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Copeia, 2007(4), pp. 788–797

A New Species of Atractus (Serpentes: Dipsadinae) from a Relictual Forest in Northeastern Brazil PAULO PASSOS, DANIEL S. FERNANDES,

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DIVA M. BORGES-NOJOSA

A new species of Atractus is described from Serra de Baturite´, an Atlantic Forest enclave in the semiarid Caatinga of the state of Ceara´, northeastern Brazil. The new species is distinguished from all congeners by the combination of 17 dorsal scale rows at midbody, long loreal, two postoculars, seven upper and lower labials, first four infralabials in contact with chinshields, seven maxillary teeth, moderate body and tail sizes, slightly bilobed and semicapitate hemipenis with lateral-tip projections, light dorsal color pattern uniformly scattered with small dark brown dots, and ventral color pattern uniformly creamish white. The new species shares most of the external morphology and hemipenis characters with A. pantostictus, differing from this taxon by general color pattern and number of subcaudal scales. Uma nova espe´cie de Atractus e´ descrita da Serra de Baturite´, um enclave de Mata Atlaˆntica na Caatinga semi-a´rida do estado do Ceara´, nordeste do Brasil. A nova espe´cie e´ distinta de todas as congeˆneres pela combinac¸a˜o de 17 fileiras de escamas dorsais no meio do corpo, loreal longa, duas po´s-oculares, sete supra e infralabiais, as quatro primeiras infralabiais em contato com mentonianas, sete dentes maxilares, corpo e cauda com tamanho moderado, hemipeˆnis levemente bilobado e semi-capitado com projec¸o˜es laterais no a´pice, padra˜o de colorido dorsal claro uniformemente salpicado com pequenas pontuac¸o˜es marrom-escuro e padra˜o de colorido ventral uniformemente creme esbranquic¸ado. A nova espe´cie compartilha a maioria dos caracteres de morfologia externa e hemipeˆnis com A. pantostictus, diferindo deste ta´xon pelo padra˜o de colorido geral e nu´mero de escamas subcaudais.

EMBERS of the Dipsadinae (5Dipsadidae sensu Vidal et al., 2007) snake genus Atractus are widely distributed in South America, occurring from Panama south to Argentina (Giraudo and Scrocchi, 2000; Myers, 2003). This genus of semi-fossorial or cryptozoical snakes currently comprises about 100 species, most of them exhibiting restricted distributions (Fernandes et al., 2000; Passos et al., 2005). The taxonomic status of many species is confused and just a few taxonomic revisions on regional scales are available (Savage, 1960; Hoogmoed, 1980; Cunha and Nascimento, 1983; Martins and Oliveira, 1993). Twenty-eight Brazilian species of Atractus are currently recognized (http:// www2.sbherpetologia.org.br/checklist/repteis.htm), three of them occurring in the Atlantic Forest in northeastern Brazil (Fernandes, 1995, 1996). Atractus guentheri is restricted to the southern portion of the state of Bahia, ranging from the municipalities of Ilhe´us south to Canavieiras (Fernandes and Argo ˆ lo, 1999). Atractus maculatus is known exclusively from the municipalities of Murici and Sa˜o Miguel dos Campos in the state of Alagoas (Fernandes et al., 2000). Atractus potschi ranges from the municipality of Maceio´ in the state of Alagoas south to Feira de Santana in the state of Bahia (Fernandes, 1995; Lima et al., 2000). Herein, we describe a new species

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from Serra de Baturite´, a mountainous Atlantic Forest enclave in the semiarid Caatinga, state of Ceara´, northeastern Brazil. MATERIALS AND METHODS The observed characters are from external morphology and hemipenis. Terminology for Atractus cephalic shields follows Savage (1960), whereas the method of counting ventral scales follows Dowling (1951). We consider three possible loreal scale conditions for the genus Atractus: short—approximately as long as high, with anterior and posterior margins with the same height; moderate—slightly longer (less than two times) than high, with anterior and posterior margins approximately with the same height; long—twice (or more) longer than high, with anterior margin higher than posterior. Hemipenis terminology follows Dowling and Savage (1960), Myers and Campbell (1981), and Zaher (1999). Techniques for hemipenis preparation follow Pesantes (1994) and Myers and Cadle (2003). Sex was determined by the presence or absence of hemipenes through a ventral incision at the base of the tail. Measurements were taken with an ocular micrometer and Zeiss (Stemi SV8) stereoscope to

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Assumptions of normality and homoscedasticity were evaluated using Kolmogorov–Smirnov and Levene’s test, respectively (Zar, 1999). In cases where characters showed insufficient variation to justify these assumptions, non-parametric tests (Mann–Whitney U-test) were performed (Zar, 1999). Atractus ronnie, new species Figures 1, 2 Holotype.—MNRJ 14194, adult female, Brazil, Ceara´ State, Serra de Baturite´, municipality of Pacoti, Granja, 04u109S, 38u559W, ca. 800 m, 10 April 1998, D. M. Borges-Nojosa.

Fig. 1. Dorsal (A) and lateral (B) views of head, and midbody (C) of holotype of Atractus ronnie (MNRJ 14194). Scale 5 5 mm.

the nearest 0.1 mm, except for snout–vent (SVL) and caudal lengths (CL), which were taken with a flexible ruler to the nearest millimeter. As segmental counts are known to be sexually dimorphic in Atractus (Savage, 1960; Passos et al., 2005) we employed an analysis of variance to verify the existence of sexual dimorphism.

Paratypes.—Twenty-four specimens (all from Ceara´ State, Serra de Baturite´): municipality of Pacoti: CHUFC 1396 and MNRJ 14195, 17 July 1989, L. W. Lima-Verde; CHUFC 2646, 18 Dec. 1997, D. M. Borges-Nojosa; CHUFC 2648, 11 Jan. 1998, D. M. Borges-Nojosa; CHUFC 2658, Sı´tio Olho d’a´gua dos Tangara´s, 9 Dec. 2005, D. M. Borges-Nojosa, J. C. L. Melo, and M. J. B. Leite; CHUFC 2481 and CHUFC 3500, Monguba, 23 Feb. 1989, D. M. Borges-Nojosa; CHUFC 3502, Sı´tio Sa˜o Jose´, 3 April 1990, D. M. Borges-Nojosa; MNRJ 14196 and MNRJ 14197, Granja, 16 Feb. 1999 and 10 April 1998, respectively, D. M. Borges-Nojosa; CHUFC 2641 and CHUFC 2647, Sı´tio Pau do Alho, 2 Nov. 1997, D. M. BorgesNojosa; CHUFC 2598, Sı´tio Xangrila´, 28 April 2005, W. C. M. Luz; CHUFC 2652–54, Cidade Pacoti, Dec. 2005, W. C. M. Luz; CHUFC 2675–76 and CHUFC 2678, Cidade Pacoti, 17 Feb.–2 March 2006, W. C. M. Luz; CHUFC 2733, Distrito de Santana, 4 April 1990, D. M. Borges-Nojosa.

Fig. 2. Dorsal (A) and ventral (B) views of Atractus ronnie (MNRJ 14195, female, paratype) in preservative, 228 mm SVL, 21 mm CL.

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Municipality of Baturite´: CHUFC 2578, Sı´tio Escuro, 11 Feb. 2005, D. M. Borges-Nojosa, P. Cascon, and J. C. L. Melo. Municipality of Mulungu: CHUFC 2645, 1991, D. M. BorgesNojosa and S. M. Corne´lio. Municipality of Guaramiranga: CHUFC 2649, Linha da Serra, 15 Aug. 1998, D. M. Borges-Nojosa; CHUFC 2651, Sı´tio Guaramiranga, 6 June 2005, D. M. Borges-Nojosa, P. Cascon, and J. C. L. Melo. Diagnosis.—Distinguished from all congeners by the following combination of characters: (1) rostral slightly visible in dorsal view; (2) head flattened in lateral view; (3) two postoculars; (4) long loreal; (5) seven upper and lower labials; (6) 17 dorsal scale rows at midbody; (7) seven maxillary teeth (five prediastemal and two postdiastemal); (8) ventral scales 154–160 in females and 134–144 in males; (9) subcaudal scales 17–20 in females and 20–25 in males; (10) dorsal ground color in preserved specimens yellowish white to reddish brown, uniformly scattered with dark brown dots; (11) ventral ground color creamish white; (12) moderate size, reaching a maximum SVL of 312 mm in females, and 220 mm in males; (13) moderate tail (8.1–10.4% of SVL) in females and (11.4–15.5% of SVL) in males; (14) hemipenis slightly bilobed, semicapitate, with lateral-tip projections. Comparisons.—Regarding the cis-Andean species of Atractus with 17 dorsal scale rows at midbody, A. ronnie differs from A. arangoi, A. badius, A. canedii, A. davidhardi, A. flammigerus, A. janethae, A. latifrons, A. lucilae, A. major, A. riveroi, A. schach, A. snethlageae, A. torquatus, A. zebrinus, and A. maculatus by having no dorsal bands or blotches (vs. dorsal color pattern with alternating black, red, or white dorsal bands or blotches). It differs from A. collaris, A. favae, A. fuliginosus, A. guentheri, A. heliobelluomini, A. limitaneus, A. univittatus (but see below), and A. zidoki by having a dorsal color pattern uniformly scattered with dark brown dots (vs. dorsal color pattern with paired white spots bordered on paravertebral region or longitudinal stripes along the vertebral, paravertebral portion or flanks). Some specimens of A. univittatus have no vertebral stripe, but A. ronnie is further distinguished from this taxon by having 20--25 subcaudals in males and 17--20 in females, and two postdiastemal teeth (vs. 28--37 and 26--34 subcaudals in males and females, respectively, and one postdiastemal tooth). Atractus ronnie is further distinguished from A. alphonsehogei, A. serranus, A. steymarki, and A. trihedrurus by having a dorsal color pattern uniformly scattered with dark brown dots and immaculate venter (vs. dorsal color

pattern uniformly dark brown or black and venter, except in A. alphonsehogei, uniformly gray or black). It is distinguished from A. edioi by having loreal scale (vs. absence of loreal scale); from A. caxiuana by loreal not contacting internasals (vs. loreal contacting internasals); from A. natans and A. emersoni by venter uniformly creamish white (vs. venter with a wide central black stripe of irregular width); from A. thalesdelemai by having two postoculars (vs. one postocular). The only other species of Atractus that share with A. ronnie a light dorsal ground color uniformly scattered with several dark brown dots and an immaculate creamish white venter are A. pantostictus and A. potschi. Atractus ronnie differs from both by lacking a barely defined (in A. pantostictus) or conspicuous (in A. potschi) dark brown collar on the occipital region. Furthermore, the new species can be distinguished from A. pantostictus by having a midbody dorsal color pattern with small dark dots (one scale length or less), and number of subcaudal scales 17–20 in females and 20–25 in males (vs. midbody dorsal color pattern with moderate to large dots, 1–2 scale length; subcaudals 21–29 in females and 27–34 in males); and from A. potschi by having 17 dorsal scale rows and flattened head in lateral view (vs. 15 dorsal scale rows and curved head in lateral view). Description of the holotype.—Adult female, SVL 275 mm; CL 26 (9.4% SVL); head length 11.7 mm (4.2% SVL) from snout tip to mouth corner; head width 7.5 mm (64% head length) taken at broadest point; interocular distance 4.3 mm; snout–orbit distance 3.2 mm (0.7 times interocular distance); head slightly distinct from body; rostral 2.0 mm wide, broader than high, sub-triangular in frontal view, slightly visible from above; internasals 1.1 mm wide, slightly broader than long; internasal suture sinistral relative to prefrontal suture; prefrontals 2.0 mm wide, as broad as long; supraocular 1.5 mm long, slightly longer than broad; frontal 3.0 mm long, as long as broad, with triangular shape dorsally; parietals 4.8 mm long, about twice longer than broad; nasal completely divided; anterior nasal slightly higher than long; posterior nasal about twice higher than long; loreal 2.1 mm long, about two times longer than high; eye diameter 1.3 mm; pupil semi-elliptical; two postoculars; upper postocular smaller than lower; temporals 2+2; upper posterior temporal elongate, about three times longer than high; seven supralabials, third to fourth contacting orbit; symphysial 1.4 mm wide, about three times broader than long, separated from chinshields by the first pair of

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infralabials; seven infralabials, first four contacting chinshields; chinshields 2.6 mm long, about twice longer than broad; three gular scales separating chinshields from first ventral; dorsal scales 17/17/17 rows, smooth, without apical pits; three preventrals; 154 ventrals; 18 divided subcaudals; cloacal plate single and slightly bilobed. Coloration of holotype in preservative.—Dorsal ground color of head almost grayish brown in dorsal and lateral views, except snout region which is mostly creamish white; supralabials creamish white, except upper portion of the third to fourth scales which are dark brown; dorsal ground color of body creamish white, uniformly scattered with small dark brown dots (1–2 scales long) barely arranged in three weakly defined longitudinal series (one on the paravertebral region and two lateral series, beginning at the level of fourth to sixth dorsal scale row) until the level of eleventh ventral scale; infralabials, gular region, venter, and tail uniformly creamish white. Coloration in life.—Dorsal ground color of head uniformly brown; supralabials yellowish, except upper portions which are generally dark brown; infralabials and gular region uniformly creamish yellow; dorsal ground color of body reddish brown uniformly scattered with dark brown dots; ventral portion of body uniformly creamish yellow. Hemipenis morphology.—Inverted organ (MNRJ 14196) extends to the level of tenth subcaudal. Fully everted and almost maximally expanded hemipenis rendered a slightly bilobed and semicapitate organ; capitular groove visible on the asulcate side, and barely defined on the sulcate and lateral sides; capitular groove located just above the level of sulcus bifurcation; sulcus spermaticus divides approximately on the medial portion of hemipenial body; branches diverge to a centrifugal position terminating on the lateral tips of the organ, which shows projections covered with spinules; sulcus spermaticus bordered by a series of spinules; intrasulcar region somewhat projected upward and covered with enlarged spines; hemipenial body uniformly covered by medium-sized hooked spines, except the most basal portion which has some dispersed spinules; basal portion of the organ naked (Fig. 3). Variation.—Atractus ronnie exhibits significant sexual dimorphism in the number of ventral (U4.2 5 0; P , 0.01; n 5 25) and subcaudal (U–4.0 5 4; P , 0.01; n 5 25) scales, and in SVL (U–0.55 16.5; P , 0.01; n 5 24). Therefore, these data are presented separately for males and females. Other characters

Fig. 3. Asulcate (A) and sulcate (B) sides from hemipenis of Atractus ronnie (MNRJ 14196). Arrows indicate lateral-tip projections. Scale 5 5 mm.

represent variation in both sexes. Largest male SVL 248 mm, tail length 37 mm; largest female SVL 312 mm, tail length 30 mm; tail 11.4–15.5% of the SVL in males, 8.1–10.4% of the SVL in females; preventrals 3 (n 5 22), 2 (n 5 2), or 3 (n 5 1); ventrals 133–144 ( x¯ 5 140.3; SD5 3.3; n 5 14) in males, 154–160 ( x¯ 5 157.4; SD 5 1.6; n 5 11) in females; subcaudals 20–25 (x¯ 5 23.1; SD 5 1.7; n 5 14) in males, 17–20 (x¯ 5 18.9; SD 5 1.1; n 5 11) in females; supralabials contacting loreal 1– 3 (n 5 1 side) or 2–3 (n 5 49 sides); infralabials contacting chinshields 1–3 (n 5 3 sides) or 1–4 (n 5 47 sides); postoculars 1 (n 5 2 sides) or 2 (n 5 48 sides); temporals 1+2 (n 5 45 sides) or 2+2 (n 5 5 sides); anterior dorsal scales rows (counted one head length from the neck) 17 (n 5 10), 16 (n 5 7), or 15 (n 5 8); dorsal scales of tail at level of the second subcaudal 8 (n 5 13), 9 (n 5 12), or 10 (n 5 1); maxillary teeth 7 (n 5 19) or 8 (n 5 1);

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Fig. 4. Geographic distribution of Atractus ronnie (triangles), A. potschi (squares), A. maculatus (circles), A. guentheri (stars), and most northern record of A. pantostictus (asterisk). Open symbols represent the respective type-localities.

hemipenis length (in situ) 7 (n 5 1), 8 (n 5 4), or 9 (n 5 1) subcaudals. The remaining characters are invariable with respect to the holotype. Distribution and natural history.—Known only from Serra de Baturite´, a crystalline mountain covered by a relictual fragment of Atlantic Rainforest in the semiarid Caatinga, state of Ceara´, northeastern Brazil in the municipalities of Pacoti, Guaramiranga, and Mulungu (Fig. 4) with altitudes ranging from 500 to 900 meters. Hoogmoed et al. (1994) and Rodrigues and Borges (1997) provide additional information on topography, vegetation, and climate of the Serra de Baturite´. All specimens were collected on the litter from primary or secondary Atlantic Forest, except for one individual that was burrowed (one-meter depth) in a banana plantation. This species was found active at several hours during the day, and specimens were observed in the field throughout most months of the year, with higher frequency from December to April. Etymology.—The specific epithet ronnie represents the nickname of our friend Dr. Ronaldo Fernandes, Curator of Herpetology at Museu Nacional, Universidade Federal do Rio de Janeiro. We dedicate this species to him for his contributions to the systematics of South American snakes, particularly the genus Atractus. DISCUSSION Savage (1960) defined three species groups of Atractus based on the similarity of characters (e.g., pholidosis, hemipenis, and color pattern). Fernandes (1996) pointed out that several

southern South American species have capitate hemipenis, but this feature does not match with the previous groups defined by Savage (1960). However, Fernandes (1996) noticed that since this character state is plesiomorphic in the tribe Dipsadini, this feature provides no support for the recognition of this assemblage as a natural group. Recently, Schargel and Castoe (2003) reported hemipenis capitation in the northern South American Atractus univittatus and A. ventrimaculatus. However, these authors suggested that this condition is very distinct from those studied by Fernandes (1996). Despite uncertain homology of these conditions, this hemipenis character has been found in several recently studied species (Hoogmoed and Prudente, 2003; Schargel and Castoe, 2003; Passos et al., 2005; Zaher et al., 2005) and might be the most widespread condition in Atractus. Besides the hemipenis capitation, the new species shares a combination of characters found only in some southern South American species of Atractus related to A. reticulatus (Fernandes, 1996; Passos et al., 2005) such as three preventrals, three gular scale rows, seven upper and lower labials, first four pairs of infralabials in contact with chinshields, and ventral color pattern generally uniformly creamish white in preserved specimens. The lateral-tip projections of the hemipenis of A. ronnie are similar to those found by Schargel and Castoe (2003) in A. poeppigi. Nevertheless, the homology of these structures must be further evaluated since the hemipenis of A. poeppigi is non-capitated. Another taxon that shows lateral-tip projections, as well as a semicapitated hemipenis, and 17 dorsal scale rows at midbody is A. pantostictus (P. Passos, unpubl. data), which might suggest a close relationship between this species and A. ronnie. Atractus pantostictus occurs in open formations of the Brazilian Cerrado from the state of Sa˜o Paulo north to the municipality of Porto Nacional, state of Tocantins (Fig. 4), while A. ronnie seems to be restricted to a relictual fragment of the Atlantic Rainforest. The new species inhabits the Serra de Baturite´, which represents an isolated fragment of Atlantic Rainforest in northeastern Brazil surrounded by the semiarid Caatinga. This relictual humid forest shows some endemic amphibians (e.g., Adelophryne baturitensis; Hoogmoed et al., 1994) and reptiles (e.g., Leposoma baturitensis; Rodrigues and Borges, 1997; Placosoma sp. and Amphisbaena sp.; D. M. Borges-Nojosa, unpubl. data), which reinforces the importance of conserving the fragments of the endangered Atlantic Rainforest biome.

PASSOS ET AL.—NEW ATRACTUS FROM NORTHEASTERN BRAZIL MATERIAL EXAMINED Institutional abbreviations are as listed at http://www.asih.org/codons.pdf, except the following institutions that had acronym alterations or were not included in these sources. Venezuela—Museo de Historia Natural, Fundacio´n La Salle (MHNLS), Caracas D.C.. Colombia—Instituto Alexander Von Humboldt (IAvH), Villa de Leyva, Boyaca´; Instituto de Ciencias Naturales, Universidad Nacional de Colombia (ICN), Bogota´ D.C.; Ecuador—Museo de Zoologia, Pontifı´cia Universidad Cato´lica de Ecuador (QCAZ), Quito; Peru—Museo de Historia Natural de la Universidad Mayor de San Marcos (MHNSM), Lima; Museo de Historia Natural de Universidad Nacional de Arequipa (MUSA), Arequipa; Bolivia—Museo de Historia Natural Noel Kempff Mercado (MNKR), Santa Cruz de la Sierra; Coleccio´n Boliviana de Fauna (CBF), La Paz; Argentina—Universidad Nacional del Nordeste (UNNEC), Corrientes; Brazil—Museu de Zoologia da Universidade Federal do Ceara´ (CHUFC), Fortaleza, CE; Museu de Zoologia da Universidade Federal de Alagoas (MUFAL), Maceio´, AL; Museu de Zoologia da Universidade Estadual de Feira de Santana (MZUEFS), Feira de Santana, BA; Instituto Vital Brazil (IVB), Nitero´i, RJ; Instituto Butantan (IBSP), Sa˜o Paulo, SP; Laborato´rio de Zoologia dos Vertebrados, Universidade Federal de Ouro Preto (LZVUFOP), Ouro Preto, MG; Museu de Cieˆncias Naturais da Pontifı´cia Universidade Cato´lica de Minas Gerais (MCNR), Belo Horizonte, MG; Museu de Histo´ ria Natural Capa˜ o da Imbuia (MHNCI), Curitiba, PR; Museu Zoolo´gico Augusto Ruschi, Universidade de Passo Fundo (CRUPF), Passo Fundo, RS; United Kingdom—Natural History Museum (NHM), London. Atractus albuquerquei. Brazil: Para´: Tome´–Ac¸u– Paragominas Highway, Vila Nova, near to Rio Timboteua, MPEG 12946 (holotype); Rondo ˆ nia: Porto Velho, Vila Cachoeirado Samuel, MNRJ 3028. Atractus alphonsehogei. Brazil: Para´: Viseu, Km 75 from Braganc¸a–Viseu Highway, Bela Vista, MPEG 14928 (holotype); Augusto Correia, Fazenda Cacoal, MPEG 9949 (paratype), Braganc¸a, Parada Bom Jesus, MPEG 2221, 8573, 8667 (paratypes); Km 224 (formerly Km 74) from Highway BR 316, MPEG 10093 (paratype); Santa Rosa, Estrada de Vigia, MPEG 12593. Atractus arangoi. Colombia: Putumayo: Mocoa, Puerto Ası´s, MLS 136 (holotype). Atractus canedii. Argentina: Anta: Salta, FML 1082 (holotype). Atractus caixuana. Brazil: Para´: Melgac¸o, Floresta Nacional de Caixuan ˜ a, MPEG 19657 (holotype), 19964, 20128 (paratypes).

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Atractus collaris. Colombia: Caqueta´: Florencia, MLS 1324, 2782. Ecuador: Napo: Pozo Petrolero Zabalo, EPN 5216; Orellana: Yasuni, QCAZ 5980; Sucumbı´os: Cayabeno, QCAZ 983, 986, 1042. Atractus davidhardi. Colombia: Amazonas: Letı´cia, ICN 10096 (holotype). Atractus elaps. Brazil: unknown locality, IBSP 20314; Amazonas: Borba, MNRJ 1523. Colombia: unknown locality, MLS 182; Amazonas: Parque Natural Nacional Amacayacu, IAvH 3211; Boyaca´: Macanal, MLS 2637; Caqueta´: unknown locality, MLS 183; Florencia, MLS 185, 187, 195, 197, 1316– 18, 1322–23, 1326–27, 1739, 2730, 2733–39; Cauca: Santa Rosa, El Carmen, IAvH 4410; Cundinamarca: Medina, MLS 192; Sasaima, MLS 2527; Guaicarano, Paratebueno, MLS 188; Meta: Acacias, MLS 191; San Juan de Arama, IAvH 929; Villaviciencio, MLS 179, 189, 193, 196, 266, 1396, 2054– 55; Rio Ocoa, S Villavicencio, MLS 190; Putumayo: unknown locality, MLS 180. Ecuador: Western Ecuador, NHM 1946.1.6.45 (holotype); Eastern Ecuador, unknown locality, EPN 6892, EPN not cataloged); Napo: Alto Napo, EPN 6856, 8686; Archidona, QCAZ 2101; Rı´o Huataracu, EPN 8687; Orellana: Balsayacu, Parque Sumaco, QCAZ 6502; Fuerte, EPN 7324; Loreto: El Tena´, EPN 8688; Parque Nacional Yasunı´, EPN 2536, QCAZ 3249, 3959; Rı´o Coca, QCAZ 440; Pastaza: Mera, EPN 1175; Montalvo: Andoas, EPN 758; Nueva Vida, Misio´n Agua Santa, QCAZ 345; Puyo, QCAZ 1277; Rı´o Bobonaza, EPN 8678–83; Rı´o Tallı´n, Alto Bobonaza, EPN 8675–77; Sarayacu–Pucayacu, EPN 8685; Sucumbı´os: Lagartococha, EPN 8689; ´ grio, EPN 5781; Shushufindi, QCAZ 3303; Lago A Erroneous localities: Pichincha, Al Occidente, EPN 8692; El Oro: Santa Rosa, EPN 8690–91. Atractus emersoni. Brazil: Amazonas: Benjamin Constant, ICN 10097 (holotype), 10098–99 (paratypes). Colombia: unknown locality, ICN not catalogued. Atractus guentheri. Brazil: Bahia: Camaca˜, MNRJ 6710; Canavieiras, NHM 1946.1.1.76 (holotype). Atractus flammigerus. Colombia: Boyaca´: Macanal, MLS 140. Atractus franciscopaivai. Colombia: Amazonas: La Pedrera, ICN 10100 (holotype), 10101–02 (paratypes). Atractus gaigeae. Ecuador: Napo: Estacio´n Biolo´ gica Sacha, EPN not catalogued; Pastaza: Bobonaza, EPN 5217; Cotosaza, EPN 8693 (paratype); Misio´n, EPN 752. Atractus heliobelluomini. Colombia: Amazonas: La Pedrera, ICN 10103 (holotype). Atractus insipidus. Venezuela: Amazonas: Poste M-1, Rio Araricapara´, Venezuela–Brazil border, MBUCV 3957 (holotype). Atractus janethae. Colombia: Amazonas: La Chorrera, ICN 10104 (holotype).

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Atractus latifrons. Brazil: unknown locality, MNRJ 20315, IBSP 20315; Amazonas: Balbina Hydroelectric Plant, CHUFC 1367; Benjamin Constant, MNRJ 729–32, 1289, 1517–20, 1522; Manaus, MNRJ 726–28; Rio Purus, MNRJ 633; Rondo ˆ nia: Samuel Hydroelectric Plant, CHUFC 1430–32. Bolivia: Beni: Rı´o San Martin, between Rı´o Blanco and Rı´o Negro MNKR 595; Santa Cruz: Guarayos, Urubicha´, MNKR 3436–39; Rı´o San Martin, MNKR 505; Nuflo de Chaves, Oquinquia, Rı´o San Martı´n, MNKR 1021; Velasco, MNKR 218, 520. Colombia: Amazonas: La Pedreira, MLS 210; La Pedreira, Rio Caqueta´, IAvH 1483; Puerto Narin ˜ o, MLS 1319–21; Rio Icara–Parana´, MLS 945; Vaupe´s: Parque Natural Nacional Chiribiquete, Corregimiento Miraflores, IAvH 4264. Peru: Loreto: Maynas, Iquitos, MHNSM 2250, 2292; Mishana, Rio Nanay, MHNSM 2590, 2616; Pebas, MNRJ 2977, 2979, 2981; Requema, MHNSM 2884. Atractus lucilae. Colombia: Amazonas: La Pedreira, Puerto Co´rdoba, ICN 10105 (holotype), 10106–07 (paratypes); Parque Natural Nacional Amacayacu, IAvH 3871; Rio Muriti–Parana´, IAvH 4093, 4097; Vaupe´s: Puerto Bogotano, Lago Taraira, Rio Apaporis, IAvH 1914. Atractus maculatus. Brazil: unknown locality, NHM 51.3.18.15 (holotype); Alagoas: Murici, Mata da Bananeira, MUFAL 474–75; Sa˜o Miguel dos Campos, Usina Ceresta MNRJ 3977. Atractus major. Bolivia: La Paz: La Paz, CBF 2321. Colombia: Amazonas: Letı´cia, MLS 2011; Caqueta´ : Rio Cuemani, Proradam, IAvH 1798; Putumayo: Puerto Caicedo, ICN 10108; Vaupe´s: Estacio´n Biolo´gica Kaparu´, Lago Taraira, Lower Rio Apaporis, IAvH 2909. Ecuador: unknown locality, EPN without number; El Oro: Santa Rosa, EPN 8734; Napo: Alto Sindi, QCAZ 3689; Boca del Coca, EPN 8699; Loreto: Alto Napo, EPN 8695, 8717; Misahiualli, QCAZ 3735; Yasuni, QCAZ 3079; Pastaza: Curaray, Arajuno, Rio Manderoyacu, EPN 6413; Pastaza: EPN 8694; Tiguino, EPN 5146; Sucu´mbios: Boca del Rı´o Cuyabeno, EPN 8697; Comuna Cofa´n Duvuno, EPN 4911; Lago Agrio, EPN 8184; Piso Tropical Oriental, EPN 8696, 8701–02; Zamora–Chinchipe: Macas, QCAZ 2178. Peru: Amazonas: Bagua, MHNSM 2454, 2457; Cusco: Paucartambo, Alto Tono, MUSA 674; Convencio´n, MHNSM 3469; Hua´nuco, MHNSM 2911; Loreto: Loreto, MHNSM 3078; Madre de Dio´s: Tambopata, MHNSM 12129, 16571; Tambopata: Sachavacayoc, MUSA 605; San Martı´n: Mariscas Ca´ceres, MHNSM 2240; San Martı´n, MHNSM 2842; Ucayali: Coronel Portillo, MHNSM 222, 2636, 3005. Atractus natans. Brazil: Amazonas: Uarinı´, Estac¸a˜o Ecolo´gica de Mamiraua´, MPEG 18838,

20213 (paratypes); Para´: Caxiuana˜, MPEG 18836 (holotype). Atractus occipitoalbus. Colombia: Putumayo: El Orito. Ecuador: Morona–Santiago: Carretera Limo´ n–Macas, QCAZ 7263–64; Rio Nepano, Mendez, EPN 8729; Napo: Rı´o Hollio, QCAZ 6268; Pastaza: Arajuno, Alto Napo, EPN 8719–20; Cabeceiras del Arajuno, EPN 8723; Rı´o Bobonaza, EPN 8724–27; Rı´o Oglan, Alto Curaray, EPN 8721–22; Puyo: Santana, EPN 6474; Sucu´mbios: La Bonita, QCAZ 2779. Atractus pantostictus. Brazil: Minas Gerais: Belo Horizonte, MHNCI 787, MNRJ 6474, 10909, IBSP 40757, 58592, MCNR 13, 27, 35, 88, 101, 129, 139, 145–48, 254, 453–54, 459, 516, 726, 929–41; Campo do Meio, IBSP 50476; Conselheiro Lafaiete, LZVUFOP 501, 614, 627; Itabirito, LZVUFOP 118, 158, 274, 282, 331, 426, 466–67, 622; Machado, IBSP 57138; Ouro Branco, LZVUFOP 421, 579–80; Ouro Preto, LZVUFOP 26, 27–29, 33, 56, 82, 382, 425; Pirapora, Fazenda Triaˆ ngulo Formoso, MNRJ 4459 (paratype); Uberlaˆndia, IBSP 54604–05; Distrito Federal: Brası´lia, Jardim Zoolo´ gico, MNRJ 4460–66; Goia´s: Alianc¸a do Norte, IBSP 43954; Cana Brava, IBSP 26711; Minac¸u, IBSP 51433–34; Sa˜o Paulo: Areais, Fazenda Vargem Grande, IBSP 40404; Barueri, IBSP 45208; Borace´ia, MZUSP 3157, 3158 (paratype); Campo Lindo, IBSP 9472, 49225; Campo Limpo Paulista, IBSP 44152, 54651, 54896; Francisco Morato, IBSP 54634; Franco da Rocha, IBSP 27305, 42093, 54844 (holotype); Jales, MZUSP 4094; Jundiaı´, MNRJ 6496, IBSP 2728, 10068–69, 42646, 42664, 43192, 45624, 46228, 49267, 54235 (paratype), 54512, 54661; Jarinu, IBSP 41427; Itaperuna da Serra, IBSP 54699; Orlaˆndia, IBSP 44537; Paranapiacaba, MZUSP 2811; Sa˜o Jose´ do Rio Preto, IBSP 40028; Sa˜o Jose´ dos Campos, IBSP 27231, 27233, 29098, 37527, 40355, 44527, 45784, 45803, 45807; Sa˜o Paulo, Perus, IBSP 54655, 54886–88; Pico do Jaragua´, IBSP 42404; Pirituba, IBSP 42485, 53545, 54641; Va´ rzea Paulista, IBSP 9862, 32501, 40855, 40857, 45167; Tocantins: Porto Nacional, Luı´s Eduardo Magalha˜es Hydroelectric Plant, IBSP 64952–64966. Atractus poeppigi. Brazil: Amazonas: Alto Rio Negro, MNRJ 10837; Borba, MNRJ 1523. Colombia: Amazonas: Letı´cia, MLS 133, 1313–15. Peru: Amazonas: Bagua, MHNSM 2380, 2447; Pasco: Cerro de Pasc¸o, Oxapampa, MHNSM 3485; San Martı´n: San Martı´n, MHNSM 3133, 3337; Tarapoto, MHNSM 3278. Atractus potschi. Brazil: Alagoas: Maceio´, IBSP 48438 (holotype); Bahia: Feira de Santana, Jaı´ba, MZUEFS 454; Feira de Santana, Faz, Brasileiro, MZUEFS 682; Teofilaˆndia, IBSP 57119; Sergipe,

PASSOS ET AL.—NEW ATRACTUS FROM NORTHEASTERN BRAZIL Salgado, MZUSP 7001, 7195–97, 7275–81 (paratypes); Sa˜o Cristo´va˜o, MNRJ 14057–58. Atractus punctiventris. Colombia: Meta: Villavicencio, MLS 254, formerly MLS 102 (holotype), 255–56. Atractus reticulatus. Argentina: Corrientes: Galarza, Santo Tome, UNNEC 7588; San Miguel, UNNEC 256–57; Formosa: Naicneck, Colonia Aborigine, UNNEC 7219. Brazil: Parana´: unknown locality, MNRJ 9820; Sa˜o Jose´ dos Pinhais, MNRJ 9086; Rio Grande do Sul: Candela´ria, MNRJ 1261; Entre Rios: CRUPF 309; Nicolau Vergueiro, CRUPF 176; Passo Fundo, CRUPF 96, 199, 213, 224–26, 249, 284, 304, 343, 376, 401, 416, 590, 686, 819, 829–30, 1064, 1204; Sa˜o Paulo: Sa˜o Paulo, MNRJ 1524. Atractus riveiroi. Venezuela: Amazonas: Marahuaca, Campo Temiche, MBUCV 7175, formerly UPR 49 [r-20] (holotype). Atractus schach. Brazil: Acre: Porto Walter, Rio Jurua´, MPEG 20376; Rio Branco, IBSP 43394; Amazonas: Manaus, IBSP 49430; Presidente Figueredo, Rio Uatuma˜, Hydroelectric Plant, MPEG 17495, 17527; Maranha˜o: Nova Vida, 25 Km from Rio Gurupi, MPEG 10347, 12255, 15790–91; Para´: Bom Jesus da Braganc¸a, MPEG 11374; Capita˜o Poc¸o, MPEG 13267; Caxiuana˜, MPEG 20071; Km 198 PA-232 Highway, MPEG 11569, 15165; Viseu, Bela Vista, MPEG 3713, 10100, 16300. Peru: Corrientes, IBSP 49433. Atractus serranus. Brazil: unknown locality, IBSP 32857; Sa˜o Paulo: Campinas, IBSP 50861; Cotia, IBSP 55698; Cubata˜o, IBSP 9706; Engenheiro Marsilack, IBSP 9075–76, 9088–89; Guarulhos, IBSP 26999; Guarulhos, Km 21 Presidente Dutra Highway, IBSP 27147, 27862; Paranapiacaba, IBSP 7200, 10589, 18645, 23518; Ribeira˜o Pires, IBSP 10136; Rio dos Campos, IBSP 9267, 9437–38, 10136; Rio Grande da Serra, IBSP 54636, 54974; Santo Amaro, Marink–Santos Highway, IBSP 4852; Santo Andre´, IBSP 53630, 55252; Santo Andre´, Km 38 Santos–Jundiaı´ Highway, IBSP 42947; Saleso´polis, Estac¸a˜o Biolo´gica da Borace´ia, MZUSP 2193; Sa˜o Joa˜o, IBSP 7002; Sa˜o Luiz do Paraitinga, IBSP 53924; Serra de Paranapiacaba, IBSP 7239 (holotype); Tapiraı´, IBSP 42963. Atractus snethlageae. Bolivia: Pando: Manuripi, Reserva Nacional de la Vida Silvestre Amazoˆnica, MNKR 3275. Brazil: unknown locality, MNRJ 9842; Amazonas: Benjamin Constant, IBSP 33369; Maranha˜o: Nova Vida, MPEG 14986, 15422 (paratype); Para´: Ananindeua, Lago Azul, MPEG 16383–85 (paratypes); Bele´m, Ilha do Mosqueiro, MPEG 2595 (paratype); Santa Ba´rbara, Benevides, MPEG 3955 (paratype); Sa˜o Joa˜o da Pratinha, MPEG 10137 (paratype); Viseu, Coloˆnia Nova, BR 316 Highway, 10 Km from Rio Gurupi, MPEG 10131 (holotype); Viseu, Bela

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Vista, MPEG 2543, 6845, 15973 (paratypes); Tucuruı´, IBSP 46454; Rondoˆnia: Ariquemas, IBSP 41530; Porto Velho, Samuel Hydroelectric Plant, CHUFC 1399. Colombia: Amazonas: ICN 10109–10110. Peru: Cajamarca: Jae´n, MHNSM 3390; San Martı´n: San Martı´n, MHNSM 3338; Corrientes, IBSP 49431–32. Ecuador: Eastern Andes, unknown locality, EPN 8718; Napo: Misahualli, QCAZ 3476–77; San Francisco de Borga, QCAZ 1320; San Rafael, El Chaco, QCAZ 1493–94. Atractus steymarki. Venezuela: Bolivar: Chimanta, Churi–Tepui, MHNLS 11004; El Dorado, MBUCV 3872. Atractus paraguayensis. Argentina: Corrientes: San Luis del Palmar, Costa Grande, UNNEC 84; Santo Tome, UNNEC 4979. Brazil: Parana´ : Pinha˜o, Rio Jorda˜o, MCP 7185, 7211, 7365; Pinha˜o, Rio Parana´, MCP 7364; Rio Grande do Sul: Carazinho, CRUPF 1180; Chapeco´, MCP 14013; Colorado, CRUPF 1196; Derrubadas, MCP 12387; Getu´lio Vargas, CRUPF 64; Ibiraquita˜, CRUPF 587; Ijuı´, MCP 13726–32; Ipira, MCP 2913; Mato Castellano, CRUPF 289, 516, 991–92, 1094; Pinheiro Machado, CRUPF 257; Planalto, MCP 5898–99; Planalto, 4u Sec¸a˜o, MCP 5915, 5997; Porto Maua´, MCP 11609, 11611, 11623; Porto Vera Cruz, MCP 11670; Santo ˆ ngelo, MCP 12516–17; Tapejara, CRUPF 477, A 814; Santa Catarina: Concordia, Entre Rios, MCP 2912; Peritiba, MCP 2939; Piratuba, MCP 2893– 94, 2897, 2902. Atractus thalesdelemai. Brazil: Rio Grande do Sul: Passo Fundo, Fazenda Corporac¸ a˜ o da Brigada Militar, MNRJ 10052 (holotype), 10053–54, 10080–81 (paratypes); Passo Fundo, Jardim Botaˆnico, CRUPF 172, 801 (paratypes); Passo Fundo, Vera Cruz, CRUPF 405. Atractus torquatus. Brazil: Amazonas: Manaus, Km 80 BR 174, MZUSP 8533–34; Manaus, Reserva Florestal Adolpho Ducke–INPA, MZUSP 8455, 9588; Novo Aira˜o, MZUSP 8205; Presidente Figueredo, Balbina Hydroelectric Plant, MPEG 17516. Colombia: Vaupe´s: ICN 10111. Venezuela: Amazonas: Frente 20, MHNLS 14488; Bolivar: El Dorado, MBUCV 1406. Atractus trihedrurus. Brazil: Parana´: Guaratuba, UHE Guaricana, MHNCI 851; Piraquara, IBSP 3067 (paratype); Santa Catarina: Campo Alegre, IBSP 32664; Campo Grande, Rio Negrinho, IBSP 32367, 32369; Sa˜o Bento do Sul, IBSP 9111, 3098 (holotype), MZUSP 9439; Sa˜o Paulo: Guapiara, IBSP 33717, 34360, 34409; Ibiu´na, IBSP 46476, 46658, 56474; Juquitiba, IBSP 33930, 44676, 46604, 53565, 54703, 62215, 62860, 68219; Miracatu, IBSP 58763; Piedade, IBSP 49752, 50280, 58413; Ribeira˜ o Pires, IBSP 31188, 42906; Tapiraı´, IBSP 42222, 46605, 52636, 56938.

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Atractus trilineatus. Brazil: Roraima: Boa Vista, Taiano region, Colo ˆ nia Coronel Mota, MPEG 479. Venezuela: Monagas: Rı´o Guarapiches, EBGG 2602; Sucre: Sabaneta del Pilor, MHNLS 13333. Atractus univittatus. Colombia: Meta: unknown locality, ICN 8264; Acacias, ICN 10651; Cubaral, ICN 10695; Lomalinda, IAvH 950, 954; Parque Natural Nacional Cueva de Los Guacharos, IAvH 1002; Restrepo, ICN 6567, 6902, MLS 2530, 2961; Villaviciencio, IAvH 2466, ICN 2699, 2716, 2996, 6129, 7104. Venezuela: Barina: Barinitas, MHNLS 16762; Carabobo, San Rafael Hydroelectric Plant, MHNLS 5622; Cojedes: San Carlos, Manrique, MHNLS 13834; Distrito Capital: Car´ vila, Canales del acas, Parque Nacional El A Naigata, MHNLS 11397; Parque Nacional El ´ vila, La Guaira´, MBUCV 2030; Miranda: GuaiA coco, MBUCV 8355. Atractus zebrinus. Brazil: unknown locality, IVB 1548, NHM 61.4.18.12, 61.4.18.13; Bahia: Porto Seguro, IBSP 57202; Espı´rito Santo: Santa Tereza, MNRJ 733–34; Minas Gerais: Bocaina de Aiuruoca, IBSP 6463; Camanducaia, IBSP 28868–69, 32453, 40106, 41443, 44794, 45431, 45620, 45622, 45691, 46286, 51491, 51683, 54818, 67697; Delfim Moreira, IBSP 57476; Extrema, IBSP 68962; Itabira, IBSP 71376; Itamonte, IBSP 43154; Liberdade, MNRJ 6497; Monte Verde, IBSP 33499, 51683, 53839, 60927, 61924; Ouro Preto, LZVUFOP 19, 27, 45, 86, 133, 369; Paraiso´polis, IBSP 71285; Sa˜o Gonc¸alo do Rio Abaixo, Peti, MNRJ 9298; Sapucaı´–Mirim, IBSP 56953, 61385, 62660–61, 66361, 70432; Parana´: Campo Largo, MHNCI 4818; Votuverava, IBSP 12893; Rio de Janeiro: Cachoeiras de Macacu´, MNRJ 7064–65; Itaboraı´, MHNCI 1295; Nova Friburgo, MNRJ 6322, 6498; Petro´polis, IVB 1203, 2485, MNRJ 4467–70, 10091; Rio de Janeiro, NHM 54.4.18.12; Tereso´polis, IBSP 41054, 41059, MNRJ 12899; Tereso´ polis, Parque Nacional da Serra dos ´ rga˜os, MNRJ 6495; Visconde de Maua´, IBSP O 48839; Santa Catarina: Peritiba, IBSP 44049; Sa˜o Paulo: Apiaı´, Serra Formosa, IBSP 52316; Borace´ia, MZUSP 2194; Campos do Jorda˜o, IBSP 7899, 44190, 50862, 54326, 68189; Cubata˜o, IBSP 45193; Cunha, IBSP 46348; Guapiara, Fazenda Oriente, IBSP 33717; Joano´polis, IBSP 55090, 57017–18, 58310; Ribeiras, Fazenda Cobalto, IBSP 43733; Santo Anto ˆ nio do Pinhal, IBSP 25020–21; Santo Anto ˆ nio do Pinhal, IBSP 21949; Santo Anto ˆ nio do Pinhal, Engenheiro Lefe´vre Road, IBSP 16435; Sa˜o Bernardo do Campo, IBSP 56207; Sa˜o Jose´ do Barreiro, IBSP 70789–91, 71018; Sa˜o Paulo, IBSP 4551; Treˆs Irma˜os Hydroeletric Plant, IVB 1519. Atractus zidoki. Brazil: Amapa´: Serra do Navio, MPEG 16437, MPEG not catalogued; Para´ : Capita˜o Poc¸o, MPEG 13265–66, 13268.

ACKNOWLEDGMENTS We are grateful to the following personnel for permission and/or facilities to examine specimens under their care: C. Aguilar and J. Santa-Gadea (MHNSM), A. Almendarı´z (EPN), F. Bilbao (EBRG), R. Casallas and A. Rodriguez (MLS), J. Ce´spedez (UNNEC), L. Coloma and S. Ron (QCAZ), M. Di-Bernardo (PUCRS), C. Ferreira (MBUCV), F. Franco (IBSP), L. Gonzales and R. Montan ˜ o (MNKR), F. Junca´ (MZUEFS), J. Lynch (ICN), A. Melgarejo (IVB), J. Moura-Leite (MHNCI), L. Nascimento (MCNR), M. Pires (LZVUFOP), D. Perico (IAvH), A. Prudente (MPEG), G. Scrochii (FML), C. Sen ˜ aris and G. Rivas (MHNLS), M. Wilkinson and C. McCarthy (NHM), H. Zaher (MZUSP), and N. Zanella (CRUPF); P. Nascimento for rendering the line art; J. Pombal, Jr. for valuable comments on the manuscript. D. Borges-Nojosa thanks Instituto Brasileiro de Meio Ambiente e dos Recursos Naturais Renova´veis (IBAMA) for the permission to collect specimens (processes 368/98-61 and 02001.000368/98-61). Financial support was provided by Conselho Nacional de Desenvolvimento Cientı´fico e Tecnolo´gico (CNPq) and Coordenac¸a˜o de Aperfeic¸oamento de Pessoal de Nı´vel Superior (CAPES). LITERATURE CITED CUNHA, O. R., AND F. P. NASCIMENTO. 1983. Ofı´dios da Amazo ˆ nia. XX. As espe´cies de Atractus Wagler, 1828, na Amazo ˆ nia Oriental e Maranha˜o (Ophidia: Colubridae). Boletim do Museu Paraense Emı´lio Goeldi 123:1–38. DOWLING, H. G. 1951. A proposed standard system of counting ventrals in snakes. British Journal of Herpetology 1:97–99. DOWLING, H. G., AND J. M. SAVAGE. 1960. A guide to the snake hemipenis: a survey of basic structure and systematic characteristics. Zoologica 45:17–28. DOWNS, F. L. 1967. Intrageneric relationships among colubrid snakes of the genus Geophis Wagler. Miscellaneous Publications, Museum of Zoology, University of Michigan 131:1–193. FERNANDES, R. 1995. A new species of snake in the genus Atractus (Colubridae: Xenodontinae) from northeastern Brazil. Journal of Herpetology 29:416–419. FERNANDES, R. 1996. Variation and taxonomy of Atractus reticulatus complex (Serpentes: Colubridae). Comunicac¸o˜es do Museu de Cieˆncias e Tecnologia da PUCRS (Se´rie Zoologia) 8:37–53. FERNANDES, R., AND J. A. S. ARGOˆLO. 1999. Rediscovery of Atractus guentheri (Wucherer, 1861) (Serpentes, Colubridae) in Southeastern Bahia, Brazil. Boletim do Museu Nacional, Nova Se´rie, Zoologia 397:1–5. FERNANDES, R., E. M. X. FREIRE, AND G. PUORTO. 2000. Geographic variation of the Brazilian Atlantic Rainforest snake Atractus maculatus (Gu¨nther,

PASSOS ET AL.—NEW ATRACTUS FROM NORTHEASTERN BRAZIL 1858), with revalidation of Rhabdosoma zebrinum Jan, 1862 (Serpentes: Colubridae). Boletim do Museu Nacional, Nova Se´rie, Zoologia 419:1–8. GIRAUDO, A. R., AND G. J. SCROCCHI. 2000. The genus Atractus (Serpentes: Colubridae) in northeastern Argentina. Herpetological Journal 10:81–90. HOOGMOED, M. S. 1980. Revision of the genus Atractus in Surinam, with the resurrection of two species (Colubridae, Reptilia). Notes on the herpetofauna of Surinam VII. Zoologische Verhandelingen 175:1–47. HOOGMOED, M. S., D. M. BORGES, AND P. CASCON. 1994. Three new species of the genus Adelophryne (Amphibia: Anura: Leptodactylidae) from Northeastern Brazil, with remarks on the other species of the genus. Zoologische Mededelingen 68:271–300. HOOGMOED, M. S., AND A. L. C. PRUDENTE. 2003. A new species of Atractus (Reptilia: Ophidia: Colubridae: Dipsadinae) from the Amazon forest region in Brazil. Zoologische Mededelingen 77:425–439. LIMA, F. M. P., I. BIONDI, AND F. A. JUNCA´. 2000. Atractus potschi. Geographical distribution. Herpetological Review 31:254. MARTINS, M., AND M. E. OLIVEIRA. 1993. The snakes of the genus Atractus Wagler (Reptilia: Squamata: Colubridae) from the Manaus region, central Amazonia, Brazil. Zoologische Mededelingen 67:21–40. MYERS, C. W. 2003. Rare snakes—five new species from eastern Panama: reviews of northern Atractus and southern Geophis (Colubridae: Dipsadinae). American Museum Novitates 3391:1–47. MYERS, C. W., AND J. E. CADLE. 2003. On the snake hemipenis, with notes on Psomophis and techniques of eversion: a response to Dowling. Herpetological Review 34:295–302. MYERS, C. W., AND J. A. CAMPBELL. 1981. A new genus and species of colubrid snake from the Sierra Madre del Sur of Guerrero, Mexico. American Museum Novitates 2708:1–20. PASSOS, P., R. FERNANDES, AND N. ZANELLA. 2005. A new species of Atractus (Serpentes: Colubridae) from southern Brazil. Herpetologica 61:209–218. PESANTES, O. 1994. A method for preparing hemipenis of preserved snakes. Journal of Herpetology 28:93–95. RODRIGUES, M. T., AND D. M. BORGES. 1997. A new species of Leposoma (Squamata: Gymnophthalmi-

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dae) from a relictual forest in semiarid Northeastern Brazil. Herpetologica 53:1–6. SAVAGE, J. M. 1960. A revision of the Ecuadorian snakes of the genus Atractus. Miscellaneous Publications, Museum of Zoology, University of Michigan 112:1–86. SCHARGEL, W. E., AND T. C. CASTOE. 2003. The hemipenis of some fossorial snakes of the genus Atractus (Colubridae: Dipsadinae) with comments on variation in the genus. Journal of Herpetology 37:718–721. VIDAL, N., A.-S. DELMAS, P. DAVID, C. CRUAUD, A. COULOUX, AND S. B. HEDGES. 2007. The phylogeny and classification of caenophidian snakes inferred from seven nuclear protein-coding genes. Comptes Rendus Biologies 330:182–187. ZAHER, H. 1999. Hemipenial morphology of the South American xenodontine snakes, with a proposal for a monophyletic Xenodontinae and a reappraisal of colubroid hemipenes. Bulletin of the American Museum of Natural History 240:1–168. ZAHER, H., I. SOUZA, D. J. GOWER, E. HINGST-ZAHER, AND N. JORGE DA SILVA. 2005. Redescription of Atractus albuquerquei (Serpentes: Colubridae: Dipsadinae), with comments on geographical distribution and intraspecific variation. Pape´is Avulsos de Zoologia 45:19–32. ZAR, J. H. 1999. Biostatistical Analysis, Fourth Edition. Prentice-Hall, Englewood Cliffs, New Jersey.

(PP) DEPARTAMENTO DE VERTEBRADOS, MUSEU NACIONAL/UFRJ, QUINTA DA BOA VISTA, 20940040, RIO DE JANEIRO, RJ, BRAZIL ; (DSF) DEPARTAMENTO DE ZOOLOGIA, INSTITUTO DE BIOLOGIA, UNIVERSIDADE FEDERAL DO RIO DE JANEIRO, ILHA DO GOVERNADOR, 21941-590, RIO DE JANEIRO, RJ, BRAZIL; AND (DMB-N) NU´CLEO REGIONAL DE OFIOLOGIA DA UFC, DEPARTAMENTO DE BIOLO´ , CAMPUS GIA, UNIVERSIDADE FEDERAL DO CEARA PICI, BLOCO 905, 60455-760, FORTALEZA, CE, BRAZIL. E-mail: (PP) [email protected]. Send reprint requests to PP. Submitted: 10 July 2006. Accepted: 1 April 2007. Section editor: S. A. Schaefer.