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Rediscovery and redescription of the Andean earth-snake Atractus wagleri (Reptilia: Serpentes: Colubridae)



Atractus wagleri was described based on a single specimen from Humbo, Department of Boyacá on the western Cordillera Oriental of Colombia, and since its original description there is no further record for the species. In the course of examination of Colombian collections the holotype of the Atractus wagleri could not be found and it is probable that it was lost, but we found three additional specimens of this poorly known snake from localities relatively close to the type locality. In this paper, we describe these specimens, and report data on meristics, morphometrics, and hemipenial variation for the species.
Accepted by D. Gower: 7 Nov. 2008; published: 5 Jan. 2009 59
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2009 · Magnolia Press
Zootaxa 1969: 5968 (2009)
Rediscovery and redescription of the Andean earth-snake Atractus wagleri
(Reptilia: Serpentes: Colubridae)
1Universidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Vertebrados, Quinta da Boa Vista, 20940-040, Rio de
Janeiro, RJ, Brazil. E-mail:
2Museo de Herpetología Universidad de Antioquia, Instituto de Biología, Universidad de Antioquia, Apartado 1226, Medellín, Antio-
quia, Colombia
Atractus wagleri was described based on a single specimen from Humbo, Department of Boyacá on the western Cordil-
lera Oriental of Colombia, and since its original description there is no further record for the species. In the course of
examination of Colombian collections the holotype of the Atractus wagleri could not be found and it is probable that it
was lost, but we found three additional specimens of this poorly known snake from localities relatively close to the type
locality. In this paper, we describe these specimens, and report data on meristics, morphometrics, and hemipenial varia-
tion for the species.
Key words: Atractus wagleri, Colombia, Cordillera Oriental, hemipenis, taxonomy
The fossorial colubrid snake genus Atractus Wagler is distributed widely in the Neotropical region, occurring
from Panama to Argentina (Giraudo & Scrocchi, 2000; Myers, 2003). Atractus is the most diverse Alethino-
phidian snake genus, with nearly 120 valid species, most of them known only from their type localities (Pas-
sos, 2008). The taxonomic status of several species remains unclear and there are many misidentified
specimens in many herpetological collections (Passos et al., 2005; Passos et al., 2007a, b; Passos & Fer-
nandes, 2008). The trans-Andean Atractus wagleri was described by Prado (1945) based on a single specimen
from Humbo in Boyacá Department of Colombia, and since its original description, there are no further
records for the species. While examining Colombian collections, we found three specimens of A. wagleri
from localities close to the type locality that we report herein.
Our aim in this study is to provide a new description of Atractus wagleri on the basis of new specimens
found in the field and already present in herpetological collections, reporting data on meristics, morphomet-
rics, and hemipenis variation for the species. In addition, we provide a new diagnosis, and comparisons
between A. wagleri and congeners occurring parapatrically in the Cordilleras Central (A. sanguineus) and Ori-
ental (A. crassicaudatus).
Material and methods
We examined Atractus specimens in the following collections: USA—United States National Museum
(USNM), Smithsonian Institution, Washington D.C. Venezuela—Colección de Vertebrados de la Universidad
60 · Zootaxa 1969 © 2009 Magnolia Press
de Los Andes (CV-ULA), Mérida; Museu de Biología de la Universidad Central de Venezuela (MBUCV),
Caracas D. C.; Museo de Historia Natural, Fundación La Salle (MHNLS), Caracas D.C.; Colección Herpeto-
logica del Laboratório de Biogeografia de la Universidad de Los Andes (ULABG), Mérida. Colom-
bia—Colégio San José (CSJ), Medellín, Antioquia; Colección Herpetologica de la Universidad de Quíndio
(UQC), Armenia, Quíndio; Colección Zoológica de la Universidad de Tolima (CZUT-R), Ibague, Tolima;
Instituto Alexander Von Humboldt (IAvH), Villa de Leyva, Boyacá; Instituto de Ciencias Naturales, Universi-
dad Nacional de Colombia (ICN), Bogotá D.C.; Museo de Herpetología Universidad de Antioquia (MHUA),
Medellín, Antioquia; Museo de la Universidad La Salle (MLS), Bogotá D.C.; Museo de Zoologia de la Uni-
versidad Javeriana (MUJ), Bogotá, D.C.; Museo de Historia Natural de Universidad Industrial de Santander
(UIS), Bucaramanga, Santander; Colección Herpetológica de la Universidad del Valle (UV-C), Cali, Valle del
Cauca. Ecuador—Escola Politecnica Nacional (EPN), Quito; Museo de Zoologia, Pontifícia Universidad
Católica de Ecuador (QCAZ), Quito. Peru—Museo de Historia Natural de la Universidad Mayor de San Mar-
cos (MHNSM), Lima; Museo de Historia Natural de Universidad Nacional de Arequipa (MUSA), Arequipa.
Brazil—Instituto Butantan (IBSP), São Paulo, SP; Museu de Zoologia da Universidade de São Paulo
(MZUSP) São Paulo, SP. England—The Natural History Museum (NHM), London. Germany— Zoologisches
Museum für Hamburg (ZMH), Hamburg. Specimens and localities are listed in Appendix and correspond to
all trans-Andean species of Atractus examined.
Terminology for Atractus cephalic shields follows Savage (1960) and ventral scale counts follow Dowling
(1951). Nomenclature regarding the loreal condition follows Passos et al. (2007b). Terminology for hemipenis
description follows Dowling and Savage (1960), as augmented by Myers and Campbell (1981) and Zaher
(1999). The hemipenes were examined in the everted condition, and when necessary they were everted using
the techniques described by Pesantes (1994). 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 analogue caliper to the
nearest 0.1 mm under a stereoscope, except for snout-vent length (SVL) and caudal length (CL), which were
taken with a flexible ruler to the nearest 1 mm.
Atractus wagleri
(Figures 1–4)
Atractus wagleri Prado, 1945; Ciencia (Mexico): 6:61.
Holotype: Female, MLS 228, from Humbo (05º36’N, 74º16’W, ca. 1000 m), Boyacá Department of Colom-
bia, collected on 1948 by Hermanos of La Salle order. Presumed lost (see discussion).
Diagnosis: Atractus wagleri is distinguished from all congeners by the combination of the following char-
acters: (1) 17/17/17 smooth dorsal scale rows; (2) two postoculars; (3) loreal long; (4) temporals 1+2; (5)
seven supralabials, third and fourth contacting orbit; (6) seven infralabials, first three contacting chinshields;
(7) six or seven maxillary teeth; (8) generally four gular scale rows; (9) four or five preventrals; (10) 174–180
ventrals in females, 157–174 in males; (11) 43–44 subcaudals in females, 46–56 in males; (12) dorsal ground
colour in preservative cream-red with an irregular vertebral black stripe connected to lateral black blotches,
constituting complete bands anteriorly and decreasing in size posteriorly; (13) venter black with paraventral
region cream; (14) moderate body size, females reaching 437 mm SVL and males 445 mm SVL; (15) moder-
ate tail length in females (13.6–15.3% SVL) and long in male (21.61% SVL); (16) hemipenis moderately
bilobed, semicapitate, and semicalyculate.
Comparisons: Among all congeners, Atractus wagleri occur parapatrically and shared a suite combina-
tion of characters with A. sanguineus: 17 dorsal scale rows, more than 170 ventrals in both sexes, more than
40 subcaudals in males, seven upper and lower labials, first three infralabials contacting chinshields, seven or
Zootaxa 1969 © 2009 Magnolia Press · 61
eight maxillary teeth, long tail in males, dorsum with blotches decreasing in size posteriorly, and venter gener-
ally most black. Atractus wagleri differs from A. sanguineus in having 43–44 subcaudals in females and para-
vertebral blotches constituting bands anteriorly (vs. 29–38 subcaudals in females and paravertebral blotches
never forming bands, except for occasionally a black collar on neck, anteriorly). The only other species of
Atractus occurring parapatrycally with A. wagleri at Cordillera Oriental of Colombia is A. crassicaudatus.
Atractus wagleri can be easily distinguished from A. crassicaudatus by having 174–180 ventrals in females,
43–44 subcaudals in females and 46–56 in males, and tail 21.61% SVL in males, 13.6–15.3% SVL in females
(vs. 147–170 ventrals in females; 21–33 subcaudals in males and 14–26 in females; tail 11.0–17.1% SVL in
males, 8.0–13.7% in females).
FIGURES 1–2. Dorsal (1) and ventral (2) views of Atractus wagleri (MHUA 14504) from San Vicenti de Chucurí. SVL
445 mm, CL 96 mm.
Description (based on 3 new specimens; 2 males, 1 female): Head twice as long as wide, flattened in
lateral view, round in dorsal view; snout truncate in lateral view, round in dorsal view; rostral subtriangular in
frontal view, broader than high, little visible in dorsal view; internasal as long as wide; internasal suture sinis-
tral with respect to prefrontal suture; prefrontal longer than wide; supraocular subtrapezoidal, slightly longer
than wide; frontal subpentagonal, broader than long; parietal about twice as long as wide; nasal divided; nos-
tril located between prenasal and postnasal; prenasal twice as high as long; postnasal about twice as high as
long, higher than prenasal; loreal long, contacting second and third supralabials; pupil round; two postoculars
of similar length; upper postocular about twice as high as lower postocular; temporals 1+2; anterior temporal
twice as long as high; upper posterior temporal elongate, three times as long as wide; seven supralabials, third
and fourth contacting orbit; second supralabial higher than first and smaller than third; third higher and sev-
enth longer than remaining supralabials; symphisial subtriangular, four times broader than long; seven infrala-
bials, first three contacting chinshields; first pair of infralabials in contact behind symphisial, preventing
symphisial/chinshields contact; chinshields three times as long as wide; three or four gular scale rows; four or
five preventrals; 17/17/17 smooth dorsal scale rows; dorsals lacking apical pits, supra-anal tubercles, and
keels; caudal spine short, robust, and rhomboid.
Maxillary arch: Arched in dorsal view, with four or five prediastemal and two or three postdiastemal
teeth; prediastemal teeth large, curved downward, decreasing gradually in size posteriorly, angular in cross
section, robust at base, and narrower at apices; maxillary diastema short; postdiastemal teeth slightly smaller
than last prediastemal tooth.
62 · Zootaxa 1969 © 2009 Magnolia Press
Colour pattern in preservative: Dorsum of head black, covered with small beige dots above internasals,
nasals, loreal, and prefrontals; background of head black to dorsal margins of supralabials; black descending
postocular stripe reaching dorsal edge of sixth and anterior ventral margin of seventh supralabial; temporal
region pigmented with irregular beige spots; median region of supralabials cream-white, except for sixth and
seventh scales; mental region cream-white, occasionally with disperse black dots; preventrals uniformly
cream; venter predominantly cream-white with irregular black dots for first 15 ventrals; venter becomes uni-
formly black before 15th ventral, except for lateral margin of scales uniformly cream-white; tail black with lat-
eral portion of subcaudals cream-white; dorsal ground colour of body black anteriorly, with cream-white
complete transverse bands (one scale long), alternated on flanks; black interspaces (three to four scales long),
reaching paraventral region; pale transverse bands of opposite sides connected middorsally only anteriorly, up
to a point level with 20th ventral; pale transverse bands increase in size at midbody (two scales long), with
black interspaces reducing gradually in size, restricted to paravertebral region (two scales long); posterior to
midbody region, dorsal ground colour becomes cream-red with black vertebral line (two scales wide), break-
ing up to for black paravertebral blotches; flanks with round black blotches (two scales long) at the level of
third and fourth scale rows; paraventral region with irregular small black blotches, constituting a barely
defined lateral stripe above first two dorsal scale rows; paraventral blotches occasionally connected to para-
vertebral ones (Figs. 1–2).
FIGURE 3. General view of Atractus wagleri (MHUA 14504) in life.
Colour pattern in life: Dorsum of head black with yellow dots above internasals, nasals, loreal, prefron-
tal, and temporal region; ventral margin of supralabials yellow; mental region and venter anteriorly cream-
Zootaxa 1969 © 2009 Magnolia Press · 63
yellow; lateral margin of ventrals cream-yellow and remaining belly uniformly black; dorsum of body black
anteriorly, with yellow transverse bands; bands gradually change to reddish orange in the median third and
uniformly red at posterior region of body; vertebral line and paravertebral blotches black; flanks and paraven-
tral region with cream-red ground colour, covered with irregular black blotches (Fig. 3).
Hemipenis (Fig. 4; everted organs n = 2): Organ moderately bilobed, semicapitate, and semicalyculate;
lobes distinct and restricted to distal portion of capitulum; lobes cylindrical with round, laterally projecting
apices; lateral projections oriented centrifugally, clearly distinct from proximal region of lobes; lobes and
capitulum uniformly covered with spinulate calyces; calyces progressively replaced by papillae toward tips of
lobes; lobes with calyces losing vertical walls and constituting spinulate transverse flounces; transverse
flounces well defined on lateral portion of asulcate side of hemipenis; asulcate side of capitulum with barely
conspicuous lobular crests; capitular crotch well marked on both sides of hemipenis; capitulum similar in size
to hemipenial body, located at the level of sulcus spermaticus bifurcation; sulcus spermaticus bifurcates in the
mid portion of the organ; branches of sulcus spermaticus centrifugal, reaching lobe apices; margins of sulcus
spermaticus stout and narrow along hemipenial body, and moderately expanded in the capitular region;
hemipenial body subcylindrical, covered with moderate hooked spines; basal naked pocket restricted to basal
portion of hemipenial body; proximal portion of hemipenis with longitudinal plicae and disperse spinules.
FIGURE 4. Sulcate (left) and asulcate (right) views of the hemipenis of Atractus wagleri (MHUA 14504). Scale =
Variation (3 new specimens plus Prado's data on female holotype): Largest male SVL 445 mm, CL 96
mm; largest female SVL 437 mm, CL 67 mm; tail 21.6% SVL in male, 13.6–15.3% (n = 2) SVL in females;
157–174 (n = 2) ventrals in males, 174–180 (n = 2) in females; 45–56 (n = 2) subcaudals in males, 43–44 (n =
2) in females; 3 (n = 1 side) or 4 (n = 5 sides) gular scale rows; 4 (n = 2) or 5 (n = 1) preventrals; 8 (n = 4
sides) or 9 (n = 2 side) dorsal scale rows in the level of second subcaudal; 6 (n = 2 sides) or 7 (n = 4 sides)
maxillary teeth; 4.5–9.0 mm midbody diameter.
64 · Zootaxa 1969 © 2009 Magnolia Press
Distribution: Western versant of Cordillera Oriental of Colombia, from Floridablanca (07º03’53’’N,
73º05’23’’W) in Santander Department to Humbo (05º36’N, 74º16’W) in Boyacá Department. Atractus
wagleri has been found in sub-Andean Tropical Forest at elevations of 740–1200 m (Fig. 5).
Remarks: Prado (1945) cited the type locality of A. wagleri as Humbo (Boyacá), Colombia. Subse-
quently, some authors cited Muzo as the proper type locality of the species (Medem, 1965; Pérez-Santos &
Moreno, 1998). This contradiction is probably because Humbo (05º36’N, 74º16’W) was a small town
included originally in the boundaries of Muzo (05º32’N, 74º06) municipality. However, the municipality of
Muzo was redefined and now does not contain Humbo, which is located instead within Quípama (05º31’N,
74º11’W) municipality (J. Lynch, pers. comm.).
FIGURE 5. Geographical distribution of Atractus sanguineus and Atractus wagleri. Open symbols correspond to
respective type localities of the species.
Prado (1945) described Atractus wagleri based on a specimen from Humbo in the Boyacá Department of
Colombia, distinguishing it from Atractus sanguineus Prado, 1944 by having a rostral slightly broader than
high, frontal as long as wide, loreal twice as long as wide, and distinct colour pattern. Unfortunately, the holo-
type of A. wagleri could not be found in the course of our complete examination of the MLS Atractus collec-
tion, and probably has been lost during a fire at Universidad La Salle in 1948 (A. Rodriguez, pers. comm.).
We report here three additional specimens of A. wagleri from localities geographically close to the type
locality (Fig. 5). Although cephalic characters earlier employed by Prado (1945) to diagnose it from A. san-
guineus falls into the range of variation of the former species, based on our sample of both taxa there are dif-
ferences in the colour pattern and number of subcaudal scales which guarantee specific recognition of A.
wagleri (see above). These differences are based on small and geographic located samples for the two taxa.
However, each species seems to be restricted to a distinct Colombian Cordillera, with the Magdalena Valley
operating as a putative barrier separating them (Fig. 5). Nonetheless, the discovery of new material of both
species in the Magdalena Valley might require the re-evaluation of the taxonomic status of Atractus wagleri.
Zootaxa 1969 © 2009 Magnolia Press · 65
We are grateful to the following personnel for permission and facilities to examine specimens under their care:
A. Acosta (MUJ), C. Aguilar and J. Santa-Gadea (MHNSM), A. Almendaríz (EPN), M. Bernal (CZUT-R), F.
Bilbao (EBRG), R. Casallas and A. Rodriguez (MLS), J. Cobos (UQC), L. Coloma and S. Ron (QCAZ), De
Keiroz and R. Williams (USNM), C. Ferreira (MBUCV), F. Franco (IBSP), J. Hallerman (ZMH), E. La Marca
(ULABG), J. Lynch (ICN), V. Paez (MHUA), A. Pascual (CV-ULA), D. Perico (IAvH), M. P. Ramírez (UIS),
C. Señaris and G. Rivas (MHNLS), M. Wilkinson and C. McCarthy (NHM), H. Zaher (MZUSP) A. Zamudio
and A. Ortíz (CSJ). We thank J Lynch (ICN), A. prudente (MPEG), and D. Gower (NHM) for constructive
criticism of the manuscript. P. Passos thanks J. Lynch (ICN) for laboratory facilities and help with the Colom-
bian collections; Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), Coordenação de
Aperfeiçoamento de Pessoal de Nível Superior (CAPES), and Fundação Carlos Chagas de Amparo à Pesquisa
do Estado do Rio de Janeiro (FAPERJ) for financial support.
Dowling, H.G. (1951). A proposed standard system of counting ventrals in snakes. British Journal of Herpetology, 1,
Dowling, H.G. & Savage, J.M. (1960). A guide to the snake hemipenis: a survey of basic structure and systematic charac-
teristics. Zoologica, 45, 17–28.
Giraudo, A.R. & ScrocchI, G.J. (2000). The genus Atractus (Serpentes: Colubridae) in Northeastern Argentina. Herpeto-
logical Journal, 10, 81–90.
Merem, F. (1965). Biliografia comentada de reptiles colombianos. Revista de la Académia Colombiana de Ciencias
Físicas, Exactas y Naturales, 12, 299–346.
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Geophis (Colubridae: Dipsadinae). American Museum Novitates, 3391, 1–47.
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Guerrero, Mexico. American Museum Novitates, 2708, 1–20.
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Thesis, Museu Nacional, Universidade Federal do Rio de Janeiro, Brazil, 671 pp.
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Atractus modestus. Herpetological Journal, 17, 1–6.
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66 · Zootaxa 1969 © 2009 Magnolia Press
Appendix I: Specimens examined.
Atractus andinus: Colombia, Antioquia, Andes: (formerly CSJ 231, now CSJ 516 holotype).
Atractus biseriatus: Colombia, Caldas, Villamaria: (MLS 145 holotype).
Atractus bocourti: Peru, Huánuco, Acomayo: (NHM 1946. 1.2.24 holotype, MHNSM 2801), Chaglla: (MHNSM
20041–43), Huancapallac: (MHNSM 20036), Molino-Panao, Pachitea: (MHNSM 3001, 3064), Panao: (MHNSM
20044), Taruco: (USNM 120809).
Atractus boulengerii: Colombia, Valle Del Cauca, Anchicayá, Bajo Anchicayá: (UV-C 6591).
Atractus carrioni: Ecuador, Loja, Loja: (EPN 8673–74, QCAZ 793, 1081–82, 1217–19, 2100), Jardin Botanico: (QCAZ
6445–46), Yangana: (QCAZ 6550), Rio Molacatus: (QCAZ 6533–34).
Atractus clarki: Colombia, Antioquia, Anori: (MHUA 14000); Chocó: Andagoya: (MLS 1213), Istmina: (MLS 1214);
Valle del Cauca: Restrepo: (ICN 10826).
Atractus crassicaudatus: Colombia, without locality: (IBSP 2443, ICN 8505, 8508–25, 8922–25, MLS 139, 152, 156,
293, 2640, MUJ 92, 355); Boyacá, Badohondo: (ICN 10693), Belen: (ICN 10709), Chiquinquirá: (MLS 2577),
Coper: (MLS 2578–79), Duitama: (ICN 10700–07), Garagoa: (ICN 10627, MUJ 315–22, 398–99, 509), Guayatá:
(IAvH 864–65), Pajarito: (IAvH 1059, ICN 2608–11, 2831–33), Pesca: (IAvH 1880), Rio Tectino: (IAvH 799),
Sogomoso: (MLS 282, 2751–52), Tunja: (MUJ 04), Ventaquemada: (MLS 2243), Villa de Leyva: (IAvH 2172–73,
3039, 3189, 4788, 4811–20, 4852, 4878, 4889, 4892–93, 4912, 4960, 4976, ICN 2792, 8332–33, 9016–19, 9027,
MLS 2021, 2564–65, 2918–20), Zetaquirá: (MUJ 05); Cundinamarca, without locality: (MUJ 482), Aguadita: (MLS
169), Albán: (IAVH 4749, ICN 10626), Bogotá: (IAvH 129, 204, 2478, ICN 1394–426, 1455, 1460–61, 2588, 2623,
2633, 2641, 3377, 4217, 4240, 6209, 6236, 6340, 6449 6490–91, 6504–05, 6509, 7100, 7102, 8260, 10806, IBSP
226, 7216–17, 10164–67, 42945, MLS 153, 164–65, 167 172, 178, 2546, 2607–09, 2614–15, 2617, 2644–45, MUJ
03, 07, 09–10, 17, 22, 24, 151, 180, 206–09, 211, 400, 609–10, 692), Arrachal: (MLS 265, 2805–13), Cerro de Suba,
La Conejera: (ICN 6336, 6577–79, 6580–81, 10692), Codazzi: (MLS 2386), San Joaquín: (MLS 2964–65), Santana:
(IAvH 4964), Cajicá: (IAvH 500), Chia: (ICN 7101, MLS 2373–77, 2382–83, 2600, 2622–23, 2830–93, 2900–08,
2935–36, MLS without number, MUJ 18, 477), Cogua: (MLS 163, 185), Cota: (MUJ 164), Facatativa: (MUJ 264,
461–62), Fontinbón: (MUJ 25), Fuquené: (MUJ 16, 20–21), Fusagasuga: (MLS 2634, MUJ 92), Guachancipá: (ICN
8261), Guachetá: (MLS 2263), Guaduas: (MUJ 01), Guasca (MLS 2626, MUJ 203–05, 215), La Calera: (MUJ 298),
La Union: (MLS 157), Machetá: (MLS 2568–70, 2653, 2921–22, 2927, 2931), Mosquera: (ICN 1453–54, 1456,
1458–59), Laguna Herrera: (IAvH 3815, ICN 859, 1277, 1457), Nemocón: (ICN 7041), Pacho: (MLS 154, 2611–12,
2616, 2923–30, MUJ 550), Pasca: (ICN 485–86, MLS 2602–04), Quetame: (ICN 4477), Represa del Sisga: (IAvH
08), Reserva Carpanta: (MLS 26), San Antonio del Tequendama: (IAvH 3038–39, MLS 150–51, 200), Sesquilá:
(MLS 2571), Sibaté: (MLS 175–76, 295), Sopo: (MLS 2624), Suesca: (MUJ 214, 649), Sumapaz: (MLS 168),
Sutatenza: (MLS 283–84, 288, 292, 1860–63, 2493–94), Tabio: (MLS 1898), Tausa (MUJ 142), Tena: (MUJ 12, 19),
Une: (MLS 160, 177, 2709–10), Usaquén: (MLS 2378–79, 2381, 2894–99, MUJ 13), Villapinzón: (ICN 2816, MLS
299), Villeta: (IAvH 1587); Meta, Cañon La Curia: (MLS 06), Lomalinda: (IAvH 967); Santander, without locality:
(MUJ 212), Bolívar: (MLS 162), Jesús Maria: (MLS 2246–48), Puente Nacional: (MLS 2629), Santa Rita: (MLS
2630). Locality probably in error: Meta, Puerto Lopez: (ICN 6500, MUJ 15).
Atractus duboisi: Ecuador, Napo: without locality: (EPN four not cataloged specimens, QCAZ 2797), Baeza: (QCAZ
1234–1241, 2103, 2759, 4110, 4156), Cantón Quijos: (EPN 1281–89, 3121), Cordillera de Guacamayos (EPN 6875,
QCAZ 3707–08, 3290), Cosanga: (QCAZ 906, 2098, 2106–07, 2759, 2798–2806, 5469), Baeza-Quito road: (QCAZ
4195, 4201), Río Hollin, Loreto road: (QCAZ 2104), Las Palmas: (QCAZ 3347–3350, 6593–95).
Atractus dunni: Ecuador, without locality: (QCAZ 219, 2884); Cotopaxi: without locality (IBSP 54328), Galapagos:
(QCAZ 1092), San Francisco de Las Pampas: (QCAZ 163, 240–47, 670, 1077, 1231–33, 1685–86, 2108–10),
Reserva Otonga, Cañon Signos: (QCAZ 4036); El Oro, Buenaventura: (EPN not cataloged); Loja: Olmedo: (QCAZ
1219); Pichincha, CERG: (QCAZ 2094), Chiriboqua: (32127–28), Mindo: (QCAZ 4151), Nanegalito: (QCAZ 638),
Tandayapa: (QCAZ 872, 1667, 2102, 2111). Localities probably in error: Piso Tropical Oriental: (EPN 8703);
Pastaza: Rio Bobonaza: (EPN 8733).
Atractus emigdioi: Venezuela, Lara: Moran, La Palma, Páramo El Jábon: (MHNLS 9299); Trujillo, Boconó, Valera-Tru-
jillo road: (ULABG 3791), Parque Nacional Guaramacal: (MHNLS 16209), Trujillo: (ULABG 4473).
Atractus eriki: Venezuela, Táchira: (CV-ULA 6117); Trujillo: Escuche: (ULABG 6710 paratype), Trujillo: (ULABG
6694 paratype), Valera: (ULABG 6693 holotype); ZULIA: Sierra de Perijá: (MBUCV not cataloged).
Atractus erythromelas: Venezuela, Mérida, Libertador: (MHNLS 902), Mérida: (NHM 1.716–17 paratypes of A. eryth-
romelas), Mucurubá: (MHNLS 276–78, 630, 902).
Atractus gigas: Ecuador, Cotopoxi, Chiribogua: (QCAZ 01), Palmeras: (QCAZ 2099), Reserva Otonga: (QCAZ 3266),
San Francisco de Las Pampas: (QCAZ 175, 179, 443, 647, 662). Locality probably in error, Piso Tropical Oriental:
(EPN 8706).
Atractus indistinctus: Colombia, Norte de Santander, Ocaña: (MLS 166 holotype, MLS 261–62, 264, 2695–96).
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Atractus iridescens: Colombia: Antioquia: San Pedro de Arama: (CSJ 563); Chocó, Nuqui: (IAvH 4539), Rio San Juan:
(MLS 1212); Nariño, Barbacoas, El Diviso, Vereda Berlin, Reserva Natural Biotopo Selva Húmeda: (ICN
Atractus lancinii: Venezuela, Aragua, Cumboto road: (EBRG 590), Maracay-Ocumare road: (EBRG 198–99, 291,
407–08, 698), Parque Nacional Pittien, Estácion Biológica Rancho Grande: (EBRG 699, 4338); Carabobo, Barbula:
(MHNLS 1750); Distrito Capital, Caracas, Parque Nacional El Ávila, Canales del Naigata: (MBUCV 2043a,b,
2044b, MHNLS 11417–18, 11797), Cerro Naigata: (MHNLS 3145), La Guaira: (MBUCV 2044a); Miranda, Gua-
caipuro: (MHNLS 6848), San Antonio de Los Altos: (MHNLS 2086, 12684), El Amarillo: (MHNLS 15150, 16788),
San Diego de Los Altos: (EBRG 1982), Santenejas: (EBRG 4088); Yaracuy, Nirgua, Santa Teresa: (MHNLS 6381).
Atractus lasallei: Colombia, without locality: (MLS 301); Antioquia, Bello: (ICN 19621), Bello, San Felix: (MHUA
14060), Belmira: (ICN 10622), El Retiro:(MHUA 14112), Guarne: (MLS 2129), La Ceja: (ICN 10713), Marinilla:
(ICN 10714), Medellín: (ICN 10618–20, MHUA 14383, MLS 2944), Medellín, Las Palmas: (ICN 1085), Medellín,
San Cristobal: (MHUA 14003), Medellín, Santa Helena: (MHUA 14194, 14221, 114368,MLS 2217), Medellín, Pie-
dras Blancas: (IAvH 970, 1008, 10012–13, 1933, 4857, MLS 204–06, 219, 223, 1782, 2829, 2946, 2958), Medellín,
San Antonio de Prado: (MHUA 14028, 14086), Rionegro: (MLS 2077, 2210), San Pedro: (IBSP 5315 holotype,
MLS 202–03, 207–09, 280, 1765, 1781, 1783, 1842–43, 1856, 1878, 1941, 2356, 2939, 2941, ICN 10628–31), San
Pedro, La Lana: (MLS 2412, 2955), Santa Rosa de Osos: (MLS 1902–04, 1946), Santo Domingo: (MLS 230), Son-
són: (ICN 10697, MLS 135). Locality probably in error, Jericó: (MLS 303).
Atractus lehmanni: Colombia, Cauca, Popayán: (ICN 1794, 2590, 2613, 10635–36, MLS 1919), Los dos Brazos:
(USNM 151633), Rio Molina: (MLS 2593), Silvia: (MLS 2595, 2681); Valle Del Cauca, Sevilla: (MHUA 1407).
Atractus loveridgei: Colombia, Antioquia, Jardin: (CSJ 566), Jericó: (IBSP 7202, 8908, 8916, 10126, MLS 213–16, 218,
220–22, 224–25, 1205). Locality probably in error San Pedro: (MLS 2355).
Atractus manizalensis: Caldas, Manizales: (IAvH 3309–10, MLS 294), Pacorá: (MLS 2216), Villamaria: (MLS 227
holotype MLS 228 paratype, MLS 146, 2461, 1999), Salamina: (MLS 173, 226, 1777, 1779–80, 2716); Quindio,
Armenia: (UQC 01, 05, 08, three specimens UQC not cataloged).
Atractus matthewi: Venezuela, Anzoategui, Macizo de Turimiquire, Cerro El Guamal: (EBRG 3952–54 paratypes of A.
matthewi, EBRG 4453 holotype of A. nororientalis, EBRG 4454 paratype of A. nororientalis, MNRJ 8127); Mona-
gas, Caripe: (MBUCV 1669); Sucre, Serrania de Turimiquire: (EBRG 3793).
Atractus melanogaster: Colombia, Caldas, without locality: (MLS 296), Pensilvania: (MLS 235, 237); Tolima, Cajama-
rca, Vereda La Palma: (ICN 10029–34), Ibague: Ibanasca: (CZUT-R 117), Pastales: (CZUT-R 10), Toche: (CZUT-R
Atractus melas: Colombia, Chocó, Quibdo: (MLS 2537); Valle del Cauca: (UV-C 8533).
Atractus michelae: Venezuela, Mérida, Canagua, Pueblos del Sur: (ULABG 2672 holotype) Táchira, Uribante, Caparo:
(CV-ULA 4445 paratype, CV-ULA 2918).
Atractus mijaresi: Venezuela, Mérida, Mucurubá, Rangel: (ULABG 4697 holotype).
Atractus modestus: Ecuador, western Ecuador (NHM 1946. 1.6.30, holotype); Azuay, Molleturo: (QCAZ 1167); Coto-
paxi, San Francisco de Las Pampas: (QCAZ 002, 201–03, 641, 1216, 2100), Pilaló: (QCAZ 6548); Morona-Santi-
ago, Plan de Milagro: (QCAZ 2013); Pichincha, without locality: (QCAZ 1134).
Atractus multicinctus:Colombia: Valle del Cauca, road to Buenaventura: (USNM 151723); Buenaventura, Queremal:
(ICN 7075).
Atractus nebularis: Colombia, Cesar, Valledupar, Nabusímaque: (ICN 10807–10); Magdalena, Montes, Cuchilla, Hierb-
abuena: (ICN 5663–65 paratypes), Santa Marta, above San Lorenzo Station: (ICN 10625), Sierra de San Lorenzo:
(ICN 2756 holotype, ICN 2757–67 paratypes).
Atractus nicefori: Colombia, Antioquia, Jardin: (MLS 2940), Jericó: (MLS 229, MLS 231–33, 239–40, 275, 279, 297,
302, 2635–37), Támesis: (MUJ 02–03).
Atractus nigricaudus: Peru, Pasco, Huáchon, Pugmaray: (MHNSM 19175, 19180, 19183, 19194), Oxapampa, San
Alberto: (MHNSM 17761), Paucartambo, Agomarca: (MHNSM 18108, 18113, 18192–93, 19047–48), Aquimarca:
(MHNSM 17811–12, 17825–27, 17842–44, 17854, 17862, 17867–68, 18015, 18101, 18103, 18050, 18105, 18575),
Mayabamba: (MHNSM 18051–54), Santa Isabel (MHNSM 18107, 18427–30), Taurapau: (MHNSM 18571),
Uchuhuerta: (MHNSM 18609–10, 19046).
Atractus nigriventris: Colombia, Boyacá, Chita: (MLS 234 holotype).
Atractus obesus: Colombia: Valle Del Cauca: Cali: Parque Nacional Natural Los Farallones: (ICN 2934).
Atractus obtusirostris: Colombia, Tolima, Ibague, Pastales: (CZUT-R 12), Toche: (CZUT-R 11), Icononzo: (ICN 2722,
6497), Juntas: (ICN 5669–71), Rio Combeyma: (ZMH-R 4428 syntype).
Atractus ochrosetrus: Venezuela, Mérida, Tovar: (ULABG 4698 holotype), Tovar-Guaraque road: (ULABG 4696 para-
Atractus oculotemporalis: Colombia, Antioquia, Jericó: (IBSP 6390 holotype).
Atractus pamplonensis: Colombia, Norte de Santander, Bochalema: (MHNLS not cataloged), Chinacotá: (MLS
2338–39), Chitagá, Chucarima; (MLS 273–74, 248, 287, 300), Cutilla: (MHUA 14163–64), El Diamante: (MLS
68 · Zootaxa 1969 © 2009 Magnolia Press
1920), Labateca: (ICN 10715–18), La Donjuana: (MLS 248), Ocaña: (MLS 277), Pamplona: (IBSP 9192 holotype,
IBSP 9190–91, 9040, 9021 paratypes, MLS 241–44, 247, 250–52, 276, 2001–02, 2364, 2369–71, 2458–60,
2688–2694, 2711–15, 2753–69, MLS without number, ICN 10719–24), Toledo: (MLS 249, 253, 2700–03).
Atractus paucidens: Ecuador, Pichincha, Santo Domingo de Los Colorados, Finca La Esperanza: (EPN 8729–32,
MZUSP 7703, USNM 232725); locality in error, Pastaza, Río Bobonaza: (USNM 232726).
Atractus roulei: Ecuador, Chimborazo: (USNM 33861); Azuay, Hierba Mala: (QCAZ 6256).
Atractus sanctaemartae: Magdalena, Ciudad Perdida: (IAvH 2290), San Sebatian de Rábago: (ICN 10711). Locality in
error, Huila, Acevedo, Parque Natural Nacional Cueva de los Guacharos: (IAvH 8302).
Atractus sanguineus: Colombia, Antioquia, Porce: (MLS 2513), San Pedro: (CSJ 518), Yarumal: (formally CSJ 232, now
CSJ 517 holotype, MLS 1784–85), Yolombo: (MHUA 1432).
Atractus tamaensis: Venezuela, Táchira, Junin, Betania: (MHNLS 8307 holotype, MHNLS 8301, 8303–06 paratypes).
Atractus taphorni: Venezuela, without locality: (IBSP 25785); Mérida, El Chorotal: La Azulita road: (CV-ULA 1838),
La Carbonera: (CV-ULA 6417), Libertador: (ULABG 3909).
Atractus trivittatus: Colombia, Boyacá, Chita: (MLS 258 holotype, MLS 286); Casanare, La Salina: (MLS 257, 290,
2638–39, 2706–07); Norte de Santander, Arboledas: (MLS 269), Gramalote: (MLS 137), La Donjuana: (MLS
Atractus variegatus: Colombia, Boyacá, Boavita: (MLS 2484–85), La Uvita: (MLS 260 holotype, MLS 217, 259, 267,
272, 278, 281, 2266, 2268–69, 2271–73, 2697).
Atractus ventrimaculatus: Venezuela, Mérida, Betania: (ULABG 2409), La Princesa: (ULABG 6701–02), Libertador, El
Valle: (MHNLS 897–901), Mérida: (NHM 1946. 1.5.15 holotype), La Mucus, Parque Nacional Sierra Nevada:
(MBUCV 2016), Pico Humbo: (EBRG 4052).
Atractus cf. vertebralis: Peru, Cusco, Urubamba, Machu-Pichu: (MHNSM 3100), Puerto Libre, Apurimac River Valley:
(USNM 538480); Huancavelica, Tayacaya: (MHNSM 2849).
Atractus vertebrolineatus: Colombia, Norte de Santander, Ocaña: (MLS 184 holotype).
Atractus vittatus: Venezuela, Aragua, withouth locality: (IBSP 41082), Colonia Tavor: El Limón road: (EBRG 700,
2959, 4059, 4092); Distrito Capital, Caracas: (MBUCV 703), El Junquito-Colonia Tavor road: (MBUCV 415), El
Limón: Las Aguaitas: (MHNLS 5159).
Atractus wagleri: Colombia, Santander, Floridablanca: (UIS-R 71), Piedecuesta, Guatigurá: Vereda Viricute: (UIS-R
281), San Vicente de Chucuri: (MHUA 14504).
Atractus werneri: Colombia, without locality: (MLS 144, 289, 483); Cundinamarca, El Colégio: (IAvH 4327), Fusa-
gasugá: (ICN 2727, MLS 2329, 2334, 2345–44, 2427, 2514, 2518, 2523, 2563, 2914–16, 2932–34, MUJ 92), La
Mesa: (MLS 161), La Vega: (IAvH 2068), San Francisco: (ICN 5738, 10696), Santandercito: (IAvH 3014, MLS
1915–16, 2118, 2020), Sasaima: (ICN 2612, MLS 236, 238), Silvania: (IAvH 145, 823–24, ICN 7268), Vereda Santa
Rita: (IAvH 17).
... Las escamas corporales son lisas sin fosetas apicales, con 17 hileras de escamas dorsales sin reducción, de 140 a 170 escamas ventrales; escama anal entera y subcaudales divididas (21 a 33 en machos y 14 a 26 en hembras) (Passos y Arredondo 2009). Maxila con 8 a 11 dientes, de longitud similar y con tendencia decreciente hacia la parte posterior de la boca (Boulenger 1894, Dunn 1944a, b, Peter y Orejas-Miranda 1970, Pérez-Santos y Moreno 1988. ...
... Atractus crassicaudatus es fácilmente distinguible de sus especies más cercanas geográficamente, A� wagleri y A� werneri, por la siguiente combinación de caracteres: A� crassicaudatus presenta 147-170 escamas ventrales (♀), 14-26 escamas subcaudales (21-33 ♂) y la cola representa el 8-13,7% (11-17% ♂) de la longitud de cuerpo, mientras que A� wagleri presenta 174-180 ventrales (♀), 43-44 subcaudales (46-56 ♂) y la cola representa el 13,6-15,3% (21,61% ♂) del cuerpo, por su parte A� werneri presenta 158-174 ventrales (♀), 21-36 subcaudales (27-37 ♂) y la cola representa el 8,7-15,2% (13,2-17,1% ♂) del cuerpo (Passos y Arredondo, 2009;Passos y Lynch, 2011); de manera que A� crassicaudatus tiene el cuerpo y la cola, proporcionalmente más corta de las tres especies. ...
... Vera-Pérez -Atractus manizalesensis in Antioquia endemic) (Passos & Arredondo, 2009;Passos et al., 2009a, b;Passos & Lynch, 2010;Uetz et al. 2019). ...
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A new locality for the Colombian endemic earth-snake Atractus manizalesensis is reported, with comments on geographical distribution of congeners from the Cauca river basin
... In recent years, some species of snakes have been rediscovered in Colombia (i.e. Atractus wagleri, Passos & Arredondo, 2009; Trilepida joshuai, Rojas-Morales & González-Durán, 2011; Coniophanes andressensis, Caicedo-Portilla, 2014;Imantodes phantasma, Medina-Rangel et al., 2018;Vanegas-Guerrero et al., 2019), which should promote the need for field studies to monitor populations of these littleknown species. The recording of species after several decades since the last known description is a relatively common case for snakes. ...
Full-text available
Dendrophidion boshelli is a poorly known and endemic snake species from the Middle Magdalena river valley in Colombia. It was described in 1944 based on a single specimen from the municipality of Caparrapí, department of Cundinamarca. Since the original description, only three additional specimens have been established. As part of the results of a herpetological monitoring in the Miel I Hydroelectric project, department of Caldas-Colombia, three additional specimens of D. boshelli were found in 2014-2015. The specimens represent the second known population of the species. We presented morphological data and pholidosis; description of the coloration in life, and a description of the habitat. Reptiles of Colombia, and the IUCN Redlist. For this reason, to promote the local conservation of this and other endemic and threatened species in the Miel I area, we suggested some actions at the local level, such as establishing a conservation area with legal status in the Middle Manso River basin, which is a tributary of the Miel I Hydroelectric project.
... In recent years, some species of snakes have been rediscovered in Colombia (i.e. Atractus wagleri, Passos & Arredondo, 2009; Trilepida joshuai, Rojas-Morales & González-Durán, 2011; Coniophanes andressensis, Caicedo-Portilla, 2014;Imantodes phantasma, Medina-Rangel et al., 2018;Vanegas-Guerrero et al., 2019), which should promote the need for field studies to monitor populations of these littleknown species. The recording of species after several decades since the last known description is a relatively common case for snakes. ...
Full-text available
Dendrophidion boshelli is a poorly known and endemic snake species from the Middle Magdalena river valley in Colombia. It was described in 1944 based on a single specimen from the municipality of Caparrapí, department of Cundinamarca. Since the original description, only three additional specimens have been established. As part of the results of a herpetological monitoring in the Miel I Hydroelectric project, department of Caldas-Colombia, three additional specimens of D. boshelli were found in 2014–2015. The specimens represent the second known population of the species. We presented morphological data and pholidosis; description of the coloration in life, and a description of the habitat. Recently, D. boshelli was included as a Critically Endangered (CR) species in the Red Book of Reptiles of Colombia, and the IUCN Redlist. For this reason, to promote the local conservation of this and other endemic and threatened species in the Miel I area, we suggested some actions at the local level, such as establishing a conservation area with legal status in the Middle Manso River basin, which is a tributary of the Miel I Hydroelectric project.
We present a catalog of type specimens deposited in the Herpetological collections at the Museo de La Salle (MLS), Bogotá, Colombia. The list includes 85 type specimens comprising 36 holotypes and 49 paratypes. Also, we include the types belonging to other institutions, corrections in the catalog numbers and localities, additions and updates to the information in the original descriptions, as well as rediscovery of material that was considered lost until now.
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We evaluated the taxonomic status of snakes from the Atractus emmeli species complex (composed by A. boettgeri, A. emmeli, A. paravertebralis, and A. taeniatus) on the basis of concordance between quantitative (meristics and morphometrics) and qualitative (pholidosis, color pattern, and hemipenis) analyses of morphological characters, in combination with ecological niche modeling and niche overlapping. We synonymize A. boettgeri, A. paravertebralis, and A. taeniatus with A. emmeli based on the congruent analytical results. We also describe a new species to accommodate the Brazilian populations from the state of Mato Grosso mainly based upon some unique states of morphological characters, including hemipenial morphology, color pattern, and meristics. We found that the new species has a distinct ecological niche compared with A. emmeli and some level of niche overlapping with A. albuquerquei. We found great differences in ecological niches of species occurring in the Cerrado versus those occurring in the Western Amazon–Andean foothills, suggesting a putative niche evolution in this group.
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The taxonomic status of the Pacific lowland Atractus is revised on the basis of meristic, morphometric, colour pattern, and hemipenial characters. Geographical variation is reported for six Atractus species (A. boulengerii, A. clarki, A. iridescens, A. melas, A. multicinctus, and A. paucidens). Atractus boulengerii is rediscovered and redescribed from a specimen from the Colombian coast. The first voucher specimens are reported for A. melas. The current status of A. microrhynchus is maintained based on the discovery of new material referrable to that species. Three new species of Atractus are described from the Pacific lowland of Colombia: A. echidna sp. nov., A. medusa sp. nov., A. typhon sp. nov. Two new Atractus species groups (multicinctus and paucidens) are proposed based on external morphology, maxillary dentition, and hemipenial characters. A new key to Pacific lowland species of Atractus is provided.
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We describe herein a new polychromatic species of the snake genus Atractus from the cloud forests of the northeastern Andes of Colombia. The new species is distinguished from all congeners by having an exclusive combination of phenotypic characters, such as: dorsal scale rows 17, loreal long, seven to ten maxillary teeth, ventrals 156–174 in females and 153–169 in males, subcaudals 20–30 in females and 23–30 in males, dorsum with variable coloration, changing from dark green to orange or red with a black nuchal band (three to four scales long) connected to a black vertebral line and two black dorsolateral continuous stripes from the occipital region to tip of the tail, venter with irregular black blotches, relatively small body size, small tail length in females and moderately long in males, hemipenis moderately bilobed, semicapitate and semicalyculate. We compared the new species with all congeners occurring along the Cordillera Oriental in Colombia, Sierra de Perijá in the Colombia/Venezuela frontier and Cordillera de Mérida in Venezuela. We discussed aspects related to polychromatism and its implication toward a robust taxonomy for the genus Atractus.
The Ground Snakes of the genus Atractus Wagler, 1828 (Dipsadidae) as currently recognized consists of over 130 species. However the genus has long been recognized as being paraphyletic, with various species groups having been assigned generic names in the past. Notwithstanding this fact, most publishing authors continue to group all within Atractus as a matter of convenience rather than an evidence-based firm belief that all should be placed within a single genus on the basis of accepted genus level affinity. This paper begins the dismemberment of the genus Atractus as known to date and formally removes three divergent species from Colombia (the wagleri group) and places them in a newly named genus according to the Zoological Code (Ride et al. 1999). Two other divergent species are each placed into monotypic (at this stage) genera, these being Atractus clarki Dunn and Bailey, 1939 and Atractus zidoki Gasc and Rodrigues, 1979. Keywords: Taxonomy; Ground Snake; Colombia; Atractus; New Genera; Shanekingus; Carstensus; Drewwilliamsus.
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Body-size is significantly correlated with the number of vertebrae (pleomerism) in multiple vertebrate lineages, indicating that somitogenesis process is an important factor dictating evolutionary change associated to phyletic allometry and, consequently, species fitness and diversification. However, the role of the evolution of extreme body sizes (dwarfism and gigantism) remains elusive in snakes, mainly with respect to postnatal ontogeny in dietary preferences associated with evolution of gigantism in many lineages. We described herein a new species in the highly diversified and species-rich genus Atractus on the basis of four specimens from the southeastern slopes of the Ecuadorian Andes. The new species is morphologically similar and apparently closely related to two other allopatric giant congeners (A. gigas and A. touzeti), from which it can be distinguished by their distinct dorsal and ventral coloration, the number of supralabial and infralabial scales, the number of maxillary teeth, and relative width of the head. In addition, we discuss on the ontogenetic trajectories hypotheses and dietary specializations related to evolution of gigantism in the goo-eaters genus Atractus.
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We present a revision of Atractus in north-eastern Argentina based on the examination of newly collected specimens and most of the material available in Argentinean museums. Four species are reported: A. snethlageae, A. paraguayensis, A. reticulatus and A. taeniatus. Atractus badius was erroneously cited as occurring in Argentina based on a specimen from Las Palmas, Chaco province which is reassigned to A. snethlageae. This record represents a considerable southern extension of the known range of the species. Atractus paraguayensis is redescribed based on three new specimens. This species was previously known only from the holotype reported from 'Paraguay' without definite locality data. Adult and juvenile colour patterns in life are described. The validity of some diagnostic characters is discussed, and new diagnostic characters are given for A. reticulatus and A. paraguayensis. All species examined showed noteworthy variation in colour pattern. Sexual dimorphism is reported in all species. The distributional patterns and phytogeographic areas occupied by each species in Argentina are discussed. We also characterize morphological variation for each and provide a key for the Argentinean species.
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A new species of Atractus is described from upper Cururu River in the Amazon Basin of Brazil. The new species differs from all currently recognized Atractus by having a dorsal colour pattern reddish brown, with first two dorsal scale rows creamish white, hemipenis slightly bilobed, with alary spines at intrasulcar region, and lateral projections of the lobes depressed in their basal portions. In addition, a discussion concerning putative close relative taxa is provided.
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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.
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A new species of Atractus is described from Serra de Baturité, an Atlantic Forest enclave in the semiarid Caatinga of the state of Ceará, 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.
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The New World xenodontine "colubrids" represent two immunologically distinct assemblages - the Central and South American lineages, neither of which has been well diagnosed to date. I follow this nomenclature and recognize the Central American lineage as containing 22 genera. This clade is supported by the synapomorphy of a sulcus spermaticus bifurcating within or at the base of the capitulum (Cadle, 1984; Myers and Cadle, 1994). The remaining xenodontines constitute a total of 68 presently recognized genera, of which 41 are placed in the subfamily Xenodontinae sensu stricto. The other 27 genera are considered incertae sedis, pending further research. The Xenodontinae sensu stricto are hypothesized as being monophyletic on the basis of the following hemipenial synapomorphies: (1) presence of enlarged lateral spines on the hemipenial body, and (2) two distinctly ornamented regions on the lobes, the asulcate surface bearing enlarged spinulate or papillate calyces (= body calyces). Some taxa recognized as Xenodontinae sensu stricto lack body calyces but have a nude area in the same topographical position (e.g., Psomophis, Tropidodryas). This pattern is viewed as the result of secondary loss. The rationale for this conclusion is based on the hypothesis that body calyces are merely enlarged capitular calyces of the asulcate/medial surfaces of the lobes. In the Xenodontinae, the body calyces are almost always separated from the calyces of the capitulum by a more or less developed overhang (except in a few genera). This overhang is generally retained on the hemipenes where the asulcate/medial surfaces of the lobes are nude (e.g., Psomophis), which supports the view that the body calyces were secondarily lost. Body calyces are also found on the surface of the hemipenial body in its asulcate side (e.g., Philodryas, Pseudablabes, Xenoxybelis). Because body calyces are interpreted as modified "capitular calyces," which are restricted to the lobular region and crotch, the presence of these structures far on the hemipenial body is here viewed as a more derived state where the body calyces extend from the lobes to the body. Various presumably monophyletic units are defined within the Xenodontinae sensu stricto. Conophis, Heterodon, and Farancia are clearly assigned to the Xenodontinae sensu stricto. The hemipenial morphology of various supra-generic "colubrid" taxa are described and compared. The variation of some hemipenial features within the colubroid radiation, as well as their bearing on the higher level phylogeny of colubroids, is investigated.
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Atractus modestus was described based on a single specimen from western Ecuador, and since its original description there have been no further records for this species. During the examination of Ecuadorian collections, we found additional specimens of this poorly known snake. In this paper, we redescribe the holotype of A. modestus, describe the hemipenis and report new specimens, localities, and data on meristic and morphometric variation in the species. We also compare and diagnose this species from all others members of this highly diverse genus.