Herpetologica, 61(2), 2005, 209–218 Ó 2005 by The Herpetologists’ League, Inc.
A NEW SPECIES OF ATRACTUS (SERPENTES: COLUBRIDAE) FROM SOUTHERN BRAZIL PAULO PASSOS1,3, RONALDO FERNANDES1, 1
AND
NOELI ZANELLA2
Departamento de Vertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, Rio de Janeiro, RJ 20940-040, Brazil 2 Instituto de Cieˆncias Biolo´gicas, Campus I, BR 285, Universidade de Passo Fundo, Passo Fundo, RS, 99001-970, Brazil
ABSTRACT: A new species of Atractus is described from grasslands of the State of Rio Grande do Sul, southern Brazil. The new species is distinguished from all congeners by the combination of a single postocular, long loreal, six supralabials, generally six infralabials, 17 dorsal scales rows, six or seven maxillary teeth, and a dorsal color pattern in preservative uniformly grayish-brown with a creamish-white temporal region. Comparisons of the new species are made with all other Atractus species, and its affinities with A. reticulatus and allied species are suggested based on the morphology of the hemipenis. Key words: Atractus thalesdelemai new species; Dipsadinae; Southern Brazil; Systematics; Taxonomy
THE XENODONTINE snake genus Atractus Wagler, 1828 is distributed widely throughout South America, occurring from Panama to Argentina (Giraudo and Scrocchi, 2002; Myers, 2003). Atractus is a genus of fossorial snakes closely related to Adelphicos and Geophis (Downs, 1967; Fernandes, 1995b; Savage, 1960) and composed of nearly 100 species, most with highly restricted distributions (Fernandes, 1995a,b). Currently, the taxonomic status of most species are confused, and there have been attempts of taxonomic revisions on regional scales only (e.g., Cunha and Nascimento, 1983; Fernandes, 1995c; Hoogmoed, 1980; Martins and Oliveira, 1993; Myers, 2003; Roze, 1961; Savage, 1960). Eight species of Atractus currently are recognized in southern South America (Fernandes, 1995a,c), and three of them (Atractus reticulates, A. taeniatus, and A. zebrinus) occur in Rio Grande do Sul, the southernmost state of Brazil (Lema, 1994). While examining Atractus specimens from Rio Grande do Sul, we discovered a number of misidentified specimens referred to as Atractus reticulatus that represent a new taxon, which is described herein. MATERIALS AND METHODS We examined more than 500 Atractus specimens from South America, acronyms and specimens examined are listed in the 3
CORRESPONDENCE: e-mail,
[email protected] 209
Appendix. Data for Atractus paraguayensis was augmented with information from Giraudo and Scrocchi (2000). Terminology for Atractus cephalic shields follows Savage (1960), whereas the method of counting ventral scales follows Dowling (1951); description of the hemipenis follows Dowling and Savage (1960), as augmented by Myers and Campbell (1981). Sex was determined by the presence or absence of hemipenes detected through a ventral incision at the base of the tail. Measurements were taken with a dial caliper to the nearest 0.1 mm, except for snout–vent (SVL) and tail lengths (TL), which were taken with a flexible ruler to the nearest millimeter. Analysis of variance (ANOVA) using segmental counts (ventral and subcaudal scales) was employed to assess the presence or absence of sexual dimorphism within each taxon (Table 1), except for A. canedii and A. paraguayensis, in which the sample size was too small to allow the use of statistical tests. Assumptions of univariate normality and homoskedasticity were evaluated using Kolmogorov-Smirnov’s test and the Levene’s test, respectively (Zar, 1999). In cases where characters showed no sufficient variation to justify such assumptions, or when the sample size was too small, nonparametric tests (MannWhitney U-test) were performed (Zar, 1999). The following characters were employed in the statistical analyses: number of ventral; subcaudal; supralabial; and infralabial scales; and number of maxillary teeth.
210
HERPETOLOGICA
[Vol. 61, No. 2
TABLE 1.—Sexual dimorphism values for segmental counts among southern South American species of Atractus. Species
Ventral scales
Subcaudal scales
A. pantostictus A. reticulatus A. serranus A. taeniatus A. thalesdelemai A. trihedrurus A. zebrinus
F1,71 5 172.60; P , 0.001 F1,247 5 536.12; P , 0.001 F1,14 5 48.57; P , 0.001 F1,42 5 101.90; P , 0.001 U1,4 = 0; P < 0.02 F1,30 5 93.04 P , 0.001 F1,39 5 42.72 P , 0.001
F1,68 5 177.17; P , 0.001 F1,246 5 408.05; P , 0.001 F1,14 5 57.16; P , 0.001 F1,39 5 135.21; P , 0.001 U1,4 = 0.5; P < 0.03 F1,30 5 58.03 P , 0.001 F1,53 5 72.16 P , 0.001
DESCRIPTION OF NEW SPECIES Atractus thalesdelemai sp. nov. Holotype.—MNRJ 10052 (Figs. 1–2), adult male, from Fazenda Corporac¸a˜o da Brigada Militar, Municipality of Passo Fundo (288 149 300 S, 528 219 270 W), State of Rio Grande do Sul, Brazil, 30 January 2001, collected by Noeli Zanella in pitfall traps. Paratypes.—Seven specimens from Municipality of Passo Fundo, State of Rio Grande do Sul: MNRJ 10053–4, two females from Fazenda Corporac¸a˜o da Brigada Militar, 26–30 November 2001, collected by Noeli Zanella in pitfall traps; MNRJ 10080–1 two males from Fazenda Corporac¸a˜o da Brigada Militar, collected by Noeli Zanella in pitfall traps, on 15 August 2001 and 25 August 2002, respectively; CRUPF 172, female from Jardim Botaˆnico, 5 August 1994 collected by Lusberta Portilho; CRUPF 405, female from Vera Cruz, 12 October 1998 collected by Rafael Bibiano; CRUPF 801, male from Jardim Botaˆnico, 12 February 2001 collected by Nicolau Pot. Diagnosis.—The new species is distinguished from all congeners by the combination of: (1) a single postocular; (2) long loreal; (3) six supralabials and generally six infralabials; (4) 17 dorsal scales rows; (5) six or seven maxillary teeth; and (6) a dorsal color pattern in preservative uniformly grayish brown with a creamish white temporal region. Refer to Table 2 for diagnostic features related to the other southern South American species of Atractus. Of all species of Atractus, the new species shares a single postocular (instead of two) with A. balzani, A. carrioni, A. elaps, A. latifrons, A. major, A. microrhynchus, A. natans, A. occiptoalbus, A. pamplonensis, A. poeppigi, A. sanctamartae, A. torquatus, and A. werneri. Of these, the new species may be distinguished from A. elaps, A. latifrons, and A. poeppigi (the
Atractus elaps species group of Savage [1960]) by having a small rostral not as large as the prefrontal (instead of large, equal of larger than the prefrontals in the A. elaps group), internasals much less than half as large as the prefrontal (instead of almost as large as the prefrontal in the A. elaps group), and prefrontal at least one and half times broader than long (instead of about as broad as long in the A. elaps group), and a long loreal much larger than the postnasal (instead of short, squarish, and about as long as the postnasal in the A. elaps species group). Furthermore, the new species can be distinguished from the remaining species by having 17 dorsal scales rows, instead of 15 in A. carrioni and A. occiptoalbus, and by having six supralabials, instead of seven or eight in A. major, A. microrhynchus, A. natans, A. pamplonensis, A. sanctamartae, A. torquatus, and A. werneri. Finally, Atractus thalesdelemai can be distinguished from A. balzani by featuring smaller tail size (maximum tail length 38 mm vs. tail length 50 mm), number of ventral scales (149–154 in males and 165–167 in females vs. 159), smaller number of subcaudals (26–30 in males and 22–26 in females vs. 32), dorsum color pattern formed by scales lighter in the posterior edge (instead of lighter in the center), and a homogenously creamish white venter (instead of yellowish speckled with brown). Compared to species that might occur in its currently known distribution, Atractus thalesdelemai can be distinguished from A. reticulatus and A. taeniatus by having 17 dorsal scales rows (instead of 15), six supralabials (instead of seven), no nuchal collar and a grayish brown reticulate dorsal pattern (instead of a nuchal collar and either a vertebral stripe pattern with gray [red in live specimens] reticulation in A. reticulatus or a vertebral streak with lateral projections sometimes creating bands in ?1
A. zebrinus
A. trihedrurus
A. thalesdelemai
A. taeniatus
A. serranus
A. reticulatus
A. paraguayensis
A. canedii A. pantostictus
Species
17
17
$ (23)
17
$ (16)
# (18)
17
17
$ (4)
# (16)
17
15
$ (10)
# (2)
15
17
$ (12)
# (4)
17
15
$ (142)
# (4)
15
# (107)
15 15
17
$ (37)
# (1) $ (3)
15 17
Dorsal scales
# (1) # (36)
Sex (n)
167 x 5 149.9 6 4.7 142–159 x 5 165.1 6 5.2 152–176 163 x 5 162.8 6 4.0 157–166 x 5 142 6 3.9 130–151 x 5 153.3 6 3.8 142–162 x 5147.5 6 1.7 146–150 x 5 157.6 6 2.7 154–163 x 5 144.2 6 2.7 141–147 x 5 157.7 6 4.7 149–165 x = 152.5 ± 2.1 151–154 x = 165.5 ± 1.0 165–167 x 5 142.1 6 4.8 132–151 x 5 154.6 6 4.3 147–160 x 5 144.1 6 4.2 136–151 x 5 154.6 6 5.6 145–170
Ventral scales
50 x 5 29.1 6 1.7 27–34 x 5 23.7 6 1.7 21–29 30 x 5 23 6 1 22–24 x 5 28.7 6 1.85 25–34 x 5 23.7 6 2.0 19–30 x 5 28.7 6 2.1 27–31 x 5 19.8 6 2.0 14–22 x 5 27.7 6 2.36 26–31 x 5 23.3 6 1.5 21–25 x = 28.5 ± 0.7 28–29 x = 23.5 ± 1.73 22–26 x 5 26 6 1.6 23–29 x 5 20.1 6 2.0 17–24 x 5 25.8 6 2.9 20–32 x 5 18.8 6 2.37 15–27
Subcaudal scales
x 5 7.1 6 0.3 7–8 x 5 6.9 6 0.1 6–7 x 5 7.0 6 0.1 7–8
7
6
6
7
x 5 7.0 6 0.1 7–8 7
x 5 7.0 6 0.2 5–8 x 5 6.9 6 0.2 6–8 7
7 7
6 x 5 6.9 6 0.2 6–7 7
Supralabial scales
7
x 5 6.9 6 0.1 6–7 7
7
x 5 6.9 6 0.2 6–7 x 5 6.9 6 0.4 6–8 x = 6.5 ± 0.7 6–7 6
7
x 5 6.9 6 0.2 6–8 x 5 6.9 6 0.2 6–8 7
x 5 7.1 6 0.1 7–8 7 7
7 7
Infralabial scales
x 5 9.2 6 0.4 9–10 x 5 8.3 6 0.7 7–9 x 5 8.3 6 1.1 7–10
x = 6.1 ± 0.2 6–7 9
x 5 8.1 6 0.2 8–9 6
x 5 7.3 6 0.5 7–8 x 5 7.3 6 0.5 6–8 x 5 9.0 6 0.4 8–10 x 5 9.0 6 0.6 8–11 8
— x 5 7.1 6 0.2 7–8 x 5 7.0 6 0.3 6–8 7 7
Maxillary teeth
2
2
2
2
1
1
2
2
2
2
2
2
2 2
2
2 2
Postocular scales
8
5
7
6
5
4
3
2
1
Dorsal color pattern
TABLE 2.—Selected diagnostic characters for male and female specimens of the species of the genus Atractus occurring in southern South America. Data for Atractus paraguayensis augmented from Giraudo and Scrocchi (2000). When possible, values indicate mean 6 SD, and range. Abbreviations for dorsal color pattern are the follow: 1 5 striped; 2 5 banded; 3 5 vertebral and lateral blotches forming stripes; 4 5 reticulated, sometimes with a vertebral line; 5 5 juveniles banded, adults uniform; 6 5 vertebral streak with lateral projections, sometimes creating bands; 7 5 uniform juveniles and adults; 8 5 transversal bands.
June 2005] HERPETOLOGICA 211
212
HERPETOLOGICA
FIG. 1.—Atractus thalesdelemai sp. nov. (holotype, MNRJ 10052): general dorsal view. SVL 265 mm, and TL 38 mm.
A. taeniatus); from A. taeniatus by having six or seven maxillary teeth (instead of 8–9); and from A. zebrinus by having six supralabials (instead of seven), six or seven maxillary teeth (instead of 8–10), and uniformly dorsal pattern (instead of banded pattern) (Table 2). Description of holotype.—An adult male, SVL 265 mm; TL 38 mm (14.3% SVL); head length 11.5 mm (4.3% SVL) from rostral scale to posterior corner of mandibles; head width 5.8 mm (50.4% head length) at broadest point; interocular distance 4.25 mm; snout–orbit distance 3 mm (0.75 times interocular distance); head slightly distinct from body; rostral 0.75 times broader than high, visible from above; internasals and prefrontals as broad as long; frontal slightly broader than long, with a triangular shape in dorsal view; parietals 1.7 times longer than wide; nasal divided; loreal 2.1 times longer than high; eyes diameter 1.5 mm; pupil round; supraorbitals longer than broad; one postocular; temporals 1 þ 2; six supralabials, 3–4 contacting orbit; symphisial twice as wide as long, separated from chinshields by first pair of infralabials; seven infralabials, 1–3 contacting chinshields; anterior chinshields about twice as long as broad; posterior chinshields absent; six maxillary
[Vol. 61, No. 2
teeth; dorsal scales 17/17/17 rows, scales smooth without apical pits; preventrals 3; ventrals 154; anal plate single; paired subcaudals 28. Color of holotype in preservative (Figs. 1–2).—Dorsum of head uniformly grayish brown; lower portion of nasal and temporals lighter posteriorly; supralabials mostly creamish white, dorsal edge of third and fourth supralabials grayish brown; venter, infralabials, gulars, first and second dorsal scales rows, and lateral region of neck uniformly creamish white; ground color of dorsum, above dorsal edge of second scale row reticulate, grayish brown; anterior portion of dorsal scales grayish brown, and posterior one grayish white. Hemipenis of holotype (Fig. 3).—Inverted organ extends to level of ninth and bifurcates at eighth subcaudal; organ capitate and bilobed, lobes distinct from base; each lobe nearly flattened on apical portion, displaying series of spinulate calyces; capitation stronger on asulcate side, shallow on sulcate side; sulcus spermaticus divides about half of organ; branches have centrolineal orientation terminating on distal region of each lobe; basal portion of hemipenial body covered by large spines, concentrated on medial portion of asulcate side; spines on asulcate side slightly larger than those on sulcate side. Variation.—The type series consists of four males and four females; largest male SVL 265 mm, tail length 38 mm; largest female SVL 381 mm, tail length 38 mm; tail 13.1–14.3% of the SVL in males and 9.1–10.8% of the SVL in females. Infralabials 6–7 ( x 5 6.12; s 5 0.35; n 5 8); infralabials contacting chinshields 1–3 (n 5 6) or 1–4 (n 5 2); preventrals 1–4 ( x5 2.75; s 5 0.88; n 5 8); ventrals 149–154 ( x 5 151; s 5 2.16; n 5 4) in males, 165–167 ( x 5 165.5; s 5 1; n 5 4) in females; subcaudals 26–30 ( x 5 28.25; s 5 1.71; n 5 2) in males, 22–26 ( x 5 23.5; s 5 1.73; n 5 4) in females; dorsal scales of the tail 9–11 ( x 5 9.87; s 5 0.64; n 5 8); hemipenis (inverted organ) extends 8–11 ( x 5 9.25; s 5 1.25; n 5 4), and bifurcation 8–10 subcaudal scales; maxillary teeth 6–7 ( x 5 6.06; s 5 0.17; n 5 8). All remaining characters examined were invariable in relation to the holotype, including coloration of adults and juveniles. Distribution (Fig. 4).—This species is known from three localities of the municipality of
June 2005]
HERPETOLOGICA
213
FIG. 2.—Atractus thalesdelemai sp. nov. (holotype, MNRJ 10052): (A) dorsal and (B) lateral views of the head, and (C) lateral view of the midbody; scales 5 5 mm.
Passo Fundo, State of Rio Grande do Sul, Southern Brazil. These three localities are in the vicinities of the urban area of the city of Passo Fundo (population 166,343) and they retain some elements of grassland environment, with field formations interspersed by Araucaria forest. The type locality vegetation is typical dirty grassland, which in addition to grasses and sedges contains shrubs (Behling, 2001), suggesting that this species may be distributed over grasslands of southern South America. Etymology.—The specific name is a noun in the genitive case, in honor of Professor Thales de Lema for his lifetime of work with the herpetofauna of southern Brazil. DISCUSSION As in many fossorial snakes, species of the genus Atractus exhibit strong sexual dimorphism in segmental counts. We found it in all
species of Atractus we examined (Table 1), in accordance to the same results found by Savage (1960) with Ecuadorian species. As these results imply that this is a common feature within the genus, we agree with Savage’s (1960) suggestion that sexes should be described and analyzed separately. Furthermore, the lack of this information may reduce the relevance of this kind of data from early authors. Atractus thalesdelemai has a combination of characters also found in A. carrioni and A. roulei, such as a single postocular, six supraand infra- labials, and a uniform color pattern. These are characters that Savage (1960) argued to be indicative of close phylogenetic relationship among these species. Nevertheless, these Ecuadorian species have 15 dorsal scales rows and a higher number of maxillary teeth (8–9 in A. carrioni and 10–11 in A. roulei), differing from the new species that has 17 dorsal scales rows and 6–7 maxillary teeth.
214
HERPETOLOGICA
[Vol. 61, No. 2
FIG. 3.—Hemipenis of Atractus thalesdelemai sp.nov. (holotype, MNRJ 10052): (A) sulcate and (B) asulcate sides; scale 5 5 mm.
As noted by Savage (1960), the presence of 15 dorsal scales rows is probably an apomorphic state in Atractus that likely has evolved independently in several lineages, whereas the higher number of maxillary teeth is an unusual character of unknown polarity in Atractus, shared by a few species (e.g., A. serranus and A. trihedrurus) (Fernandes, 1995c). Some Andean species of Atractus (A. boettgeri and A. occipitoalbus) share some rare features with the new species. Atractus occipitoalbus occasionally has a single postocular, and A. boettgeri has only six supralabials. Savage (1960) considered A. occipitoalbus to be more closely related to A. trilineatus species group with 17 dorsal scale rows, based mainly on similarities of color pattern, whereas he considered A. boettgeri to be of uncertain relationship. Furthermore, A. boettgeri, A. carrioni, A. occipitoalbus, and A. roulei were included in the A. trilineatus species group by Savage (1960), defined by possessing noncapitate hemipenes, a putative synapomorphic state in the tribe Dipsadini (Fernandes, 1995c). Another species, Atractus balzani shares at least two rare character states
(single postocular, six supralabials) with A. thalesdelemai, which can be distinguished from it by several features (see diagnosis). However, a more complete comparison is difficult because of the brevity of the original description (Boulenger, 1898) that omits the sex of the holotype and do not provide an illustration. Furthermore, A. balzani is only known from the holotype, collected in the Bolivia, ca. 2000 km from the type locality of A. thalesdelemai. Atractus thalesdelemai has a capitate hemipenis with distal calyces. Fernandes (1995c) and Fernandes et al. (2000) noticed that Atractus maculatus, A. reticulatus, A. serranus, A. taeniatus, A. trihedrurus, and A. zebrinus share the same character. Recently, Schargel and Castoe (2003) found the same condition in A. univittatus and A. ventriaculatus. However, Schargel and Castoe (2003) pointed out that the hemipenes they described are very different from those studied by Fernandes (1995c), as they have spinulate flounces (A. ventrimaculatus) or papillate calyces (A. univittatus), versus spinulate calyces in the capitulum. Nevertheless, those species share an unusual
June 2005]
HERPETOLOGICA
215
FIG. 4.—Geographical distribution of Atractus thalesdelemai sp. nov. Type locality 5 open circle.
Dipsadine state character, which, as pointed out by Fernandes (1995b), is a plesiomorphic state in the subfamily and, as such, cannot be used as evidence of monophyly for this assemblage. It is possible that these snakes might represent a basal grade in relation to all other species of Atractus, in which the hemipenis has no capitation, a character state interpreted as apomorphic by Fernandes (1995c). In fact, A. reticulatus was included by Fernandes (1995b) in his analysis, which suggested a basal position to this species within the genus. Finally, despite the poorly known phylogenetic relationships of the genus, we propose the association of the new species to this apparently basal stock of Atractus based in the unusual hemipenial morphology for the genus. RESUMO Uma nova espe´cie de Atractus e´ descrita dos campos do Estado do Rio Grande do Sul, sul
do Brasil. A nova espe´cie pode ser distinta de todas as suas congeˆneres pela seguinte combinac¸a˜o de caracteres: uma po´s-ocular, loreal longa, seis supralabiais e geralmente seis infralabiais, 17 fileiras de escamas dorsais, seis ou sete dentes maxilares e um padra˜o uniforme de colorac¸a˜o dorsal marrom acinzentado com a regia˜o temporal branco creme esbranquic¸ado. Comparac¸~oes sa˜o feitas com todas as demais espe´cies de Atractus do sul da Ame´rica do Sul e a afinidade filogene´tica da espe´cie nova com A. reticulatus e espe´cies afins e´ sugerida com base na morfologia do hemipeˆnis. Acknowledgments.—We thank: C. McCarthy and M. Wilkinson (BMNH); S. P. Carvalho e Silva (DZUFRJ); G. Scrocchi and S. Kretzschmar (FML); F. L. Franco (IB); M. Leita˜o de Arau´jo (MCN); M. Di Bernardo (MCP); R. S. Be´rnils, S. A. A. Morato and J. C. Moura Leite (MHNCI); A. M. Ramos Costa and H. Zaher (MZUSP); H. Grillitsch and F. Tiedemann (NMW); A. G. Kluge and G. Schneider (UMMZ); and R. Gu¨nther (ZMB) for allowing us to examine the specimens under their care. We are grateful to P. R. Nascimento for rendering the line art. We thank
216
HERPETOLOGICA
L. F. B. de Oliveira and A. Giraudo for calling our attention to relevant literature, and U. Caramaschi, D. S. Fernandes, J. P. Pombal, Jr., and M. Raposo for comments on the manuscript. Financial support to P. Passos and R. Fernandes was provided by Conselho Nacional de Desenvolvimento Cientı´fico e Tecnolo´gico (CNPq), and Fundac¸a˜o Carlos Chagas Filho de Amparo a` Pesquisa do Estado do Rio de Janeiro (FAPERJ).
LITERATURE CITED BEHLING, H. 2001. South and southeastern Brazilian grasslands during the late Quaternary times: a synthesis. Paleogeography, Paleoclimatology, Paleoecology 2710:1–9. BOULENGER, G. A. 1898. A list of the reptiles and batrachians collected by the late Prof. L. Balzan in Bolivia. Annali del Museo Civico di Storia Naturale di Genova serie 2,XIX:128–133. CUNHA, O. R., AND F. P. NASCIMENTO. 1983. As espe´cies de Atractus Wagler, 1828, na Amazoˆnia oriental e Maranha˜o. (Ophı´dia, Colubridae). Boletim do Museu Paraense Emı´lio Goeldi, Nova Se´rie, Zoologia 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. 1995a. A new species of snake in the genus Atractus (Colubridae: Xenodontinae) from Northeastern Brazil. Journal of Herpetology 29:416–419. ———. 1995b. Phylogeny of the Dipsadine Snakes. Ph.D. Dissertation, University of Texas, Arlington, Texas, U.S.A. ———. 1995c. Variation and taxonomy of Atractus reticulatus complex (Serpentes: Colubridae). Comunicac¸~oes do Museu de Cieˆncias e Tecnologia da PUCRS, Se´rie Zoologia 8:37–53. FERNANDES, R., E. M. X. FREIRE, AND G. PUORTO. 2000. Geographic variation of the Brazilian Atlantic Rain Forest snake Atractus maculatus (Gu¨nther, 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. ———. 2002. Argentinian snakes: an annotated checklist. Smithsonian Herpetological Information Service 132: 1–53. 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. LEMA, T. 1994. Lista comemtada dos re´pteis ocorrentes no Rio Grande do Sul, Brasil. Comunicac¸~oes do Museu de Cieˆncias e Tecnologia da PUCRS, Se´rie Zoologia 7:41–150. LEVITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E. DAWSON. 1985. Standards in herpetology and ichthyology: Part 1. Standard symbolic codes for institutional
[Vol. 61, No. 2
resource collections in herpetology and ichthyology. Copeia 1985:802–832. MARTINS, M., AND 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 No. 3391:1–47. 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. ROZE, J. A. 1961. El genero Atractus (Serpentes: Colubridae) en Venezuela. Acta Biologica Venezuelica 3:103–119. SAVAGE, J. M. 1960. A revision of the Ecuadorian snakes of the colubrid genus Atractus. Miscellaneous Publications, Museum of Zoology, University of Michigan 112:1–86. SCHARGEL, W. E., AND T. A. CASTOE. 2003. The hemipenes of some snakes of the semifossorial genus Atractus, with comments on the variation in the genus. Journal of Herpetology 37:718–721. ZAR, J. H. 1999. Biostatistical Analysis. Prentice Hall, Upper Sadle River, New Jersey, U.S.A. Accepted: 28 January 2005 Associate Editor: Christopher Shei
APPENDIX Specimens Examined We examined Atractus specimens in the following collections (acronyms follow Leviton et al., 1985): Colec¸a˜o Zoolo´gia Grego´rio Bondar (CZGB), Centro de Pesquisas do Cacau, Ilhe´us; Departamento de Zoologia, Universidade Federal do Rio de Janeiro (DZUFRJ), Rio de Janeiro; Fundacio´n Miguel Lillo (FML), San Miguel de Tucuma´n; Instituto Butantan (IB), Sa˜o Paulo; Museu de Cieˆncias Naturais, Fundac¸a˜o Zoobotaˆnica do Rio Grande do Sul (MCN), Porto Alegre; Museu de Cieˆnciais Naturais, Pontificia Universidade Cato´lica de Minas Gerais (MCNR), Belo Horizonte; Museu de Cieˆncias e Tecnologia, Pontificia Universidade Cato´lica do Rio Grande do Sul (MCP), Porto Alegre; Museu de Histo´ria Natural Capa˜o da Imbuia (MHNCI), Curitiba; Museu Nacional, Universidade Federal do Rio de Janeiro (MNRJ), Rio de Janeiro; Museu de Zoologia, Universidade Federal de Alagoas (MUFAL), Maceio´; Museu Zoolo´gico Augusto Ruschi, Universidade de Passo Fundo (CRUPF), Passo Fundo; Museu de Zoologia, Universidade de Sa˜o Paulo (MZUSP), Sa˜o Paulo; Museum fur Naturkunde, Universitat Humboldt (ZMB), Berlin; Museum of Zoology, University of Michigan (UMMZ), Ann Arbor; Naturhistorisches Museum Wien (NMW), Vienna; Natural History Museum (BMNH), London. Atractus boettgeri (n 5 1): BOLIVIA: JUNGAS: (BMNH 1946.1.6.29 [holotype]). Atractus canedii (n 5 1): ARGENTINA: ANTA: Salta: (FML 1082 [holotype]).
June 2005]
HERPETOLOGICA
Atractus elaps (n 5 3): BRAZIL: Unknown locality: (IB 20314). AMAZONAS: Borba: (MNRJ 1523). ECUADOR: Western Ecuador: (??)(BMNH 1946.1.6.45 [holotype]. Atractus latifrons (n 5 18): BRAZIL: Unknown locality: (MNRJ 20315). AMAZONAS: Benjamin Constant: (MNRJ 729–732, 1289, 1517–20, 1522); Manaus: (MNRJ 726–28). PERU: PEVAS: (MNRJ 2977, 2979, 2981). Atractus maculatus (n 5 5): BRAZIL: Unknown locality: (BMNH 1946.6.46 [holotype]); ALAGOAS: Murici, Mata da Bananeira: (MUFAL 474–75); Sa˜o Miguel dos Campos: (MNRJ 3977). BAHIA: Porto Seguro: (IB 57202). Atractus major (n 5 3): ECUADOR: PASTAZA: (BMNH 1946.9.7.59–60 [paratypes], Canelos: (BMNH 1946.9.7.27 [holotype]). Atractus pantostictus (n 5 124): BRAZIL: MINAS GERAIS: Belo Horizonte: (MHNCI 787, MNRJ 6474, IB 40757, 58592, MCNR 13, 27, 35, 88, 101, 129, 139, 145–47, 148, 254, 453–54, 459, 726, 929–40); Campo do Meio: (IB 50476); Machado: (IB 57138); Pirapora, Fazenda Triaˆngulo Formoso: (MNRJ 4459, paratype); Uberlaˆndia: (IB 54604–05). DISTRITO FEDERAL: Brası´lia, Jardim Zoolo´gico: (MNRJ 4460–66). GOIA´S: Alianc¸a do Norte: (IB 43954); Cana Brava: (IB 26711); Minac¸u: (IB 51433– 34). SA˜O PAULO: Areais, Fazenda Vargem Grande: (IB 40404); Barueri: (IB 45208); Borace´ia: (MZUSP: 3157, MZUSP 3158 [paratype]); Campo Lindo: (IB 9472, 49225); Campo Limpo Paulista: (IB 44152, 54651, 54896); Francisco Morato: (IB 54634); Franco da Rocha: (IB 27305, 42093, 54844 [holotype]); Jales: (MZUSP 4094); Jundiaı´: (MNRJ 6496, IB 2728, 10068–69, 42646, 42664, 43192, 45624, 46228, 49267, 54235 [paratype], 54512, 54661; Jarinu´: (IB 41427); Itaperuna da Serra: (IB 54699); Orlaˆndia: (IB 44537); Paranapiacaba: (MZUSP 2811); Sa˜o Jose´ do Rio Preto: (IB 40028); Sa˜o Jose´ dos Campos: (IB 27231, 27233, 29098, 37527, 40355, 44527, 45784, 45803, 45807); Sa˜o Paulo: Perus: (IB 54655, IB 54886–88); Pico do Jaragua´: (IB 42404), Pirituba: (IB 42485, 53545, 54641); Va´rzea Paulista: (IB 9862, 32501, 40855, 40857, 45167). TOCANTINS: Porto Nacional, Hydroeletric Plant Rescue Luı´s Eduardo Magalha˜es: (IB 64952–64966). Atractus paraguayensis (n 5 1): PARAGUAY: Unknown locality: (NMW 23443 [holotype]). Atractus poeppigi (n 5 1): BRAZIL: AMAZONAS: Alto Rio Negro: (MNRJ MNRJ 10837). Atractus reticulatus (n 5 249): BRAZIL: PARANA´: Unknown locality: (MNRJ 9820); Apucarana: (MHNCI 3455); Arauca´ria: (MHNCI 1351, 3707, 4357, 4471, IB 11980–83, 11985, 18369); Boqueira˜o: (IB 27564); Castro: (IB 4495, 4497, 4499, 4511, 7837, 7876, 8329, 9647, 10307, 10586); Parque Florestal do Caxambu: (MHNCI 604); Curitiba: (MCN 3350–52), (MCP 3639), (MHNCI 259, 262, 296, 299–300, 397–400, 742, 802, 917, 1117, 1320, 1588, 1685, 2183, 2210, 2783, 2833, 3443, 4193, 4588, 4601–02, 4993–94); Guarapuava: (MHNCI 3312–13); Itarare´: (IB 40324, 45742); Jaguariaı´va: (MHNCI 3345); Joaquim Murtinho: (IB 13631); Lapa: (IB 10360, MHNCI 2070); Nova Restinga: (IB 15007); Palmeira: (MHNCI 298, 1184); Ponta Grossa: (MHNCI 1936); Porto Amazonas: (MHNCI 4553–55); Piraı´ do Sul: (IB 8981, 9399); Sa˜o Jose´ dos Pinhais: (IB 43633, MHNCI 2379–80, 2451, 2731, MNRJ 9086). RIO GRANDE DO SUL: Bossoroca, Fazenda Santa Catarina: (MCN 2842); Candela´ria: (MNRJ 1261); Canoas: Capa˜o do Corvo: (MCP 02), Vila Matias Velho: (MCP 1281); Cachoeirinha: (MCP 2009); Entre Rios
217
do Sul: (CRUPF 309); Esteio, Polo Petroquı´mico: (MCN 9438); General Camara´, Boca da Picada: (MCP 2611); Gravataı´: (MCN 2954, 2999–3001, 1183, 1858); Guaı´ba: (MCP 3241), Mariana Pimentel: (MCP 2530, 2538); Mato Castelhano: (CRUPF 991); Nicolau Vergueiro: (CRUPF 173, 176); Passo Fundo: (CRUPF 96, 199, 213, 224–26, 249, 284, 304, 309, 343, 376, 401, 416, 590, 686, 819, 829–30, 1064, 1204); Pelotas, Instituto de Pesquisas Agra´rias do Sul: (MCN 1329); Porto Alegre: (MCN 03, 416, 1013, 1015–16, 1794, 2766, 3517, 4145, 5405, 5407, 5409, 5411–12, 6060, 6095, 6392, 6489, MCP 01, 06, 886, 1186, 1205, 1304, 1458, 1634–35, 1876, 1907, 2047, 2171, 2470, 2502, 3079, 3479, 3495, 3745, 3921, 4382, 4551, 4663, 4730, 4788–95, 4842); Porto Xavier: (MCP 3306); Santa Maria: (MCN 3055, 4039, 7315), Camobi: (MCP 3873); Santo Antoˆnio: (MCP 03–05); Sa˜o Borja, Barreiro: (MCP 3144); Sa˜o Leopoldo: (MCN 5410), Sı´tio Laurindo: (MZUSP 5054); Sa˜o Lourenc¸o do Sul: (BMNH 1946.1.2.7 [holotype]); Viama˜o: (MCN 4096). SANTA CATARINA: Campos Novos: (MCP 2848–49); Rio Vermelho: (MZUSP 9417). SA˜O PAULO: Guaruja´: MZUSP (10577); Guianases: (MZUSP 4096); Ibiu´na: (IB 45169); Itarare´, Fazenda Santa Maria do Espinho: (IB 40324, 45742); Jarinu: (MZUSP 9496); Juquitiba: (IB 32556); Mogi da Cruzes: (IB 9452); Osasco: (IB 2731); Pissaguera: (MZUSP 1282, 1856); Santo Andre´: (IB 45738), Campestre: (IB 30452); Sa˜o Bernardo do Campo: (IB 45678); Sa˜o Caetano do Sul: (IB 4873); Sa˜o Miguel Paulista: (IB 27510, 33104); Sa˜o Paulo: (IB 1208, 1258, 3506–07, 5322, 5361, 8023, 5366, 9456, 20718, 34301, 45674, 54637, 54663, MCN 5406, MNRJ 1524, MZUSP 1226, 1230–34, 1661, 2008, 2624, 2633–34, 2638, 2643, 3159, 4167, 4153, 4164, 4204–05, UMMZ 109058); Sa˜o Vicente: (MZUSP 4582–85). Atractus serranus (n 5 38): BRAZIL: Unknown locality: (IB 32857). SA˜O PAULO: Alto da Serra: (IB 31188, 42947); Borace´ia: (MZUSP 2193); Campinas: (IB 50861), Capa˜o Bonito, Sı´tio Guarapiava: (IB 32664); Cotia: (IB 55698); Cubata˜o: (IB 9706); Engenheiro Marsilack: (IB 9075–76, 9088–89); Guapiara, Fazenda Oriente: (IB 34360, 34409); Guarulhos: (IB 26999), Km 21 of Presidente Dutra Highway: (IB 27147, 27862); Paranapiacaba: (IB 10589, 18645, 23518), Campo Grande (7200); Ribeira˜o Pires: (IB 10136); Rio dos Campos: (IB 9267, 9437–38, 10136); Rio Grande da Serra: (IB 54636, 54974); Santo Amaro, Marink-Santos Highway: (IB 4852); Santo Andre´: (53630, 55252), Km 38 of the Santos-Jundiaı´ Highway: (IB 42947); Sa˜o Joa˜o:(IB 7002); Sa˜o Luiz do Paraitinga: (IB 53924); Serra de Paranapiacaba: (IB 7239 [holotype]); Tapiraı´: (IB 42222, 42963, 52636). Atractus taeniatus (n 5 42): BRAZIL: PARANA´: Pinha˜o, Rio Jorda˜o (MCP 7185, 7211, 7364–65). RIO GRANDE DO SUL: Carazinho: (CRUPF 1180); Colorado: (CRUPF 1196); Derrubadas, Parque Florestal Estadual do Turvo: (MCP 12387); Getu´lio Vargas: (CRUPF 64); Girua´: (IB 10380); Ibirapuita˜: (CRUPF 587); Ijuı´: (MCP 13726–32); Mato Castelhano: (CRUPF 289, 516, 992, 1094); Pinheiro Machado: (CRUPF 257); Planalto: (MCP 5898–99, 5915, 5997); Porto Maua´: (MCP 11609, 11611, 11623); Porto Vera Cruz (MCP 11670); Santo Aˆngelo: (IB 9552, MCP 12516– 17); Tapejara: (CRUPF 417, 814). SANTA CATARINA: Chapeco´: (MCP 14013); Concordia, Entre Rios: (MCP 2912); Ipira: (MCP 2913); Peritiba: (MCP 2939); Piratuba (MCP 2893–94, 2902). Atractus torquatus (n 5 6): BRAZIL: AMAZONAS: Manaus, Reserva INPA-WWF: (MZUSP 8455, 9588);
218
HERPETOLOGICA
Km 80 of the Br 174 Highway, Fazenda Dimona: (MZUSP 8533–34); Novo Aira˜o: (MZUSP 8205). PARA´: Alto Cumina˜: (MNRJ 7888). Atractus trihedrurus (n 5 18): BRAZIL: PARANA´: Guaratuba, UHE Guaricana: (MHNCI 851); Piraquara: (IB 3067 [paratype]). SANTA CATARINA: Campo Alegre: (IB 32664); Campo Grande, Rio Negrinho: (IB 32367, 32369); Sa˜o Bento do Sul: (IB 9111, IB 3098 [holotype], MZUSP 9439). SA˜O PAULO: Ibiu´na: (IB 46476, 564474); Juquitiba: (IB 33930, 53565, 54703, 62215); Piedade: (IB 49752, 50280); Ribeira˜o Pires: (IB 31188, 42906). Atractus univittatus (n 51): VENEZUELA: ANZOA´TEGUI: Cerro Laguna (MNRJ 8127). Atractus ventrimaculatus (n 5 1): VENEZUELA: MERIDA: Merida: (BMNH 1946.1.5.15 [paratype]). Atractus zebrinus (n 5 42): BRAZIL: Unknown Locality: (BMNH 61.4.18.12, 61.4.18.13). BAHIA: Porto
[Vol. 61, No. 2
Seguro: (IB 57202). ESPI´RITO SANTO: Santa Tereza: (MNRJ 733–34). MINAS GERAIS: Camanducaia: (IB 40106, 45431, 45620, 45622, 45691, 51491, 51683); Liberdade: (MNRJ 6497); Sa˜o Gonc¸alo do Rio Abaixo, Peti: (MNRJ 9298). PARANA´: Campo Largo: (MHNCI 4818); Votuverava: (IB 12893). RIO DE JANEIRO: Cachoeiras de Macacu: (MNRJ 7064–65); Itaboraı´: (MHNCI 1295); Nova Friburgo: (MNRJ 6498, ZMB 7448); Petro´polis: (MNRJ 4467–70, DZUFRJ 497, 563); Rio de Janeiro: (BMNH 54.4.18.12, ZMB 6006); Tereso´p´ rga˜os: (MNRJ 6495). olis, Parque Nacional da Serra dos O SANTA CATARINA: Peritiba: (IB 44049). SA˜O PAULO: Apiaı´: (IB 52316); Borace´ia: (MZUSP 2194); Campos do Jorda˜o: (IB 54326); Cubata˜o: (IB 45193); Engenheiro Lefe´vre: (IB 16435); Guapiara, Fazenda Oriente: (IB 33717); Ribeira, Fazenda Cobalto: (IB 43733); Tapiraı´: (IB 42222, 46605, 52636, 56938).
