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A New Species of Scincella (Squamata: Scincidae) from the Cuatro Cie´negas Basin, Coahuila, Mexico Uri Omar Garcı´a-Va ´ zquez1, Luis Canseco-Ma´rquez1, and Adria´n Nieto-Montes de Oca1 A new species of Scincella, previously confused with S. lateralis, is described from the Cuatro Cie´negas Basin, Coahuila, Mexico. The new species differs from all congeners in North and Middle America by possessing two dark, narrow ventrolateral stripes on each side (vs. dark, narrow ventrolateral stripes on each side absent in the other species); from all congeners in North and Middle America, except S. lateralis, by having three or more pairs of nuchal scales (vs. fewer than three pairs of nuchals in the other species) and the first nuchal in contact with the tertiary temporal row (vs. first nuchal usually in contact with the upper secondary temporal in the other species); and from S. lateralis by having longer limbs (limbs overlapping by 1–15 scales when adpressed against body, hindlimb length/SVL ratio 0.30–0.42, ” x = 0.36; vs. limbs separated by 2–23 scales when adpressed against body, hindlimb length/SVL ratio 0.24–0.37, ” x = 0.30, in S. lateralis) and usually more scales around midbody (28–30, ” x = 29.0, n = 15; vs. 24–29, ” x = 26.4, n = 28, in S. lateralis). The new species is geographically closest, and morphologically most similar, to S. lateralis. Se describe una nueva especie de Scincella, previamente confundida con S. lateralis, de la Cuenca de Cuatro Cie´negas, Coahuila, Me´xico. La nueva especie difiere de todos sus conge´neres en Norte y Centroame´rica por poseer dos lı´neas ventrolaterales oscuras y angostas en cada lado (vs. lı´neas ventrolaterales oscuras y angostas ausentes en las otras especies); de todos sus conge´neres en Norte y Centroame´rica, excepto S. lateralis, por poseer tres o ma ´ s pares de escamas nucales (vs. menos de tres pares de escamas nucales en las otras especies) y la primera escama nucal en contacto con la hilera temporal terciaria (vs. primera escama nucal usualmente en contacto con la escama temporal secundaria superior en las otras especies), y de S. lateralis por poseer extremidades ma ´ s largas (extremidades que se superponen de 1 a 15 escamas cuando se pliegan contra el cuerpo, cociente longitud de la extremidad posterior/longitud hocico-cloaca 0.30–0.42, ” x= 0.36; vs. extremidades separadas por 2–23 escamas cuando se pliegan contra el cuerpo, cociente longitud de la extremidad posterior/longitud hocico-cloaca 0.24–0.37, ” x = 0.30, en S. lateralis) y usualmente ma ´ s hileras longitudinales de escamas alrededor de la mitad del cuerpo (28–30, ” x = 29.0, n = 15; vs. 24–29, ” x = 26.4, n = 28, en S. lateralis). La nueva especie es geogra ´ ficamente ma ´ s cercana, y morfolo ´ gicamente ma´s similar, a S. lateralis.
A
LTHOUGH the scincid lizard genus Scincella includes 23 species in eastern Asia, only three species are recognized in North and Middle America (Mittleman, 1950; Ouboter, 1986; Eremchenko, 2003). Scincella lateralis occurs from northern North Carolina in the United States (Smith, 1946) to central Coahuila and northern Durango in Mexico (Smith et al., 1997). Scincella gemmingeri, with two subspecies, occurs from southern Nuevo Leo´n (Martı´n del Campo, 1953) to northeastern Oaxaca, Mexico (Smith and Taylor, 1950). Scincella silvicola, also with two subspecies, occurs from east-central Coahuila (Garcı´a-Va´zquez et al., 2005) to northwestern Oaxaca and southern Veracruz, Mexico (Smith and Taylor, 1950). Although S. lateralis was one of the first species of scincid lizards to be described from America, the species was not reported from the Cuatro Cie´negas Basin, Coahuila, until a personal communication by McCoy, reporting a Leiolopisma laterale ‘‘derived form, distinctive at racial or specific level (that) occurs at Cuatro Cie´negas, Coahuila,’’ was cited by Morafka (1977). A few years later, McCoy (1984) reported an undescribed subspecies of Scincella lateralis (‘‘S. lateralis ssp.’’) endemic to the Cuatro Cie´negas Basin, but did not describe it. Overall, few specimens of S. lateralis have been reported from Mexico. In addition to the specimens from the Cuatro Cie´negas Basin, two specimens were recently reported from northern Durango by Smith et al. (1997);
1
however, only brief information about the morphology and color pattern of these specimens was provided. Our examination of several specimens of Scincella from the Cuatro Cie´negas Basin suggested that they represent an undescribed species. Subsequent field work and examination of additional specimens of Scincella from Mexico and the United States supported this assessment. Because neither Morafka (1977) nor McCoy (1984) mentioned whether the specimens from the Cuatro Cie´negas Basin examined by McCoy were deposited in some herpetological collection, we were unable to examine them. However, because of their geographic provenance, we assume that they represent the same undescribed species. Herein, we describe this species on the basis of a sample of 15 specimens. MATERIALS AND METHODS A total of 15 specimens from the Cuatro Cie´negas Basin, Coahuila, were examined. Ten of these specimens were museum specimens, whereas the other five were collected during field work conducted in the basin in March 2003 and June 2004. These specimens were fixed in 10% buffered formalin, preserved in 70% ethanol, and deposited in the herpetological collection of the Museo de Zoologı´a of the Facultad de Ciencias, Universidad Nacional Auto´noma de Me´xico (MZFC). The 15 specimens of Scincella from the
Museo de Zoologı´a and Departamento de Biologı´a Evolutiva, Facultad de Ciencias, Universidad Nacional Auto´noma de Me´xico, Apdo. Postal 70-153, Me´xico 04510, D.F., Me´xico; E-mail: (UOGV)
[email protected]; (LCM)
[email protected]; and (ANMO) anietomontesdeoca@ me.com. Send reprint requests to UOGV. Submitted: 12 February 2009. Accepted: 15 March 2010. Associate Editor: D. Kizirian. DOI: 10.1643/CH-09-045 F 2010 by the American Society of Ichthyologists and Herpetologists
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Cuatro Cie´negas Basin were compared with representatives of all of the species and subspecies of the genus in North and Middle America, including the type series of Scincella g. gemmingeri, S. g. forbesorum, S. s. silvicola, and S. s. caudaequinae. Nomenclature of scales follows Taylor (1935). The diagnosis was based on the specimens examined and relevant literature (Taylor, 1937; Mittleman, 1950; Smith, 1951; Greer, 1974). Scale counts were performed with the aid of a dissecting microscope. Measurements were taken with calipers to the nearest 0.01 mm. In the description of the holotype, the maximum width and maximum length of some scales are given (in that order) in parentheses. Bilateral characters were scored on both the left and right sides. When the condition of a given meristic character was not identical on both sides, the conditions on the left and right sides are given as left/right. Color descriptions and codes (in parentheses) follow Smithe (1975). For each of the remaining specimens examined, bilateral characters were scored only on the left side, except for the supraoculars and nuchals, which were scored on both the left and right sides (conditions given as left/right). Recognition of this species is based on the phylogenetic species concept proposed by Eldredge and Cracraft (1980), Nelson and Platnick (1981), and Cracraft (1983), among others. Scincella kikaapoa, new species Figures 1–2, Tables 1–2 Scincella lateralis.—Morafka, 1977:73.–McCoy, 1984:50 (in part). Holotype.—MZFC 17667, adult male, Mexico, Coahuila, 4 km SE Cuatro Cie´ negas, Poza El Mojarral, 26u55911.90N, 100u06953.20W, 739 m, 6 July 2005, A. Contreras. Paratypes.—All (n 5 14) from Mexico, Coahuila, Cuatro Cie´negas Basin: MZFC 17668, same locality as the holotype; MZFC 17664–17666, 13 km S Cuatro Cie´negas, Poza Azul; MZFC 2012, KU 47088–47089, 12.5 km SE Cuatro Cie´negas, Poza Churince; UIMNH 43231–43236, 48328, 9.17 km SE Cuatro Cie´negas. Diagnosis.—Scincella kikaapoa can be distinguished from all congeners in North and Middle America by having two dark, narrow ventrolateral stripes separated from each other by a narrow ventrolateral pale line on each side (vs. dark ventrolateral stripes on each side absent in S. s. caudaequinae, S. s. silvicola, S. lateralis, and S. gemmingeri). In addition, S. kikaapoa may be distinguished from all congeners in North and Middle America, except S. lateralis, by having three or more pairs of nuchals (vs. fewer than three pairs of nuchals in the other species) and the first nuchal separated from the upper secondary temporal by the tertiary temporal row (in 100% of the specimens, n 5 15; vs. first nuchal in contact with the upper secondary temporal in $92% of the specimens in S. gemmingeri and S. silvicola; n 5 81 and 102, respectively; Table 1), and from S. lateralis by having longer limbs (limbs overlapping by 1–15 scales when adpressed against body, hindlimb length/SVL ratio 0.30–0.42, x 5 0.36; vs. limbs separated by 2–23 scales when adpressed against body, hindlimb length/SVL ratio 0.24–0.37, x 5 0.30, in S. lateralis; Table 1) and usually more scales around midbody (28–30, x 5 29.0, n 5 15, in S. kikaapoa; vs. 24–29, x 5 26.4, n 5 28, in S. lateralis).
Fig. 1. Scincella kikaapoa, holotype, MZFC 17667. Head scales in dorsal view (top), left lateral view (middle), and ventral view (bottom). Scale bar represents 1 mm.
Description of holotype.—Length from tip of snout to vent (SVL) 42.38 mm; distance from tip of snout to anterior border of orbit 2.83 mm; distance from tip of snout to anterior insertion of forelimb 15.10 mm; trunk length from axilla (posterior point of contact between forelimb and body wall with limb held at right angle to body) to groin (anterior point of contact between hindlimb and body wall with limb held at right angle to body) 23.14 mm; greatest head width (between angles of jaws) 5.32 mm; head length (from tip of snout to posterior margin of ear) 9.18 mm; greatest body width 5.76 mm. Tail partly regenerated, 72.60 mm in length (total length 114.98 mm). Limbs pentadactyl, long, overlap by four scales when adpressed against body (hindlimb length/SVL ratio 0.30), longest fingers reaching posterior corner of eye; straightened limbs measuring (from insertion to tip of longest digit) about 9.54 mm forelimb (22% SVL), 14.30 mm hindlimb (34% SVL).
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Fig. 2. Scincella kikaapoa in life, MZFC 17667, holotype.
Snout acutely rounded in dorsal view, rounded in lateral view; rostral approximately twice as wide as high, in broad contact with frontonasal posteriorly. Frontonasal slightly wider than long, in contact with prefrontals posteriorly, its
posterior margin with a medial lobe between prefrontals. Prefrontals about as wide as long, with posterior margin rounded, in contact with anterior and posterior loreals, first superciliary, and (very narrowly on left side, widely on right
UTA 55544 (see Material Examined) was not included. a
S. g. forbesorum (n = 9) S. g. gemmingeri (n = 72) S. s. silvicola (n = 75) S. lateralis (n = 28a) S. s. caudaequinae (n = 27) Character/taxon
Table 1. Variation in Selected Characters in North and Middle American Scincella.
Nuchals left/right 0–3/0–3 2–4/2–4 0–3/0–3 0–5/0–5 0–3/1–2 3–4/3–5 Upper tertiary temporal–parietal contact (%) 8 94 4 8 0 100 Longitudinal scale rows around midbody x¯ 5 28.87 (27–30) x¯ 5 26.42 (24–29) x¯ 5 30.42 (28–34) x¯ 5 26.75 (24–30) x¯ 5 27.77 (26–30) x¯ 5 29.00 (28–30) Hindlimb length/SVL ratio x¯ 5 0.34 (0.27–0.39) x¯ 5 0.30 (0.24–0.37) x¯ 5 0.37 (0.30–0.41) x¯ 5 0.31 (0.23–0.38) x¯ 5 0.33 (0.29–0.38) x¯ 5 0.36 (0.30–0.42) Subdigital lamellae on fourth toe x¯ 5 18.06 (16–19) x¯ 5 16.32 (15–18) x¯ 5 19.12 (17–21) x¯ 5 15.75 (13–19) x¯ 5 17.55 (16–19) x¯ 5 17.93 (18–20) Dark ventrolateral stripes Absent Absent Absent Absent Absent Present
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side) first supraocular; minimum distance between prefrontals 0.54 mm. Frontal large, elongate, kite-shaped (1.49 mm 3 3.10 mm), in contact with frontonasal, prefrontals, first two and three supraoculars on left and right side, respectively, and frontoparietals. Frontoparietals large, not fused; frontoparietal on left and right sides in lateral contact with second, third, and fourth and with third, fourth, and fifth supraoculars, respectively. Supraoculars large, 4/5 (first supraocular on right side presumably abnormally divided). Interparietal small (1.43 mm 3 2.15 mm), slightly elongate, kite-shaped, enclosed by frontoparietals and parietals; pineal eye situated near posterior end of interparietal. Parietals large, elongate, obliquely oriented, in narrow contact with each other behind interparietal; in contact with last supraocular, last superciliary, uppermost postsubocular, upper secondary temporal, uppermost tertiary temporal, first nuchal, and (broadly and narrowly on left and right side, respectively) second left nuchal. Three pairs of nuchals; those of first pair about 2.5–3.0 times as wide as long, separated from each other by second left nuchal; those of second and third pairs noticeably wider, in medial contact with each other. Nasal rhomboidal, slightly longer than high, with naris centrally situated; nasals widely separated by frontonasal. Anterior loreal about twice as high as wide. Posterior loreal slightly higher than wide, wider than anterior loreal, on left side; partially fused with preocular and presumably fused with anteriormost presubocular on right side, forming a large scale nearly twice as wide as high, much wider than anterior loreal. One small preocular, partially fused with posterior loreal on right side. Superciliaries 8/9 (third scale in right series divided longitudinally into two); first superciliary large, in contact with first supraocular, prefrontal, posterior loreal, and preocular (posterior loreal partially fused with preocular and presumably fused with anteriormost presubocular on right side); second to sixth and fourth to seventh superciliaries lower than first and first, second, and third superciliaries on left and right sides, respectively; last superciliary about as large as penultimate superciliary, in contact with parietal, last supraocular, seventh superciliary, posterior postocular, and uppermost postsubocular. Upper ciliaries 7/6; lower ciliaries 9/10; lower eyelid movable, scaly, with elongate, translucent median window. Subocular row incomplete, separated by fifth supralabial below center of eye. Presuboculars three on left side, anteriormost one much larger than the other ones; two, small, on right side (anteriormost presubocular presumably fused with preocular and posterior loreal). Postsuboculars 4/4, arranged in an oblique row extending posterodorsally to parietal. Postoculars 2/2; anterior one in contact with orbit on left side, separated from orbit by two tiny scales on right side. Temporals 1-2-3; primary temporal large, separated from parietal by uppermost postsubocular and upper secondary temporal; lower secondary temporal slightly smaller than primary temporal, overlapping a much larger upper secondary temporal; lower tertiary temporal large, about twice as high as wide, and remaining tertiary temporals slightly higher than wide on left side; lower tertiary temporal moderate in size, about as high as wide, and remaining tertiary temporals distinctly larger, about twice as high as wide, on right side. Nuchals separated from upper secondary temporal by tertiary temporal row. Supralabials seven. Infralabials 7/6. Mental more than twice as wide as long. Postmental longer than mental, in contact with first two infralabials. Three pairs of enlarged chin scales, with first
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Table 2. Variation in Selected Morphometric Characters in the Type Series of Scincella kikaapoa, Excluding the Holotype (n = 14). SVL = snout–vent length; HW = head width; HL = head length; EW = ear width; FL = forelimb length; HIL = hindlimb length; DL = fourth toe length; TL = trunk length.
Specimen/character KU 47088 KU 47089 MZFC 2012 MZFC 17664 MZFC 17665 MZFC 17666 MZFC 17668 UIMNH 43231 UIMNH 43232 UIMNH 43233 UIMNH 43234 UIMNH 43235 UIMNH 43236 UIMNH 48328 x¯ 1 SE
SVL
HW
HL
EW
FL
HIL
DL
TL
43.85 35.67 35.70 35.52 43.28 41.42 39.21 48.96 40.91 39.80 40.60 39.75 41.57 29.09 39.84 4.58
4.67 4.16 4.80 4.52 5.24 5.08 4.36 6.29 4.90 4.99 4.68 4.77 5.16 4.50 4.87 0.49
7.53 6.72 7.20 7.08 8.04 8.07 7.23 7.95 7.38 7.60 7.58 6.87 7.75 6.10 7.43 0.59
1.34 1.09 0.80 1.04 1.10 1.00 0.92 1.43 1.33 1.24 1.10 1.02 1.36 0.69 1.09 0.21
9.39 11.99 9.40 9.70 10.08 14.34 9.56 9.40 9.90 10.19 9.94 9.04 11.08 9.09 10.15 1.05
12.78 10.86 14.00 11.72 14.38 11.01 13.07 11.19 13.71 15.27 15.01 11.91 14.20 12.27 13.10 1.42
4.18 5.56 4.60 5.09 4.90 5.70 4.50 4.83 5.20 5.70 5.00 4.73 5.73 4.06 4.96 0.50
24.91 19.59 20.10 19.47 22.08 22.28 21.62 23.27 24.16 20.92 25.39 23.09 23.15 16.43 21.97 2.36
pair in broad contact, second pair separated by one scale, third pair separated by three scales. External ear opening a large obliquely oriented oval (smaller than eye opening), without lobules or spines. Scales smooth and unstriated, in 31 rows around neck just anterior to forelimb; in 30 rows around midbody; in 20 rows around base of tail. Dorsals in 67 transverse rows (from end of nuchals to level above vent), nearly equal in size to ventrals. Scales of limbs smooth and unstriated as on body. Outer posterior tubercle of wrist enlarged. Tubercles under sole approximately 53, variable in size; series bordering heel slightly enlarged. Supradigital scales in one row; upper and lower ungual sheath scales short, covering only base of claw; subdigital lamellae rounded. Digits long. Subdigital lamellae on manus I 4/4 II 8/8 III 11/11 IV 12/12 V 7/8. Subdigital lamellae on pes I 5/5 II 9/8? (digit incomplete) III 14/14 IV 18/18 V 10/10. Median pair of enlarged precloacal scales (nearly twice as large as adjacent ventrals), each overlapping part of smaller outer precloacal scale on each side. Subcaudals on base of tail nearly as large as ventrals, wider on regenerated part of tail. Color of holotype in preservative.—Dorsum brownish olive (29), slightly darker or olive (30) on head and tail; separated on either side from dark, broad dorsolateral stripe by narrow, ill-defined pale line. Dorsolateral stripe dark grayish brown (20); extending posteriorly from naris throughout rest of head, trunk, and tail; narrow anterior to orbit, occupying only dorsal three-fourths of loreal region; wider posterior to orbit, occupying two scale rows at level of axilla, one and two half scale rows at level of midbody, and two scale rows at level of groin; with some small, indistinct, pale scattered flecks throughout its extension. Dorsolateral stripe defined ventrally by strong, narrow (about one-half scale row at level of midbody) pale neutral gray (86) midlateral stripe extending posteriorly from along supralabials, above forelimb and hindlimb, then along tail. Midlateral stripe separated ventrally from pale ventral surface by two narrow, dark grayish brown (20) ventrolateral stripes separated from each other by narrow (one-half scale row at level of midbody), pale line; upper ventrolateral stripe extending
posteriorly from mid ear throughout rest of trunk and tail; ill defined on neck; encompassing one and one-half scale rows at level of axilla and one scale row at levels of midbody and groin; slightly blurred by numerous small, indistinct, pale scattered flecks throughout its extension; lower ventrolateral stripe distinct on posterior two-thirds of trunk, gradually disappearing towards axilla, dim on posterior end of neck and base of tail; encompassing only tips of two adjacent scale rows at level of midbody. Small, irregular dark spots scattered mostly on supralabials, infralabials, and postlabials comprising only remaining marks on head. Ventral surface pale neutral gray (86), gradually becoming dark neutral gray (83) posteriorly on tail. Variation.—Frontonasal separated from frontal (prefrontals in medial contact) in ten specimens; in contact (prefrontals separated) in four. Variation in selected meristic characters in type series (including holotype) as follows: supraoculars 4–4/4–5 (4/4 [n 5 14], 4/5 [n 5 1]), x 5 4.0/4.07; nuchals 2– 5/3–4 (3/3 [n 5 8]; 3/4 [n 5 2]; 4/4 [n 5 4]; 5/4 [n 5 1]), x 5 3.4/3.3; infralabials 6–7 (6 [n 5 2]; 7 [n 5 13]), x 5 6.8; dorsals in 61–71 transverse rows (61 [n 5 2]; 63 [n 5 4]; 65 [n 5 3]; 66 [n 5 1]; 67 [n 5 3]; 71 [n 5 2]), x 5 64.6; and lamellae under fourth toe 16–20 (16 [n 5 1]; 17 [n 5 5]; 18 [n 5 5]; 19 [n 5 3]; 20 [n 5 1]), x 5 17.9. Measurements for all of the specimens of the type series are provided in Table 2. Geographic distribution.—Scincella kikaapoa appears to be allopatric with respect to the other species of Scincella (Fig. 3). The nearest records of Scincella to the type-locality of S. kikaapoa are those of S. lateralis from northern Coahuila (municipality of Melchor Muzquiz), ca. 114 km (straight line) to the north, and northern Durango (Smith et al., 1997), ca. 233 km (straight line) to the southwest. Excluding S. lateralis, the nearest records of Scincella to the Cuatro Cie´negas Basin are those of Los Lirios, municipality of Arteaga, Coahuila (S. silvicola caudaequinae; Garcı´a-Va´zquez et al., 2005), and Cola de Caballo, municipality of Santiago, Nuevo Leo´n (typelocality of S. s. caudaequinae), to the southeast. The other Mexican Scincella, S. g. gemmingeri, S. g. forbesorum, and S. s. silvicola, occur much further to the southeast.
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Etymology.—The specific name kikaapoa is an indeclinable word from the language of the Kikapue, the native people that inhabits the center of Coahuila, that means ‘‘those who walk on the land.’’ DISCUSSION
Fig. 3. Geographic distribution of Scincella kikaapoa. Discontinuous lines represent state limits. The triangle represents the type-locality of this species. The squares and circles represent the closest localities of Scincella silvicola caudaequinae and S. lateralis, respectively. Scale bar represents 50 km.
Ecology.—The Cuatro Cie´negas Basin is a unique region because of its remarkable aquatic features, which include lagunas, playas and associated gypsum dunes, rivers, subterranean tubes, artesian wells, pozos, and marshes (Pinkava, 1979, 1984). The vegetation on the basin floor has been divided into several major zones: basin zacato´n grasslands, aquatic and semi-aquatic habitats (sedges and marshes), gypsum dune assemblages, and a transition zone of shrubs and trees between the grasslands and the slopes at the foot of the steep limestone sierras that surround the region (Pinkava, 1979, 1984). McCoy (1984) regarded Scincella kikaapoa (his Scincella lateralis spp.) as a semiaquatic species that lives in marshes, sedge mats, and other such habitats peripheral to water courses and lagunas, able to survive seasonal or temporary habitat drying, and capable of dispersal across areas of discontinuous aquatic habitat. The type series of S. kikaapoa was collected in the shores of ditches and small ponds in the basin. These shores were covered by low (height usually less than 30 cm) halophilic vegetation (Fig. 4). The vegetation of the aquatic and semiaquatic habitats in the basin was described by Pinkava (1979). The climate at the type-locality is temperate (mean annual temperature 5 16–22uC; mean temperature of the coldest and warmest months 12uC and 28uC, respectively) and arid, with annual seasonal precipitation averaging less than 200 mm and a rainy season that extends from May through December (INEGI, 1994). Conservation.—The conservation status of the lagoons in the Cuatro Cie´negas Basin has long been a matter of concern (i.e., Pinkava, 1987; Breunig, 2006). In a visit to the typelocality (Poza El Mojarral) two years after the holotype and a paratype of S. kikaapoa were collected, it was noted that the extension of the pond had reduced and some of the ditches connected to it had dried. In a recent visit (December 2009), additional dry ditches were observed in the region. No Scincella were found.
The evidence for the diagnosability and therefore specific status of Scincella kikaapoa seems conclusive. The presence in this species of two dark, narrow ventrolateral stripes separated from each other by a narrow ventrolateral pale line on each side is a unique character for North and Middle American Scincella. Furthermore, S. kikaapoa possesses a unique combination of other characters among the North and Middle American species of the genus (Table 1). Smith et al. (1997) assigned two specimens of Scincella from northern Durango (14.5 km SSW of Picardias, 1342 m) to S. lateralis. We visited this area in March 2003, June 2004, and September 2007; however, no specimens were found. Examination of specimens of S. lateralis from Texas, Florida, and northern Coahuila, in addition to those reported by Smith et al. (1997) from Durango, revealed no tendency whatsoever towards the color pattern or scalation of S. kikaapoa. The habitat of S. kikaapoa also is unique among the habitats occupied by Scincella in North and Middle America. Whereas S. kikaapoa has been considered a semiaquatic species (McCoy, 1984) and lives in the short halophilic vegetation surrounding the lagoons in the Cuatro Cie´negas Basin, the other species of Scincella inhabit mainly forested areas, including oak, pine, pine–oak, cloud, rain, and temperate and tropical deciduous forests (Garcı´a-Va´zquez, 2003). Because the Durango population is located approximately 300 km from the closest record known of S. lateralis, we suggest that assignment of this population to S. lateralis should be regarded as tentative until additional material from this population becomes available. MATERIAL EXAMINED Institutional abbreviations for museum and collections follow Leviton et al. (1985) or are listed at http://www.asih.org/ codons.pdf, except for the following institutions: EBUAP (Escuela de Biologı´a, Beneme´rita Universidad Auto´noma de Puebla), ITAH (Instituto Tecnolo´gico Agropecuario de Hidalgo), MZFC (see above), and UNAM-LT (Estacio´n Biolo´gica Los Tuxtlas, Universidad Nacional Auto´noma de Me´xico). Scincella gemmingeri forbesorum: Me´xico: Hidalgo: UIMNH 27399 (holotype), UIMNH 27400, FMNH 100418, 100528–31 (paratypes), La Placita, 10 mi S Jacala; ITAH 998, Tepehuaca´n de Guerrero; MZFC 7586, Tlahuiltepa, Tolantongo. Scincella gemmingeri gemmingeri: Me´ xico: Hidalgo: UTA 31011–12, ITAH 107, 223, Tlanchinol; ENCB 12330–31, Barranca de Mextitlan; UTA 11989–91, 11995, Lolotla; ITAH 741–45, Calnali, Atempa; MZFC 3449, Zacualtipan, Molango; UTA 11992, Zacualtipan, Xochicoatlan; MZFC 7743, San Bartolo Tutotepec. Oaxaca: MZFC 4502–04, Santiago Comaltepec, Puerto Antonio; MZFC 13458, San Jose´ Tenango; AMH 64 (EBUAP uncatalogued), San Isidro Buenos Aires; UTA 12087, Sierra de Jua´rez, Buena Vista; UTA 8429, Sierra de Jua´rez, Cerro Pelo´n; UTA 30314–17, MZFC 10545, Sierra de Jua´rez, La Esperanza; UTA 12013, 30331, Sierra de Jua´rez, Los Metates; JAC 21635 (MZFC uncatalogued), JAC 21637 (UTA uncatalogued), JAC 21638 (MZFC uncatalogued), Sierra de Jua´rez, San Mateo Yetla; UTA 8440–41,
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Fig. 4. Habitat of Scincella kikaapoa at the type locality.
12011, 12014, 12088–90, Sierra de Jua´rez, Vista Hermosa; MZFC 7058, 16 km E Rodulfo Figueroa; UTA 12095–97, Rodulfo Figueroa, Cerro Bau´l; MZFC 9956, San Miguel Chimalapa; UTA 5679, 10 mi W Cerro Bau´l; UTA 8558,
12005–06, 12008–10, 12091–92, 30329–30, JAC 21140 (UTA uncatalogued), JAC 21543 (MZFC uncatalogued), Sierra Mixe, Totontepec. Puebla: UOGV 112, 115–16 (EBUAP uncatalogued), Apulco, Zacapoaxtla; LCM 1154 (EBUAP
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uncatalogued), Tahitic, Zacapoaxtla; IBH 6896–97, 6898 (2 specimens), Tepango de Rodrı´guez. Veracruz: USNM 6331 (holotype), Orizaba; UNAM-LT 3534, Cuidad Mendoza; UNAM-LT 3551, 4020, Ixhuatla´n del Cafe´; JAC 22563 (UTA uncatalogued), JAC 22567 (MZFC uncatalogued), La Perla, Metlac; MZFC 5579, Mextitlan, San Andre´s Tuxtla; UNAM-LT 1354, Ocotal Chico, Pajapan; UNAM-LT 3329, Petlalcala, San Andre´s Tuxtla. Scincella lateralis: United States: Florida: CAS 214306, Citrus Co.; UF 41295–97, NE Lake Rosalie, Osceola; UF 41318, Winter Haven, Lake Shipp, Polk; UF 41367, S Medart, Wakulla. Texas: UF 1023–25, 10 mi E College Station, Brazos; UTA 11118–22, Cherokee, Houston; UTA 14905–10, 14912, El Llano, Houston; UF 1019–21, NW of Normangee, Leon. Me´xico: Coahuila: FMNH 28663, Melchor Muzquiz; UTA 55544, Sierra del Carmen, Rancho Alejandro Garza. Durango: UF 105868–69, 14.5 km SSW Picardias. Scincella silvicola caudaequinae: Me´xico: Nuevo Leo´n: KU 192606, Can ˜ o´n la Huasteca; KU 92614–15, Paraje los Osos, Santiago; MZFC 11220, FMNH 11220, 30698, 30701, 36993– 95, Santiago; KU 87736–41, Zaragoza. Quere´taro: MZFC 8830, 9423, Jalpan de Serra; MZFC 9804, 9831–32, Neblinas, Landa de Matamoros; MZFC 9805, Rı´o Tancuilin. San Luis Potosı´: UIMNH 10132, FMNH 100414–16 (paratypes), 10 mi W El Naranjo. Tamaulipas: MZFC 8514, Go´mez Farı´as. Scincella silvicola silvicola: Me´xico: Hidalgo: CHH 6, 10 (ITAH uncatalogued), Acuatempa, Huejutla; ITAH 336, Tlaplexco. Oaxaca: EBUAP 1794–95, 6 km W Concepcio´n Pa´palo; MZFC 6035, IBH 11194–98, Cuicatla´n; EBUAP 725, MZFC 8592, 8595, 8604, Quiotepec, Cuicatla´n; EBUAP 726, Santa Marı´a Texcatitla´n; JAC 22686 (UTA uncatalogued), 22688 (MZFC uncatalogued), Tutepetongo, Cuicatla´n. Puebla: JAC 22499 (UTA uncatalogued), Cuetzalan; EBUAP 1542, Cuetzalan, Cuichat; MZFC 6449, Jonotla; EBUAP 1535, LCM 1160 (MZFC uncatalogued), Cuetzalan, Las Hamacas; UOGV 72 (MZFC uncatalogued), LCM 1165 (MZFC uncatalogued), Presa La Soledad, Ayotoxco de Guerrero; IFP 212, 226, 237, 251, 265, 292, 313, 331, 410, 430, 432, 441–42 (EBUAP uncatalogued), EBUAP 1541, 1444–45, 1547–48, 1452, Cuetzalan, San Andre´s Tzicuilan; EBUAP 1533–34, Cuetzalan, Tazipehualt; EBUAP 1550, Cuetzalan, Tzanaco; EBUAP 1530–32, Cuetzalan, Tzinacapan; EBUAP 1527, Cuetzalan, Xalpana; EBUAP 1529, Cuetzalan, Yancualtipan. Veracruz: FMNH 100008 (holotype), FMNH 104678 (paratype), near San Lorenzo, 10 mi SE Co´rdoba; UNAM-LT 3527, Amatla´n de Los Reyes; MZFC 182, La Isla, Nautla; UNAM-LT 2916, La´zaro Ca´rdenas, Catemaco; MZFC 603, Mocambo; ENCB 7545, Palma Sola; IBH 410, Rı´o Quetzalapan, Catemaco; UNAM-LT 3069, 3550, 3554, 3576–77, Volca´n San Martı´n, San Andre´s Tuxtla; MZFC 7373, IBH 50569, 50597, Santa Martha, Catemaco; UNAM-LT 1103, 3529, Estacio´n Biolo´gica UNAM; ENCB 6511, Xico; UNAM-LT 188, Zacatal, San Andre´s Tuxtla. ACKNOWLEDGMENTS Support for field work was provided by grants to A. NietoMontes de Oca from CONACyT (number P47590-Q) and the Direccio´n General de Apoyo al Personal Acade´mico, Universidad Nacional Auto´noma de Me´xico (PAPIIT grant numbers IN210707 and IN224009). For loan of specimens, we thank J. Campbell and E. Smith (University of Texas at Arlington), L. Trueb and J. Simmons (University of Kansas), K. de Queiroz (USNM), J. B. Ladonski and A. Resetar
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(FMNH), G. Pe´rez Higareda (Estacio´n de Biologı´a Tropical ‘‘Los Tuxtlas,’’ Universidad Nacional Auto´noma de Me´xico), P. Lavı´n (Universidad Auto ´ noma de Tamaulipas), J. C. Lo´pez (Escuela Nacional de Ciencias Biolo´gicas, Instituto Polite´cnico Nacional), H. Eliosa (Escuela de Biologı´a, Universidad Auto´noma de Puebla), and F. Mendoza (Instituto Tecnolo´gico Agropecuario de Hidalgo). We thank J. Aguilar Lo ´ pez, M. Feria Ortiz, A. Contreras Arquieta, A. Mendoza Herna´ndez, R. Mendoza Paz, J. Ramos Alvarez, and C. Yanez Arenas for their invaluable help in collecting the specimens of the type series during 2004 and 2005 and/or for comparative material. LITERATURE CITED Breunig, L. 2006. Conservation in context: establishing natural protected areas during Mexico’s neoliberal reformation. Unpubl. Ph.D. diss., The University of Arizona, Tucson, Arizona. Cracraft, J. 1983. Species concepts and speciation analysis. Current Ornithology 1:159–187. Eldredge, N., and J. Cracraft. 1980. Phylogenetic Patterns and the Evolutionary Process. Method and Theory in Comparative Biology. Columbia University Press, New York. Eremchenko, V. K. 2003. Generic and specific redefinition and redescription of the North-Vietnam skink Scincella melanosticta (Boulenger, 1887). Izvestiya Vuzov, Bishkek 1–2:20–28. Garcı´a-Va´zquez, U. O. 2003. Revisio´n taxono´mica del ge´nero Scincella (Lacertilia: Scincidae) de Me´xico. Unpubl. B.S. thesis, Universidad Auto´noma de Puebla, Puebla, Me´xico Garcı´a-Va´zquez, U. O., D. Lazcano-Villareal, M. C. Garcı´ade la Pen ˜ a, and G. Castan ˜ eda. 2005. Geographic distribution: Scincella silvicola caudaequinae. Herpetological Review 36:337. Greer, A. E., Jr. 1974. The generic relationships of the scincid lizard genus Leiolopisma and its relatives. Australian Journal of Zoology, Supplementary Series 31:1–34. INEGI (Instituto Nacional de Estadı´stica, Geografı´a e Informa´tica). 1994. Carta Topogra´fica 1:50000, Cuatro Cie´negas G13B59, Coahuila. Instituto Nacional de Estadı´stica, Geografı´a e Informa´tica, Distrito Federal, Me´xico. Leviton, A. E., R. H. Gibbs, Jr., E. Heal, and C. E. Dawson. 1985. Standards in herpetology and ichthyology: part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985:802–832. Martı´n del Campo, R. 1953. Contribucio´n al conocimiento de la herpetologı´a de Nuevo Leo´n. Universidad 11:53–77. McCoy, C. J. 1984. Ecological and zoogeographic relationships of amphibians and reptiles of the Cuatro Cie´negas Basin. Journal of the Arizona–Nevada Academy of Science 19:49–59. Mittleman, M. B. 1950. The generic status of Scincus lateralis Say, 1823. Herpetologica 6:17–19. Morafka, D. J. 1977. A biogeographical analysis of the Chihuahuan Desert through its herpetofauna. Biogeographica 9:1–313. Nelson, G., and N. I. Platnick. 1981. Systematics and Biogeography. Columbia University Press, New York. Ouboter, P. E. 1986. A revision of the genus Scincella (Reptilia: Sauria: Scincidae) of Asia, with some notes on its evolution. Zoologische Verhandelingen 229:1–66. Pinkava, D. J. 1979. Vegetation and flora of the Bolson of Cuatro Cienegas region, Coahuila, Mexico, I. Boletı´n de la Sociedad Bota´nica de Me´xico 38:35–44.
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Smith, H. M., and E. H. Taylor. 1950. An annotated checklist and key to the reptiles of Mexico exclusive of the snakes. Bulletin of the United States National Museum 199:1–253. Smithe, F. B. 1975. Naturalist’s Color Guide. American Museum of Natural History, New York. Taylor, E. H. 1935. A taxonomic study of the cosmopolitan scincoid lizards of the genus Eumeces with an account of the distribution and relationships of its species. The Kansas University Science Bulletin 23:1–643. Taylor, E. H. 1937. Two new lizards of the genus Leiolopisma from Mexico, with comments on another Mexican species. Copeia 1937:5–11.
