Herpetology Notes, volume 12: 591-602 (2019) (published online on 11 June 2019)

Herpetofauna of three “Brejos de Altitude” in the interior of the state of Pernambuco, northeastern Brazil

Marco Antonio de Freitas1,*, Arthur Diesel Abegg2,5, Daniel da Silva Araújo3, Hugo Estevam de Almeida Coelho4, Weverton dos Santos Azevedo5, Márcio Frazão Chaves6, Conrado Mario da Rosa7, and Geraldo Jorge Barbosa de Moura8

Abstract. “Brejos de altitude” are ecoregions found at high altitudes, generally over 600 meters above sea level, located in both mountainous and plateau regions distributed throughout the Brazilian Northeast. These relict altitudinal forests display unique biotic evolutionary characteristics mainly due to the geological time they have spent in isolation, serving as “islands” for biological specialization, with many registered endemic species (six species of amphibians and 22 reptiles of the northeast region). Studies providing basic herpetofauna data are important to understand the structure and composition of these communities. In this context, the present study aimed to compile an inventory of the herpetofauna of three brejos de altitude in the state of Pernambuco, Northeastern Brazil. A total of 82 species were recorded in all three study areas, 30 amphibians (two orders, eight families and 15 genera) and 52 reptiles (two orders, 19 families and 42 genera). Four species of lizards found in the present study are endangered, and one tortoise and one snake species are data deficient (DD).

Keywords. Altitudinal forests, Caatinga, Inventory

Introduction

South America presents a wide variety of topographies 1 Instituto Chico Mendes de Conservação da Biodiversidade and climates, resulting in a remarkably rich and diverse (ICMBio), ESEC Murici, Rua Marino Vieira de Araújo 32, herpetofauna (Duellman, 1979). However, many species Cidade Alta, CEP 57820-000, Murici, AL, Brazil. may disappear before being catalogued by researchers 2 Universidade de São Paulo (USP), Instituto de Biociências (Dirzo and Raven, 2003). Inventories rank among the (IB-USP), Rua do Matão, 14, São Paulo, 05508-090, São main stages in the process required to describe, map Paulo, Brazil. 3 SOWITEC do Brasil Energias Alternativas Ltda. Avenida Luís and measure biodiversity for conservation planning Viana, nº 6.462 Edifício Wall Street Empresarial Torre East, actions (Margules and Pressey, 2000), and are important Salas 1.507 a 1.523, Paralela CEP: 41730-101, Salvador BA, for understanding richness and endemism and in the Brazil. detection of the presence of threatened species and their 4 Rua Prof. Carlos Ott, nº 79, Alameda Shopping, Sala 13, Stella requirements regarding habitat structure and quality. Maris, CEP40.000-000, Salvador, BA, Brazil. 5 According to Leivas et al. (2015), local species Instituto Butantan, Laboratório Especial de Coleções inventories are one of the main factors responsible for Zoológicas, Avenida Vital Brasil, 1.500, Butantã, São Paulo – SP, 05503-900, Brazil. the increasing number of described species. The same 6 Centro de Educação e Saúde, unidade acadêmica de Biologia authors also state that these types of study are particularly e Química, Universidade Federal de Campina Grande. Olho essential in high diversity countries. Consequently, D´água da Bica, CEP 58175-000, Cuité, PB, Brazil. countries with high diversity rates and low knowledge 7 Universidade Federal de Santa Maria, Programa de Pós- on this diversity should invest urgent efforts in order to Graduação em Biodiversidade Animal, CCNE, Laboratório generate biological knowledge (Marques and Lamas, de Herpetologia, Av. Roraima 1000, 97105-00, Santa Maria, 2006), as these data are important for species and habitat Rio Grande do Sul, Brazil. 8 Universidade Federal Rural de Pernambuco, Laboratório de conservation. Estudos Herpetológicos e Paleoherpetológicos, Rua Dom The Brazilian Northeast is still poorly understood, Manoel de Medeiros, s/n, Dois Irmãos, CEP 52171-900, with few published inventories in specific areas, notably Recife, PE, Brazil. in altitudinal forests. Only a total of 43 humid enclaves * Corresponding author. E-mail: [email protected] 592 Marco Antonio de Freitas et al.

and Santos, 2004). Thus, these regions are climatically isolated from the rest of the Caatinga located in the northeastern semi-arid Brazilian regions (Tabarelli and Santos, 2004). These relict altitudinal forests display unique evolutionary biota characteristics, mainly due to the geological time they have spent in isolation, serving as “islands” for biological specialization, with many registered endemic species (Vanzolini, 1981; Borges- Nojosa and Caramaschi, 2003; Rodrigues 2003; Freitas and Silva 2007; Pereira-Filho and Montingelli, 2011; Freitas, 2015). Aiming to promote a refinement of the brejos de altitude herpetofauna distribution and richness records Figure 1. Map of Northeastern Brazil, indicating the areas in the state of Pernambuco, Brazil, the results of the assessed in the present study. Areas in brown represent inventories carried out in three different localities are altitudes from 500 meters a.s.l. presented here.

Material and methods comprising altitudinal forests are known in the states of Three areas were inventoried at different times (Figure Pernambuco, Paraíba, Rio Grande do Norte and Ceará 1), applying a variety of sampling and methodologies, (Pereira-Filho and Montingelli, 2011). In Pernambuco, including the use or non-use of traps. In addition, quick these altitudinal forests, or “brejos de altitude”, are inventories obtained from consulting activities regarding considered disjunctive enclaves of tropical evergreen wind farm licensing in the Sertânia and Arcoverde forests inserted in the Caatinga area (Andrade-Lima, regions were also used. The taxonomic nomenclature 2007). applied herein follows Costa and Bérnils (2018) for Brejos de altitude are found at high altitudes, generally reptiles and Segalla et al. (2016) for amphibians, with over 600 meters above sea level, in both mountainous modifications made by Dubois (2017). and plateau regions distributed throughout the Brazilian Belo Jardim (-8.3365S, -36.4235W) (Fig. 2A): The Northeast (Tabarelli and Santos, 2004; Guedes et al., Fazenda Vale do Tabocas is a seasonal altitudinal forest 2014). These areas exert a physical barrier on the humid fragment comprising almost 250 hectares, between 600 winds and lower clouds from the Atlantic Ocean, which and 1100 meters in elevation. The area is characterized condense and provide orographic rainfall. These rains as in early and medium regeneration stages. Field play a significant role in the formation of highly humid sampling was carried out in May 2016 and was vegetation and results in different degrees of seasonality concentrated between 900 and 1050 meters in altitude. throughout the year throughout this biome (Tabarelli Three lines of pitfall traps with seven 40-liter buckets

Figure 2. Brejos de altitude areas sampled in the present study. (A) Belo Jardim, (B) Sertânia and (C) Arcoverde. Photographs by Marco Antonio de Freitas. Herpetofauna of three “Brejos de Altitude” in northeastern Brazil 593 were installed. The buckets were connected by a five- the participation of five people, obtaining both visual meter long plastic canvas, resulting in three lines of a and auditory records. In addition, occasional records forty-meter linear barrier. The lines were installed as were also considered. The collected amphibians were follows: one on the forest edge, one in an open canopy euthanized by applying lidocaine ointment (lidocaine) forest area and one in the interior of the fragment, on the ventral region and the reptiles through overdosing totalling 1512 hours-buckets sampling effort, over five with ether (Callefo, 2002; Franco and Salomão, 2002). trapping days. Additionally, daily active searches were All specimens were fixed with 10% formalin and then performed in the area, beginning at 6 pm and ending preserved in 70% ethanol. The collected material was at 10 pm, totalling 400 hours of sampling effort, with deposited at the Laboratory of Herpetological and

Figure 3. (A) Adenomera sp.; (B) Boana crepitans; (C) Boana faber; (D) Dendropsophus branneri; (E) Dendropsophus decipiens; (F) Dendropsophus oliveirai; (G) Dendropsophus sp.; (H) Dermatonotus muelleri; (I) Leptodactylus macrosternum; (J) Leptodactylus troglodytes; (K) Odontophrynus carvalhoi; (L) Trachycephalus atlas. All photos were taken in Belo Jardim. Photographs by Marco Antonio de Freitas. 594 Marco Antonio de Freitas et al.

Paleoherpetological Studies (LEHP) belonging to the effort, with the participation of three people, obtaining Federal Rural University of Pernambuco), in Recife both visual and auditory records. No traps were laid. (Appendix 1), under collection license SISBIO 53914- No specimens were collected in these two locations, 1. as these they were quick environmental consulting Sertânia (-8.0756S, -37.2693W) and Arcoverde inventories, and no collection license was issued. Thus, (-8.4184S, -37.0585W) (Fig. 2B-C): These two only photographic records were obtained. municipalities are located on the Borborema Plateau. The sampled areas are characterized by mountain ranges with Results and Discussion altitudes that surpass 800 meters in altitude, containing arboreal Caatinga vegetation in its lower portions and Species composition - A total of 82 species were deciduous forests in its higher portions. Two campaigns recorded at the three brejos de altitude in the state of comprising 10 days each were carried out, one in May Pernambuco, 30 amphibians (two orders, 8 families and of 2013 and one in October of 2014, with three people 14 genera) (Figs. 3-4, Table 1) and 52 reptiles (2 orders, in the field. The active search methodology was applied 19 families and 42 genera) (Figs. 5-7, Table 2). The area to observe amphibians and reptiles. Additionally, daily displaying the greatest overall richness was Sertânia active searches were performed in the area, beginning (total: 66; amphibians: 20; reptiles: 46) followed by at 6 pm and ending at 10 pm, and beginning at 7 am Belo Jardim (total: 61; amphibians: 25; reptiles: 36) and and ending at 11 am totalling 480 hours of sampling Arcoverde (total: 57; amphibians: 16; reptiles: 41).

Figure 4. (A) Physalaemus cuvieri; (B) Pithecopus nordestinus; (C) Pleurodema diplolister; (D) Pristimantis ramagii; (E) Proceratophrys renalis; (F) Scinax eurydice; (G) Scinax pachycrus; (H) Rhinella jimi; (I) Scinax x-signatus. All photos were taken in Belo Jardim. Photographs by Marco Antonio de Freitas and Daniel Araújo. Herpetofauna of three “Brejos de Altitude” in northeastern Brazil 595

Table 1. Amphibian species recorded in the brejos de altitude in interior Pernambuco, Brazil. *Species endemic to the northeast region. Table 1. Amphibian species recorded in the brejos de altitude in interior Pernambuco, Brazil. *Species endemic to the northeast region.

ORDER/Family/Species (Author, year) Belo Jardim Sertânia Arcoverde ANURA

Bufonidae

  Rhinella crucifer (Wied-Neuwied, 1821) Rhinella jimi (Stevaux, 2002)    Rhinella granulosa (Spix, 1824)    Craugastoridae

Pristimantis ramagii (Boulenger, 1888) 

Hylidae

Dendropsophus branneri (Cochran, 1948)    Dendropsophus decipiens (Lutz, 1925) 

Dendropsophus oliveirai (Bokermann, 1963)*    Dendropsophus minutus (Peters, 1872) 

Dendropsophus sp. 

Boana crepitans (Wied-Neuwied, 1824)      Boana faber (Wied-Neuwied, 1821) Boana albomarginata (Spix, 1824) 

Boana raniceps Cope, 1862 

Scinax eurydice (Bokermann, 1968) 

Scinax pachycrus (Miranda-Ribeiro, 1937)*    Scinax x-signatus (Spix, 1824)    Phyllomedusidae

Pithecopus nordestinus (Caramaschi, 2006)*    Leptodactylidae

Leptodactylus vastus (A. Lutz, 1930)    Leptodactylus mystacinus (Burmeister, 1861) 

Leptodactylus macrosternum Miranda-Ribeiro, 1926*    Leptodactylus troglodytes A. Lutz, 1926*    Adenomera sp. 

Physalaemus kroyeri (Reinhardt and Lütken, 1862 "1861")    Physalaemus cuvieri Fitzinger, 1826    Pleurodema diplolister (Peters, 1870)    Microhylidae

Dermatonotus muelleri (Boettger, 1885) 

Odontophrynidae

Proceratophrys cristiceps (Müller, 1884 “1883”)*  

Proceratophrys renalis (Miranda-Ribeiro, 1920) 

Odontophrynus carvalhoi Savage and Cei, 1965 

GYMNOPHIONA

Siphonopidae

Siphonops paulensis Boettger, 1892  596 Marco Antonio de Freitas et al.

Three reptiles, Psychosaura agmosticha, Stenolepis and two Crocodylia are recorded, totalling 126 species ridleyi, Thamnodynastes almae, and three amphibians, (SEMAS, 2017). Of these, 52 species were found in Scinax eurydice, Boana faber and Dendropsophus the present study, equivalent to 41.2% of the state’s decipiens are new records for the area in the state of total richness. It should be noted, however, that some Pernambuco (Freitas, 2015). reptile species were not included in this list until Brazil is the most species-rich country in the world now (SEMAS, 2017), such as Phrynops tuberosus concerning amphibians (Segalla et al., 2016), and yet (Peters,1870), Amphisbaena lumbricalis Vanzolini, the true number of species is still underestimated. 1996, Hemidactylus mabouia (Moreau de Jonnès, 1818) These animals are considered by some researchers as , Ophiodes sp. and Thamnodynastes phoenix Franco, the most threatened group of vertebrates in the world, Trevine, Montingelli and Zaher, 2017, which were due to habitat loss and fragmentation (Gibbons et observed in the present study. al. 2000, Becker et al. 2007, Loyola et al. 2007), and When comparing the total richness of the studied disease (Lips, 2016). Two Gymnophiona species and 85 community (82 spp.), with herpetofauna surveys carried anuran species are recorded for the state of Pernambuco out in different dry regions of the Caatinga biome, a (SEMAS, 2015). The 30 species found in the three relatively higher number of species is noted (Table brejos de altitude represent approximately 34% of the 3). This difference may be associated to the applied total species recorded for this state. collection methodology, total sampling effort and due to In relation to the reptile fauna of Pernambuco, eight historical local flora and fauna formation processes. The Testudines, 41 lizards, eight Amphisbaenia, 67 snakes latter may be particularly important, as the process of

Figure 5. (A) Ameivulla ocellifera; (B) Brasiliscincus heathi; (C) Chelonoides carbonarius; (D) Diploglossus lessonae; (E) Enyalius bibronii; (F) Gymnodactylus geckoides; (G) Hemydactylus brasilianus; (H) Iguana iguana; (I) Kinosternon scorpioides. Photos were taken in Belo Jardim, Sertânia and Arcoverde. Photographs by Marco Antonio de Freitas and Daniel Araújo. Herpetofauna of three “Brejos de Altitude” in northeastern Brazil 597 altitude heath formation resulted in humid forest remnants coloration pattern variations and populations from high isolated after the South America forest expansion during north-eastern areas have not yet been satisfactorily the Quaternary. Although it is located in the Caatinga described taxonomically (WSA, pers. obs). It is thus biome, it does not share the same fauna and flora as of paramount importance that future revision studies these surrounding, dry environments, emphasized by concerning this genus include brejos de altitude samples the presence of endemics, predominantly forest species, (Pereira-Filho and Montingelli, 2011; Marques, et al., including six species of amphibians and 22 reptiles of 2017). In addition, it was not possible to unequivocally the northeast region (Table 1, 2). (Vanzolini, 1981; determine species-level classification for two amphibian Borges-Nojosa and Caramaschi, 2003; Freitas, 2015; taxa (Adenomera sp. and Dendropsophus sp.) based Borges-Nojosa et al., 2016). on external morphology alone, which may constitute Since the brejos de altitude constitutes relict forest new species and deserve further attention. Finally, one formations in semi-arid landscapes, they may play reptile (Ophiodes sp.) is known to be a new species, a role as potential “continental islands”, promoting occurring along forested areas in both southeastern and speciation through isolation (Borges-Nojosa et al., northeastern Brazil, previously confused with Ophiodes 2016). As a result, remarkable interpopulational striatus (Borges-Martins, 1998). variations (morphological, ecological and/or genetic), According to the recent assessment regarding the risk or even new species, may evolve in these areas. In this of fauna extinction in the state of Pernambuco (SEMAS, context, the Tantilla genus in Brazil presents significant 2017), three lizard species reported herein are threatened

Figure 6. (A) Ligodactylus klugei; (B) Mesoclemmys tuberculata; (C) Phyllopezus periosus; (D) Phyllopezus pollicaris; (E) Psychosaura agmosticha; (F) Vanzosaura multiscutata; (G) Tropidurus semitaeniatus; (H) Stenolepys rydley; (I) Tropidurus hispidus. Photos were taken in Belo Jardim, Sertânia and Arcoverde. Photographs by Marco Antonio de Freitas and Daniel Araújo. 598 Marco Antonio de Freitas et al.

Table 2. Reptile species recorded in the brejos de altitude in interior Pernambuco, Brazil. *Species endemic to the northeast region. Table 2. Reptile species recorded in the brejos de altitude in interior Pernambuco, Brazil. * Species endemic to the northeast region.

ORDER/Family/Species (Author, year) Belo Jardim Sertânia Arcoverde TESTUDINES

Chelidae

Mesoclemmys tuberculata (Lüderwaldt, 1926)*    Phrynops tuberosus (Peters, 1870)    Testudinidae

Chelonoidis carbonarius (Spix, 1824)  

Kinosternidae

Kinosternon scorpioides (Linnaeus, 1766)    SQUAMATA

Amphisbaenidae

Amphisbaena alba Linnaeus, 1758    Amphisbaena lumbricalis Vanzolini, 1996* 

Tropiduridae

Tropidurus hispidus (Spix, 1825)    Tropidurus semitaeniatus (Spix, 1825)*    Gekkonidae

Hemidactylus mabouia (Moreau de Jonnès, 1818)    Hemidactylus brasilianus (Amaral, 1935)*    Lygodactylus klugei (Smith, Martin and Swain, 1977)*  

Teiidae

Salvator merianae Duméril and Bibron, 1839    Ameiva ameiva (Linnaeus, 1758)    Ameivula ocellifera (Spix, 1825)*    Phyllodactylidae

Phyllopezus pollicaris (Spix, 1825)    Phyllopezus periosus Rodrigues, 1986* 

Gymnodactylus geckoides Spix, 1825*    Enyalius bibronii Boulenger, 1885*  

Polychrotidae

Polychrus acutirostris Spix, 1825  

Iguanidae

Iguana iguana (Linnaeus, 1758)    Diploglossidae

Ophiodes sp. 

Diploglossus lessonae Peracca, 1890* 

Gymnophthalmidae

  Cercosaura ocellata Wagler, 1830 Stenolepis ridleyi Boulenger, 1887* 

Micrablepharus maximiliani (Reinhardt and Luetken, 1862)  

Vanzosaura multiscutata (Amaral, 1933)*   

Herpetofauna of three “Brejos de Altitude” in northeastern Brazil 599

Table 2. Continued.

ORDER/Family/Species (Author, year) Belo Jardim Sertânia Arcoverde Mabuyidae

Brasiliscincus heathi (Schmidt and Inger, 1951)*    Psychosaura agmosticha (Rodrigues, 2000)*  

Leptotyphlopidae

Epictia borapeliotes (Vanzolini, 1996)*    Boidae

Boa constrictor Linnaeus, 1758    Epicrates assisi Machado, 1945*    Colubridae

Chironius flavolineatus (Jan, 1863)    Oxybelis aeneus (Wagler in Spix, 1824)    Spilotes pullatus (Linnaeus, 1758)    Tantilla cf. melanocephala (Linnaeus, 1758)  

Dipsadidae

Apostolepis cearensis Gomes, 1915* 

Pseudoboa nigra (Duméril, Bibron and Duméril, 1854) 

Boiruna sertaneja Zaher, 1996* 

Leptodeira annulata (Linnaeus, 1758) 

Erythrolamprus poecilogyrus (Wied, 1824)    Erythrolamprus viridis (Günther, 1862)    Oxyrhopus trigeminus Duméril, Bibron and Duméril, 1854    Thamnodynastes almae Franco and Ferreira, 2003*  

Thamnodynastes phoenix Franco, Trevine, Montingelli and    Zaher, 2017* Thamnodynastes sertanejo Bailey, Thomas and Silva-Jr,   2005* Philodryas nattereri Steindachner, 1870    Philodryas olfersii (Liechtenstein, 1823)    Xenodon merremii (Wagler in Spix, 1824) 

Elapidae

Micrurus ibiboboca (Merrem, 1820)    Viperidae

Bothrops leucurus Wagler in Spix, 1824 

Bothrops erythromelas Amaral, 1923*  

Crotalus durissus Linnaeus, 1758   

with extinction, while Cercosaura ocellata, observed in deficient (DD): the tortoise Chelonoidis carbonarius Belo Jardim and Sertânia swamps, and Stenolepis ridleyi and the snake Micrurus ibiboboca (SEMAS, 2017). observed in Belo Jardim, are classified as vulnerable These records are particularly noteworthy and deserve (VU). In addition, two other species are listed as special attention, as the areas sampled in the present 600 Marco Antonio de Freitas et al.

Figure 7. (A) Bothrops erythromelas; (B) Bothrops leucurus; (C) Crotalus durissus; (D) Micrurus ibiboboca; (E) Oxyrhopus trigeminus; (F) Tantilla cf. melanocephala; (G) Thamnodynastes almae; (H) Thamnodynastes phoenix; (I) Thamnodynastes sertanejo. Photos were taken in Belo Jardim, Sertânia and Arcoverde. Photographs by Marco Antonio de Freitas and Daniel Araújo.

Table 3. Total richness of amphibians and reptiles found in different studies in the Caatinga biome, compared to the richness found in the brejos de altitudeTable 3.in Totalthe interior richness of of Pernambuco, amphibians andBrazil. reptiles found in different studies in the Caatinga biome, compared to the richness found in the brejos de altitude in the interior of Pernambuco, Brazil.

Source Vegetation State Municipality/ Amphibians Reptiles Total Locality This study Brejo de altitude Pernambuco Sertânia 20 46 66 This study Brejo de altitude Pernambuco Belo Jardim 25 36 61 Pedrosa et al., (2014) Arboreous Caatinga Pernambuco Parque Nacional do 21 37 58 Catimbau This study Brejo de altitude Pernambuco Arcoverde 16 41 57 Pereira et al., (2015) Caatinga seca Pernambuco Serrita 23 31 54 Garda et al., (2013) Caatinga baixa Bahia Raso da Catarina 21 32 53 Ecological Station Arzabe et al., (2005) Arboreous Caatinga/ Paraíba Curibataú 21 31 52 Mata seca Caldas et al., (2016) Caatinga aberta Rio Grande do Seridó Ecological 19 22 41 Norte Station Cavalcanti et al., (2014) Arboreous Caatinga Piauí Serra da Capivara 8 29 37 National Park

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Segalla, M.V., Caramaschi, U., Cruz, C.G., Garcia, P.C.A., Grant, Appendix. Voucher list. T., Haddad, C.F.B., Garcia, P.C.A., Berneck, B.V.M., Langone, J.A. (2016): Brazilian amphibians: List of species. Herpetologia Siphonops annulatus (CHPUFRPE 4535); Brasileira 5 (2): 34–46. Odontophrynys carvalhoi (CHPUFRPE 4975); Rhinella SEMAS, Secretaria de meio ambiente e sustentabilidade (2015): granulosa (CHPUFRPE 4976); Pithecopus nordestinus Resolução SEMAS n° 1 de 09 de janeiro de 2015. Lista Estadual (CHPUFRPE 4542); Boana faber (CHPUFRPE 4977); Oficial de Espécies da Fauna Ameaçadas de Extinção – Anfíbios. Scinax eurydice (CHPUFRPE 4978); Scinax pachychrus Pernambuco, SEMAS. (CHPUFRPE 4979-80); Scinax x-signatus (CHUNB SEMAS, Secretaria de meio ambiente e sustentabilidade (2017): Resolução SEMAS n° 1 de 15 de maio de 2017. Lista Estadual 58517-18); Physalaemus cuvieri (CHPUFRPE Oficial de Espécies da Fauna Ameaçadas de Extinção – Répteis. 4981-82); Rhinella jimi (4983-85); Leptodactylus Pernambuco, SEMAS. troglodytes (CHPUFRPE 4986); Adenomera sp. Tabarelli, M., Santos, A.M.M. (2004): Uma breve descrição sobre (13498-99-13500); Leptodactylus vastus (CHUNB a história natural dos Brejos Nordestinos. Brejos de Altitude em 14400); Pseudopaludicola sp. (CHUNB 58513-16); Pernambuco e Paraíba, História Natural, Ecologia e Conservação Rhinella crucifer (CHPUFRPE 4987); Dendropsophus (Porto, K.C., Cabral, J.J.P., Tabarelli, M., eds.). Ministério do oliveirai (CHPUFRPE 4988-90); Dendropsophus Meio Ambiente 9: 17–24. Vanzolini, P.E. (1981): A quase-historical approach to the natural sp. (CHPUFRPE 4991-92); Pristimantis ramagii history of differentiation of reptiles in the tropical geographic (CHPUFRPE 4993), (CHUNB 58960); Proceratophrys isolated. Papéis Avulsos de Zoologia 34 (19): 189–204. renalis (CHPUFRPE 4996-4614); Bothrops leucurus (4533); Crotalus durissus (CHPUFRPE 4934); Philodryas nattereri (CHPUFRPE 4536); Oxyrhopus trigeminus (CHPUFRPE 4537); Thamnodynastes phoenix (CHUNB13765); Brasiliscincus heathi (CHUNB 09297); Hemidactylus brasiliana (CHUNB 11160-61-13491-96); Phyllopezus pollicaris (CHUNB 11162-67-13497); Gymnodactylus geckoides (CHUNB 11190-98-13566-80); Tropidurus hispidus (CHUNB 13464-67-13490-13539-60); Tropidurus semitaeniatus (CHUNB 04330-39); Ameiva ameiva (CHUNB 13589); Stenolepis ridleyi (CHPUFRPE 4994-95).

Accepted by Christoph Liedtke