Zootaxa 3920 (3): 443–452 www.mapress.com /zootaxa / Copyright © 2015 Magnolia Press
Article
ISSN 1175-5326 (print edition)
ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3920.3.3 http://zoobank.org/urn:lsid:zoobank.org:pub:143FCD6B-F614-4422-832C-190114C28FEB
A new species of Amerotyphlops from Northeastern Brazil, with comments on distribution of related species ROBERTA GRABOSKI1, 2, 5, GENTIL ALVES PEREIRA FILHO1, ARIANE AUXILIADORA ARAÚJO DA SILVA3, 4, ANA LÚCIA DA COSTA PRUDENTE4 & HUSSAM ZAHER1 1
Museu de Zoologia da Universidade de São Paulo, CP 42494, São Paulo, 04218-070 São Paulo, Brasil Programa de Pós Graduação em Zoologia, Universidade Estadual Paulista Júlio de Mesquita Filho, Av. 24 A, 1515, Bairro Bela Vista, Rio Claro, 13506-900, São Paulo, Brasil 3 Programa de Pós-Graduação em Zoologia, Universidade Federal do Pará, Museu Paraense Emílio Goeldi. Av. Perimetral, 1901,Terra Firme, CP 399, Belém, 66017-970 Pará, Brasil. 4 Laboratório de Herpetologia, Museu Paraense Emílio Goeldi, CP 399, Av. Perimetral, 1901, Terra Firme, Belém, 66077-530, Pará, Brasil 5 Corresponding author. E-mail: [email protected] 2
Abstract We describe a new species of Amerotyphlops from an upland forest enclave in the state of Paraíba, Northeastern Brazil. The new species is distinguished from the other seven South American species of Amerotyphlops by the combination of the following characters: nasal suture incomplete; rostral scale oval and yellowish cream with some dark brown spots; four supralabial scales; three infralabial scales; rows of scales around the body 18/18/18; middorsal scales from 204 to 225; dorsum with twelve to thirteen rows of scales dark brown and belly with four to five rows of scales immaculate yellowish cream; caudal spine dark brown; subcaudal scales 8–10 in female and 11–13 in males; maximum total length 233 mm. The new species is morphologically similar to A. amoipira and A. paucisquamus, sharing 18/18/18 rows of scales around the body and a small overlap of counts of middorsal scales. Key words: Atlantic rainforest, Typhlopidae, taxonomy, Amerotyphlops paucisquamus, Amerotyphlops amoipira
Introduction The genus Amerotyphlops was recently proposed by Hedges et al. (2014) for the clade of typhlopids that currently comprises 14 species found in the New World. Thirteen species are distributed on the mainland, from Mexico to northern Argentina, and one insular species occur in the West Indian (Hedges et al. 2014; Pyron & Wallach 2014). From the 13 mainland species, seven are restricted to South America: Amerotyphlops reticulatus (Linnaeus 1758); A. lehneri (Roux 1926), A. brongersmianus (Vanzolini 1972), A. minuisquamus (Dixon & Hendricks 1979), A. paucisquamus (Dixon & Hendricks 1979); A. yonenagae (Rodrigues, 1991), and A. amoipira (Rodrigues & Juncá 2002). Amerotyphlops lehneri occurs exclusively in the state of Falcon, northern Venezuela (Roux 1926; Shreve 1947; Dixon & Hendricks 1979; Rivas et al. 2012; Wallach et al. 2014), while A. yonenagae and A. amoipira are endemic to the predominantly psammophilous environment of dunes along the São Francisco River, both occurring in the state of Bahia, Northeastern Brazil (Rodrigues 1991; Rodrigues & Juncá 2002; Wallach et al. 2014). Amerotyphlops paucisquamus is endemic in the Atlantic forest of Northeastern Brazil, from state of Pernambuco and Paraíba (Dixon & Hendricks 1979; Rodrigues 1991; Freire 2001; Rodrigues & Juncá 2001; França et al. 2012; Wallach et al. 2014). The three remaining species are widely distributed in South America: A. minuisquamus is distributed throughout the Amazon basin, in Peru, western Brazil, Venezuela, Colombia and the Guyana Shield (Roze 1956; Dixon & Hendricks 1979; Caicedo-Portilla 2011; Wallach et al. 2014); A. reticulatus occurs in both cis- and trans-Andean lowlands of Colombia (Caicedo-Portilla 2011), and throughout the Amazon basin and
Accepted by P. Passos: 16 Jan. 2015; published: 23 Feb. 2015
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adjacent ecotonal regions southern of the Amazon forest, including gallery forests in the Cerrado biome (Cunha & Nascimento 1978, 1993; Dixon & Hendricks 1979; França et al. 2006; França & Venâncio 2010; Ávila & Kawashita-Ribeiro 2011; Wallach et al., 2014); A. brongersmianus is widely distributed along the open formations of South America (Cerrado, Chaco, and Caatinga) and savanna enclaves within the Amazon and Atlantic forests (Dixon & Hendrix 1979; Rodrigues 1991; Loebmann 2008; Santana et al. 2008; Martins et al. 2010; Arruda et al. 2011; Wallach et al., 2014). Here, we describe a new species of Amerotyphlops from an upland rainforest (locally known as “Brejo de Altitude”) in the state of Paraíba, Northeastern Brazil. This new species is morphologically similar to A. amoipira and A. paucisquamus.
Material and methods We examined 303 specimens from six South American species of Amerotyphlops (Appendix 1). Amerotyphlops lehneri was the only South American species for which no specimens were available for this study; we thus used the available information in the literature (Roux, 1926; Shreve,1947; Dixon & Hendricks, 1979). All specimens examined are deposited in the following institutions (acronyms in parentheses): Instituto Butantan, São Paulo, São Paulo (IBSP); Instituto Nacional de Pesquisas da Amazônia, Manaus, Amazonas (INPA); Museu de Ciências e Tecnologia da PUC-RS, Porto Alegre, Rio Grande do Sul (MCP-PUCRS); Museu de Zoologia da Universidade de São Paulo, São Paulo, São Paulo (MZUSP); Museu de Zoologia da Universidade Estadual de Campinas, Campinas, São Paulo (ZUEC-REP); Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Rio de Janeiro (MNRJ); Museu Paraense Emílio Goeldi, Belém, Pará (MPEG); Universidade Federal da Bahia, Salvador, Bahia (UFBA); Universidade Federal da Paraíba, João Pessoa, Paraíba (CHUFPB); Museu de Zoologia João Moojen, Universidade Federal de Viçosa, Viçosa, Minas Gerais (MZUFV); Museu de Zoologia Professor Mello Leitão, Santa Teresa, Espírito Santo (MBML); Universidade Federal do Mato Grosso, Cuiabá, Mato Grosso (UFMT); Laboratório de Anfíbios e Répteis, Universidade Federal do Rio Grande do Norte, Natal, Rio Grande do Norte (AAGARDA). We measured the total length (TTL) to the nearest 1 mm by stretching carefully the specimen along a graduated ruler. We measured the cephalic scales, midbody diameter (MBD), and tail length (TL) to the nearest 0.1 mm with a digital caliper. We followed Wallach (2003) for terminology on cephalic scales, and Thomas & Hedges (2007) and Dixon & Hendrix (1979) for measurements and scale counts, except for middorsal scales, which were counted along the paravertebral row and did not include the rostral scale and terminal spine. We counted the transversal scale rows in three different places along the body: 1) at the level of the 20th scale posterior to the symphysial, 2) at mid body, 3) at the level of the 10th scale anterior to the cloacal shield. We determined sex by making a small incision at the base of the tail to access the presence or absence of hemipenes. We generated a distribution map through the software ArcGIS v10.2.2 (ESRI 1999). Geographical coordinates were obtained from the Species Link 2013 online database, based on the institutions database, or directly taken with GPS devices at the localities. The software Google Earth (Google Inc. 2013) was used, whenever possible, to refine the coordinates. We performed an univariate analysis of variance (ANOVA) to assess the possible presence of secondary sexual dimorphism in several meristic variables (subcaudal, middorsal and ventral scales), and two continuous variables (TTL and TL). Assumptiom of univariate normality and homoscedasticity were performed with KolmogorovSmirnov’s and Levene’s tests, respectively. Characters that showed sexual dimorphism are shown separately for males and females. Only adult individuals, determined following Ávila et al. (2006), were employed in the analysis of sexual dimorphism to avoid the allometric effect by continuous variables. Only one juvenile (MZUSP 21274) from the 18 specimens examined (eight females and 10 males), was removed from the analysis.
Amerotyphlops arenensis sp. nov. Fig. 1 Holotype. An adult male, MZUSP 20042, collected by Gentil P. Filho on 19th November 2008 from the Reserva
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Ecológica Mata do Pau Ferro (06° 58'12'' S, 35° 42'15'' W; ca. 600 m), municipality of Areia, state of Paraíba, Brazil. Paratypes. Eighteen specimens all collected in the type locality by Gentil P. Filho. MZUSP 20037–38 collected in 16th October 2008, MZUSP 20039 collected in 17th October 2008, MZUSP 20040–41 collected in 19th November 2008, MZUSP 20136 collected in 19th November 2008, MZUSP 20043–45 collected in 17th December 2008, MZUSP 20046 collected in 18th December 2008, MZUSP 20047 and MZUSP 21273 collected in 21st January 2009, MZUSP 20048 collected in 28th February 2009, MZUSP 20049–50 and MZUSP 21274 collected in 27th March 2009, MZUSP 20051 collected in 23rd April 2009, and MZUSP 20052 collected in 21st May 2009. Diagnosis. Amerotyphlops arenensis is distinguished from all other South American species by the following combination of characters: nasal suture incomplete; rostral scale oval; supralabial scales four; infralabial scales three; rows scales around the body 18/18/18; middorsal scales 204 to 225; rows of dorsal scales dark brown 12–13th; rows of ventral scales yellowish cream and immaculate 4–5; caudal spine dark brown; subcaudal scales 8–10 in female and 11–13 for males; and maximum TTL 233 mm.
FIGURE 1. Lateral (A), ventral (B), and dorsal (C) views of the head of the holotype of Amerotyphlops arenensis sp. nov. (MZUSP 20042). Scale Bar = 1mm. NEW AMEROTYPHLOPS FROM NORTHEASTERN BRAZIL
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FIGURE 2. General view in life of the paratype (MZUSP 20038) of Amerotyphlops arenensis. Photographed by Gentil P. Filho.
The new species differs from Amerotyphlops lehneri by having an incomplete nasal suture (vs. complete nasal suture); from A. brongersmianus, A. reticulatus, and A. minuisquamus by having 18/18/18 rows scales around the body (vs. 18/16/14, 18/18/14, 20/18/14 or 20/18/15 in A. minuisquamus; 20/20/18 or 20/20/20 in A. brongersmianus and A. reticulatus); from A. yonenagae by having less than 250 middorsal scales (vs. more than 250 middorsal); from A. amoipira by having highly pigmented cephalic scales with a dark brown dorsum (vs. few pigmented cephalic scales, creamish brown dorsum with a fine darker brown paravertebral line concentrated in the anterior part of the body) and from A. paucisquamus by having a largest number of middorsal, between 204–225 (vs. fewer number of middorsal, between 162–209). Description of the holotype. Adult male, TTL 191 mm, TL 7 mm, TTL/MBD 28.08, and TTL/TL 27.28. Head slightly depressed dorsoventrally, not distinct from neck. Snout round in dorsal and ventral views. Rostral oval, longer than wide, narrow at anteroposterior region and wider at medial region; visible in dorsal view, extending ventrodorsally without reaching the imaginary transverse line between anterior borders of eyes. Rostral contacting nasal (anterior and posterior) dorsolaterally, and first supralabial and anterior nasal scales ventrally. Nasal suture incomplete, only partially dividing the anterior and posterior portions of nasal scale. Suture begins in the upper edge of second supralabial, pass through nostril, but fails to reach rostral. Anterior nasal in contact with first infralabial and upper edge of second infralabial. Posterior nasal longer than wide, contacting upper margin of second supralabial and preocular. Supralabials four, fourth twice longer than third supralabial. Infralabials three, third largest. Eye diameter 0.91 mm; eyes not visible in ventral view, located dorsolaterally, close to suture between preocular and ocular scales, completely covered by ocular scale. Ocular scales contacting frontal. Body cylindrical and robust. Midbody diameter 6.80 mm. Dorsal and ventral scales cycloid, wider than long, highly imbricated and arranged in diagonal series; scale rows around the body 18/18/18. Middorsal scales 213. Cloacal plate rounded, bordered anteriorly by three rows of scales and posteriorly by five rows of scales. Subcaudal scales 13, excluding the terminal spine. Terminal spine large, stout base and dark brown. Coloration of the holotype in life. Dorsum (12–13 rows of scales) dark brown, venter (4–5 rows of scales) light cream. Dorsal and ventral portions of snout pinkish, with a few dark brown spots, covering both rostral and nasal scales (only in the upper half of snout). Symphysial region pinkish and immaculate. Dorsal head scales (supraoculars, frontal, posfrontal, parietals, and occipitals) and dorsal portions of lateral head scales (ocular, nasal, and lower nasal) predominantly dark brown, and ventral portions pinkish. Cloacal plate light cream and terminal spine dark brown. After six years preserved in 70% ethanol, the coloration of the holotype remained mostly the same. All regions in light cream and pinkish changed to yellowish cream (Fig. 2). Variation. The ANOVA revealed secondary sexual dimorphism only for the number of subcaudal scales (F= 63.43, p < 0.001) and TL (F=47.61, p < 0.001) among the verified variables. Number of subcaudal scales 11–13 (mean = 11.8, SD= 0.63, N= 11) in males and 8–10 (mean = 9.2, SD= 0.70, N= 8) in females. Tail length 4.42–6.77 % of TTL (N= 10) in males and 3.71–4.91% of TTL (N= 8) in females. Largest female and male with 233 mm
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TTL and 5.23 mm of TL (MZUSP 20043) and 215 mm of TTL and 7.19 of TL (MZUSP 20044), respectively. MBD 5.41–8.04 (mean = 7.21, SD= 0.77, N=10) in males and 5.85–9 (mean = 8; SD = 1.12; N= 8) in females; number of middorsal scales 204–221 (mean = 214.1, SD= 4.70, N= 11) in males and 207–225 (mean = 217.25, SD= 7.10, N= 8) in females. The color patterns of the paratypes are similar to that found in the holotype. However, two individuals (11%) lack dark spots in their rostral scale while the remaining individuals (88%) retain dark spots in their rostral scale. Table 1 shows additional morphometric characters and scale patterns found in the new species and in two morphologically similar species distributed in the Northeastern Brazil (A. amoipira and A. paucisquamus). TABLE 1. Variation for some selected characters of morphologically similar species of Amerotyphlops from Northeast region of Brazil. The values displayed in the table represent (in this sequence) range, mean, standard deviations, and available sample size. Abbreviations are as following: ED = eye diameter, HR = head radius, HWE = head width, IN = internasal distance, INORB = interorbital distance, RL = rostral length, RW = rostral width, TL = tail length, TTL: total length. Character
A. arenensis sp. nov.
A. amoipira
A. paucisquasmus
ED
0.93 ± 0.09 (0.75–1.05) 10 0.90 ± 0.10 (0.77–1.10) 8
0.83 ± 0.09 (0.69–0.98) 8 0.80 ± 0.24 (0.62–0.97) 2
0.80 ± 0.15 (0.43–0.99) 12 0.85 ± 0.19 (0.52–1.29) 12
HR
3 ± 0.10 (2.78–3.13) 10 3.17 ± 0.21 (2.77–3.42) 8
2.52 ± 0.39 (1.95–3.04) 8 2.77 ± 0.68 (2.28–3.35) 2
2.62 ± 0.26 (2.05–2.93) 12 2.69 ± 0.26 (2.27–3.14) 12
HWE
5.15 ± 0.33 (4.43–5.47) 10 5.47 ± 0.42 (4.80–6.02) 8
3.97 ± 0.58 (3.25–5.13) 8 4.04 ± 0.60 (3.62–4.46) 2
4.11± 0.42 (3.17–4.64) 12 4.24 ± 0.42 (3.58–5.00) 12
IN
2.47 ± 0.14 (2.16–2.65) 10 2.58 ± 0.18 (2.35–2.80) 8
2.19 ± 0.30 (1.74–2.74) 8 2.14 ± 0.33 (1.91–2.37) 2
2.14 ± 0.25 (1.54–2.50) 12 2.18 ± 0.18 (1.86–2.61) 14
INORB
3.29 ± 0.13 (3.05–3.45) 10 3.41± 0.24 (3–3.86) 8
2.23 ± 0.25 (1.81–2.56) 8 2.23 ± 0.20 (2.09–2.36) 2
2.58 ± 0.26 (1.94–2.88) 12 2.48 ± 0.19 (2.09–2.76) 12
RL
2.84 ± 0.15 (2.59–3.09) 10 3.05 ± 0.22 (2.71–3.41) 8
2.19 ± 0.29 (1.82–2.75) 8 2.23 ± 0.50 (1.88–2.59) 2
2.40 ± 0.22 (1.90–2.76) 12 2.36 ± 0.22 (2.11–2.78) 12
RW
1.66 ± 0.08 (1.57–1.80) 10 1.73 ± 0.14 (1.56–2.04) 8
1.41 ± 0.22 (1.15–1.83) 8 1.38 ± 0.14 (1.28–1.47) 2
1.30 ± 0.13 (1.03–1.45) 12 1.31 ± 0.13 (1.09–1.57) 12
TTL/TL
30.50 ± 2.27 (27.02–33.89) 10 42.21 ± 2.70 (38.02–45.98) 8
36.03 ± 3.59 (32.76–43.45) 8 45.95 ± 3.83 (43.23–48.65) 2
33.86 ± 7.05 (27.49–54.72) 12 39.17 ± 3.32 (32.64–45.58) 12
Etymology. The specific epithet is derived from the Latin name “arena”, a reference to the name of the type locality of the new species, the municipality of Areia. Distribution and habitat. Amerotyphlops arenensis is only known from the Reserva Ecológica Mata do Pau Ferro (REMPF), situated at 5 km from the municipality of Areia, state of Paraíba, Brazil (Fig. 3). This region is considered one of the largest upland forest fragments of the state of Paraíba (about 600 hectares). The prevalent phytophysiognomy in the REMPF correspond to a sub-montane seasonal semi-decidual forest (Fig. 4) (Barbosa et al. 2004), ranging from 400–600 meters above sea level, with an annual average of temperature and rainfall of 22° C and 1.400 mm, respectively (Mayo & Fevereiro 1981). The REMPF has several different levels of successional forests, presenting in some parts a high and dense forest with large height trees (approximately 35 meters), with a well-defined canopy structure, woodland and understory well established and preserved (Barbosa et al. 2004). All specimens were captured with the aid of pitfall traps into the most preserved forest.
Discussion Despite the significant advances in our knowledge of the South American snake fauna, the genus Amerotyphlops remains elusive, with several species still known only from their type series (Dixon & Hendricks 1979; Rodrigues 1991). Amerotyphlops amoipira, A. yonenagae, and A. paucisquamus represent species morphologically similar with distributions restricted to the São Francisco sand dunes and coastal Atlantic forest of northeastern Brazil (Dixon & Hendricks 1979; Rodrigues 1991; Rodrigues & Juncá 2002). Our analysis revealed the presence of a fourth species in Northeastern Brazil, A. arenensis that is apparently restricted to the upland forests of the state of NEW AMEROTYPHLOPS FROM NORTHEASTERN BRAZIL
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Paraíba. Currently, A. arenensis is still known only from its type-locality but, similarly to A. paucisquamus and A. amoipira, future surveys along the northeastern region may likely extend its range of distribution. New geographic records for the later two species are discussed in more detail below, along with some of the more conspicuous features that help distinguish these species. Amerotyphlops amoipira was described from the sand dunes of the São Francisco River, in the municipality of Ibiraba, state of Bahia. Recently, Fernandes et al. (2010) extended its geographic distribution to Fazenda Santa Maria da Vereda, municipality of Bonito de Minas, state of Minas Gerais. Brito & Freire (2012) also extended the distribution of A. amoipira to some fragments of Atlantic forest; dunes and sandbank areas (sandy habitats with herbaceous and shrubby vegetation) in the states of Alagoas (Piaçabuçu, Coruripe and São José da Laje municipalities) and Rio Grande do Norte (municipality of Nísia Floresta). We further extend the distribution of A. amoipira eastward from the type locality (approximately 650 km airline), with five additional specimens from the municipality of Barra dos Coqueiros, along the sandy coast of the State of Sergipe (Fig. 3). All individuals show an incomplete nasal suture, 18/18/18 scales around the body, 209 to 230 middorsal scales and a light brown coloration that is only slightly pigmented.
FIGURE 3. Geographical distribution of Amerotyphlops arenensis (star), A. amoipira (triangles), A. paucisquamus (squares), and A. yonenagae (circle). Open symbols correspond to new records and closed symbols to type localities. For A. paucisquamus we plotted the more precise locality of the paratype.
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FIGURE 4. General view of the Reserva Mata do Pau-Ferro, state of Paraíba, Northeastern Brazil. Photographed by Gentil P. Filho.
Amerotyphlops paucisquamus was described based only on two specimens, with the holotype from the state of Pernambuco, while the paratype has a more specific locality, municipality of Recife (Dixon & Hendricks 1979). Rodrigues (1991) reported nine specimens of A. paucisquamus from the municipality of João Pessoa, in the state of Paraíba. França et al. (2012) also reported 50 specimens of A. paucisquamus from Reserva Biológica Guaribas, between the municipalities of Rio Tinto and Mamanguape, state of Paraíba. During the course of this research, several previously unidentified specimens (Amerotyphlops spp.) from Northeastern Brazil were identified as A. paucisquamus. These records extend significantly the distribution of the species (see Fig. 3). All specimens show an incomplete nasal suture, 18/18/18 scales around the body, fewer numbers of middorsal scales (162–209), a brownish-pigmented dorsum, and light brownish ventral scales. All specimens were found in fragments of the predominantly coastal semi-deciduous Atlantic Forest of Northeastern Brazil. Amerotyphlops yonenagae is currently reported to be endemic to the municipality of Santo Inácio, state of Bahia, where it occurs only in the sandy dune system along the São Francisco River. Here we report a new record from the Estação Ecológica Raso da Catarina, municipality of Paulo Afonso, state of Bahia, which extends the distribution of the species of approximately 500 km southwest within the Caatinga biome (Fig. 3). The single individual shows an incomplete nasal suture, 18/18/18 scales around the body, 250 middorsal scales, and a pale cream dorsum with dark paravertebral scales.
Acknowledgements We are grateful to Miguel T. Rodrigues (IB-USP), Francisco Franco (IBSP), Paulo Roberto Manzani and Luis Felipe de Toledo Ramos Pereira (ZUEC), Paulo Passos (MNRJ), Glaucia F. Pontes (MCP-PUCRS), Rejane Lira (UFBA), Frederico França (UFPB), Renato Feio (MZUFV), Marcos André de Carvalho (UFMT), Richard Vogt (INPA), Gustavo Vieira (CHUFPB), and Adrian Garda (UFRN) for allowing us to examine specimens under their care; Felipe G. Grazziotin and Juan C. Arredondo reviewed an earlier draft of the manuscript; and Luciana Moreira Lobo produced the line drawings. RG was supported by a Master fellowship from the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP; process 08/52285-0). GAPF is supported by a postdoctoral fellowship from the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP; process 2012/19753-6). AAAS was
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supported by a Master fellowship from the Conselho Nacional de Desenvolvimento Científico e Programa de Fomento a Pós-graduação (CAPES-PROF; process 15001016022P6). ALCP was supported by grants from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq; processes 308950/2011-9 and 562171/ 2010-0). This research was supported by grants from the Fundação de Amparo à Pesquisa do Estado de São Paulo (BIOTA/FAPESP; process 11/50206-9) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq; processes 565046/2010-1 and 303545/2010-0) to HZ. We are also grateful to Paulo Passos and Van Wallach for reviewing the manuscript.
References Arruda, M.P., Almeida, C.H.L.N., Rolim, D.C. & Maffei, F. (2011) First record in midwestern region of São Paulo, Brazil of Typhlops brongersmianus Vanzolini, 1976 (Squamata: Typhlopidade). Check List, 7 (4), 571–573. Ávila, R.W., Ferreira, W.L. & Souza, V.B. (2006) Biology of the blindsnake Typhlops brongersmianus (TYPHLOPIDAE) in a semideciduous forest from central Brazil. Herpetological Journal, 16, 403–405. Ávila, R.W. & Kawashita-Ribeiro, R.A. (2011) Herpetofauna of São João da Barra hydroeletric plant state of Mato Grosso, Brazil. Check List, 7 (6), 750–755. Barbosa, M.R.V., Agra, M.F., Sampaio, E.V.S., Da Cunha, J.P. & Andrade, L.A. (2004) Diversidade florística na Mata do PauFerro, Areia, Paraíba. In: Porto, K.C., Cabral, J.J.P. & Tabarelli, M. (Eds.), Brejos de Altitude em Pernambuco e Paraíba: História Natural, Ecologia e Conservação. Série Biodiversidade 9. Ministério do Meio Ambiente, Brasília, pp. 111–121. Brito, P.S. & Freire, E.M.X. (2012) New records and geographic distribution map of Typhlops amoipira Rodrigues and Juncá, 2002 (Typhlopidae) in the Brazilian Rainforest. Check List, 8 (6), 1347–1349. Caicedo-Portilla, J.R. (2011) Sexual dimorphism and geographic variation of the blind snake Typhlops reticulatus (Scolecophidia: Typhlopidae) and distribution of species of genus in Colombia. Caldasia, 33 (1), 221–234. Dixon, J.R. & Hendricks, F.S. (1979) The wormsnakes (Family Typhlopidae) of the Neotropics, exclusive of the Antilles. Zoologische Verhandelingen, 173, 1–39. Esri (1999) ArcGIS (version 10.2.2) Environmental Systems Research Institute. Inc, New York. Fernandes, V.D., Ribeiro, M.M., Santos Dayrell, J.S., Santana, D.J. & Rocha Lima, L.H. (2010) Reptilia, Squamata, Serpentes, Typhlopidae, Typhlops amoipira Rodrigues and Juncá: range extension and new state record. Check List, 6 (2), 268–269. França, F.G.R. & Venâncio, N.M. (2010) Reptiles and amphibians of poorly known region in southwest Amazonia. Biotemas, 23 (3), 71–84. http://dx.doi.org/10.5007/2175-7925.2010v23n3p71 França, F.G.R., Mesquita, D.O. & Colli, G. (2006) A checklist of snakes from Amazonian savannas in Brazil, housed in the coleção herpetológica da Universidade de Brasília, with new distribution records. Sam Noble Oklahoma Occasional Papers, 17, 1–13. França, R.C., Germano, C.E.S. & França, F.G.R. (2012) Composition of a snake assemblage inhabiting an urbanized area in the Atlantic forest of Paraíba State, Northeast Brazil. Biota Neotropica, 12 (3), 183–195. http://dx.doi.org/10.1590/S1676-06032012000300019 Freire, E.M.X. (2001) Composição, Taxonomia, Diversidade e Considerações Zoogeográficas sobre a Fauna de Lagartos e Serpentes de Remanescentes da Mata Atlântica do Estado de Alagoas, Brazil. PhD Thesis, Universidade Federal do Rio de Janeiro, Rio de Janeiro, 144 pp. Google Inc. (2013) Software Google Earth. Available from: http://www.google.com/earth/index.html (accessed 04 November 2013) Hedges, S.B., Marion, A.B., Lipp, K.M., Marin, J. & Vidal, N. (2014) A taxonomic framework for typhlopid snakes from the Caribbean and other regions (Reptilia, Squamata). Caribbean Herpetology, 49, 1–61. Loebmann, D. (2008) Geographic distribution. Typhlops brongersmianus. Herpetological Review, 32, 244. Martins, A.R., Silveira, A.L. & Bruno, S.F. (2010) New records of Typhlops brongersmianus (Serpentes, Typhlopidae) in southeastern Brazil. Herpetology Notes, 3, 247–248. Mayo, S.J. & Fevereiro V.P.B. (1982) Mata do Pau-Ferro, a Pilot Study of the Brejo Forest. Royal Botanic Gardens (BethamMoxon Trust) in association with Chrurchil Memorial Trust, Great Britain, 29 pp. Pyron, R.A. & Wallach, V. (2014) Systematic of the blindsnakes (Serpentes: Scolecophidia: Typhlopoidea) based on molecular and morphological evidence. Zootaxa, 3829 (1), 1–81. http://dx.doi.org/10.11646/zootaxa.3829.1.1 Rivas, G., Molina, C.R., Ugueto, G.N., Barros, T.R., Barrio-Amorós, C.L. & Kok, P.J.R. (2012) Reptiles of Venezuela: an updated and commented checklist. Zootaxa, 3211, 1–64. Rodrigues, M.T. (1991) Herpetofauna das dunas interiores do Rio São Francisco, Bahia, Brasil. IV. Uma nova espécie de Typhlops (Ophidia, Typhlopidae). Papéis Avulsos de Zoologia, 37 (22), 343–346. http://dx.doi.org/10.1590/s0031-10492002001300001 Rodrigues, M.T. & Juncá, F.A. (2002) Herpetofauna of the quartenary sand dunes of the middle Rio São Francisco: Bahia:
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Brazil. VII. Typhlops amoipira sp. nov., a possible relative of Typhlops yonenagae (Serpents, Typhlopidae). Papéis Avulsos de Zoologia, 42 (13), 325–333. http://dx.doi.org/10.1590/S0031-10492002001300001 Roux, J. (1926) Notes d'erpétologie Sud-Américaine. Revue Suisse de Zoologie, 33, 291–299. Roze, J.A. (1956) Ofidios coleccionados por la expedición Franco Venezolana al alto Orinoco 1951-1952. Boletin del Museo de Ciencias Naturales, 1, 179–195. Santana, G.G., Vieira, W.L.S., Pereira-Filho, G.A., Delfim, F.R., Lima, Y.C.C. & Vieira, K.S. (2008) Herpetofauna em um fragmento de Floresta Atlântica no estado da Paraíba, região Nordeste do Brasil. Biotemas, 21 (1), 75–84. http://dx.doi.org/10.5007/2175-7925.2008v21n1p75 Shreve, B. (1947) On Venezuelan reptiles and amphibians collected by Dr. H.G. Kugler. Bulletin of Museum of Comparative Zoology, 99, 519–537. SpeciesLink (2013) Rede SpeciesLink. Centro de Referência em Informação Ambiental (CRIA), Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP). Available from: http://www.splink.org.br/ (accessed 5 November 2013) Thomas, R. & Hedges, S.B. (2007) Eleven new species of the genus Typhlops (Serpentes: Typhlopidae) from Hispaniola and Cuba. Zootaxa, 1400, 1–26. Uetz, P. & Hošek, J. (2013) The Reptile Database, Available from: http://www.reptile-database.org (accessed 17 November 2013) Vanzolini, P.E. (1972) Typhlops brongersmai spec. nov. from the coast of Bahia, Brasil (Serpentes, Typhlopidae). Zoologische Mededelingen, 47 (3), 27–29. Wallach, V. (2003) Scolecophidia Miscellanea. Hamadryad, 27 (2), 227–245. Wallach, V., Williams, K.L. & Boundy, J. (2014) Snakes of the World: a Catalogue of Living and Extinct Species. CRC Press, Boca Raton, 1209 pp.
APPENDIX 1. Specimens examined. Amerotyphlops amoipira (N = 10). BRAZIL: SERGIPE: Aracajú: Barra dos Coqueiros: MZUSP 17464, MZUSP 17645, MZUSP 17466–68; BAHIA: Ibiraba: MZUSP 12299–301 (paratypes), MZUSP 12303 (paratype), MZUSP 12298 (holotype). Amerotyphlops brongersmianus (N = 185): BRAZIL: ALAGOAS: Campo Alegre: MZUSP 17271–72, Passo do Camaragibe: MZUSP 17729–31; BAHIA: Ilhéus: MZUSP 5218 (holotype), MZUSP 14666, MZUSP 14674, MZUSP 14678, MZUSP 14685, MZUSP 14687, MZUSP 14696, MZUSP 14702, Laje: IBSP 74025, Sapiranga: IBSP 55110, Uruçuca: MZUSP16737; CEÁRA: Ubajara: IBSP 76845–47; ESPÍRITO SANTO: Aracruz: MBML 158, Linhares: IBSP 76577, Presidente Kennedy: MNRJ 18743–45, MNRJ 18747, São Mateus: MZUSP 11597, Vitória: UFV 389; GOIÁS: Pequena Central Hidrelétrica Mosquitão: MZUSP 17675–77, Pequena Central Hidrelétrica Santa Edwirges: MZUSP 17776–80, Petrolina de Goiás: MZUSP 12491–93, Rio Verde: MZUSP 3850, Serra da Mesa: MZUSP 11037–38; MARANHÃO: Alcântara: MZUSP 17221; MATO GROSSO DO SUL: Anaurilândia: IBSP 63025–26, Bataguaçú: MZUSP 11689–98, Brasilândia: IBSP 62644, Campo Grande: MZUSP 10100, Corumbá: MZUSP 4440, MZUSP 4442–43, Ribas do Rio Pardo: IBSP 74589–90, Rio Pardo: Hidrelétrica Sérgio Motta: MZUSP 13986, Santa Rita do Pardo: IBSP 64080–81, MZUSP 11675, Três Lagoas: IBSP 75138, MZUSP 4401; MATO GROSSO: Alto Taquari: UFMT 6090, UFMT 6235; Araputanga: UFMT 5920–21, UFMT 4179, Aripuanã: UFMT 4241, UFMT 4245, UFV 1429–30, UFV 1440–41, UFV 1689–91, Cáceres: MZUSP 11460, MZUSP 11458; Chapada dos Guimarães (Usina Hidrelétrica Manso): MZUSP 14767, MZUSP 14772, MZUSP 14778, MZUSP 14781, MZUSP 14785–86, MZUSP 14795, MZUSP 14797, MZUSP 14812–14, MZUSP 14972, Nova Lacerda: UFMT 6236, UFMT 6239–41, Poconé: UFMT 5946, UFMT 5948–49, ZUEC 893, ZUEC 1159, São José do Rio Claro: UFMT 5876, Sapezal: UFMT 5896–97, Vale de São Domingos: UFMT 6237–38, UFMT 6242–43; MINAS GERAIS: Frutal: MZUSP 2720, Resplendor: UFV 1782, Volta Grande: UFV 1291; PARÁ: Alter do Chão: Lago Caapiranda: INPA H10677–79, INPA H10681–83; Bragança: MPEG 6344, MPEG 8459, MPEG 9972, Curuçá: MPEG 21483, Ourém: MPEG 1266, Tucuruí: MPEG 16804, Vitória do Xingu: Usina Hidrelétrica Belo Monte: MPEG 19819, PARAÍBA: João Pessoa: MZUSP 10672, Mata do Buraquinho: UFPB 4470–71, Rio Tinto: Reserva Biológica Guaribas: CHUFPB 11088–90, CHUFPB 11092, CHUFPB 11094, CHUFPB 11095; RIO GRANDE DO SUL: Arroio Teixeira: MCP 6723, Capão da Canoa: MCP 12079, Terra de Areia: MCP 12495, Tramandaí: MCP 0974, SANTA CATARINA: Florianópolis: MCP 7722, SÃO PAULO: Araçatuba: IBSP 8964, Barretos: IBSP 4753, Cachoeira de Emas: MZUSP 2455, Ilha Solteira: IBSP 33860–61, IBSP 36487, IBSP 36754, IBSP 38324–25, Indiaporã: IBSP 41837, IBSP 41842, IBSP 41847, IBSP 41851, Pereira Barreto: IBSP 54106–07, Primavera: IB 63061, Teodoro Sampaio: MZUSP 14557, MZUSP 15325; TOCANTINS: Aguiarnópolis: IBSP 75304–05, Babaçulândia: MPEG 23641, MPEG 23643, Guaraí: MZUSP 12682–84, MZUSP 13867, Mateiros: MZUSP 14978–80, Palmas: MZUSP 11508–10, MZUSP 11512; Usina Hidrelétrica Luiz Eduardo Magalhães: MZUSP 14758, MZUSP 14898–901, MZUSP 14903–04. PARAGUAY: ASSUNCIÓN: Baía de Assunción: IBSP 10117–19. Amerotyphlops minuisquamus (N = 18). BRAZIL: AMAZONAS: Manaus: MZUSP 5233 (paratype): São Raimundo: MZUSP 7638: Campus do Instituto Nacional de Pesquisas da Amazônia: MZUSP 8038, Presidente Figueiredo: IBSP 51726. GUYANA: POTARO-SIPARUNI: Vicinity of Kuribrong River: MZUSP 21446–51, MZUSP 21453–60.
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Amerotyphlops paucisquamus (N = 28). BRAZIL: ALAGOAS: Campo Alegre: MZUSP 17273–74, MZUSP 17289, CEARÁ: Caucáia: IBSP 76985; PARAÍBA: João Pessoa: MZUSP 8309–12, MZUSP 8914–15, MZUSP 8980, Rio Tinto: Reserva Biológica Guaribas: CHUFPB 11096–106; PERNAMBUCO: Cabo de Santo de Agostinho: Reserva Florestal do Sistema Gurjaú: MNRJ 17056; Recife: IBSP 17219 (paratype); MARANHÃO: São Luís: Usina Termoelétrica Ponta da Madeira: MNRJ 16956, MNRJ 16958, MNRJ 16967; RIO GRANDE DO NORTE: Goianinha: AAGARDA 2786. Amerotyphlops reticulatus (N = 23). BRAZIL: AMAPÁ: Oiapoque: MPEG 197; AMAZONAS: Boa Vista do Amanã: Reserva de Desenvolvimento Sustentável Amanã: INPA H10425–26, Manaus: Campus do Instituto de Pesquisas da Amazônia: INPA H14847, Urucará: MPEG 23526; MATO GROSSO: Nova Ubiratã: UFMT 5345, Sinop: UFMT 5790; MARANHÃO: Humberto de Campos: MNRJ 7851, São Luís: MNRJ 16980–81; PARÁ: Almerim: Monte Dourado road: MPEG 21434–38, Barcarena: MPEG 23640, Curionópolis: MPEG 20738, Ilha do Marajó: Chaves: MPEG 23581–82, RONDÔNIA: Espigão D'Oeste: Fazenda Jaburu: INPA H12096, Porto Velho: IBSP 52913, IBSP 53123; RORAIMA: Caracaraí: Parque Nacional Viruá: INPA H21663. Amerotyphlops yonenagae (N = 20). BRAZIL: BAHIA: Santo Inácio: MZUSP 10086 (holotype), MZUSP 10087 (paratype), MZUSP 10541, MZUSP 12497–500, MZUSP 22219–MTR 22223, MTR 11375,Gentio do Ouro: Vila de Santo Inácio: Lagoa de Itaparica: UFBA 1705–06, UFBA 001708–11, Paulo Afonso: Estação Ecológica Raso da Catarina: CHUFPB 11087.
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Article
ISSN 1175-5326 (print edition)
ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3920.3.3 http://zoobank.org/urn:lsid:zoobank.org:pub:143FCD6B-F614-4422-832C-190114C28FEB
A new species of Amerotyphlops from Northeastern Brazil, with comments on distribution of related species ROBERTA GRABOSKI1, 2, 5, GENTIL ALVES PEREIRA FILHO1, ARIANE AUXILIADORA ARAÚJO DA SILVA3, 4, ANA LÚCIA DA COSTA PRUDENTE4 & HUSSAM ZAHER1 1
Museu de Zoologia da Universidade de São Paulo, CP 42494, São Paulo, 04218-070 São Paulo, Brasil Programa de Pós Graduação em Zoologia, Universidade Estadual Paulista Júlio de Mesquita Filho, Av. 24 A, 1515, Bairro Bela Vista, Rio Claro, 13506-900, São Paulo, Brasil 3 Programa de Pós-Graduação em Zoologia, Universidade Federal do Pará, Museu Paraense Emílio Goeldi. Av. Perimetral, 1901,Terra Firme, CP 399, Belém, 66017-970 Pará, Brasil. 4 Laboratório de Herpetologia, Museu Paraense Emílio Goeldi, CP 399, Av. Perimetral, 1901, Terra Firme, Belém, 66077-530, Pará, Brasil 5 Corresponding author. E-mail: [email protected] 2
Abstract We describe a new species of Amerotyphlops from an upland forest enclave in the state of Paraíba, Northeastern Brazil. The new species is distinguished from the other seven South American species of Amerotyphlops by the combination of the following characters: nasal suture incomplete; rostral scale oval and yellowish cream with some dark brown spots; four supralabial scales; three infralabial scales; rows of scales around the body 18/18/18; middorsal scales from 204 to 225; dorsum with twelve to thirteen rows of scales dark brown and belly with four to five rows of scales immaculate yellowish cream; caudal spine dark brown; subcaudal scales 8–10 in female and 11–13 in males; maximum total length 233 mm. The new species is morphologically similar to A. amoipira and A. paucisquamus, sharing 18/18/18 rows of scales around the body and a small overlap of counts of middorsal scales. Key words: Atlantic rainforest, Typhlopidae, taxonomy, Amerotyphlops paucisquamus, Amerotyphlops amoipira
Introduction The genus Amerotyphlops was recently proposed by Hedges et al. (2014) for the clade of typhlopids that currently comprises 14 species found in the New World. Thirteen species are distributed on the mainland, from Mexico to northern Argentina, and one insular species occur in the West Indian (Hedges et al. 2014; Pyron & Wallach 2014). From the 13 mainland species, seven are restricted to South America: Amerotyphlops reticulatus (Linnaeus 1758); A. lehneri (Roux 1926), A. brongersmianus (Vanzolini 1972), A. minuisquamus (Dixon & Hendricks 1979), A. paucisquamus (Dixon & Hendricks 1979); A. yonenagae (Rodrigues, 1991), and A. amoipira (Rodrigues & Juncá 2002). Amerotyphlops lehneri occurs exclusively in the state of Falcon, northern Venezuela (Roux 1926; Shreve 1947; Dixon & Hendricks 1979; Rivas et al. 2012; Wallach et al. 2014), while A. yonenagae and A. amoipira are endemic to the predominantly psammophilous environment of dunes along the São Francisco River, both occurring in the state of Bahia, Northeastern Brazil (Rodrigues 1991; Rodrigues & Juncá 2002; Wallach et al. 2014). Amerotyphlops paucisquamus is endemic in the Atlantic forest of Northeastern Brazil, from state of Pernambuco and Paraíba (Dixon & Hendricks 1979; Rodrigues 1991; Freire 2001; Rodrigues & Juncá 2001; França et al. 2012; Wallach et al. 2014). The three remaining species are widely distributed in South America: A. minuisquamus is distributed throughout the Amazon basin, in Peru, western Brazil, Venezuela, Colombia and the Guyana Shield (Roze 1956; Dixon & Hendricks 1979; Caicedo-Portilla 2011; Wallach et al. 2014); A. reticulatus occurs in both cis- and trans-Andean lowlands of Colombia (Caicedo-Portilla 2011), and throughout the Amazon basin and
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adjacent ecotonal regions southern of the Amazon forest, including gallery forests in the Cerrado biome (Cunha & Nascimento 1978, 1993; Dixon & Hendricks 1979; França et al. 2006; França & Venâncio 2010; Ávila & Kawashita-Ribeiro 2011; Wallach et al., 2014); A. brongersmianus is widely distributed along the open formations of South America (Cerrado, Chaco, and Caatinga) and savanna enclaves within the Amazon and Atlantic forests (Dixon & Hendrix 1979; Rodrigues 1991; Loebmann 2008; Santana et al. 2008; Martins et al. 2010; Arruda et al. 2011; Wallach et al., 2014). Here, we describe a new species of Amerotyphlops from an upland rainforest (locally known as “Brejo de Altitude”) in the state of Paraíba, Northeastern Brazil. This new species is morphologically similar to A. amoipira and A. paucisquamus.
Material and methods We examined 303 specimens from six South American species of Amerotyphlops (Appendix 1). Amerotyphlops lehneri was the only South American species for which no specimens were available for this study; we thus used the available information in the literature (Roux, 1926; Shreve,1947; Dixon & Hendricks, 1979). All specimens examined are deposited in the following institutions (acronyms in parentheses): Instituto Butantan, São Paulo, São Paulo (IBSP); Instituto Nacional de Pesquisas da Amazônia, Manaus, Amazonas (INPA); Museu de Ciências e Tecnologia da PUC-RS, Porto Alegre, Rio Grande do Sul (MCP-PUCRS); Museu de Zoologia da Universidade de São Paulo, São Paulo, São Paulo (MZUSP); Museu de Zoologia da Universidade Estadual de Campinas, Campinas, São Paulo (ZUEC-REP); Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Rio de Janeiro (MNRJ); Museu Paraense Emílio Goeldi, Belém, Pará (MPEG); Universidade Federal da Bahia, Salvador, Bahia (UFBA); Universidade Federal da Paraíba, João Pessoa, Paraíba (CHUFPB); Museu de Zoologia João Moojen, Universidade Federal de Viçosa, Viçosa, Minas Gerais (MZUFV); Museu de Zoologia Professor Mello Leitão, Santa Teresa, Espírito Santo (MBML); Universidade Federal do Mato Grosso, Cuiabá, Mato Grosso (UFMT); Laboratório de Anfíbios e Répteis, Universidade Federal do Rio Grande do Norte, Natal, Rio Grande do Norte (AAGARDA). We measured the total length (TTL) to the nearest 1 mm by stretching carefully the specimen along a graduated ruler. We measured the cephalic scales, midbody diameter (MBD), and tail length (TL) to the nearest 0.1 mm with a digital caliper. We followed Wallach (2003) for terminology on cephalic scales, and Thomas & Hedges (2007) and Dixon & Hendrix (1979) for measurements and scale counts, except for middorsal scales, which were counted along the paravertebral row and did not include the rostral scale and terminal spine. We counted the transversal scale rows in three different places along the body: 1) at the level of the 20th scale posterior to the symphysial, 2) at mid body, 3) at the level of the 10th scale anterior to the cloacal shield. We determined sex by making a small incision at the base of the tail to access the presence or absence of hemipenes. We generated a distribution map through the software ArcGIS v10.2.2 (ESRI 1999). Geographical coordinates were obtained from the Species Link 2013 online database, based on the institutions database, or directly taken with GPS devices at the localities. The software Google Earth (Google Inc. 2013) was used, whenever possible, to refine the coordinates. We performed an univariate analysis of variance (ANOVA) to assess the possible presence of secondary sexual dimorphism in several meristic variables (subcaudal, middorsal and ventral scales), and two continuous variables (TTL and TL). Assumptiom of univariate normality and homoscedasticity were performed with KolmogorovSmirnov’s and Levene’s tests, respectively. Characters that showed sexual dimorphism are shown separately for males and females. Only adult individuals, determined following Ávila et al. (2006), were employed in the analysis of sexual dimorphism to avoid the allometric effect by continuous variables. Only one juvenile (MZUSP 21274) from the 18 specimens examined (eight females and 10 males), was removed from the analysis.
Amerotyphlops arenensis sp. nov. Fig. 1 Holotype. An adult male, MZUSP 20042, collected by Gentil P. Filho on 19th November 2008 from the Reserva
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Ecológica Mata do Pau Ferro (06° 58'12'' S, 35° 42'15'' W; ca. 600 m), municipality of Areia, state of Paraíba, Brazil. Paratypes. Eighteen specimens all collected in the type locality by Gentil P. Filho. MZUSP 20037–38 collected in 16th October 2008, MZUSP 20039 collected in 17th October 2008, MZUSP 20040–41 collected in 19th November 2008, MZUSP 20136 collected in 19th November 2008, MZUSP 20043–45 collected in 17th December 2008, MZUSP 20046 collected in 18th December 2008, MZUSP 20047 and MZUSP 21273 collected in 21st January 2009, MZUSP 20048 collected in 28th February 2009, MZUSP 20049–50 and MZUSP 21274 collected in 27th March 2009, MZUSP 20051 collected in 23rd April 2009, and MZUSP 20052 collected in 21st May 2009. Diagnosis. Amerotyphlops arenensis is distinguished from all other South American species by the following combination of characters: nasal suture incomplete; rostral scale oval; supralabial scales four; infralabial scales three; rows scales around the body 18/18/18; middorsal scales 204 to 225; rows of dorsal scales dark brown 12–13th; rows of ventral scales yellowish cream and immaculate 4–5; caudal spine dark brown; subcaudal scales 8–10 in female and 11–13 for males; and maximum TTL 233 mm.
FIGURE 1. Lateral (A), ventral (B), and dorsal (C) views of the head of the holotype of Amerotyphlops arenensis sp. nov. (MZUSP 20042). Scale Bar = 1mm. NEW AMEROTYPHLOPS FROM NORTHEASTERN BRAZIL
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FIGURE 2. General view in life of the paratype (MZUSP 20038) of Amerotyphlops arenensis. Photographed by Gentil P. Filho.
The new species differs from Amerotyphlops lehneri by having an incomplete nasal suture (vs. complete nasal suture); from A. brongersmianus, A. reticulatus, and A. minuisquamus by having 18/18/18 rows scales around the body (vs. 18/16/14, 18/18/14, 20/18/14 or 20/18/15 in A. minuisquamus; 20/20/18 or 20/20/20 in A. brongersmianus and A. reticulatus); from A. yonenagae by having less than 250 middorsal scales (vs. more than 250 middorsal); from A. amoipira by having highly pigmented cephalic scales with a dark brown dorsum (vs. few pigmented cephalic scales, creamish brown dorsum with a fine darker brown paravertebral line concentrated in the anterior part of the body) and from A. paucisquamus by having a largest number of middorsal, between 204–225 (vs. fewer number of middorsal, between 162–209). Description of the holotype. Adult male, TTL 191 mm, TL 7 mm, TTL/MBD 28.08, and TTL/TL 27.28. Head slightly depressed dorsoventrally, not distinct from neck. Snout round in dorsal and ventral views. Rostral oval, longer than wide, narrow at anteroposterior region and wider at medial region; visible in dorsal view, extending ventrodorsally without reaching the imaginary transverse line between anterior borders of eyes. Rostral contacting nasal (anterior and posterior) dorsolaterally, and first supralabial and anterior nasal scales ventrally. Nasal suture incomplete, only partially dividing the anterior and posterior portions of nasal scale. Suture begins in the upper edge of second supralabial, pass through nostril, but fails to reach rostral. Anterior nasal in contact with first infralabial and upper edge of second infralabial. Posterior nasal longer than wide, contacting upper margin of second supralabial and preocular. Supralabials four, fourth twice longer than third supralabial. Infralabials three, third largest. Eye diameter 0.91 mm; eyes not visible in ventral view, located dorsolaterally, close to suture between preocular and ocular scales, completely covered by ocular scale. Ocular scales contacting frontal. Body cylindrical and robust. Midbody diameter 6.80 mm. Dorsal and ventral scales cycloid, wider than long, highly imbricated and arranged in diagonal series; scale rows around the body 18/18/18. Middorsal scales 213. Cloacal plate rounded, bordered anteriorly by three rows of scales and posteriorly by five rows of scales. Subcaudal scales 13, excluding the terminal spine. Terminal spine large, stout base and dark brown. Coloration of the holotype in life. Dorsum (12–13 rows of scales) dark brown, venter (4–5 rows of scales) light cream. Dorsal and ventral portions of snout pinkish, with a few dark brown spots, covering both rostral and nasal scales (only in the upper half of snout). Symphysial region pinkish and immaculate. Dorsal head scales (supraoculars, frontal, posfrontal, parietals, and occipitals) and dorsal portions of lateral head scales (ocular, nasal, and lower nasal) predominantly dark brown, and ventral portions pinkish. Cloacal plate light cream and terminal spine dark brown. After six years preserved in 70% ethanol, the coloration of the holotype remained mostly the same. All regions in light cream and pinkish changed to yellowish cream (Fig. 2). Variation. The ANOVA revealed secondary sexual dimorphism only for the number of subcaudal scales (F= 63.43, p < 0.001) and TL (F=47.61, p < 0.001) among the verified variables. Number of subcaudal scales 11–13 (mean = 11.8, SD= 0.63, N= 11) in males and 8–10 (mean = 9.2, SD= 0.70, N= 8) in females. Tail length 4.42–6.77 % of TTL (N= 10) in males and 3.71–4.91% of TTL (N= 8) in females. Largest female and male with 233 mm
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TTL and 5.23 mm of TL (MZUSP 20043) and 215 mm of TTL and 7.19 of TL (MZUSP 20044), respectively. MBD 5.41–8.04 (mean = 7.21, SD= 0.77, N=10) in males and 5.85–9 (mean = 8; SD = 1.12; N= 8) in females; number of middorsal scales 204–221 (mean = 214.1, SD= 4.70, N= 11) in males and 207–225 (mean = 217.25, SD= 7.10, N= 8) in females. The color patterns of the paratypes are similar to that found in the holotype. However, two individuals (11%) lack dark spots in their rostral scale while the remaining individuals (88%) retain dark spots in their rostral scale. Table 1 shows additional morphometric characters and scale patterns found in the new species and in two morphologically similar species distributed in the Northeastern Brazil (A. amoipira and A. paucisquamus). TABLE 1. Variation for some selected characters of morphologically similar species of Amerotyphlops from Northeast region of Brazil. The values displayed in the table represent (in this sequence) range, mean, standard deviations, and available sample size. Abbreviations are as following: ED = eye diameter, HR = head radius, HWE = head width, IN = internasal distance, INORB = interorbital distance, RL = rostral length, RW = rostral width, TL = tail length, TTL: total length. Character
A. arenensis sp. nov.
A. amoipira
A. paucisquasmus
ED
0.93 ± 0.09 (0.75–1.05) 10 0.90 ± 0.10 (0.77–1.10) 8
0.83 ± 0.09 (0.69–0.98) 8 0.80 ± 0.24 (0.62–0.97) 2
0.80 ± 0.15 (0.43–0.99) 12 0.85 ± 0.19 (0.52–1.29) 12
HR
3 ± 0.10 (2.78–3.13) 10 3.17 ± 0.21 (2.77–3.42) 8
2.52 ± 0.39 (1.95–3.04) 8 2.77 ± 0.68 (2.28–3.35) 2
2.62 ± 0.26 (2.05–2.93) 12 2.69 ± 0.26 (2.27–3.14) 12
HWE
5.15 ± 0.33 (4.43–5.47) 10 5.47 ± 0.42 (4.80–6.02) 8
3.97 ± 0.58 (3.25–5.13) 8 4.04 ± 0.60 (3.62–4.46) 2
4.11± 0.42 (3.17–4.64) 12 4.24 ± 0.42 (3.58–5.00) 12
IN
2.47 ± 0.14 (2.16–2.65) 10 2.58 ± 0.18 (2.35–2.80) 8
2.19 ± 0.30 (1.74–2.74) 8 2.14 ± 0.33 (1.91–2.37) 2
2.14 ± 0.25 (1.54–2.50) 12 2.18 ± 0.18 (1.86–2.61) 14
INORB
3.29 ± 0.13 (3.05–3.45) 10 3.41± 0.24 (3–3.86) 8
2.23 ± 0.25 (1.81–2.56) 8 2.23 ± 0.20 (2.09–2.36) 2
2.58 ± 0.26 (1.94–2.88) 12 2.48 ± 0.19 (2.09–2.76) 12
RL
2.84 ± 0.15 (2.59–3.09) 10 3.05 ± 0.22 (2.71–3.41) 8
2.19 ± 0.29 (1.82–2.75) 8 2.23 ± 0.50 (1.88–2.59) 2
2.40 ± 0.22 (1.90–2.76) 12 2.36 ± 0.22 (2.11–2.78) 12
RW
1.66 ± 0.08 (1.57–1.80) 10 1.73 ± 0.14 (1.56–2.04) 8
1.41 ± 0.22 (1.15–1.83) 8 1.38 ± 0.14 (1.28–1.47) 2
1.30 ± 0.13 (1.03–1.45) 12 1.31 ± 0.13 (1.09–1.57) 12
TTL/TL
30.50 ± 2.27 (27.02–33.89) 10 42.21 ± 2.70 (38.02–45.98) 8
36.03 ± 3.59 (32.76–43.45) 8 45.95 ± 3.83 (43.23–48.65) 2
33.86 ± 7.05 (27.49–54.72) 12 39.17 ± 3.32 (32.64–45.58) 12
Etymology. The specific epithet is derived from the Latin name “arena”, a reference to the name of the type locality of the new species, the municipality of Areia. Distribution and habitat. Amerotyphlops arenensis is only known from the Reserva Ecológica Mata do Pau Ferro (REMPF), situated at 5 km from the municipality of Areia, state of Paraíba, Brazil (Fig. 3). This region is considered one of the largest upland forest fragments of the state of Paraíba (about 600 hectares). The prevalent phytophysiognomy in the REMPF correspond to a sub-montane seasonal semi-decidual forest (Fig. 4) (Barbosa et al. 2004), ranging from 400–600 meters above sea level, with an annual average of temperature and rainfall of 22° C and 1.400 mm, respectively (Mayo & Fevereiro 1981). The REMPF has several different levels of successional forests, presenting in some parts a high and dense forest with large height trees (approximately 35 meters), with a well-defined canopy structure, woodland and understory well established and preserved (Barbosa et al. 2004). All specimens were captured with the aid of pitfall traps into the most preserved forest.
Discussion Despite the significant advances in our knowledge of the South American snake fauna, the genus Amerotyphlops remains elusive, with several species still known only from their type series (Dixon & Hendricks 1979; Rodrigues 1991). Amerotyphlops amoipira, A. yonenagae, and A. paucisquamus represent species morphologically similar with distributions restricted to the São Francisco sand dunes and coastal Atlantic forest of northeastern Brazil (Dixon & Hendricks 1979; Rodrigues 1991; Rodrigues & Juncá 2002). Our analysis revealed the presence of a fourth species in Northeastern Brazil, A. arenensis that is apparently restricted to the upland forests of the state of NEW AMEROTYPHLOPS FROM NORTHEASTERN BRAZIL
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Paraíba. Currently, A. arenensis is still known only from its type-locality but, similarly to A. paucisquamus and A. amoipira, future surveys along the northeastern region may likely extend its range of distribution. New geographic records for the later two species are discussed in more detail below, along with some of the more conspicuous features that help distinguish these species. Amerotyphlops amoipira was described from the sand dunes of the São Francisco River, in the municipality of Ibiraba, state of Bahia. Recently, Fernandes et al. (2010) extended its geographic distribution to Fazenda Santa Maria da Vereda, municipality of Bonito de Minas, state of Minas Gerais. Brito & Freire (2012) also extended the distribution of A. amoipira to some fragments of Atlantic forest; dunes and sandbank areas (sandy habitats with herbaceous and shrubby vegetation) in the states of Alagoas (Piaçabuçu, Coruripe and São José da Laje municipalities) and Rio Grande do Norte (municipality of Nísia Floresta). We further extend the distribution of A. amoipira eastward from the type locality (approximately 650 km airline), with five additional specimens from the municipality of Barra dos Coqueiros, along the sandy coast of the State of Sergipe (Fig. 3). All individuals show an incomplete nasal suture, 18/18/18 scales around the body, 209 to 230 middorsal scales and a light brown coloration that is only slightly pigmented.
FIGURE 3. Geographical distribution of Amerotyphlops arenensis (star), A. amoipira (triangles), A. paucisquamus (squares), and A. yonenagae (circle). Open symbols correspond to new records and closed symbols to type localities. For A. paucisquamus we plotted the more precise locality of the paratype.
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FIGURE 4. General view of the Reserva Mata do Pau-Ferro, state of Paraíba, Northeastern Brazil. Photographed by Gentil P. Filho.
Amerotyphlops paucisquamus was described based only on two specimens, with the holotype from the state of Pernambuco, while the paratype has a more specific locality, municipality of Recife (Dixon & Hendricks 1979). Rodrigues (1991) reported nine specimens of A. paucisquamus from the municipality of João Pessoa, in the state of Paraíba. França et al. (2012) also reported 50 specimens of A. paucisquamus from Reserva Biológica Guaribas, between the municipalities of Rio Tinto and Mamanguape, state of Paraíba. During the course of this research, several previously unidentified specimens (Amerotyphlops spp.) from Northeastern Brazil were identified as A. paucisquamus. These records extend significantly the distribution of the species (see Fig. 3). All specimens show an incomplete nasal suture, 18/18/18 scales around the body, fewer numbers of middorsal scales (162–209), a brownish-pigmented dorsum, and light brownish ventral scales. All specimens were found in fragments of the predominantly coastal semi-deciduous Atlantic Forest of Northeastern Brazil. Amerotyphlops yonenagae is currently reported to be endemic to the municipality of Santo Inácio, state of Bahia, where it occurs only in the sandy dune system along the São Francisco River. Here we report a new record from the Estação Ecológica Raso da Catarina, municipality of Paulo Afonso, state of Bahia, which extends the distribution of the species of approximately 500 km southwest within the Caatinga biome (Fig. 3). The single individual shows an incomplete nasal suture, 18/18/18 scales around the body, 250 middorsal scales, and a pale cream dorsum with dark paravertebral scales.
Acknowledgements We are grateful to Miguel T. Rodrigues (IB-USP), Francisco Franco (IBSP), Paulo Roberto Manzani and Luis Felipe de Toledo Ramos Pereira (ZUEC), Paulo Passos (MNRJ), Glaucia F. Pontes (MCP-PUCRS), Rejane Lira (UFBA), Frederico França (UFPB), Renato Feio (MZUFV), Marcos André de Carvalho (UFMT), Richard Vogt (INPA), Gustavo Vieira (CHUFPB), and Adrian Garda (UFRN) for allowing us to examine specimens under their care; Felipe G. Grazziotin and Juan C. Arredondo reviewed an earlier draft of the manuscript; and Luciana Moreira Lobo produced the line drawings. RG was supported by a Master fellowship from the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP; process 08/52285-0). GAPF is supported by a postdoctoral fellowship from the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP; process 2012/19753-6). AAAS was
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supported by a Master fellowship from the Conselho Nacional de Desenvolvimento Científico e Programa de Fomento a Pós-graduação (CAPES-PROF; process 15001016022P6). ALCP was supported by grants from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq; processes 308950/2011-9 and 562171/ 2010-0). This research was supported by grants from the Fundação de Amparo à Pesquisa do Estado de São Paulo (BIOTA/FAPESP; process 11/50206-9) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq; processes 565046/2010-1 and 303545/2010-0) to HZ. We are also grateful to Paulo Passos and Van Wallach for reviewing the manuscript.
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Brazil. VII. Typhlops amoipira sp. nov., a possible relative of Typhlops yonenagae (Serpents, Typhlopidae). Papéis Avulsos de Zoologia, 42 (13), 325–333. http://dx.doi.org/10.1590/S0031-10492002001300001 Roux, J. (1926) Notes d'erpétologie Sud-Américaine. Revue Suisse de Zoologie, 33, 291–299. Roze, J.A. (1956) Ofidios coleccionados por la expedición Franco Venezolana al alto Orinoco 1951-1952. Boletin del Museo de Ciencias Naturales, 1, 179–195. Santana, G.G., Vieira, W.L.S., Pereira-Filho, G.A., Delfim, F.R., Lima, Y.C.C. & Vieira, K.S. (2008) Herpetofauna em um fragmento de Floresta Atlântica no estado da Paraíba, região Nordeste do Brasil. Biotemas, 21 (1), 75–84. http://dx.doi.org/10.5007/2175-7925.2008v21n1p75 Shreve, B. (1947) On Venezuelan reptiles and amphibians collected by Dr. H.G. Kugler. Bulletin of Museum of Comparative Zoology, 99, 519–537. SpeciesLink (2013) Rede SpeciesLink. Centro de Referência em Informação Ambiental (CRIA), Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP). Available from: http://www.splink.org.br/ (accessed 5 November 2013) Thomas, R. & Hedges, S.B. (2007) Eleven new species of the genus Typhlops (Serpentes: Typhlopidae) from Hispaniola and Cuba. Zootaxa, 1400, 1–26. Uetz, P. & Hošek, J. (2013) The Reptile Database, Available from: http://www.reptile-database.org (accessed 17 November 2013) Vanzolini, P.E. (1972) Typhlops brongersmai spec. nov. from the coast of Bahia, Brasil (Serpentes, Typhlopidae). Zoologische Mededelingen, 47 (3), 27–29. Wallach, V. (2003) Scolecophidia Miscellanea. Hamadryad, 27 (2), 227–245. Wallach, V., Williams, K.L. & Boundy, J. (2014) Snakes of the World: a Catalogue of Living and Extinct Species. CRC Press, Boca Raton, 1209 pp.
APPENDIX 1. Specimens examined. Amerotyphlops amoipira (N = 10). BRAZIL: SERGIPE: Aracajú: Barra dos Coqueiros: MZUSP 17464, MZUSP 17645, MZUSP 17466–68; BAHIA: Ibiraba: MZUSP 12299–301 (paratypes), MZUSP 12303 (paratype), MZUSP 12298 (holotype). Amerotyphlops brongersmianus (N = 185): BRAZIL: ALAGOAS: Campo Alegre: MZUSP 17271–72, Passo do Camaragibe: MZUSP 17729–31; BAHIA: Ilhéus: MZUSP 5218 (holotype), MZUSP 14666, MZUSP 14674, MZUSP 14678, MZUSP 14685, MZUSP 14687, MZUSP 14696, MZUSP 14702, Laje: IBSP 74025, Sapiranga: IBSP 55110, Uruçuca: MZUSP16737; CEÁRA: Ubajara: IBSP 76845–47; ESPÍRITO SANTO: Aracruz: MBML 158, Linhares: IBSP 76577, Presidente Kennedy: MNRJ 18743–45, MNRJ 18747, São Mateus: MZUSP 11597, Vitória: UFV 389; GOIÁS: Pequena Central Hidrelétrica Mosquitão: MZUSP 17675–77, Pequena Central Hidrelétrica Santa Edwirges: MZUSP 17776–80, Petrolina de Goiás: MZUSP 12491–93, Rio Verde: MZUSP 3850, Serra da Mesa: MZUSP 11037–38; MARANHÃO: Alcântara: MZUSP 17221; MATO GROSSO DO SUL: Anaurilândia: IBSP 63025–26, Bataguaçú: MZUSP 11689–98, Brasilândia: IBSP 62644, Campo Grande: MZUSP 10100, Corumbá: MZUSP 4440, MZUSP 4442–43, Ribas do Rio Pardo: IBSP 74589–90, Rio Pardo: Hidrelétrica Sérgio Motta: MZUSP 13986, Santa Rita do Pardo: IBSP 64080–81, MZUSP 11675, Três Lagoas: IBSP 75138, MZUSP 4401; MATO GROSSO: Alto Taquari: UFMT 6090, UFMT 6235; Araputanga: UFMT 5920–21, UFMT 4179, Aripuanã: UFMT 4241, UFMT 4245, UFV 1429–30, UFV 1440–41, UFV 1689–91, Cáceres: MZUSP 11460, MZUSP 11458; Chapada dos Guimarães (Usina Hidrelétrica Manso): MZUSP 14767, MZUSP 14772, MZUSP 14778, MZUSP 14781, MZUSP 14785–86, MZUSP 14795, MZUSP 14797, MZUSP 14812–14, MZUSP 14972, Nova Lacerda: UFMT 6236, UFMT 6239–41, Poconé: UFMT 5946, UFMT 5948–49, ZUEC 893, ZUEC 1159, São José do Rio Claro: UFMT 5876, Sapezal: UFMT 5896–97, Vale de São Domingos: UFMT 6237–38, UFMT 6242–43; MINAS GERAIS: Frutal: MZUSP 2720, Resplendor: UFV 1782, Volta Grande: UFV 1291; PARÁ: Alter do Chão: Lago Caapiranda: INPA H10677–79, INPA H10681–83; Bragança: MPEG 6344, MPEG 8459, MPEG 9972, Curuçá: MPEG 21483, Ourém: MPEG 1266, Tucuruí: MPEG 16804, Vitória do Xingu: Usina Hidrelétrica Belo Monte: MPEG 19819, PARAÍBA: João Pessoa: MZUSP 10672, Mata do Buraquinho: UFPB 4470–71, Rio Tinto: Reserva Biológica Guaribas: CHUFPB 11088–90, CHUFPB 11092, CHUFPB 11094, CHUFPB 11095; RIO GRANDE DO SUL: Arroio Teixeira: MCP 6723, Capão da Canoa: MCP 12079, Terra de Areia: MCP 12495, Tramandaí: MCP 0974, SANTA CATARINA: Florianópolis: MCP 7722, SÃO PAULO: Araçatuba: IBSP 8964, Barretos: IBSP 4753, Cachoeira de Emas: MZUSP 2455, Ilha Solteira: IBSP 33860–61, IBSP 36487, IBSP 36754, IBSP 38324–25, Indiaporã: IBSP 41837, IBSP 41842, IBSP 41847, IBSP 41851, Pereira Barreto: IBSP 54106–07, Primavera: IB 63061, Teodoro Sampaio: MZUSP 14557, MZUSP 15325; TOCANTINS: Aguiarnópolis: IBSP 75304–05, Babaçulândia: MPEG 23641, MPEG 23643, Guaraí: MZUSP 12682–84, MZUSP 13867, Mateiros: MZUSP 14978–80, Palmas: MZUSP 11508–10, MZUSP 11512; Usina Hidrelétrica Luiz Eduardo Magalhães: MZUSP 14758, MZUSP 14898–901, MZUSP 14903–04. PARAGUAY: ASSUNCIÓN: Baía de Assunción: IBSP 10117–19. Amerotyphlops minuisquamus (N = 18). BRAZIL: AMAZONAS: Manaus: MZUSP 5233 (paratype): São Raimundo: MZUSP 7638: Campus do Instituto Nacional de Pesquisas da Amazônia: MZUSP 8038, Presidente Figueiredo: IBSP 51726. GUYANA: POTARO-SIPARUNI: Vicinity of Kuribrong River: MZUSP 21446–51, MZUSP 21453–60.
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Amerotyphlops paucisquamus (N = 28). BRAZIL: ALAGOAS: Campo Alegre: MZUSP 17273–74, MZUSP 17289, CEARÁ: Caucáia: IBSP 76985; PARAÍBA: João Pessoa: MZUSP 8309–12, MZUSP 8914–15, MZUSP 8980, Rio Tinto: Reserva Biológica Guaribas: CHUFPB 11096–106; PERNAMBUCO: Cabo de Santo de Agostinho: Reserva Florestal do Sistema Gurjaú: MNRJ 17056; Recife: IBSP 17219 (paratype); MARANHÃO: São Luís: Usina Termoelétrica Ponta da Madeira: MNRJ 16956, MNRJ 16958, MNRJ 16967; RIO GRANDE DO NORTE: Goianinha: AAGARDA 2786. Amerotyphlops reticulatus (N = 23). BRAZIL: AMAPÁ: Oiapoque: MPEG 197; AMAZONAS: Boa Vista do Amanã: Reserva de Desenvolvimento Sustentável Amanã: INPA H10425–26, Manaus: Campus do Instituto de Pesquisas da Amazônia: INPA H14847, Urucará: MPEG 23526; MATO GROSSO: Nova Ubiratã: UFMT 5345, Sinop: UFMT 5790; MARANHÃO: Humberto de Campos: MNRJ 7851, São Luís: MNRJ 16980–81; PARÁ: Almerim: Monte Dourado road: MPEG 21434–38, Barcarena: MPEG 23640, Curionópolis: MPEG 20738, Ilha do Marajó: Chaves: MPEG 23581–82, RONDÔNIA: Espigão D'Oeste: Fazenda Jaburu: INPA H12096, Porto Velho: IBSP 52913, IBSP 53123; RORAIMA: Caracaraí: Parque Nacional Viruá: INPA H21663. Amerotyphlops yonenagae (N = 20). BRAZIL: BAHIA: Santo Inácio: MZUSP 10086 (holotype), MZUSP 10087 (paratype), MZUSP 10541, MZUSP 12497–500, MZUSP 22219–MTR 22223, MTR 11375,Gentio do Ouro: Vila de Santo Inácio: Lagoa de Itaparica: UFBA 1705–06, UFBA 001708–11, Paulo Afonso: Estação Ecológica Raso da Catarina: CHUFPB 11087.
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