Brejos Nordestinos, Tabuleiros, and urban areas (França et al. 2012, Rodrigues et al. 2015, Sampaio et al. 2018). Chironiusflavolineatus feeds on amphibians (Pinto et al. 2008, Rodrigues et al. 2016). Its litter can range from 3 to 8 eggs (Dixon et al. 1993, Hamdan and Fernandes 2015).
Dendrophidionatlantica Freire, Caramaschi & Gonçalves, 2010 – A small-sized terrestrial species (average SVL = 366 mm; N = 24), with diurnal activity (Marques et al. 2019). Dendrophidionatlantica is endemic to the PEC and occurs in the states of Alagoas, Pernambuco and Paraíba (Fig. 8E), being found in Forest (Freire et al. 2010, Pereira Filho et al. 2017, Barbosa et al. 2019). Dendrophidionatlantica feeds on amphibians (Marques et al. 2019). Its litter can have 3 eggs (Lima et al. 2019).
Drymarchoncorais (Boie, 1827) – A large terrestrial species (average SVL = 1288 mm; N = 7), with diurnal activity (Marques et al. 2019). It presents a wide distribution, being registered in the Amazon Forest, Cerrado, Caatinga and Pantanal (Cunha and Nascimento 1993, Strussmann and Sazima 1993, Guedes et al. 2014, Marques et al. 2015, 2019). In the PEC it occurs in all states (Fig. 8F), being found in Forest, Tabuleiros and urban areas (Rodrigues et al. 2015, Mesquita et al. 2018). Drymarchoncorais feeds on amphibians, amphisbaenians, lizards, snakes, birds and mammals (Prudente et al. 2014). Its litter can range from 3 to 15 eggs (Prudente et al. 2014).
Drymoluberdichrous (Peters, 1863) – A small-sized terrestrial species (average SVL = 348 mm; N = 15), with diurnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Amazon Forest, and Caatinga (Cunha and Nascimento 1993, Guedes et al. 2014, Marques et al. 2019). In the PEC it occurs in the states of Alagoas and Paraíba (Fig. 8F), being found in Forest, Brejos Nordestinos, Tabuleiros and urban areas (Rodrigues et al. 2015, Pereira Filho et al. 2017, Mesquita et al. 2018, França and França 2019). Drymoluberdichrous feeds on lizards and amphibians (Martins and Oliveira 1998, Borges-Nojosa and Lima 2001). Its litter can range from 2 to 6 eggs (Martins and Oliveira 1998, Fraga et al. 2013).
Leptophisahaetulla (Linnaeus, 1758) – An arboreal and terrestrial, moderate-sized species (average SVL = 582 mm; N = 42), with diurnal activity (Marques et al. 2019). This species occurs in Atlantic Forest, Amazon Forest, Caatinga, Cerrado, Pantanal, and Pampas (Strussmann and Sazima 1993, Bérnils et al. 2007, Guedes et al. 2014, Marques et al. 2015, 2019). In the PEC it can be found in all states (Fig. 8G) in Forest, Brejos Nordestinos and urban areas (Pereira Filho and Montingelli 2011, França and França 2019). Leptophisahaetulla feeds on amphibians and lizards (Albuquerque et al. 2007). Its litter can range from 3 to 12 eggs (Vitt and Vangilder 1983, Mesquita et al. 2009).
Oxybelisaeneus (Wagler, 1824) – An arboreal, moderate-sized species (average SVL = 780 mm; N =46), with diurnal activity (Marques et al. 2019). It presents a wide distribution, being found in the Atlantic Forest, Amazon Forest, Caatinga, Cerrado, and Pantanal (Cunha and Nascimento 1993, Marques et al. 2005, 2015, 2019, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 8H), being found in Forest, Brejos Nordestinos, Tabuleiros, and urban areas (Pereira Filho and Montingelli 2011, Rodrigues et al. 2015, França and França 2019). Oxybelisaeneus feeds on lizards, amphibians, and occasionally fishes (Henderson 1982, Hetherington 2006, Grant and Lewis 2010, Mesquita et al. 2013, Franzini et al. 2018). Its litter can range from 4 to 9 eggs (Vitt and Vangilder 1983, Mesquita et al. 2009, Fraga et al. 2013).
Palusophisbifossatus (Raddi, 1820) – A moderate-sized terrestrial species (average SVL = 801 mm; N = 5), with diurnal activity (Marques et al. 2019). It presents a wide distribution, occurring in the Atlantic Forest, Amazon Forest, Caatinga, Cerrado, Pampas, and Pantanal (Cunha and Nascimento 1993, Strussmann and Sazima 1993, Lema 2003, Bérnils et al. 2007, Guedes et al. 2014, Marques et al. 2019). In the PEC it occurs in all states (Fig. 8G), being found in Forest and Brejos Nordestinos (Pereira Filho and Montingelli 2011, Pereira Filho et al. 2017). Palusophisbifossatus feeds on amphibians, mammals, and lizards (Leite et al. 2007). Its litter can range from 4 to 24 eggs (Costa et al. 2010).
Spilotespullatus (Linnaeus, 1758) – A large, semi-arboreal species (average SVL = 1442 mm; N = 21), with diurnal activity (Marques et al. 2019). It presents a wide distribution, being found in the Atlantic Forest, Amazon Forest, Caatinga, Cerrado, and Pantanal (Cunha and Nascimento 1993, Marques et al. 2005, 2015, 2019, Guedes et al. 2014). In the PEC, it occurs in the states of Alagoas, Pernambuco and Paraíba (Fig. 8I), being found in Forest, Brejos Nordestinos, Tabuleiros, and urban areas (Pereira Filho et al. 2017, Mesquita et al. 2018, França and França 2019). Spilotespullatus feeds on mammals and birds (Silva et al. 2010, Marques et al. 2014). Its litter can range from 2 to 5 eggs (Hauzman et al. 2005, Fraga et al. 2013).
Spilotessulphureus (Wagler, 1824) – A moderate-sized semi-arboreal species (average SVL = 911 mm; N =20), with diurnal activity (Marques et al. 2019). It presents a wide distribution, being found in the Atlantic Forest, Amazon Forest, Caatinga and Cerrado (Cunha and Nascimento 1993, Guedes et al. 2014, Marques et al. 2015, 2019). In the PEC, it occurs in the states of Alagoas and Paraíba (Fig. 8I), being found in Forest and urban areas (Morais et al. 2018). Spilotessulphureus feeds on mammals and birds (Beebe 1946, Cunha and Nascimento 1993, Rufino and Bernardi 1999). Its litter can range from 7 to 15 eggs (Good 1989, Fraga et al. 2013, Morais et al. 2018).
Tantillamelanocephala (Linnaeus, 1758) – A small-sized fossorial species (average SVL = 233 mm; N = 172), with diurnal and nocturnal activity (Marques et al. 2019). It presents a wide distribution, occurring in the Atlantic Forest, Amazon Forest, Caatinga, Cerrado, Pampas, and Pantanal (Cunha and Nascimento 1993, Marques et al. 2005, 2015, 2019, Bérnils et al. 2007, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 8H), being found in Forest, Brejos Nordestinos, Tabuleiros, and restingas (Pereira Filho and Montingelli 2011, Mesquita et al. 2018, Sampaio et al. 2018). Tantillamelanocephala feeds on arthropods. Its litter can range from 1 to 3 eggs (Mesquita et al. 2009, Fraga et al. 2013)
Dipsadidae Bonaparte, 1838
Apostolepiscearensis Gomes, 1915 – A small-sized fossorial species (average SVL = 329 mm; N = 44), with diurnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Caatinga and Cerrado (Guedes et al. 2014, Marques et al. 2015, Mesquita et al. 2018). In the PEC it occurs in the states of Rio Grande do Norte, Paraíba and Pernambuco (Fig. 9A), being found in Forest, Tabuleiros, and urban areas (Mesquita et al. 2018, França and França 2019). Apostolepiscearensis feeds on small elongated reptiles (Mesquita et al. 2009, Amorim et al. 2015, Marques et al. 2019).
Geographic distribution records for snakes of the Pernambuco Endemism Center (PEC). AApostolepislongicaudata and A.cearensisBAtractuscaete, A.maculatus and A.potschiCBoirunasertaneja and Caaeteboia sp. DDipsasmikanii and D.neuwiediED.sazimai and D.variegataFEchinantheracephalomaculata and E.cephalostriataGErythrolamprusalmadensis, E.taeniogaster, E.miliaris, and E.aesculapiiHE.poecilogyrus, E.viridis and E.reginae. IHelicopsangulatus and H.leopardinus.
https://binary.pensoft.net/fig/485068
Apostolepislongicaudata Gomes, 1921 – A small-sized fossorial species (average SVL = 235 mm; N = 8), with diurnal activity (Marques et al. 2019). This species occurs in the Cerrado, Caatinga and Floresta Atlântica (Curcio et al. 2011, França et al. 2012). In the PEC it occurs only in a conservation unit (Reserva Biológica Guaribas) located in the state of Paraíba (Fig. 9A), being found in Forest. Apostolepislongicaudata feeds on small elongated reptiles (Marques et al. 2019). We found two eggs in a female.
Atractuscaete Passos, Fernandes, Bérnils & Moura-Leite, 2010 – A small-sized fossorial and cryptozoic species (average SVL = 376 mm, N = 1), with nocturnal activity (Passos et al. 2010, Marques et al. 2019). This species is endemic to the PEC and occurs only in the state of Alagoas (Fig. 9B), being found in Forest areas. Atractuscaete feeds mostly on earthworms (Passos et al. 2010).
Atractusmaculatus (Günther, 1858) – A small-sized fossorial and cryptozoic species (average SVL = 326 mm; N = 5), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest and Caatinga (Passos et al. 2010, Abegg et al. 2017a). In the PEC it occurs in the states of Alagoas and Pernambuco (Fig. 9B), being found in Forest and urban areas, when close to forests. Atractusmaculatus feeds mostly on earthworms (Passos et al. 2010).
Atractuspotschi Fernandes, 1995 – A small-sized fossorial and cryptozoic species (average SVL = 312 mm, N = 1), with nocturnal activity (Passos et al. 2010, Marques et al. 2019). This species occurs in the Atlantic Forest and Caatinga (Guedes et al. 2014). In the PEC it occurs in the state of Alagoas (Fig. 9B), being found in Forest (Passos et al. 2010). Atractuspotschi feeds mostly on earthworms (Passos et al. 2010).
Boirunasertaneja Zaher, 1996 – A large terrestrial species (average SVL = 1358 mm; N = 2), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest and Caatinga (Guedes et al. 2014, Pereira Filho et al. 2017). In the PEC it can be found in the states of Pernambuco and Alagoas (Fig. 9C), in Tabuleiros and Forest (Rodrigues et al. 2015, Pereira Filho et al. 2017). Boirunasertaneja eats snakes, lizards and mammals (Vitt and Vangilder 1983, Gaiarsa et al. 2013). Its litter can range from 4 to 14 eggs (Vitt and Vangilder 1983, Gaiarsa et al. 2013).
Caaeteboia sp. – A small to moderate-sized terrestrial species (average SVL = 411 mm; N = 2), with diurnal activity (personal observation). This species is endemic to the PEC and occurs only in the states of Pernambuco and Paraíba (Fig. 9C), being found in Forest.
Dipsasmikanii Schlegel, 1837 – A small-sized terrestrial species (average SVL = 302 mm; N = 72), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Cerrado, Caatinga and Pantanal (Marques et al. 2005, 2015, 2019, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 9D), being found in Forest, Brejos Nordestinos, Tabuleiros and urban areas (França et al. 2012, Pereira Filho et al. 2017, Sampaio et al. 2018). Dipsasmikanii feeds on mollusks (Laporta-Ferreira et al. 1986). Its litter can range from 3 to 10 eggs (Pizzatto et al. 2008).
Dipsasneuwiedi (Ihering, 1911) – A small-sized terrestrial species (average SVL = 369 mm; N = 17), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest and Caatinga (Guedes et al. 2014, Marques et al. 2019). In the PEC it occurs in the states of Alagoas, Pernambuco and Paraíba (Fig. 9D), being found in Forest, Brejos Nordestinos and urban areas (Pereira Filho et al. 2017). Dipsasneuwiedi feeds on mollusks (Laporta-Ferreira et al. 1986). Its litter can range from 4 to 12 eggs (Pizzatto et al. 2008).
Dipsassazimai Fernandes, Marques & Argôlo, 2010 – A small-sized arboreal and terrestrial species (average SVL = 299 mm; N = 1), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest and Caatinga (Fernandes et al. 2010, Guedes et al. 2014). In the PEC it occurs in the states of Alagoas and Pernambuco (Fig. 9E), being found in Forest. Dipsassazimai feeds on mollusks (Fernandes et al. 2010).
Dipsasvariegata (Duméril, Bibron & Duméril, 1854) – A small to moderate size arboreal and terrestrial species (average SVL = 464 mm; N = 4), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest and Amazon Forest (Cunha and Nascimento 1993, Marques et al. 2019). In the PEC it occurs only in the state of Alagoas (Fig. 9E), being found in Forest. Dipsasvariegata feeds on mollusks (Marques et al. 2019).
Echinantheracephalomaculata Di Bernardo, 1994 – A small to moderate size terrestrial species (average SVL = 297 mm, N = 2), with diurnal activity (Di-Bernardo 1994, Marques et al. 2019). This species is endemic to the PEC and occurs only in the states of Alagoas and Pernambuco (Fig. 9F), being found in Forest (Roberto et al. 2015, Freitas et al. 2019b). Echinantheracephalomaculata feeds on amphibians (Marques et al. 2019).
Echinantheracephalostriata Di Bernardo, 1996 – A moderate-sized terrestrial species, with diurnal activity (Di-Bernardo 1996, Marques et al. 2019). This species only occurs in the Atlantic Forest (Marques et al. 2019). In the PEC it occurs in the state of Alagoas (Fig. 9F), being found only in the Reserva Biológica de Pedra Talhada (Roberto et al. 2015). In the report of this species for the PECRoberto et al. (2015) provide a photo and a voucher (URCA-H 4103). Echinantheracephalostriata feeds on amphibians (Marques et al. 2009).
Erythrolamprusaesculapii (Linnaeus, 1758) – A moderate-sized terrestrial species (average SVL = 562 mm; N = 7), with diurnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Amazon forest, Caatinga, Cerrado and Pantanal (Cunha and Nascimento 1993, Marques et al. 2005, 2015, 2017a, 2019). In the PEC it occurs in the states of Alagoas and Pernambuco (Fig. 9G), being found in Forest and urban areas. Erythrolamprusaesculapii feeds on snakes and lizards (Marques and Puorto 1992). Its litter can range from 1 to 8 eggs (Marques 1996a).
Erythrolamprusalmadensis (Wagler, 1824) – A small-sized semi-aquatic species (average SVL = 298 mm; N = 4), with diurnal activity (Marques et al. 2019). This species has a wide distribution, occurring in the Atlantic Forest, Amazon forest, Caatinga, Cerrado, Pantanal and Pampas (Dixon 1989, França et al. 2006, Bérnils et al. 2007, Guedes et al. 2014). In the PEC it occurs in the states of Paraíba and Rio Grande do Norte (Fig. 9G), being found in Forest (Pereira Filho et al. 2017, França and França 2019). Erythrolamprusalmadensis feeds on amphibians (Bernarde and Abe 2010, Rodrigues et al. 2016). Its litter can have five eggs.
Erythrolamprusmiliaris (Linnaeus, 1758) – A small-sized semi-aquatic species (average SVL = 382 mm; N = 7), with diurnal and nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Amazon forest, Caatinga and Cerrado (Cunha and Nascimento 1993, Nogueira et al. 2010, Marques et al. 2017a, 2019). In the PEC it occurs in the states of Alagoas, Pernambuco and Paraíba (Fig. 9G), being found in Forest and Brejos Nordestinos. Erythrolamprusmiliaris feeds on amphibians and fish (Marques et al. 2019). Its litter can range from 1 to 30 eggs (Pizzatto and Marques 2006).
Erythrolampruspoecilogyrus (Wied-Neuwied, 1825) – A small-sized terrestrial species (average SVL = 313 mm; N = 35), with diurnal and nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Caatinga, Cerrado, Pantanal and Pampas (Marques et al. 2005, 2015, 2019, Bérnils et al. 2007, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 9H), being found in Forest, Brejos Nordestinos, Mangroves, Tabuleiros and urban areas (França et al. 2012, Pereira Filho et al. 2017, Mesquita et al. 2018). Erythrolampruspoecilogyrus feeds on amphibians and lizards (Prieto et al. 2012). Its litter can range from 3 to 17 eggs (Vitt and Vangilder 1983, Mesquita et al. 2009). In Figure 4N we show a juvenile that is in the process of changing its coloration to the adult stage. This species has a different color pattern in the region (Pereira Filho et al. 2017) if compared to other populations located more southwards.
Erythrolamprusreginae (Linnaeus, 1758) – A small-sized semi-aquatic species (average SVL = 355 mm; N = 4), with diurnal activity (Marques et al. 2019). This species occurs in the Atlantic and Amazon forests, Caatinga, Cerrado and Pantanal (Cunha and Nascimento 1993, Marques et al. 2005, 2015, 2019, Guedes et al. 2014). In the PEC it occurs in the states of Alagoas and Pernambuco (Fig. 9H), being found in Forest. Erythrolamprusreginae feeds on amphibians, lizards, and fish (Martins and Oliveira 1998, Albarelli and Santos-Costa 2010, Silva et al. 2010, Rodrigues et al. 2016). Its litter can range from 1 to 4 eggs (Arzamendia 2016, Marques et al. 2016)
Erythrolamprustaeniogaster (Jan, 1863) – A small-sized semi-aquatic species (average SVL = 364 mm; N = 45), with diurnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Amazon forest, Caatinga, Cerrado and Pantanal (Cunha and Nascimento 1993, Marques et al. 2005, 2015, 2019, Guedes et al. 2014). In the PEC it occurs in the states of Alagoas, Pernambuco and Paraíba (Fig. 9G), being found in Forest, Tabuleiros, Restingas and urban areas (Rodrigues et al. 2015, Pereira Filho et al. 2017, Mesquita et al. 2018, Sampaio et al. 2018). Erythrolamprustaeniogaster feeds on amphibians and fish (Cunha and Nascimento 1993, Rodrigues et al. 2016). Its litter can range from 7 to 10 eggs (Cunha and Nascimento 1993).
Erythrolamprusviridis (Günther, 1862)– A small-sized terrestrial species (average SVL = 243 mm; N = 21), with diurnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest and Caatinga (Guedes et al. 2014, Marques et al. 2019). In the PEC it occurs in all states (Fig. 9H), being found in Forest, Brejos Nordestinos and urban areas (Pereira Filho and Montingelli 2011, Pereira Filho et al. 2017). Erythrolamprusviridis feeds on amphibians and lizards (Vitt and Vangilder 1983, Mesquita et al. 2009). Its litter can range from 2 to 7 eggs (Vitt and Vangilder 1983, Mesquita et al. 2009).
Helicopsangulatus (Linnaeus, 1758) – A small to moderate sized aquatic species (average SVL = 413 mm; N = 236), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic and Amazon forests, Caatinga, Cerrado and Pantanal (Cunha and Nascimento 1993, Marques et al. 2005, 2015, 2019, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 9I), being found in Forest, Mangroves, Restingas and urban areas (França et al. 2012, Pereira Filho et al. 2017, Sampaio et al. 2018). Helicopsangulatus feeds on fish and amphibians. Its litter can range from 1 to 21 eggs (Braz et al. 2016).
Helicopsleopardinus (Schlegel, 1837) – A small-sized aquatic species (average SVL = 324 mm; N = 9), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Amazon Forest, Caatinga, Cerrado, Pantanal and Pampas (Strussmann and Sazima 1993, Marques et al. 2005, 2015, 2019, Bérnils et al. 2007, Guedes et al. 2014, Rodrigues et al. 2016). In the PEC it occurs in the states of Alagoas and Pernambuco (Fig. 9I), being found in Forest, Restingas and urban areas. Helicopsleopardinus feeds on fish and amphibians (Ávila et al. 2006). Its litter can range from 3 to 31 eggs (Scartozzoni and Almeida-Santos 2006, Braz et al. 2016).
Hydrodynastesgigas (Duméril, Bibron & Duméril, 1854) – A large aquatic and terrestrial species (average SVL = 1296 mm; N =10), with diurnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Amazon Forest, Cerrado, Pantanal and Pampas (Lema 2003, Marques et al. 2005, 2015, 2019, Rodrigues et al. 2016). In the PEC it occurs in the states of Paraíba and Rio Grande do Norte (Fig. 10A), being found in Forest and Restingas (Pereira Filho et al. 2017, Sampaio et al. 2018). Hydrodynastesgigas feeds on fish, amphibians, mammals and snakes (López and Giraudo 2004). Its litter can range from 14 to 42 eggs (Vogel 1958, Fraga et al. 2013).
Geographic distribution records for snakes of the Pernambuco Endemism Center (PEC). AHydrodynastesgigas and LygophisdilepisBImantodescenchoa and LeptodeiraannulataCOxyrhopusguibei and O.petolariusDO.trigeminusEPhilodryasnattereri, P.olfersii and P.patagoniensisFPhimophisguerini and PseudoboanigraGPsomophisjoberti, Sibonnebulatus and SiphlophiscompressusHTaeniophallusaffinis and T.occipitalisIThamnodynastesalmae, T.hypoconia, T.pallidus and T.phoenix.
https://binary.pensoft.net/fig/485069
Imantodescenchoa (Linnaeus, 1758) – An arboreal, moderate-sized species (average SVL = 633 mm; N = 23), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Amazon Forest, Caatinga, Cerrado and Pantanal (Cunha and Nascimento 1993, Marques et al. 2005, 2015, 2019, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 10B), being found in Forest and Tabuleiros (Rodrigues et al. 2015, Mesquita et al. 2018). Imantodescenchoa feeds on amphibians and lizards (Martins and Oliveira 1998, Sousa et al. 2014). Its litter can range from 1 to 7 eggs (Martins and Oliveira 1998, Pizzatto et al. 2008, Fraga et al. 2013, Sousa et al. 2014).
Leptodeiraannulata (Linnaeus, 1758) – A moderate-sized arboreal and terrestrial species (average SVL = 576 mm; N = 6), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Amazon Forest, Caatinga, Cerrado and Pantanal (Ávila and Morais 2007, Guedes et al. 2014, Marques et al. 2015, 2019). In the PEC it occurs in the states of Alagoas and Pernambuco (Fig. 10B), being found in Forest, Brejos Nordestinos, and Restingas (Pereira Filho and Montingelli 2011, Roberto et al. 2015). Leptodeiraannulata feeds on amphibians and lizards (Moura 1999, Mesquita et al. 2013, Santos-Silva et al. 2014). Its litter can range from 3 to 13 eggs (Petzold 1969, Pizzatto et al. 2008).
Lygophisdilepis Cope, 1862 – A small-sized terrestrial species (average SVL = 356 mm; N = 9), with diurnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Caatinga and Cerrado (Guedes et al. 2014, Marques et al. 2015, Mesquita et al. 2018). In the PEC it occurs in the states of Pernambuco, Paraíba and Rio Grande do Norte (Fig. 10A), being found in Forest, Brejos Nordestinos, and urban areas (Pereira Filho and Montingelli 2011, França et al. 2012, Mesquita et al. 2018). Lygophisdilepis feeds on amphibians (Mesquita et al. 2009). Its litter can range from 4 to 6 eggs (Mesquita et al. 2009).
Oxyrhopusguibei Hoge & Romano, 1977 – A small sized terrestrial species (average SVL = 442 mm; N = 10), with diurnal and nocturnal activity (Marques et al. 2017). This species occurs in the Atlantic Forest, Caatinga, Cerrado and Pantanal (Marques et al. 2005, 2015, 2019, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 10C), being found in Forest, Brejos Nordestinos, and Tabuleiros (Pereira Filho and Montingelli 2011, Mesquita et al. 2018). Oxyrhopusguibei feeds on mammals and lizards (Andrade and Silvano 1996, Barbo et al. 2011). Its litter can range from 3 to 20 eggs (Pizzatto and Marques 2002).
Oxyrhopuspetolarius (Linnaeus, 1758) – A small size terrestrial species (average SVL = 423 mm; N = 36), with nocturnal activity (Marques et al. 2017). This species occurs in the Atlantic Forest, Amazon Forest, Caatinga, Cerrado and Pantanal (Marques et al. 2005, 2015, 2019, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 10C), being found in Forest, Brejos Nordestinos, Tabuleiros, and urban areas (Pereira Filho et al. 2017, Mesquita et al. 2018, Sampaio et al. 2018, França and França 2019). Oxyrhopuspetolarius feeds on lizards, mammals, birds and amphibians (Alencar et al. 2013). Its litter can range from 2 to 12 eggs (Lynch 2009, Gaiarsa et al. 2013).
Oxyrhopustrigeminus Duméril, Bibron & Duméril, 1854 – A small-sized terrestrial species (average SVL = 360 mm; N = 237), with nocturnal activity (Marques et al. 2017). This species occurs in the Atlantic Forest, Caatinga, Cerrado and Pantanal (Marques et al. 2005, 2015, 2019, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 10D), being found in Forest, Brejos Nordestinos, Restingas, Tabuleiros, and urban areas (Pereira Filho and Montingelli 2011, Sampaio et al. 2018, França and França 2019). Oxyrhopustrigeminus feeds on lizards, mammals, and birds (Vitt and Vangilder 1983, Mesquita et al. 2009, Alencar et al. 2012). Its litter can range from 6 to 9 eggs (Vitt and Vangilder 1983, Mesquita et al. 2009).
Philodryasnattereri Steindachner, 1870 – A moderate-sized terrestrial or semi-arboreal species (average SVL = 712 mm; N = 76), with diurnal activity (Marques et al. 2017). This species occurs in the Atlantic Forest, Caatinga, Cerrado, Pantanal (Marques et al. 2005, 2015, Guedes et al. 2014, Mesquita et al. 2018). In the PEC it occurs in all states (Fig. 10E), being found in Forest, Brejos Nordestinos, Tabuleiros, and urban areas (França et al. 2012, Pereira Filho et al. 2017, Sampaio et al. 2018). Philodryasnattereri feeds on lizards, mammals, amphibians, snakes, and birds (Mesquita et al. 2011b). Its litter can range from 4 to 13 eggs (Vitt and Vangilder 1983, Mesquita et al. 2009).
Philodryasolfersii (Lichtenstein, 1823) – A moderate-sized terrestrial or semi-arboreal species (average SVL = 562 mm; N = 123), with diurnal activity (Marques et al. 2017). This species occurs in the Atlantic Forest, Caatinga, Cerrado, Pantanal and Pampas (Marques et al. 2005, 2015, 2019, Bérnils et al. 2007, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 10E), being found in Forest, Brejos Nordestinos, Tabuleiros, Mangroves and urban areas (Pereira Filho and Montingelli 2011, França et al. 2012, Pereira Filho et al. 2017, Sampaio et al. 2018). Philodryasolfersii feeds on amphibians, lizards, birds and mammals (Hartmann and Marques 2005). Its litter can range from 1 to 16 eggs (Vitt and Vangilder 1983, Fowler et al. 1998, Mesquita et al. 2009).
Philodryaspatagoniensis (Girard, 1858) – A small to moderate sized terrestrial species (average (average SVL = 436 mm; N = 68), with diurnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Caatinga, Cerrado, Pantanal and Pampas (Marques et al. 2005, 2015, 2019, Bérnils et al. 2007, Guedes et al. 2014). In the PEC it occurs in the states of Pernambuco, Paraíba, and Rio Grande do Norte (Fig. 10E), being found in Forest, Tabuleiros, Restingas, and urban areas (França et al. 2012, Pereira Filho et al. 2017, Sampaio et al. 2018). Philodryaspatagoniensis feeds on amphibians, lizards, mammals, birds, and snakes (Hartmann and Marques 2005). Its litter can range from 3 to 19 eggs (Fowler et al. 1998).
Phimophisguerini (Duméril, Bibron & Duméril, 1854) – A small to moderate sized terrestrial species (average SVL = 497 mm; N = 15), with nocturnal activity (Marques et al. 2017). This species occurs in the Atlantic Forest, Caatinga, Cerrado, Pampas and Pantanal (Lema 2003, Marques et al. 2005, 2015, Guedes et al. 2014, Mesquita et al. 2018). In the PEC it occurs in the states of Alagoas and Paraíba (Fig. 10F), being found in Forest and Tabuleiros (Rodrigues et al. 2015, Pereira Filho et al. 2017). Phimophisguerini feeds on lizards and mammals (Alencar et al. 2013). Its litter can range from 3 to 7 eggs (Gaiarsa et al. 2013).
Pseudoboanigra (Duméril, Bibron & Duméril, 1854) – A moderate-sized terrestrial species (average SVL = 543 mm; N = 64), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Caatinga, Cerrado and Pantanal (Marques et al. 2005, 2015, 2019, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 10F), being found in Forest, Brejos Nordestinos, Tabuleiros, and urban areas (Pereira Filho and Montingelli 2011, França et al. 2012, Pereira Filho et al. 2017, Mesquita et al. 2018). Pseudoboanigra feeds on lizards, mammals, and snakes (Alencar et al. 2012). Its litter can range from 3 to 24 eggs (Orofino et al. 2010, Gaiarsa et al. 2013).
Psomophisjoberti (Sauvage, 1884) – A small-sized terrestrial species (average SVL = 285 mm; N = 11), with diurnal activity (Marques et al. 2017). This species occurs in the Atlantic Forest, Amazon Forest, Caatinga and Cerrado (Guedes et al. 2014, Marques et al. 2015, Rodrigues et al. 2016, Mesquita et al. 2018). In the PEC it occurs only in the state of Paraíba (Fig. 10G), being found in Forest and urban areas (França et al. 2012, Pereira Filho et al. 2017). Psomophisjoberti feeds on amphibians and lizards (Strussmann and Sazima 1993, Rodrigues et al. 2016). Its litter can have 7 eggs (Mesquita et al. 2009, 2011a).
Sibonnebulatus (Linnaeus, 1758) – A small-sized arboreal species (average SVL = 377 mm; N = 21), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Amazon Forest and can also be found on relict moist forests in Caatinga (Cunha and Nascimento 1993, Guedes et al. 2014, Marques et al. 2019). In the PEC it occurs in all states (Fig. 10G), being found in Forest, Tabuleiros, and urban areas (França et al. 2012, Rodrigues et al. 2015). Sibonnebulatus feeds on mollusks (Duellman 2005). Its litter can have 5 eggs (Boos 2001).
Siphlophiscompressus (Daudin, 1803) – A moderate-sized arboreal and terrestrial species (average SVL = 527 mm; N = 13), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest and Amazon Forest (Cunha and Nascimento 1993, Marques et al. 2019). In the PEC it occurs in the states of Alagoas, Pernambuco, and Paraíba (Fig. 10G), being found in Forest and Tabuleiros (Roberto et al. 2015, Rodrigues et al. 2015, Pereira Filho et al. 2017). Siphlophiscompressus feeds mainly on lizards, but may also feed on snakes (Martins and Oliveira 1998, Alencar et al. 2013). Its litter can range from 3 to 12 eggs (Martins and Oliveira 1998, Fraga et al. 2013, Gaiarsa et al. 2013).
Taeniophallusaffinis (Günther, 1858) – A small-sized cryptozoic species (average SVL = 172 mm; N = 9), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest and Caatinga (Guedes et al. 2014, Marques et al. 2019). In the PEC it occurs in the states of Alagoas, Pernambuco, and Paraíba (Fig. 10H), being found in Forest, Brejos Nordestinos, and Tabuleiros (Rodrigues et al. 2015, Pereira Filho et al. 2017). Taeniophallusaffinis feeds on lizards, amphibians, amphisbaenians, and mammals (Sousa and Cruz 2000, Barbo and Marques 2003, Zacariotti and Gomes 2010, Gomes 2012). Its litter can range from 5 to 7 eggs (Amaral 1978).
Taeniophallusoccipitalis (Jan, 1863) – A small-sized cryptozoic species (average SVL = 272 mm; N = 63), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Caatinga, Cerrado and Pampas (Bérnils et al. 2007, Guedes et al. 2014, Marques et al. 2015, 2019). In the PEC it occurs in all states (Fig. 10H), being found in Forest, Brejos Nordestinos, Tabuleiros, and urban areas (Pereira Filho and Montingelli 2011, Rodrigues et al. 2015, Pereira Filho et al. 2017, França and França 2019). Taeniophallusoccipitalis feeds on lizards, amphibians, and snakes (Balestrin and Di-Bernardo 2005, Gomes 2012). Its litter can have two eggs.
Thamnodynastesalmae Franco & Ferreira, 2003 – A moderate-sized arboreal and terrestrial, with nocturnal activity (Franco and Ferreira 2002, Marques et al. 2019). This species occurs in the Atlantic Forest and Caatinga (Guedes et al. 2014, Marques et al. 2019). In the PEC it occurs only in Brejos Nordestinos in the state of Pernambuco (Fig. 10I) (Freitas et al. 2019a). Thamnodynastesalmae feeds on amphibians and lizards (Marques et al. 2017a).
Thamnodynasteshypoconia (Cope, 1860) – A moderate-sized arboreal and terrestrial, with nocturnal activity (Marques et al. 2017a). This species occurs in the Atlantic Forest, Caatinga, Cerrado and Pampas (Bérnils et al. 2007, Guedes et al. 2014, Marques et al. 2015, 2019). In PEC it occurs only in the Parque Estadual Mata do Pau-Ferro, state of Paraíba, a Brejo Nordestino (Fig. 10I) (Pereira Filho et al. 2017). Thamnodynasteshypoconia feeds on amphibians and lizards (Bellini et al. 2013). Its litter can range from 4 to 13 hatchlings (Bellini et al. 2013).
Thamnodynastespallidus (Linnaeus, 1758) – A small-sized arboreal and terrestrial (average SVL = 325 mm; N = 92), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Amazon Forest and Caatinga (Bailey et al. 2005, Guedes et al. 2014, Marques et al. 2019). In the PEC it occurs in the states of Alagoas, Pernambuco and Paraíba (Fig. 10I), being found in Forest and Tabuleiros (Rodrigues et al. 2015, Pereira Filho et al. 2017). Thamnodynastespallidus feeds on amphibians (Guedes et al. 2014, Protázio et al. 2017). Its litter can range from 3 to 6 hatchlings (Cunha and Nascimento 1981, Araújo et al. 2018).
Thamnodynastesphoenix Franco, Trevine, Montingelli & Zaher, 2017 – A small to moderate size arboreal and terrestrial, with nocturnal activity (Franco et al. 2017, Marques et al. 2017a). This species occurs in the Atlantic Forest, Caatinga and Cerrado (Guedes et al. 2014, Franco et al. 2017, Freitas et al. 2019a). In the PEC it occurs only in Brejos Nordestinos of the state of Pernambuco (Fig. 10I) (Freitas et al. 2019a). Thamnodynastesphoenix feeds on amphibians (Pergentino and Ribeiro 2017).
Xenodonmerremii (Wagler, 1824) – A small to moderate size species (average SVL = 446 mm; N = 97), with diurnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Caatinga, Cerrado, Pampas Pantanal (Marques et al. 2005, 2015, 2019, Bérnils et al. 2007, Guedes et al. 2014). In the PEC it occurs in all states (Fig. 11A), being found in Forest, Brejos Nordestinos, Tabuleiros, and urban areas (Pereira Filho and Montingelli 2011, França et al. 2012, Rodrigues et al. 2015). Xenodonmerremii feeds on amphibians (Vitt and Vangilder 1983, Mesquita et al. 2009). Its litter can range from 4 to 30 eggs (Gaiarsa et al. 2013).
Geographic distribution records for snakes of the Pernambuco Endemism Center (PEC). AXenodonmerremii and X.rabdocephalusBXenopholisscalaris and X.undulatusCMicruruscorallinus, M.ibiboboca and M.potyguaraDEpictiaborapeliotesEAmerotyphlopsamoipira and A.arenensisFA.brongersmianus and A.paucisquamusGBothropsbilineatus and B.erythromelasHB.leucurus and B.muriciensisICrotalusdurissus and Lachesismuta.
https://binary.pensoft.net/fig/485070
Xenodonrabdocephalus (Wied-Neuwied, 1824) – A moderate-sized terrestrial species (average SVL = 630 mm; N = 2), with diurnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Amazon Forest and Cerrado (Cunha and Nascimento 1993, Marques et al. 2015, 2019). In the PEC it occurs in the states of Alagoas and Pernambuco (Fig. 11A), being found in Forest. Xenodonrabdocephalus feeds on amphibians (Martins and Oliveira 1998). Its litter can range from 6 to 8 eggs (Martins and Oliveira 1998).
Xenopholisscalaris (Wucherer, 1861) – A small-sized terrestrial species (average SVL = 167 mm; N = 10), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest and Amazon Forest (Marques et al. 2015, 2019, França et al. 2019). In the PEC it occurs in the states of Alagoas and Pernambuco (Fig. 11B), being found in Forest. Xenopholisscalaris feeds on amphibians (Martins and Oliveira 1998, Bernarde and Abe 2010). Its litter can range from 2 to 3 eggs (Martins and Oliveira 1998).
Xenopholisundulatus (Jensen, 1900) – A small-sized terrestrial species (average SVL = 268 mm; N = 2), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Caatinga and Cerrado (Guedes et al. 2014, Marques et al. 2015, 2019). In the PEC it occurs in the states of Alagoas, Pernambuco, and Paraíba (Fig. 11B), being found in Forest and Brejos Nordestinos (Pereira Filho et al. 2017). Xenopholisundulatus feeds on amphibians (Cunha and Nascimento 1993, Kokobum and Maciel 2010). Its litter can have 3 eggs (Costa et al. 2013).
Elapidae Boie, 1827
Micruruscorallinus (Merrem, 1820) – A small to moderate size cryptozoic species (average SVL = 465 mm; N =1), with diurnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest (Marques et al. 2019). In the PEC it occurs only in the state of Rio Grande do Norte (Fig. 11C), being found in Forest. Micruruscorallinus feeds on amphisbaenians, lizards, snakes, and caecilians (Marques and Sazima 1997). Its litter can range from 2 to 12 eggs (Azevedo 1961, Marques 1996b).
Micrurusibiboboca (Merrem, 1820) – A moderate-sized cryptozoic species (average SVL = 533 mm; N =391), with diurnal and nocturnal activity (Marques et al. 2017). This species occurs in the Atlantic Forest and Caatinga (Marques et al. 2017a, 2019). In the PEC it occurs in all states (Fig. 11C), being found in Forest, Brejos Nordestinos, Tabuleiros, and urban areas (Pereira Filho and Montingelli 2011, França et al. 2012, Rodrigues et al. 2015, Pereira Filho et al. 2017). Micrurusibiboboca feeds on amphisbaenians, snakes, and lizards (Vitt and Vangilder 1983, Mesquita et al. 2009). We found 9 to 14 vitellogenic follicles in females.
Micruruspotyguara Pires, Da Silva Jr, Feitosa, Prudente, Preira-Filho & Zaher, 2014 – A moderate-sized cryptozoic species (average SVL = 523 mm; N = 14), with diurnal and nocturnal activity (Marques et al. 2019). Micruruspotyguara is endemic to the PEC, occurring in the states of Pernambuco, Paraíba, and Rio Grande do Norte (Fig. 11C), being found in Forest, Tabuleiros, and urban areas (Pires et al. 2014, Rodrigues et al. 2015, França and França 2019).
Leptotyphlopidae Stejneger, 1891
Epictiaborapeliotes (Vanzolini, 1996) – A small-sized fossorial species (average SVL = 111 mm; N = 34), with diurnal and nocturnal activity (Guedes et al. 2014). This species occurs in the Atlantic Forest and Caatinga (Guedes et al. 2014, Marques et al. 2019). In the PEC it occurs in the states of Pernambuco, Paraíba, and Rio Grande do Norte (Fig. 11D), being found in Forest, Brejos Nordestinos, and in Restingas (Pereira Filho et al. 2017, Sampaio et al. 2018, Freitas et al. 2019a). Epictiaborapeliotes feeds on arthropods (Marques et al. 2019).
Typhlopidae Merrem, 1890
Amerotyphlopsamoipira (Rodrigues & Juncá, 2002) – A small-sized fossorial species (average SVL = 146 mm; N = 3), with nocturnal activity (Marques et al. 2017). This species occurs in the Caatinga and Atlantic Forest (Brito and Freire 2012). In the PEC it occurs in the states of Alagoas and Rio Grande do Norte (Fig. 11E), being found in Restinga (Brito and Freire 2012). Amerotyphlopsamoipira feeds on arthropods (Marques et al. 2017a).
Amerotyphlopsarenensis Graboski, Pereira Filho, Silva, Costa Prudente & Zaher, 2015 – A small-sized fossorial species (average SVL = 148 mm; N = 13). This species occurs in the Atlantic Forest and Caatinga (Graboski et al. 2015, 2019). In the PEC it occurs in the states of Alagoas, Pernambuco and Paraíba (Fig. 11E), being found in Forest and Brejos Nordestinos (Roberto et al. 2012, Graboski et al. 2015). We found 7 to 8 vitellogenic follicles in females.
Amerotyphlopsbrongersmianus (Vanzolini, 1976) – A small-sized fossorial species (average SVL = 212 mm; N = 120), with nocturnal activity (Marques et al. 2019). This species occurs in all Brazilian biomes (Graboski et al. 2019). In the PEC it occurs in the states of Alagoas, Pernambuco and Paraíba (Fig. 11F), being found in Forest and Tabuleiros (Pereira Filho et al. 2017, Sampaio et al. 2018). This species occurs in the Atlantic Forest (Marques et al. 2019). Amerotyphlopsbrongersmianus feeds on ant larvae (Avila et al. 2006). Its litter can range from 4 to 5 eggs (Avila et al. 2006).
Amerotyphlopspaucisquamus (Dixon, 1979) – A small-sized fossorial species (average SVL = 133 mm; N =153), with nocturnal activity (Marques et al. 2019). This species is endemic to the PEC, occurring in all states (Fig. 11F), being found in Forest and Tabuleiros (Rodrigues et al. 2015, Pereira Filho et al. 2017). We found four eggs in one female and another individual laid three eggs after being collected.
Viperidae Laurenti, 1768
Bothropsbilineatus (Wied-Neuwied, 1821) – A small to moderate sized arboreal species (average SVL = 495 mm; N = 5), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest and Amazon Forest (Bernarde et al. 2011, Marques et al. 2019). In PEC occurs only in Alagoas state (Fig. 11G), being found in Forest. Bothropsbilineatus feeds on mammals, amphibians, birds, snakes, and lizards (Cunha and Nascimento 1993, Martins et al. 2002, Turci et al. 2009). Its litter can range from 4 to 16 hatchlings (Dixon and Soini 1986, Campbell and Lamar 2004, Grego et al. 2012, Almeida et al. 2019).
Bothropserythromelas Amaral, 1923 – A small to moderate size terrestrial species (average SVL = 445 mm; N = 3), with nocturnal activity (Marques et al. 2017). This species occurs in the Caatinga, but can also be found in transitional areas with the Atlantic Forest (Guedes et al. 2014). In the PEC it occurs in the states of Pernambuco and Rio Grande do Norte (Fig. 11G), being found in Forest. Bothropserythromelas feeds on arthropods when juveniles, and frogs, lizards, and mammals when adults (Martins et al. 2002). Its litter can range from 2 to 21 hatchlings (Barros et al. 2014, Reis et al. 2015).
Bothropsleucurus Wagler, 1824 – A moderate-sized terrestrial species (average SVL = 589 mm; N =207), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest (Marques et al. 2019). In the PEC it occurs in the states of Alagoas, Pernambuco and Paraíba (Fig. 11H), being found in Forest, Brejos Nordestinos, Tabuleiros, mangroves, and urban areas when near forest areas (Pereira Filho and Montingelli 2011, Rodrigues et al. 2015, Pereira Filho et al. 2017, França and França 2019). Bothropsleucurus feeds on amphibians, lizards, snakes, birds, and mammals. Its litter can range from 5 to 7 hatchlings (Lira-da-Silva et al. 1994).
Bothropsmuriciensis Ferrarezzi & Freire, 2001 – A moderate-sized terrestrial species (average SVL = 512 mm; N = 6), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest (Marques et al. 2019). This species is endemic to the PEC, occurring only in the Estação Ecológica de Murici (Fig. 11H), located in the state of Alagoas, being found in Forest. See Freitas et al. (2012) for additional information on this species. As observed in other congenerics, it probably feeds on anurans and small mammals.
Crotalusdurissus Linnaeus, 1758 – A moderate-sized terrestrial species (average SVL = 790 mm; N = 13), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest, Caatinga, Cerrado, Pampas, and Pantanal (Marques et al. 2005, 2015, 2019, Bérnils et al. 2007, Guedes et al. 2014). In the PEC it occurs in the states of Alagoas, Pernambuco, and Paraíba (Fig. 11I), being found in Forest, Brejos Nordestinos, and Restingas (Lira-da-silva et al. 2009, Pereira Filho and Montingelli 2011). Crotalusdurissus feeds on mammals (Vitt and Vangilder 1983, Strussmann and Sazima 1993, Rodrigues et al. 2016). Its litter can range from 21 to 31 hatchlings (Vitt and Vangilder 1983).
Lachesismuta (Linnaeus, 1766) – A large size terrestrial species (average SVL = 1217 mm; N = 4), with nocturnal activity (Marques et al. 2019). This species occurs in the Atlantic Forest and Amazon Forest (Cunha and Nascimento 1993, Marques et al. 2019). In the PEC it occurs in the states of Alagoas, Pernambuco and Paraíba (Fig. 11I), being found in Forest (Pereira Filho et al. 2017). Lachesismuta feeds on mammals (Cunha and Nascimento 1993, Martins and Oliveira 1998). Its litter can range from 1 to 18 eggs (Martins and Oliveira 1998, Souza 2007, Alves et al. 2014).
Discussion
Our results show a broad view of PEC’s snake fauna, including distribution data, natural history, and diversity. According to Marques et al. (2019), about 142 species of snakes occur in the Brazilian Atlantic Forest, the 78 species recorded in the PEC represent 51.3% of this total, which we can consider a high richness. In addition, new species are still being discovered in this region, for example, the species D.atlantica (Freire et al. 2010), M.potyguara (Pires et al. 2014), and A.arenensis (Graboski et al. 2015) have been described in the last ten years and at least one new species (Caaeteboia sp.) is being described at the moment (Pereira Filho et al. 2017).
The mixed composition of snake species that inhabit the Atlantic Forest located north of the São Francisco River can be considered a remarkable characteristic of this fauna (Pereira Filho et al. 2017). We can highlight that the main difference between the PEC and other portions of the biome is due to the large number of species of open areas and also of wide distribution that are present in this region. The PEC shares more species with the Caatinga and the Cerrado (74.3% and 56.4% of the shared species, respectively) than with the southern and southeastern regions of the Atlantic Forest (30% of the species are shared). This may be due to the fact that the PEC presents different physiognomic features, such as patches of Tabuleiros, which are natural enclaves of savannah found even in the middle of forests and which may provide adequate conditions for the establishment of populations of species from open areas (Mesquita et al. 2018). In addition, the proximity to the Caatinga may also have favoured the penetration and establishment of these populations (Pereira Filho et al. 2017). These arguments are supported by historical factors that are based on the expansion and retraction of the boundaries of dry and open habitat ecoregions, due to climatic fluctuations over geological time, which have reached coastal areas of northeastern Brazil (Ab’Saber 1977, Pennington et al. 2006). Thus, species considered previously endemic to the Caatinga, for example, E.borapeliotes and E.assisi (Guedes et al. 2014), and species considered endemic to the Cerrado, for example C.flavolineatus (Nogueira et al. 2010), are also abundant in the PEC.
Most reptiles are considered habitat specialists, which means that many species can only survive in one or a few distinct environments (Martins and Molina 2008). In the PEC, the great majority of snake species were found in forest areas and 26 species were collected only in this environment. Due to the occupation of the area for agriculture and urbanization, most of the forest in the PEC was lost or reduced to small fragments, mostly smaller than ten hectares, which represent less than 2% of the original coverage of the Center (Ranta et al. 1998, Tabarelli et al. 2005). This is especially worrying because species that do not use the surrounding matrix as part of their area of use or that cannot use these environments to move between the fragments, can become extinct regionally as the populations are becoming isolated, making them unviable in the long term, due to the reduced population size (Nunney and Campbell 1993). On the other hand, some species seem to be generalists in terms of habitat and can be found in different physiognomies of the PEC and even urban areas, as is the case of B.constrictor, P.olfersii, B.leucurus and O.trigeminus.
Most snake species found in the PEC mainly use soil as substrate, as well as snakes in other regions of Brazil, such as the Caatinga (Guedes et al. 2014), Atlantic Forest (Marques et al. 2017b), Cerrado (França and Braz 2013), Pantanal (Strussmann and Sazima 1993) and Amazon (Martins and Oliveira 1998, Bernarde and Abe 2006). However, PEC also harbours a great variety of semi-arboreal and arboreal species, which is a characteristic of forest biomes, such as the Atlantic Forest and Amazon (Martins and Oliveira 1998, Argôlo 2004, Marques and Sazima 2004, Bernarde and Abe 2006).
More than half of PEC snakes feed on lizards or amphibians. These types of prey are commonly found in the snake diet, although other vertebrates like mammals, birds, and snakes are also important preys (Bernarde and Abe 2006, Hartmann et al. 2009, Mesquita et al. 2009). Some species of the PEC are generalists, as boids and snakes of the genus Philodryas and Oxyrhopus. Snakes belonging to the genera Apostolepis, Dipsas, and Atractus have specialized diet, feeding on snakes, mollusks and earthworms, respectively, as well as the genera Xenodon and Xenopholis, which are specialists in amphibians. (Vitt and Vangilder 1983, Laporta-Ferreira et al. 1986, Cunha and Nascimento 1993, Martins and Oliveira 1998, Mesquita et al. 2009, Bernarde and Abe 2010, Fernandes et al. 2010, Kokobum and Maciel 2010).
It is important to emphasize that the PEC presents at least seven endemic species (A.caete, A.maculatus, B.muriciensis, Caaeteboia sp., D.atlantica, E.cephalomaculata, and M.potyguara) of which basic information on natural history and ecology are scarce. Most of these species have a very restricted distribution, have been little recorded in nature and consequently are poorly represented in scientific collections. For example, B.muriciensis has only nine records and was found only in a single location (Freitas et al. 2012), the E.cephalomaculata has seven known records and was found only in four locations (Freitas et al. 2019b) and Caaeteboia sp., which has only three records and should be a new species for the region (Pereira Filho et al. 2017). Moreover, some species have confused taxonomy, such as M.ibiboboca and D.neuwiedi, being a complex of different taxa. Some of these taxa could figure as endemic species in PEC in the future. Besides the endemic species, other PEC species deserve special attention due to the absence of information on natural history and ecology, for being rare in the region and for presenting a restricted distribution in the PEC, for example, L.trefauti, A.potschi, D.sazimai, D.variegata, E.cephalostriata, and A.arenensis.
The conservation status of PEC snake species is still little known. Of the 78 species registered in the region, only 25 species have been evaluated by the IUCN (International Union for Conservation of Nature) to date. On the Brazilian list of threatened species, some PEC species are present, they are: A.amoipira, A.caete, and B.muriciensis as “endangered” and A.paucisquamus and E.cephalomaculata as “vulnerable” (ICMBio 2018). Given the high richness of snake species, the number of endemic species and the fragmented conditions of the region’s forests, regional conservation efforts need to be intensified, because few forests north of the São Francisco River are formally protected, and the majority are small, which means that many species in the region may be threatened with extinction (Ranta et al. 1998, Uchoa Neto and Tabarelli 2002, Tabarelli et al. 2006a).
In general, many studies still need to be developed in the PEC region, so that we can better understand the snake fauna of this region. Fauna inventories in areas that are not well sampled, population dynamics studies and distribution patterns are important for better conservation planning of PEC snake species.
Acknowledgements
RCF thanks FAPESB for a PhD scholarship (BOL0353/2016) and CNPq for Doutorado Sanduíche no Exterior (SWE) scholarship (208442/2017-0). FGRF thanks the financial support from CNPq (Universal grant 404671/2016-0). MS acknowledges funding from CNPq (Productivity grant 304999/2015-6) and CAPES/Alexander von Humboldt Foundation (BEX 0585/16-5). We further thank Fausto Erritto Barbo, an anonymous reviewer and the editor Robert Jadin for detailed and constructive comments that greatly improved our manuscript.
ReferencesAb’SaberAN (1977) Espaços ocupados pela expansão dos climas secos na América do Sul, por ocasião dos períodos glaciais quaternários.3: 1–19.AbeggADFreitasMAMouraGJB (2017a) First confirmed record of Atractusmaculatus (Serpentes, Dipsadidae) from the state of Pernambuco, northeastern Brazil.13: 1–3. https://doi.org/10.15560/13.2.CL2080AbeggADFreitasMAMouraGJB (2017b) New records of Liotyphlopstrefauti Freire, Caramaschi & Argôlo, 2007 (Squamata: Anomalepididae).10: 345–347.AlbarelliLPPSantos-CostaMC (2010) Feeding ecology of Liophisreginaesemilineatus (Serpentes: Colubridae: Xenodontinae) in Eastern Amazon, Brazil.27: 87–91. https://doi.org/10.1590/S1984-46702010000100013AlbuquerqueNRGalattiUDi-BernardoM (2007) Diet and feeding behaviour of the Neotropical parrot snake (Leptophisahaetulla) in northern Brazil.41: 1237–1243. https://doi.org/10.1080/00222930701400954AlencarLRVGaldinoCABNascimentoLB (2012) Life History Aspects of Oxyrhopustrigeminus (Serpentes: Dipsadidae) from Two Sites in Southeastern Brazil.46: 9–13. https://doi.org/10.1670/09-219AlencarLRVGaiarsaMPMartinsM (2013) The evolution of diet and microhabitat use in Pseudoboine snakes.8: 60–66. https://doi.org/10.2994/SAJH-D-13-00005.1AlmeidaBJLAlmeidaMSMCavalcanteCSBernardePS (2019) Bothropsbilineatusbilineatus (Two-striped Forest Pitviper) Reproduction.50: 385–386.AlvesFQArgôloAJSCarvalhoGC (2014) Reproductive biology of the bushmaster Lachesismuta (Serpentes: Viperidae) in the Brazilian Atlantic Forest.13: 99–109. https://doi.org/10.11606/issn.2316-9079.v13i2p99-109AmaralA (1978) Serpentes do Brasil: iconografia colorida. 2a edição. Melhoramentos e ED-USP, São Paulo.AmorimDMSilvaMCQuirinoTFRobertoIJÁvilaRW (2015) Apostolepiscearensis (Burrowing snake). Diet.46: 265–266.Andrade-LimaD (1982) Present day forest refuges in Northeastern Brazil. In: PranceGT (Ed.) Biological diversification in the tropics., 245–251.AndradeROSilvanoRAM (1996) Comportamento alimentar e dieta da "falsa-coral" Oxyrhopusguibei Hoge & Romano (Serpentes, Colubridae) Revista Brasileira de Zoologia 13: 143–150. https://doi.org/10.1590/S0101-81751996000100014AraujoDSD (1992) Vegetation types of sandy coastal plains of tropical Brazil: A first approximation. In: Seeliger U (Ed.) Coastal Plant Communities of Latin America. Academic Press, New York, 392. https://doi.org/10.1016/B978-0-08-092567-7.50027-1AraújoPFrançaRCNascimentoFSLaranjeirasDOFrançaFGR (2019) New records and range expansion of Chironiuscarinatus (Linnaeus, 1758) (Serpentes, Colubridae) from the state of Paraíba, northeast Brazil.15: 927–932. https://doi.org/10.15560/15.5.927AraújoPFSilvaWMFrançaRCFrançaFGR (2018) A case of envenomation by neotropical opisthoglyphous snake Thamnodynastespallidus (Linnaeus, 1758) (Colubridae: Dipsadinae: Tachymenini) in Brazil.60: 11–14. https://doi.org/10.1590/s1678-9946201860038ArgôloAJS (2004) Editus, Ilhéus, 260 pp.ArzamendiaV (2016) New Southern record of Erythrolamprusreginae (Linnaeus, 1758) (Serpentes: Dipsadidae), a vulnerable species in Argentina.12: 1–4. https://doi.org/10.15560/12.5.1976AvilaRWFerreiraVLSouzaVB (2006) Biology of the blindsnake Typhlopsbrongersmianus (Typhlopidae) in a semideciduous forest from Central Brazil.16: 403–405.ÁvilaRWMoraisDH (2007) Notes on the ecology of the colubrid snake Leptodeiraannulata in the Pantanal, Brazil.38: 278–280.ÁvilaRWFerreiraVLArrudaJAO (2006) Natural History of the South American Water Snake Helicopsleopardinus (Colubridae: Hydropsini) in the Pantanal, Central Brazil.40: 274–279. https://doi.org/10.1670/113-05N.1AzevedoACP (1961) Notas sobre cobras-corais (Serpentes- Elapidae) III a IV.18: 1–23.BaileyJRThomasRASilva JrNJ (2005) A revision of the South American snake genus Thamnodynastes Wagler, 1830 (Serpentes, Colubridae, Tachymenini). I. Two new species of Thamnodynastes from Central Brazil and adjacent areas, with a redefinition of and neotype designation of Thamnodynastespallidus (Linnaeus, 1758).4: 83–101. https://doi.org/10.11606/issn.2316-9079.v4i2p83-101BalestrinRLDi-BernardoM (2005) Ophiophagy in the colubrid snake Echinantheraoccipitalis (Jan, 1863) from southern Brazil.41: 221–222.BarboFEMarquesOAV (2003) Do aglyphous colubrid snakes prey on live amphisbaenids able to bite? Phyllomedusa 2: 113–114. https://doi.org/10.11606/issn.2316-9079.v2i2p113-114BarboFEMarquesOAVSawayaRJ (2011) Diversity, Natural History, and Distribution of Snakes in the Municipality of São Paulo.6: 135–160. https://doi.org/10.2994/057.006.0301BarbosaVNAmaralJMSLimaLFLFrançaRCFrançaFGRSantosEM (2019) A case of melanism in Dendrophidionatlantica Freire, Caramaschi & Gonçalves, 2010 (Colubridae) from northeastern Brazil.12: 109–111.BarrosVARojasCAAlmeida-SantosSM (2014) Reproductive Biology of Bothropserythromelas from the Brazilian Caatinga.2014: 1–11. https://doi.org/10.1155/2014/680861BeebeW (1946) Field notes on the snakes of Kartabo, British Guiana, and Caripito, Venezuela.31: 11–52.BelliniGArzamendiaVGiraudoAR (2013) Ecology of Thamnodynasteshypoconia in Subtropical – Temperate South America.69: 67–79. https://doi.org/10.1655/HERPETOLOGICA-D-12-00027BernardePSAbeAS (2006) A snake community at Espigão do Oeste, Rondônia, southwestern Amazon, Brazil. South American Journal of Herpetology 1: 102–113. https://doi.org/10.2994/1808-9798(2006)1[102:ASCAED]2.0.CO;2BernardePSAbeAS (2010) Hábitos alimentares de serpentes em Espigão do Oeste, Rondônia, Brasil Introdução Material e Métodos Resultados.10: 167–173. https://doi.org/10.1590/S1676-06032010000100017BernardePSCostaHCMachadoRASão-PedroVDA (2011) Bothriopsisbilineatabilineata (Wied, 1821) (Serpentes: Viperidae): new records in the states of Amazonas, Mato Grosso and Rondônia, northern Brazil. Check List 7: 343. https://doi.org/10.15560/7.3.343BérnilsRSGiraudoARCarreiraSCechinSZ (2007) Répteis das porções subtropical e temperada da região neotropical.35: 101–136.BoosHEA (2001) The snakes of Trinidad and Tobago. Texas A&M University Press, 270 pp.Borges-NojosaDMLimaDC (2001) Dieta de Drymoluberdichrous (Peters, 1863) dos brejos-de-altitude do estado do Ceará, Brasil (Serpentes: Colubridae). Boletim do Museu Nacional.468: 1–5.BrazHBScartozzoniRRAlmeida-SantosSM (2016) Reproductive modes of the South American water snakes: A study system for the evolution of viviparity in squamate reptiles.263: 33–44. https://doi.org/10.1016/j.jcz.2016.04.003BritoPSFreireEMX (2012) New records and geographic distribution map of Typhlopsamoipira Rodrigues and Juncá, 2002 (Typhlopidae) in the Brazilian Rainforest.8: 1347–1349. https://doi.org/10.15560/8.6.1347CampbellJALamarWW (2004) Cornell University Press, Ithaca, 425 pp.Coimbra-FilhoAFCâmaraIG (1996) FBCN, Rio de Janeiro, 86 pp.CostaHCClaraMGurgelC (2013) Xenopholisundulatus (Serpentes: Xenodontinae): Reprodução e alimentação em cativeiro.2: 36–38.CostaHCPantojaDLPontesJLFeioRN (2010) Serpentes do Município de Viçosa, Mata Atlântica do Sudeste do Brasil Material e Métodos.10: 352–376. https://doi.org/10.1590/S1676-06032010000300033CunhaORNascimentoFP (1981) Ofídios da Amazônia. XII – Observações sobre a viviparidade em ofídios do Pará e Maranhão (Ophidia: Aniliidae, Boidae, Colubridae e Viperidae. Boletim do Museu Paraense Emilio Goeldi. Nova Série Zoologia: 24.CunhaORNascimentoFP (1993) Ofídios da Amazônia. As cobras da região leste do Pará.9: 1–188.CurcioFFSales NunesPMHarveyMBRodriguesMT (2011) Redescription of Apostolepislongicaudata (Serpentes: Xenodontinae) with comments on its hemipenial morphology and natural history.67: 318–331. https://doi.org/10.1655/HERPETOLOGICA-D-10-00043.1Di-BernardoM (1994) Uma nova espécie de Echinanthera Cope, 1894 (Serpentes, Colubridae) do nordeste do Brasil.2: 75–81.Di-BernardoM (1996) A new species of the neotropical snake genus Echinanthera COPE 1894 from southeastern Brazil (Serpentes, Coluridae).27: 120–126.DixonJR (1989) A key and checklist to the neotropical snake genus Liophis with country lists and maps.79: 1–40. https://doi.org/10.5479/si.23317515.79.1DixonJRSoiniP (1986) Milwaukee Public Museum The reptiles of the upper Amazon Basin, Iquitos Region, Peru. Milwaukee, 91 pp.DixonJRWiestJACeiJM (1993) Revision of the Neotropical Snake Genus Chironius Fitzinger (Serpentes, Colubridae).47: 155–167.DuellmanWE (2005) Cornell University Press, Ithaca and London, 472 pp.FernandesDSMarquesOAVArgôloAJS (2010) A new species of Dipsas Laurenti from the Atlantic Forest of Brazil.66: 57–66. https://doi.org/10.11646/zootaxa.2691.1.4FowlerIRSalomãoMDGJordãoRS (1998) A description of the female reproductive cycle in four species from the Neotropical colubrid snake Philodryas (Colubridae, Xenodontinae).28: 71–78.FragaRLimaAPPrudenteALCMagnussonWE (2013) Guia de Cobras da região de Manaus- Amazonia Central. Editora Inpa.FrançaFGRBrazVS (2013) Diversity, activity patterns, and habitat use of the snake fauna of Chapada dos Veadeiros National Park in Central Brazil.13: 74–84. https://doi.org/10.1590/S1676-06032013000100008FrançaFGRMesquitaDOColliGR (2006) A checklist of snakes from Amazonian Savannas in Brazil, housed in the coleção herpetológica da Universidade de Brasília, with new distribution records. Occasional Papers, Oklahoma Museum of Natural History: 1–13.FrançaRCFrançaFGR (2019) Spatial patterns of snake diversity in an urban area of north-east Brazil.29: 274–281. https://doi.org/10.33256/hj29.4.274281FrançaRCGermanoCESFrançaFGR (2012) Composition of a snake assemblage inhabiting an urbanized area in the Atlantic Forest of Paraíba State, Northeast Brazil.12: 183–195. https://doi.org/10.1590/S1676-06032012000300019FrançaRCMoraisMSRFreitasMAMouraGJBFrançaFGR (2019) A new record of Xenopholisscalaris (Wucherer, 1861) (Dipsadidae) for the state of Pernambuco, Brazil.12: 57–59.FrancoFLFerreiraTG (2002) Descrição de uma nova espécie de Thamnodynastes Wagler, 1830 (Serpentes, Colubridae) do nordeste brasileiro, com comentários sobre o gênero. Phyllomedusa: Journal of Herpetology 1: 57. https://doi.org/10.11606/issn.2316-9079.v1i2p57-74FrancoFLTrevineVCMontingelliGGZaherH (2017) A new species of Thamnodynastes from the open areas of central and Northeastern Brazil (Serpentes: Dipsadidae: Tachymenini).53: 339–350.FranziniLDPedroCKBCavalcantiLBDQMesquitaDO (2018) Predation of Hemidactylusmabouia (Sauria: Gekkonidae) by a vine snake Oxybelisaeneus (Serpentes: Colubridae) in an Atlantic Forest fragment, Northeastern Brazil.2: 67–70. https://doi.org/10.29215/pecen.v2i1.587FreireEMXCaramaschiUArgôloAJS (2007) A new species of Liotyphlops (Serpentes: Anomalepididae) from the Atlantic Rain Forest of Northeastern Brazil.26: 19–26. https://doi.org/10.11646/zootaxa.1393.1.2FreireEMXCaramaschiUGonçalvesU (2010) A new species of Dendrophidion (Serpentes: Colubridae) from the Atlantic Rain Forest of Northeastern Brazil.2719: 62–68. https://doi.org/10.11646/zootaxa.2719.1.5FreitasMAFrançaDPFGraboskiRUhligVVeríssimoD (2012) Notes on the conservation status, geographic distribution and ecology of Bothropsmuriciensis Ferrarezzi & Freire, 2001 (Serpentes, Viperidae).8: 338–343.FreitasMAAbeggADAraújoDSCoelhoHEAAzevedoWSChavesMFRosaCMMouraGJB (2019a) Herpetofauna of three “Brejos de altitude” in the interior of the state of Pernambuco, northeastern Brazil.12: 591–602.FreitasMABarbosaGGBernardinoKPDomingosJFilhoP (2019b) First records of the rare snake Echinantheracephalomaculata Di-Bernardo, 1994 in the state of Pernambuco, Brazil (Serpentes: Dipsadidae).12: 1005–1009.GaiarsaMPAlencarLRVMartinsM (2013) Natural History of Pseudoboine Snakes.53: 261–283. https://doi.org/10.1590/S0031-10492013001900001GoldbergSR (2007) Note on Reproduction of Whipsnakes, Genus Chironius (Serpentes: Colubridae), from Costa Rica.42: 148–149.GomesC (2012) História natural das serpentes dos gêneros Echinanthera e Taeniophallus (Echinantherini). MSc Dissertation, Universidade Estadual Paulista.GoodM (1989) Pseustessulphureus. Reproduction. Herpetological Review 20: 73.GraboskiRPereira FilhoGASilvaAAACosta PrudenteALZaherH (2015) A new species of Amerotyphlops from Northeastern Brazil, with comments on distribution of related species.3920: 443–452. https://doi.org/10.11646/zootaxa.3920.3.3GraboskiRArredondoJCGrazziotinFGSilvaAAAPrudenteALCRodriguesMTBonattoSLZaherH (2019) Molecular phylogeny and hemipenial diversity of South American species of Amerotyphlops (Typhlopidae, Scolecophidia).48: 139–156. https://doi.org/10.1111/zsc.12334GrantPBCLewisTR (2010) Predation attempt by Oxybelisaeneus (Wagler) (Mexican Vine- snake) on Basiliscusplumifrons (Cope).5: 19–22.GregoKFFernandesWCroceAPVasconcellosDRSant’AnnaSSCoragemJT (2012) Bothriopsisbilineatasmaragdinus (green jararaca) reproduction. Herpetological Review 43: 492.GuedesTBNogueiraCMarquesOAV (2014) Diversity, natural history, and geographic distribution of snakes in the Caatinga, Northeastern Brazil.3863: 1–93. https://doi.org/10.11646/zootaxa.3863.1.1HamdanBFernandesDS (2015) Taxonomic revision of Chironiusflavolineatus (Jan, 1863) with description of a new species (Serpentes: Colubridae).4012: 97–119. https://doi.org/10.11646/zootaxa.4012.1.5HartmannPAMarquesOAV (2005) Diet and habitat use of two sympatric species of Philodryas (Colubridae), in south Brazil.26: 25–31. https://doi.org/10.1163/1568538053693251HartmannPAHartmannMTMartinsM (2009) Ecology of a snake assemblage in the Atlantic Forest of southeastern Brazil.49: 343–360. https://doi.org/10.1590/S0031-10492009002700001HauzmanECostaACORScartozonniRR (2005) Spilotespullatus (Tiger Ratsnake). Reproduction. Herpetological Review 36: 328.HendersonRW (1982) Trophic Relationships and Foraging Strategies of some New World Tree Snakes (Leptophis, Oxybelis, Uromacer).3: 71–80. https://doi.org/10.1163/156853882X00185HetheringtonTE (2006) Oxybelisaeneus (Brown Vinesnake). Diet.37: 94–95.IBGE (2004) Mapa de biomas e vegetação do Brasil. Instituto Brasileiro de Geografia e Estatística – IBGE. https://www.ibge.gov.br/geociencias/informacoes-ambientais/estudos-ambientais/15842-biomas.html?=&t=downloads [November 24, 2019]KokobumMNCMacielMN (2010) Scinaxfuscovarius NCN). Predation.41: 480–481.Laporta-FerreiraILSalomãoMGSawayaP (1986) Biologia de Sibynomorphus (Colubridae – Dipsadinae) – Reprodução e Hábitos Alimentares.46: 793–799.LeitePTNunesSFCechinSZ (2007) Dieta e uso de habitat da jararaca-do-brejo, Mastigodryasbifossatus Raddi (Serpentes, Colubridae) em domínio subtropical do Brasil.24: 729–734. https://doi.org/10.1590/S0101-81752007000300025LemaTD (2003) Editora da Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre, 485 pp.LimaLFLAmaralJMSBarbosaVNSantosEM (2019) Dendrophidionatlantica/Freire, Caramaschi e Gonçalves, 2010 – Contribuições sobre reprodução.48: 1–4. https://doi.org/10.5380/abpr.v48i(1-2).69161Lira-da-SilvaRMMiseYFCasais-e-SilvaLLUlloaJHamdanB (2009) Serpentes de importância médica do nordeste do Brasil.79: 7–20.Lira-da-SilvaRMCasais-e-SilvaLLQueirozIBNunesTB (1994) Contribuição á biologia de serpentes da Bahia, Brasil. I. Vivíparas.11: 187–193. https://doi.org/10.1590/S0101-81751994000200002LópezMSGiraudoAR (2004) Diet of the large water snake Hydrodynastesgigas (Colubridae) from northeast Argentina María.25: 178–184. https://doi.org/10.1163/1568538041231148LynchJD (2009) Snakes of the genus Oxyrhopus (Colubridae: Squamata) in Colombia: Taxonomy and geographic variation.49: 319–337. https://doi.org/10.1590/S0031-10492009002500001MarquesOAVSazimaI (2004) História Natural dos Répteis da Estação Ecológica JuréiaItatins. In: MarquesOAVDulebaW (Eds) , Estação Ecológica JuréiaItatins., 257–277.MarquesOAV (1996a) Biologia reprodutiva da cobra-coral Erythrolamprusaesculapii Linnaeus (Colubridae), no Sudeste do Brasil.13: 747–753. https://doi.org/10.1590/S0101-81751996000300022MarquesOAV (1996b) Reproduction, seasonal activity and growth of the coral snake, Micruruscorallinus (Elapidae), in the southeastern Atlantic forest in Brazil.17: 277–285. https://doi.org/10.1163/156853896X00441MarquesOAVPuortoG (1992) Dieta e comportamento alimentar de Erythrolamprusaesculapii, uma serpente ofiófaga.54: 253–259.MarquesOAVSazimaI (1997) Diet and feeding behavior of the coral snake, Micruruscorallinus, from the Atlantic Forest of Brazil.5: 88–93.MarquesOAVEterovicASazimaI (2019) Ponto A, Cotia, 318 pp.MarquesOAVEterovicAStrüssmannCSazimaI (2001) Serpentes da Mata Atlântica: guia ilustrado para a Serra do Mar. Holos. Ribeirão Preto, 184 pp.MarquesOAVEterovicAStrüssmannCSazimaI (2005) Serpentes do Pantanal: guia ilustrado. Holos. Ribeirão Preto, 184 pp.MarquesOAVEterovicANogueiraCSazimaI (2015) Serpentes do Cerrado: guia ilustrado. Holos. Ribeirão Preto, 248 pp.MarquesOAVEterovicAGuedesTBSazimaI (2017a) Ponto A, Cotia, 240 pp.MarquesOAVPereiraDNBarboFEGermanoVJSawayaRJ (2009) Os Répteis do Município de São Paulo: diversidade e ecologia da fauna pretérita e atual Métodos.9: 139–150. https://doi.org/10.1590/S1676-06032009000200014MarquesOAVMuniz-Da-SilvaDFBarboFECardosoSRTMaiaDCAlmeida-SantosSM (2014) Ecology of the Colubrid Snake Spilotespullatus from the Atlantic Forest of Southeastern Brazil.70: 407–416. https://doi.org/10.1655/HERPETOLOGICA-D-14-00012MarquesRRödderDSoléMTinôcoMS (2017b) Diversity and habitat use of snakes from the coastal Atlantic rainforest in northeastern Bahia, Brazil.53: 34–43.MarquesRMebertKFonsecaÉRödderDSoléMTinôcoMS (2016) Composition and natural history notes of the coastal snake assemblage from Northern Bahia, Brazil.611: 93–142. https://doi.org/10.3897/zookeys.611.9529MartinsMOliveiraME (1998) Natural history of snakes in forests of the Manaus region, Central Amazonia, Brazil.6: 78–150.MartinsMMolinaFB (2008) Panorama geral dos répteis ameaçados do Brasil. In: MachadoABDrummondGPagliaA (Eds) Livro vermelho da fauna brasileira ameaçada de extinção., 327–334.MartinsMMarquesOAVSazimaI (2002) Ecological and phylogenetic correlates of feeding habits in neotropical pitvipers of the genus Bothrops. In: SchuettGWHoggrenMDouglasMEGreeneHW (Eds) Biology of the Vipers., 307–328. http://jararacailhoa.org/conservacaoinsularis/bothrops_feeding.pdfMesquitaDOAlvesBCFPedroCKBLaranjeirasDOCaldasFLSPedrosaIMMCRodriguesJBDrummondLOCavalcantiLBQNogueira-CostaPFrançaRCFrançaFGR (2018) Herpetofauna in two habitat types (tabuleiros and Stational Semidecidual Forest) in the Reserva Biológica Guaribas, northeastern Brazil.11: 455–474.MesquitaPCMDBritoWBorges-NojosaDM (2011a) Natural History Notes: Psomophisjoberti (NCN). Reproduction. Herpetological Review 42: 302.MesquitaPCMDPassosDCBorges-NojosaDMBeyerraCH (2009) Apostolepiscearensis (Gomes’ Burrowing snake) diet. Herpetological Review 40: 440.MesquitaPCMDBorges-NojosaDMPassosDCBezerraCH (2011b) Ecology of Philodryasnattereri in the Brazilian semi-arid region.21: 193–198.MesquitaPCMDPassosDCBorges-NojosaDMCechinSZ (2013) Ecologia e história natural das serpentes de uma área de Caatinga no nordeste Brasileiro.53: 99–113. https://doi.org/10.1590/S0031-10492013000800001ICMBio [Instituto Chico Mendes de Conservação da Biodiversidade/MMA–Ministério do Meio Ambiente] (2018) Livro Vermelho da Fauna Brasileira Ameaçada de Extinção: Volume IV–Répteis.MoraisMSRFrançaRCDelfimFRFrançaFGR (2018) Eggs and hatchling morphometry of Spilotessulphureus (Wagler in Spix, 1824) (Serpentes: Colubridae: Colubrinae: Colubroidea: Caenophidia) from northeastern Brazil.11: 441–444.MorellatoLPCHaddadCFB (2000) Introduction: The Brazilian Atlantic Forest.32: 786–792. https://doi.org/10.1111/j.1744-7429.2000.tb00618.xMouraGJBSantosEMSOliveiraMABCabralMCC (2011) Brasília, Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Renováveis, 443 pp.MouraJM (1999) Leptodeiraannulata (Culebra Desteñida, Banded Cat-eyed Snake). Diet. Herpetological Review 30: 102.MyersNMittermeierRMittermeierCDaFonescaGKentJ (2000) Biodiversity hotspots for conservation priorities. Conservation Biology 403: 853. https://doi.org/10.1038/35002501NogueiraCColliGRCostaGMachadoRB (2010) Diversidade de répteis Squamata e evolução do conhecimento faunístico no Cerrado. In: DinizIRMarinho-FilhoJMachadoRBCavalcantiRB (Eds) Cerrado: conhecimento científico quantitativo como subsídio para ações de conservação., 329–372.NunneyLCampbellKA (1993) Assessing minimum viable population size: Demography meets population genetics.8: 234–239. https://doi.org/10.1016/0169-5347(93)90197-WOrofinoRDPPizzattoLMarquesOAV (2010) Reproductive biology and food habits of Pseudoboanigra (Serpentes: Dipsadidae) from the Brazilian cerrado.9: 53–61. https://doi.org/10.11606/issn.2316-9079.v9i1p53-61PassosPFernandesR (2008) Revision of the Epicratescenchria Complex (Serpentes: Boidae). Herpetological Monographs: 1–30. https://doi.org/10.1655/06-003.1PassosPFernandesRBérnilsRSMoura-LeiteJC (2010) Taxonomic revision of the Brazilian Atlantic Forest Atractus (Reptilia: Serpentes: Dipsadidae).2364: 1–63. https://doi.org/10.11646/zootaxa.2364.1.1PenningtonRTLewisGPRatterJA (2006) An overview of the plant diversity, biogeography and conservation of neotropical savannas and seasonally dry forests. In: T. PR, Ratter JA, Lewis GP (Eds) Neotropical savannas and seasonally dry forests: Plant biodiversity, biogeography and conservation. CRC Press, Boca Raton, 1–29. https://doi.org/10.1201/9781420004496-1Pereira FilhoGAVieiraWLSAlvesRRNFrançaFGR (2017) Serpentes da Paraíba: Diversidade e Conservação. João Pessoa, 316 pp.Pereira FilhoGAMontingelliGG (2011) Check list of snakes from the Brejos de Altitude of Paraíba and Pernambuco, Brazil.11: 145–151. https://doi.org/10.1590/S1676-06032011000300011PergentinoHESRibeiroLB (2017) Anurophagy by the snake Thamnodynastesphoenix (Squamata: Dipsadidae: Tachymenini) in dry forested areas of Northeastern Brazil.10: 597–600.PetzoldHG (1969) Observations on the reproductive biology of the American ringed snake Leptodeiraannulata at East Berlin Zoo.9: 54–56. https://doi.org/10.1111/j.1748-1090.1969.tb02613.xPintoRRFernandesROtavioAVM (2008) Morphology and diet of two sympatric colubrid snakes, Chironiusflavolineatus and Chironiusquadricarinatus (Serpentes: Colubridae).29: 149–160. https://doi.org/10.1163/156853808784125027PiresMGSilva JRNJFeitosaDTPrudenteALCFilhoGAPZaherH (2014) A new species of triadal coral snake of the genus Micrurus Wagler, 1824 (Serpentes: Elapidae) from northeastern Brazil.3811: 569–584. https://doi.org/10.11646/zootaxa.3811.4.8PizzattoLMarquesOAV (2002) Reproductive biology of the false coral snake Oxyrhopusguibei (Colubridae) from southeastern Brazil.23: 495–504. https://doi.org/10.1163/15685380260462392PizzattoLMarquesOAV (2006) Interpopulational variation in sexual dimorphism, reproductive output, and parasitism of Liophismiliaris (Colubridae) in the Atlantic forest of Brazil.27: 37–46. https://doi.org/10.1163/156853806776052128PizzattoLMarquesOAV (2007) Reproductive Ecology of Boine Snakes with Emphasis on Brazilian Species and a Comparison to Pythons. South American Journal of Herpetology 2: 107–122. https://doi.org/10.2994/1808-9798(2007)2[107:REOBSW]2.0.CO;2PizzattoLMarquesOAVFacureK (2010) Food habits of Brazilian boid snakes: Overview and new data, with special reference to Corallushortulanus.30: 533–544. https://doi.org/10.1163/156853809789647121PizzattoLCantorMOliveiraJLMarquesOAVCapovillaVMartinsM (2008) Reproductive Ecology of Dipsadine Snakes, with emphasis on South American species.64: 168–179. https://doi.org/10.1655/07-031.1PranceGT (1982) Columbia University Press, New York, 714 pp.PrietoYAGiraudoARLópezMS (2012) Diet and Sexual Dimorphism of Liophispoecilogyrus (Serpentes, Dipsadidae) from the Wetland Regions of Northeast Argentina.46: 402–406. https://doi.org/10.1670/10-228ProtázioASProtázioASConceiçãoLCRibeiroACCruzSJ (2017) Thamnodynastespallidus (Serpentes: Dipsadidae) predation on Boanasemilineata (Anura: Hylidae) in fragment of Atlantic Forest, northeastern Brazil.10: 521–523.Prudente AL daCMenksACSilvaFMMaschioGF (2014) Diet and reproduction of the western indigo snake Drymarchoncorais (serpentes: Colubridae) from the Brazilian Amazon.7: 99–108.RantaPBlomTNiemeläJJoensuuESiitonenM (1998) The fragmented Atlantic rain forest of Brazil: Size, shape and distribution of forest fragments.7: 385–403. https://doi.org/10.1023/A:1008885813543ReisPMAGCoehloRDFMenezesLMNRibeiroLB (2015) Contribution to the reproductive biology of Bothropserythromelas (Squamata: Viperidae) in the semiarid region of Brazil.46: 327–331.RibeiroMCMetzgerJPMartensenACPonzoniFJHirotaMM (2009) The Brazilian Atlantic Forest: How much is left, and how is the remaining forest distributed? Implications for conservation.142: 1141–1153. https://doi.org/10.1016/j.biocon.2009.02.021RobertoIJÁvilaRWMelgarejoAR (2015) Répteis (Testudines, Squamata, Crocodylia) da Reserva Biológica de Pedra Talhada. In: StuderANusbaumerLSpichigerR (Eds) Biodiversidade da Reserva Biológica de Pedra Talhada (Alagoas, Pernambuco – Brasil).68: 357–375.RobertoIJOliveiraCRAraujo FilhoJAOliveiraHFÁvilaRW (2012) The herpetofauna of the Serra do Urubu mountain range: a key biodiversity area for conservation in The brazilian atlantic forest.57: 347–373. https://doi.org/10.11606/0031-1049.2017.57.27RochaLAAleixoAAllenGAlmedaFBaldwinCCBarclayMVBerumenML (2014) Specimen collection: An essential tool.344: 814–815. https://doi.org/10.1126/science.344.6186.814RodriguesGMMaschioGFPrudenteALC (2016) Snake assemblages of Marajó Island, Pará state, Brazil.33: 1–13. https://doi.org/10.1590/S1984-4689zool-20150020RodriguesJBGamaSCAPereira FilhoGAFrançaFGR (2015) Composition and ecological aspects of a snake assemblage on the savanna enclave of the Atlantic Forest of the Guaribas Biological Reserve in Northeastern Brazil.10: 143–156. https://doi.org/10.2994/SAJH-D-15-00016.1RufinoNBernardiJAR (1999) Natural History Notes. Pseustessulphureussulphureus. Diet.30: 103–104.SampaioILRSantosCPFrançaRCPedrosaIMMCSoléMFrançaFGR (2018) Ecological diversity of a snake assemblage from the Atlantic Forest at the south coast of Paraíba, northeast Brazil.2018: 107–125. https://doi.org/10.3897/zookeys.787.26946Santos-SilvaCRAndradeISAraújoMLNBarrosLCSGomesLFerrariSF (2014) Predation of six anuran species by the banded cat-eyed snake, Leptodeiraannulata (Serpentes: Dipsadidae), in the Caatinga scrub of northeastern Bahia, Brazil.7: 123–126.ScartozzoniRRAlmeida-SantosSM (2006) Helicopsleopardinus (Water snake): Reproduction. Herpetological Bulletin: 30–40.SeigelRA (1993) Summary: future research on snakes, or how to combat” lizard envy.”. In: SeigelRACollinsJT (Eds) Snakes: Ecology and Behavior., 395–402.SilvaJMCCasteletiCHM (2003) Status of the biodiversity of the Atlantic Forest of Brazil. In: Galindo-Leal C, Câmara IG (Eds) The Atlantic Forest of South America: Biodiversity Status, Threats and Outlook.Silva JrNJ (2016) Editora da PUC, Goiânia, 417 pp.SilvaMVSouzaMBBernardePS (2010) Riqueza e dieta de serpentes no Estado do Acre, Brasil.12: 165–176.SousaBMCruzCAG (2000) Echinantheraaffinis (NCN). Diet.31: 178–178.SousaKRMPrudenteALCMaschioGF (2014) Reproduction and diet of Imantodescenchoa (Dipsadidae: Dipsadinae) from the Brazilian Amazon.31: 8–19. https://doi.org/10.1590/S1984-46702014000100002SouzaRCG (2007) Reproduction of the Atlantic Bushmaster (Lachesismutarhombeata) for the first time in captivity.42: 41–43.StrussmannCSazimaI (1993) The snake assemblage of the Pantanal at Poconé, Western Brazil: Faunal composition and ecological summary.28: 157–168. https://doi.org/10.1080/01650529309360900TabarelliMMarinsJFSilvaJMC (2002) La biodiversidad brasileña, amenazada.308: 42–49.TabarelliMSiqueira FilhoJASantosAMM (2006a) A Floresta Atlântica ao Norte do Rio São Francisco. In: PôrtoCK (Ed.) Diversidade Biológica e Conservação da Floresta Atlântica ao Norte do Rio São Francisco., 25–40.TabarelliMMeloMDLiraOC (2006b) Nordeste; Piauí; Ceará; Rio Grande do Norte; Paraíba; Pernambuco e Alagoas: O Pacto Murici. In: CampaniliMProchnowM (Eds) Mata Atlântica: uma rede pela floresta., 149–164.TabarelliMPintoLPSilvaJMCHirotaMMBedêLC (2005) Desafios e oportunidades para a conservação da biodiversidade na Mata Atlântica brasileira. Megadiversidade, 132–138.TurciLCBAlbuquerqueSBernardePSMirandaDB (2009) Uso do hábitat, atividade e comportamento de Bothriopsisbilineatus e de Bothropsatrox (Serpentes: Viperidae) na floresta do Rio Moa, Acre, Brasil.9: 197–206. https://doi.org/10.1590/S1676-06032009000300020Uchoa NetoCAMTabarelliM (2002) Diagnóstico e estratégia de conservação do Centro de Endemismo Pernambuco. Centro de Pesquisas Ambientais do Nordeste – CEPAN: 1–69.VittLJVangilderLD (1983) Ecology of a Snake Community in Northeastern Brazil.4: 273–296. https://doi.org/10.1163/156853883X00148VogelZ (1958) Surucucu do Pantanal.11: 178–181.ZacariottiRLGomesCA (2010) Diet of the black-headed forest racer Taeniophallusaffinis Günther, 1858 in the Brazilian Atlantic forest.3: 11–12.