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Distinguishing characters and variation in Aspidoscelis neomexicana (Squamata: Teiidae): a little known diploid parthenogenetic whiptail lizard in Texas.

Abstract.--In 1966, diploid parthenogenetic Cnemidophorus neomexicanus = Aspidoscelis neomexicana (Lowe & Zweifel, 1952) was reported by another worker from an inexact site in Presidio County, Texas, based on a specimen in the University of Kansas Natural History Museum (KU 40264). Nevertheless, doubt concerning the presence of A. neomexicana in this part of Texas has been expressed for many years. Between 1988 and 2008, numerous visits by JEC to precisely documented sites near the Rio Grande south of Candelaria, Presidio County, >225 airline km southeast of the City of El Paso, resulted in collection of 29 specimens identified as A. neomexicana (New Mexico Whiptail). Those designations, which were initially based on the same qualitative diagnostic characters used in the original description of the species in 1952, are herein verified based on statistical comparisons of means for nine meristic characters for the sample of putative A. neomexicana from Presidio County and for samples of the species from El Paso and Hudspeth counties, Texas. Relevant samples of the same and different species of whiptail lizards from Hidalgo County in extreme southwestern New Mexico were also compared with the Texas samples to detect both intraspecific geographic variation and interspecific differentiation with reference to A. neomexicana in Texas.

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The first report (Axtell 1966) of Cnemidophorus neomexicanus (= Aspidoscelis neomexicana; New Mexico Whiptail) in the herpetofauna of Presidio County, Texas, which borders the Rio Grande, was based on specimen KU 40264 in the University of Kansas Natural History Museum. Subsequently, Cordes et al. (1989), Cordes & Walker (2009), and Walker et al. (2013) also reported on specimens of the species from some combination of El Paso, Hudspeth, and Presidio counties, emphasizing locality records and aspects of habitat utilization. However, they did not include data on distinguishing characters and variation in support of their identifications. Dixon (2013) listed I 1 species of whiptail lizards (for application of Aspidoscelis in the Family Teiidae see Reeder et al. 2002) in Texas including five gonochoristic species (Cnemidophorus = Aspidoscelis gularis, A. inornota, A. marmorata, A. scalaris, and A. sexlineata) and six parthenogenetic species (A. dixoni, A. exsanguis, A. laredoensis, A. neomexicana, A. tesselata, and A. uniparens). However, Dixon (2013) included this statement of uncertainty on the Texas distribution of diploid hybrid-derived A. neomexicana described as Cnemidophorus meomexicanus by Lowe & Zweifel (1952) from Socorro County, New Mexico: "Museum records from Culberson and Presidio counties are questionable." Therefore, to remove the uncertainty expressed by Dixon (2013) on the status of the species in Presidio County, this report verifies the identity of specimens listed as parthenogenetic A. neomexicana by Cordes et al. (1989) and Cordes & Walker (2009) from El Paso, Hudspeth, and Presidio counties using comparisons with each other, with samples of diploid parthenogenetic A. tesselata pattern class E from Texas, and with samples of A. neomexicana and gonochoristic A. inornata from New Mexico.

Materials and Methods

Aspects of scutellation character states and statistically analyzed meristic characters were compared using 27 of 29 specimens of A. neomexicana and 23 of 25 specimens of A. tesselata E from sites south of Candelaria, Presidio County (Table 1), to verify indentities initially based on diagnostic characters of color pattern and morphology. Also used in meristic comparisons for assessment of geographic variation and interspecific distinctions were 10 specimens of A. neomexicana from neighboring El Paso and Hudspeth counties, Texas, 24 of A. neomexicana and 18 of A. inornata from Hidalgo County, New Mexico, and nine of A. neomexicana from Grant County, New Mexico (Table 1). With the exception of the specimens from El Paso and Hudspeth counties from collections at Sul Ross State University (SRSU) and University of Texas at El Paso (UTEP), all other specimens were collected by JEC under authority of appropriate Texas and New Mexico permits and presently bear numbers of the University of Arkansas "Department of Zoology" (UADZ) herpetology collection (Table 1).

Scutellation, meristic, and color pattern characters referenced in this study were identified and described in Tables 2-4. With a few exceptions they included characters examined by Cole et al. (1988) and Manning et al. (2005) in other samples of A neomexicana from New Mexico. The identities of specimens in the American Museum of Natural History verified by electrophoresis, karyotypes, and meristic data as A. neomexicana and A. inornata by Cole et al. (1988) were from the same areas of Hidalgo County where JEC collected the UADZ specimens of these species used in this study.

Statistical analyses were performed on an institutional PC using University of Arkansas licensed JMP software (Version 10; SAS Institute, Inc., Cary, North Carolina, 2012) to generate a mean [+ or -] 1 standard error and range of variation for each character and ratio analyzed. Multiple sample means for a character were compared for significant differences ([alpha] = 0.05) using Tukey posthoc tests in JMP to preserve alpha.

Results and Discussion

Color patterns.--Aspidoscelis neomexicana was distinguishable from other species of whiptail lizards in Texas and elsewhere by distinctive dorsal and ventral color patterns throughout ontogeny (Fig. 1). This description was based on all 29 specimens (SVL 38-72 mm) from sites near the Rio Grande between 1.5 and 3.5 km south of Candelaria, Presidio County (Table 1). Dorsally, the upper and lower lateral fields above and below the lateral stripes were gray and gray-brown, respectively, whereas the dorsolateral and vertebral fields were pale brown (Table 2; Fig. 1). The lateral and dorsolateral stripes were cream-white to pale yellow and essentially straight-margined, whereas the paravertebral stripes were yellow-tan and wavy (Table 2; Fig. 1). The highly diagnostic yellow-tan vertebral (= middorsal) stripe typically had a 1-2 mm bifurcation on the parietal scales from which it was relatively straight and narrow on the neck and then irregularly margined (= wavy) to the base of the tail (Fig. 1). In two specimens of 61 mm SVL (UADZ 3807) and 60 mm SVL (UADZ 3808), part of the vertebral "stripe" resembled two strands in a twisted cord. Lateral and dorsolateral stripes extended onto the base of the tail (Fig. 1), the distal two-thirds of the tail being gray-green in life. In specimens of A. neomexicana, the stripes did not undergo fragmentation, nor were prominent light-colored vertical bars present between the stripes (Fig. 1), as were present in syntopic congener in Presidio County, diploid parthenogenetic A. tesselata E (Common Checkered Whiptail). Examples of specimens of A. neomexicana with well-defined rounded beige spots in the upper and lower lateral fields were included in Fig. 1 (UADZ 4698, gravid female of 67 mm SVL; UADZ 4700, 72 mm SVL), whereas only indistinct spots were present in UADZ 4722 (66 mm SVL). Distinct spots were not present in juveniles from Presidio County. In contrast, presence of straight stripes and a complete absence of spots at all ontogenetic stages of pattern development characterize A. inornata (Little Striped Whiptail) which also occurs in Presidio County; however, it was not observed at the riverine sites from which A. neomexicana was collected. Ventrally, live individuals of A. neomexicana were gray with a blue to green hue.

In 24 specimens (52-76 mm SVL; Table 1) of A. neomexicana from Hidalgo County, New Mexico, greater contrast between the pale colored stripes and spots and darker intervening fields were more apparent than in the 29 specimens from Presidio County.

Scutellation.--Specimens of the diploid parthenogenetic species A. neomexicana and A. tesselata E from sites of syntopy in Presidio County, sorted into samples based on the foregoing color pattern characters, were also distinctive in the following scutellation character states (see Table 2) described for the former species followed by the latter species: postantebrachial scales granular (versus scales very slightly enlarged) and scales bordering the edge of the gular fold small (versus scales enlarged). In all 23 specimens of A. tesselata E examined for the character, the circumorbital scale series terminated on each side of the head at the level of the second supraocular sutures and thus were designated incomplete (Table 3).

Scutellation data for the Presidio County sample of 27 specimens of A. neomexicana examined for the circumorbital scale character revealed unanticipated individual and geographical variation, compared with the less variable Hidalgo County sample of the species. In the Presidio County sample, only six (22.2%) specimens had complete series on both sides of the head (Table 3), whereas of the remaining specimens six (22.2%) had a complete series on one side of the head and 15 (55.6%) had incomplete series on both sides of the head. In 24 specimens of A. neomexicana from Hidalgo County and nine from Grant County, extreme southwestern New Mexico, 30 (90.9%; Table 3) specimens had complete circumorbital series on both sides of the head as expected in this species (Lowe & Zweifel 1952). Eighteen specimens of A. inornata from syntopic contacts with A. neomexicana in Hidalgo County had incomplete circumorbital series that extended no farther anteriorly than the midpoints of the third supraocular scales which provided evidence of the diagnostic utility of the character in distinguishing these species in New Mexico (Table 3). Also, most specimens (91.7%) of A. neomexicana from Hidalgo County lacked the noticeably smaller scales centrally located along the guiar fold that were present in all specimens of the species from Presidio County. Manning et al. (2005), based on 96 specimens of A. neomexicana from the Conchas Lake area, San Miguel County, New Mexico, reported two preanal scales immediately anterior to the vent in 34 (35.4%) and one in 62 (64.6%). In the Presidio County sample 29 specimens all (96.6%) except one had two preanal scales. In summary, these analyses revealed the presence of unanticipated differences in anterior extent of the circumorbital series, size of mesoptychial scales near midpoint of the guiar fold, and number of preanal scales between Texas and New Mexico arrays of A. neomexicana.

A comparison of statistical data from samples of A. neomexicana and A. tesselata E from the sites near the Rio Grande south of Candelaria, Presidio County, sorted by subjective characters revealed the presence of mutually exclusive ranges of variation in the GAB character and nearly so in the GOR character, as well as significant differences in the PV/GAB, SDL, and ILS (see Table 4).

A comparison of samples of A. neomexicana and A. inomata from sites of syntopy in Hidalgo County, New Mexico, sorted into samples based on the foregoing color pattern differences, were also distinctive with reference to the following scutellation character states (Table 2) described for the former species followed by the latter species: granular postantebrachial scales (versus very slightly enlarged); small mesoptychial scales bordering the edge of the guiar fold (versus variable size); and circumorbital scale series either complete or nearly so (versus typically only to middle of third supraocular scales). The samples of these species also had mutually exclusive ranges of variation in the GAB and GOR characters, as well as significant differences in the PV, FP, SDL, COS, LSG, and ILS (Tabic 4).

Comparisons of allopatric samples of A. neomexicana, two from Texas and two from New Mexico, revealed interstate differences in meristic characters as well as the aforementioned anterior extent of the circumorbital scale series. However, statistical comparisons of the four samples revealed the following (Table 4): three samples were significantly different in the FP [A. n. (PC-TX), A. n. (E/HC-TX, and A. n. (HC-NM)]; none were significantly different in the PV and LSG; and the HC-NM New Mexico sample was significantly different from both Texas samples that were alike in the GAB, GOR, SDL, and ILS.

Maximum snout vent length (SVL).--The maternal progenitor of both of these syntopic diploid parthenogenetic species in Presidio County was A. marmorata (Eastern Marbled Whiptail); however, the maximum SVL of A. neomexicana and A. tesselata E from the Candelaria sites were 72 and 94 mm, respectively.

Conclusions.--Several genetic studies have characterized allodiploid parthenogenetic A. neomexicana in New Mexico as having low clonal diversity, indicating a recent origin via hybridization of A. marmorata x A. inomata (Parker & Selander 1984; Cole et al. 1988; Dessauer & Cole 1989; Manning et al. 2005). Nevertheless, this study identified intriguing geographical differences in morphology (e.g., number of preanal scales, anterior extent of circumorbital series, and size of central mesoptychial scales anterior to guiar fold) and color pattern (e.g., contrast between pale stripes and dark fields). Samples from Texas utilized in this study form part of an unusual distribution pattern in which arrays of a parthenogenetic specics are disjunctively represented in Arizona (Persons & Wright 1999) and Utah (Oliver & Wright 2007) from inferred human introductions, and in New Mexico (Dessauer et al. 2000; Burkett et al. 2004; Manning et al. 2005; Cordes et al. 2011) and Texas (Axtell 1966; Cordes & Walker 2009; Walker et al. 2013) from natural occurrences. For different interpretations of the geographic origins of arrays of the species in New Mexico see Leuck et al. (1981) and Oliver & Wright (2007).

The only study bearing on the genetic relationship of Texas arrays of A. neomexicana to those elsewhere was that of Cordes et al. (1990) in which skin patches exchanged between a lizard (UADZ 3908; 65 mm SVL; Table 1) from south of Candelaria, Presidio County, and one from Conchas Lake, San Miguel County, New Mexico, were mutually accepted. Mutual histocompatibility between these individuals was inferred by Cordes et al. (1990) to indicate genetic homogeneity between the two arrays of parthenogenetic A. neomexicana which would be expected only if they were the descendants, many generations removed, from a single hybrid individual. In conclusion, a sample of lizards from near the Rio Grande south of Candelaria, the area of origin of one of the lizards used in the histocompatibility experiment, is conclusively representative of an array of A. neomexicana herein verified to exist in Presidio County as first reported by Axtell (1966) based on one specimen.

Acknowledgments

All UADZ voucher specimens of Aspidoscelis used in this study were collected in accordance with the provisions of New Mexico Game and Fish Department permit 1850 and Texas Parks and Wildlife Department permits SP061 and SPR-1090-298 issued to JEC.

Literature Cited

Axtell, R.W. 1966. Geographical distribution of the unisexual whiptail Cnemidophorus neomexicanus (Sauria: Teiidae)-present and past. Herpetologica 22:241-253.

Burkett, D. W., M. Hartsough & N. Swink. 2004. Geographic distribution. Aspidoseclis (= Cnemidophorus) neomexicana. Herpetol. Rev. 35:408.

Cole, C. J., 11. C. Dessauer & G. F. Barrowclough. 1988. Hybrid origin of a unisexual species of whiptail lizard, Cnemidophorus neomexicanus. in western North America: new evidence and a review. Am. Mus. Novitat. 2905:1-38.

Cordes, J. E. & J .M. Walker. 2009. Parthenogenetic Aspidoscelis neomexicana (Sauria: Teiidae) and syntopic congeners in Presidio County, Texas. Southwest. Nat. 54:226-230.

Cordes, J. E., J. M. Walker & R. M. Abuhteba. 1990. Genetic homogeneity in geographically remote populations of parthenogenetic Cnemidophorus neomexicanus (Sauria: Teiidae). Texas J. Sci. 42:303-305.

Cordes, J. E., J. M. Walker & G. J. Manning. 201 1. Geographic Distribution. Aspidoscelis neomexicana (New Mexico Whiptail). Herpetol. Rev. 42:568.

Cordes, J. E., J. M. Walker, J. F. Scudday & R. M. Abuhteba. 1989. Distribution and habitat of the parthenogenetic whiptail lizard. Cnemidophorus neomexicanus (Sauria: Teiidae), in Texas. Texas J. Sci. 41:425-428.

Dessauer, H. C. & C. J. Cole. 1989. Diversity between and within nominal forms of unisexual teiid lizards. In R.M. Dawley & J.P. Bogart (editors). Evolution and ecology of unisexual vertebrates: 49-71. Albany, NY: New York State Mus. Bull. 466.

Dessauer, H. C., C. J. Cole & C. R. Townsend. 2000. Hybridization among western whiptail lizards (Cnemidophorus tigris) in southwestern New Mexico: population genetics, morphology, and ecology in three contact zones. Bull. Am. Mus. Nat. Hist. 246:1 148.

Dixon. J. R. 2013. Amphibians and reptiles of Texas: with keys, taxonomic synopses, bibliography, and distribution maps. Texas A & M University Press. College Station.

Leuck, B. E., E. E. Leuck II & R. T. B. Sherwood. 1981. A new population of New Mexico whiptail lizards, Cnemidophorus neomexicanus (Teiidae). Southwest. Nat. 26:72-74.

Lowe, C. H. & R. G. Zweifel. 1952. A new species of whiptailed lizard (genus Cnemidophorus) from New Mexico. Bull. Chicago. Acad. Sci. 9:229-247.

Manning, G. J., C. J. Cole. FL C. Dessauer & J. M. Walker. 2005. Hybridization between parthenogenetic lizards (Aspidoscelis neomexicana) and gonochoristic lizards (Aspidoscelis sexlineata viridis) in New Mexico: ecological, morphological, cytological, and molecular context. Am. Mus. Novitat. 3492:1-56.

Oliver, G. V. & J. W. Wright. 2007. The New Mexico Whiptail, Cnemidophorus neomexicanus (Squamata: Teiidae), in the Great Basin of north central Utah. Western North Am. Nat. 67:461-467.

Parker, E. D., Jr. & R. K. Selander. 1984. Low clonal diversity in the parthenogenetic lizard Cnemidophorus neomexicanus (Sauria: Teiidae). Herpetologica 40:245-252.

Persons, T. & J. W. Wright. 1999. Discovery of Cnemidophorus neomexicanus in Arizona. Herpetol. Rev. 30:207-209.

Reeder, T. W., C. J. Cole & H. C. Dessauer. 2002. Phylogenetic relationships of whiptail lizards of the genus Cnemidophorus (Squamata, Teiidae): a test of monophyly. reevaluation of karyotypic evolution, and review of hybrid origins. Am. Mus. Novitat. 3365:1-61.

Walker, J. M., J. E. Cordes, D. W. Burkett & J. F. Scudday. 2013. Aspidoscelis neomexicana (New Mexico Whiptail). Habitat. Herpetol. Rev. 44:318-319.

JMW at: jmwalker@ uark.edu

James M. Walker (1), James E. Cordes (2) and James R. Dixon (3)

(1) Department of Biological Sciences, University of Arkansas, Fayetteville, Arkansas 72701

(2) Division of Sciences and Mathematics, Louisiana State University Eunice, Eunice, Louisiana 70535

(3) Curator Emeritus of Amphibians and Reptiles, Texas Cooperative Wildlife Collection. Texas A & M University, College Station, Texas 77843 (Deceased)

Caption: Figure 1. Specimens of diploid parthenogenetic Aspidoscelis neomexicana (Lowe & Zweifel, 1952) collected by JEC on 13 July 1992 from Presidio County, 1.5-3.9km S of Candelaria via Texas Farm Road 170 (30.103889[degrees]N, 104.667222[degrees]W). Left, UADZ 4698 adult female of 67 mm SVL; right, UADZ 4700 adult female of 72 mm SVL.
Table 1. Specimens of Aspidoscelis neomexicana, A. tesselata pattern
class E, and A. inornata examined for comparative data used in this
study (UADZ = University of Arkansas "Department of Zoology;" SRSU =
Sul Ross State University; and UTEP = University of Texas at El Paso).

                  State
Species (N)       (Visits)   Localities

A. neomexicana    Texas      Presidio Co.: 1.5-3.9 km S
(29)              (18)       Candelaria via TX FR 170,
                             30.103889[degrees]N,
                             104.667222[degrees]W

A. neomexicana    Texas      El Paso Co.: vicinities Hueco
(9)               (6)        Tanks State Park, Horizon
                             Lake, Hueco Mountains

A. neomexicana    Texas      Hudspeth Co.: between
(2)               (1)        Horizon Lake and National
                             Compressor Station

A. neomexicana    New        Hidalgo Co.: 21.3 km NW jct
(24)              Mexico     US Hwys 90 and 70, on 70,
                  (7)        32.515000[degrees]N,
                             108.876667[degrees]W

A. neomexicana    New        Grant Co.: 2.88 km N of
(9)               Mexico     Hidalgo/Grant boundary near
                  (2)        New Mexico Hwy 464

A. tesselata E    Texas      Presidio Co.: 1.5-3.9 km S
(25)              (16)       Candelaria via TX FR 170,
                             30.103889[degrees]N,
                             104.667222[degrees]W

A. inornata       New        Hidalgo Co.: 21.3 kmNWjct
(18)              Mexico     US Hwys 90 and 70 on 70.
                  (7)        32.515000[degrees]N,
                             108.876667[degrees]W

Species (N)        Specimens Examined

A. neomexicana     UADZ 3466, 3792-3794, 3801(29)
3803, 3807-3810, 3823-3826,
                   3848, 3881, 3898-3899, 3908,
                   3910, 3915, 4621, 4698, 4700,
                   4722, 5128, 8259, 8380

A. neomexicana     SRSU 609, 1695-1696, 4156,
(9)                4161-4162, 4195, 10793-10794

A. neomexicana     SRSU 4185-4186
(2)

A. neomexicana     UADZ 6079-6080, 6082, 6180(24)
6183, 6188-6189, 6195-6196,
                   6456-6458, 6490-6496, 69066908

A. neomexicana     UADZ 7070-7074, 7093-7096
(9)

A. tesselata E     UADZ 3795, 3798-3800, 3812,
(25)               3827-3828, 3847, 3858-3860,
                   3875-3878, 3911, 3921, 4619,
                   4699, 4702, 5120-5121, 5130,
                   8258, 8379

A. inornata        UADZ 6090-6091. 6173, 6175(18)
6178,6193-6194, 6459-6462
                   6497-6498, 6909-6911

Table 2. Characters referenced qualitatively in this report pertaining
to variation in scutellation and color pattern in samples of specimens
of diploid parthenogenetic Aspidoscelis neomexicana from Texas and New
Mexico, diploid parthenogenetic A. tesselata from Texas, and
gonochoristic A. inornata from New Mexico.

Character                  Description

Anterior Extent of         Complete or incomplete series medial to
Circumorbital Scales       right and left sides of head supraoculars on

Size of Mesoptychial       Expressed as small or abruptly enlarged
Scales                     fold bordering the guiar

Size of Central            Expressed as either evenly small or
Mesoptychials              in center of guiar fold unusually smaller
                           scales

Size of Postantebrachial   Expressed as small, moderately enlarged, or
Scales                     distal part of forelimb enlarged on

Number of Preanal          Expressed as one or two scales anterior to
Scales                     vent

Lateral Stripes            Pale stripe on each side superior to lateral
                           row of ventral scales

Dorsolateral Stripes       Pale stripe on each side superior to lateral
                           stripe

Paravertebral Stripes      Pale stripe on each side superior to
                           dorsolateral stripe

Vertebral Stripe           Pale midsagittal stripe between the
                           paravertebral stripes

Lower Lateral Fields       Dark field on each side between lateral row
                           and lateral stripe of ventral scales

Upper Lateral Fields       Dark field on each side between lateral and
                           stripes dorsolateral

Dorsolateral Fields        Dark field on each side between dorsolateral
                           paravertebral stripes and

Vertebral Field            Dark field between paravertebral stripes
                           divided by vertebral stripe longitudinally

Dorsal Spots               Expressed as present or absent lightfields
                           and/or on stripes colored rounded
                           areas in

Table. 3. Comparison of the anterior extent of the circumorbital scale
series located on each side between the supraocular and median head
scales, designated by previous workers as a critical distinguishing
character of A. neomexicana (A. n.), in samples of the species from
Texas and New Mexico compared with A. tesselata pattern class E (A. t.
E) from Texas and A. inomata (A. i.) from New Mexico (LC-RC = left
complete-right complete; LC-RI left complete-right incomplete; LI-RC =
left incomplete-right complete; LI-RI = left incomplete-right
incomplete).

State: County       N     Species        LC-RC

TX: Presidio Co.   27      A. n.       6 (22.2%)

TX: El Paso/       10      A. n.         4 (40%)
Hudspeth Cos.

NM: Hidalgo Co.    24      A. n.      21 (87.5%)

NM: Grant Co.       9      A. n.      9 (100.0%)

TX: Presidio Co.   23     A. t. E         --

NM: Hidalgo Co.    18      A. i.          --

State: County        LC-RI        LI-RC         LI-RI

TX: Presidio Co.   3 (11.1%)    3 (11.1%)    15 (55.6%)

TX: El Paso/         4 (40%)      1 (10%)       1 (10%)
Hudspeth Cos.

NM: Hidalgo Co.        --           --        3 (12.5%)

NM: Grant Co.          --           --        --

TX: Presidio Co.       --           --        23 (100%)

NM: Hidalgo Co.        --           --        18 (100%)

Table 4. Data for meristic characters (above mean [+ or -] SE and
below range and N) for samples of Aspidoscelis neomexicana (A. n.)
from Presidio (PC-TX) and El Paso-Hudspeth (E-HC-TX) counties, Texas,
and Hidalgo (HC-NM) and Grant (GC NM) counties, New Mexico, A.
tesselata pattern class E (A. t. E) from Presidio (PC'-TX) County,
Texas, and A. inornata (A. i.) from Hidalgo (HC-NM) County, New
Mexico. Only means [+ or -] SE for the same character with all
different superscripted letters are significantly different (P =
0.05). Abbreviations for sample repositories are: UADZ = University of
Arkansas Department of Zoology; SRSU = Sul Ross State University; and
UTEP = University of Texas at El Paso.

                                                  A.n.
                                                 (E/HC-
Character                       A. n.              TX)
Abbreviation:                  (PC-TX)            SRSU,
Description                     UADZ              UTEP

GAB: granular scales        77.6 [+ or -]     76.1 [+ or -]
around midbody from           0.65 (C)            0.601
one lateral row of
ventral scales to               73-86             73-79
opposite row of lateral         (27)              (10)
scales

GOR: granular scales       193.7 [+ or -]    195.8 [+ or -]
between occipital and         0.62 (B)          l.82 (B)
first row of caudal
scales                         187-199           188-205
                                (27)              (10)

PV: granular scales        12.1 [+ or -]     11.7 [+ or -]
between paravertebral         0.17 (AB)         0.45 (AB)
stripes at midbody
                                10-14             10-14
                                (27)              (10)

PV/GAB x 100: % of         15.7 [+ or -]     15.3 [+ or -]
granular scales around        0.22 (A)          0.64 (AB)
midbody located
between paravertebral           13.1-             12.9-
stripes at midbody            18.9(27)          18.6(10)

FP: femoral pores          39.2 [+ or -]     37.3 [+ or -]
summed from both              0.25 (B)          0.52 (C)
hind limbs
                                36-42             35-41
                                (27)              (10)

SDL: subdigital             31.8 [+ or -]     31.7 [+ or -]
lamellae of the fourth        0.19 (C)          0.30 (C)
toe of the
left pes                        30-34             30-33
                                (27)              (10)

COS: circumorbital          19.9 [+ or -]     21.2 [+ or -]
scales summed from            0.47 (B)          0.68 (A)
between supraocular
and median head                 16-24             16-24
scales from both sides          (27)              (10)

LSG: lateral                40.5 [+ or -]     39.5 [+ or -]
supraocular granules          0.96 (AB)         0.82 (AB)
summed from
anterior to third               31-52             37-45
supraocular sutures             (27)              (10)
from both sides

ILS: interlabial scales,    29.7 [+ or -]     27.3 [+ or -]
summed from both              1,06 (C)          1.61 (C)
sides
of head                         21-44             18-36
                                (27)              (10)

                                A. n.             A. n.
Character                       (HC-              (GC-
Abbreviation:                    NM)               NM)
Description                     UADZ              UADZ

GAB: granular scales        80.7 [+ or -]     79.0 [+ or -]
around midbody from           0.44 (B)          1.05 (BC)
one lateral row of
ventral scales to               75-83             73-83
opposite row of lateral         (24)               (9)
scales

GOR: granular scales       207.9 [+ or -]    210.6 [+ or -]
between occipital and         1 ,52 (A)         3.03 (A)
first row of caudal
scales                         193-220           190-221
                                (24)               (9)

PV: granular scales        11.6 [+ or -]     12.0 [+ or -]
between paravertebral         0.15 (B)          0.33 (AB)
stripes at midbody
                                11-13             10-13
                                (24)               (9)

PV/GAB x 100: % of          14.4 [+ or -]     15.0 [+ or -]
granular scales around        0.16 (BC)         047 (ABC)
midbody located
between paravertebral           13.2-             12.8-
stripes at midbody            15.6(24)           16.6(9)

FP: femoral pores           40.8 [+ or -]     40.2 [+ or -]
summed from both              0.31 (A)          0.28 (AB)
hind limbs
                                38-43             39-41
                                (24)               (9)

SDL: subdigital             34.3 [+ or -]     33.9 [+ or -]
lamellae of the fourth        0.20 (B)          0.26 (B)
toe of the
left pes                        33-36             33-35
                                (24)               (9)

COS: circumorbital          24.0 [+ or -]     23.9 [+ or -]
scales summed from            0.32 (A)          0.42 (A)
between supraocular
and median head                 20-26             22-26
scales from both sides          (24)               (9)

LSG: lateral                43.8 [+ or -]     42.8 [+ or -]
supraocular granules          1.11 (A)          0.94 (A)
summed from
anterior to third               30-55             39-48
supraocular sutures             (24)               (9)
from both sides

ILS: interlabial scales,    35.4 [+ or -]     31.3 [+ or -]
summed from both              0.90 (B)          1.00 (BC)
sides
of head                         29-46             27-36
                                (24)               (9)

                                                  A. i.
Character                      A. t. E            (HC-
Abbreviation:                  (PC-TX)             NX)
Description                     UADZ              UADZ

GAB: granular scales        99.4 [+ or -]     61.7 [+ or -]
around midbody from           0.54 (A)           0.8 (D)
one lateral row of
ventral scales to              94-105             57-69
opposite row of lateral         (23)              (18)
scales

GOR: granular scales       211.1 [+ or -]    151.0 [+ or -]
between occipital and         1,09 (A)          1,56 (C)
first row of caudal
scales                         197-219           139-160
                                (23)              (18)

PV: granular scales         12.8 [+ or -]     8.3 [+ or -]
between paravertebral         0.25 (A)          0.35 (C)
stripes at midbody
                                10-15             6-12
                                (23)              (18)

PV/GAB x 100: % of          12.8 [+ or -]     13.5 [+ or -]
granular scales around        0.23 (D)          0.51 (CD)
midbody located
between paravertebral           10.8-             11.2-
stripes at midbody            15.1 (23)         19.0(18)

FP: femoral pores           39.8 [+ or -]     31.5 [+ or -]
summed from both              0.32 (AB)         0.53 (D)
hind limbs
                                37-43             28-35
                                (23)              (18)

SDL: subdigital             36.0 [+ or -]     27.7 [+ or -]
lamellae of the fourth        0.26 (A)          0.39 (D)
toe of the
left pes                        34-39             26-32
                                (23)              (18)

COS: circumorbital          19.3 [+ or -]     11.2 [+ or -]
scales summed from            0.32 (B)          0.60 (C)
between supraocular
and median head                 14-21             9-19
scales from both sides          (23)              (18)

LSG: lateral                39.6 [+ or -]     25.1 [+ or -]
supraocular granules          0.66 (B)          1.26 (C)
summed from
anterior to third               34-47             16-41
supraocular sutures             (23)              (18)
from both sides

ILS: interlabial scales,    43.7 [+ or -]     21.5 [+ or -]
summed from both              0.64 (A)          1.14 (D)
sides
of head                         38-50             14-33
                                (23)              (18)
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Article Details
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Author:Walker, James M.; Cordes, James E.; Dixon, James R.
Publication:The Texas Journal of Science
Article Type:Report
Geographic Code:1USA
Date:Feb 1, 2016
Words:4593
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