Ameiva ameiva complex in Venezuela hmon-25-00-113-170-e

HERPETOLOGICAL M O N O G R A P H S

REVISION OF AMEIVA AMEIVA LINNAEUS (SQUAMATA: TEIIDAE) IN VENEZUELA: RECOGNITION OF FOUR SPECIES AND STATUS OF INTRODUCED POPULATIONS IN SOUTHERN FLORIDA, USA GABRIEL N. UGUETO1,3

2

AND

MICHAEL B. HARVEY2

1 11111 Biscayne Boulevard, No. 556, Miami, FL 33181, USA Department of Biological Sciences, Broward College, 3501 Southwest Davie Road, Davie, FL 33314, USA

Herpetological Monographs, 25, 2011, 113–170 E 2011 by The Herpetologists’ League, Inc.

REVISION OF AMEIVA AMEIVA LINNAEUS (SQUAMATA: TEIIDAE) IN VENEZUELA: RECOGNITION OF FOUR SPECIES AND STATUS OF INTRODUCED POPULATIONS IN SOUTHERN FLORIDA, USA GABRIEL N. UGUETO1,3

AND

MICHAEL B. HARVEY2

1

2

11111 Biscayne Boulevard, No. 556, Miami, FL 33181, USA Department of Biological Sciences, Broward College, 3501 Southwest Davie Road, Davie, FL 33314, USA

ABSTRACT: The common, terrestrial teiid lizards of the Ameiva ameiva complex have a broad Neotropical distribution from Costa Rica and Panama to Paraguay and northern Argentina. These lizards have also been reported from the Lesser Antilles, Trinidad, Tobago, Providencia, and the Swan Islands. The group has a complicated taxonomic history and has not been comprehensively reviewed. With the use of external morphology, we reviewed this complex in Venezuela. We conclude that A. ameiva (sensu lato) in Venezuela comprises at least four evolutionary species and recommend elevating A. ameiva (sensu stricto), A. atrigularis, and A. praesignis to species. On the other hand, we synonymize A. ameiva fischeri, A. a. ornata, and A. a. vogli, with A. praesignis; we synonymize A. a. laeta and A. a. petersii with A. ameiva; and we synonymize A. a. melanocephala with A. atrigularis. We designate a lectotype of A. atrigularis. Populations from savannahs of southern Monagas represent a new species described in this work. Both A. ameiva and A. praesignis exist as apparently viable breeding populations in southern Florida, USA. RESUMEN: El lagarto teiido Ameiva ameiva, es una especie comu´n con una distribucio´n muy amplia por el neotro´pico, desde Costa Rica y Panama´ hasta Paraguay y el norte de Argentina. La especie tambie´n esta presente en las antillas menores, Trinidad, Tobago, Providencia y la Islas del Cisne. La historia taxono´mica de A. ameiva ha sido controvertida, cao´tica y no ha sido revisada recientemente. En el presente trabajo, revisamos esta especie en Venezuela y examinamos la morfologı´a externa y coloracio´n de varias poblaciones correspondientes con las varias subespecies sen˜aladas para el paı´s. Se concluye que Ameiva ameiva (sensu lato) esta conformada por al menos cuatro especies evolucionarias y recomendamos elevar A. ameiva (sensu stricto), A. atrigularis y A. praesignis a especies. Consideramos a A. ameiva fischeri, A. a. ornata y A. a. vogli como sino´nimos de A. praesignis; tambien consideramos A. a. laeta y A. a. petersii sino´nimos de A. ameiva y A. a. melanocephala sino´nimo de A. atrigularis. Designamos un lectotipo de A. atrigularis. Las poblaciones de las sabanas del sur del estado Monagas resultaron ser una especie nueva la cual describimos en este trabajo. Adicionalmente, aclaramos la identidad taxono´mica de las poblaciones de Ameiva introducidas en los Estados Unidos de Norte Ame´rica. Reportamos varias poblaciones introducidas de A. ameiva y una de A. praesignis en el sureste del estado de la Florida.

Key words: Ameiva ameiva; Ameiva atrigularis; Ameiva pantherina; Ameiva praesignis; Monagas; New Species; South America; Squamata; Systematics; Teiidae; Venezuela

were based on color pattern of adult specimens with few or no additional corroborating characters (e.g., Donoso-Barros, 1968; Peters and Donoso-Barros, 1970). Some authors adopted a more conservative view and submerged different subspecies within a monotypic, widespread A. ameiva (e.g., Vanzolini, 1986).

The genus Ameiva Meyer is currently recognized as a paraphyletic group with respect to other cnemidophorine teiids (Reeder et al., 2002). This genus contains terrestrial, medium to large, diurnal lizards distributed from Mexico to Peru west of the Andes, and to northern Argentina east of that mountain chain. Ameiva also extends throughout the Antilles and the Bahamas (Savage, 2002).

SUBSPECIES and species concepts have been the subject of intense discussion among biologists in the last century, especially during the last few decades (e.g., de Queiroz, 1998).

Herpetologists have long suspected that subspecies of many widespread squamates actually represent evolutionary species. Recent partition of Bothropoides neuwiedi, Epicrates cenchria, and Micrurus frontalis (Jorge da Silva and Sites, 1999; Passos and Fernandes, 2008; Silva and Rodrigues, 2008) reveals that these suspicions are well founded. Most previous reviews of Ameiva ameiva (Linnaeus, 1758) recognized several subspecies, probably because the most useful diagnostic characters 3

CORRESPONDENCE: e-mail, gabrieluguetto@yahoo.com 113

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Among the 33 species currently included in this genus (Uetz, 2010), A. ameiva has the broadest distribution. In Central America it occurs from extreme southwestern Costa Rica to central Panama; in South America, its range extends from northern Colombia and Venezuela to southeastern Brazil, Paraguay, and northern Argentina. Ameiva ameiva has also been reported on the Caribbean islands of Providencia, Trinidad, Tobago, and the southern Lesser Antilles (Savage, 2002). Historically, this species occurred on the Swan Islands, where it apparently went extinct before 1914 (Dunn and Saxe, 1950). Additionally, it has been introduced to several localities of southeastern Florida in the United States (Smith and Krysko, 2007).

The taxonomic history of A. ameiva is chaotic. Most confusion surrounding this species complex was generated in the 19th and early 20th centuries. Until the last two decades, many authors regarded A. ameiva as a polytypic taxon and recognized the following subspecies: A. a. ameiva (Linnaeus, 1758) occurring in southern Venezuela, Guyana, Suriname, French Guiana and northern Brazil; A. a. atrigularis Garman (1887) from the islands of Trinidad and Tobago; A. a. fischeri Peters and Donoso-Barros (1970) from Sabana Larga and Sierra de Santa Marta in northern Colombia; A. a. fulginosa Cope (1862) from the Swan Islands and the Colombian island of Providencia; A. a. laeta Cope (1862) occurring in southeastern Brazil; A. a. melanocephala Barbour and Noble (1915) occurring in north-central and northeastern Venezuela, including Isla de Margarita; A. a. ornata Mu¨ller and Hellmich (1940) from the Llanos of Fusagasuga´ in Colombia; A. a. petersii Cope (1868) occurring in the upper Amazon Basin; A. a. praesignis (Baird and Girard, 1852) occurring mostly in open and arid formations in Panama, northern Colombia and northwestern Venezuela; A. a. tobagana Cope (1879) from the Lesser Antilles; A. a. vogli Mu¨ller (1929) from the Llanos of western and central Venezuela (Donoso-Barros, 1968; Dunn and Saxe, 1950; Peters and Donoso-Barros, 1970; Staton and Dixon, 1977; Tuck and Hardy, 1973). However, the status of the different subspecies was unstable, in part because of the lack of

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sufficient comparative material and varying interpretations of geographic and ontogenetic variation in size, squamation, and color pattern. In spite of the striking geographic color variation exhibited by A. ameiva in the northern part of its distribution, most herpetologists during the last three decades have followed Vanzolini (1986), who considered the subspecies of A. ameiva to be invalid and biologically unmeaningful. Ameiva ameiva is now usually regarded as a widespread, morphologically and ecologically variable species.

Based on material from international museum collections, which provided thorough geographic coverage for this study, we examined morphological variation within A. ameiva in Venezuela and tested whether sufficient evidence exists for recognizing separate taxa in the country. The purpose of this article is to report the results of these tests, present a key for the species of the genus Ameiva in Venezuela, and propose a new taxonomy that clarifies the complicated history of A. ameiva in South America.

MATERIALS AND METHODS We measured snout–vent length (SVL, from tip of snout to cloacal opening), head length (from tip of snout to anterior margin of ear opening), head width (widest part of head), head depth (highest part of head), snout length (from tip of snout to anterior corner of eye), tibia length (from middle of tibio-femoral articulation to middle of tibiotarsal articulation), foot length (from tibiotarsal articulation to tip of fourth toe), hand length (from radio-carpal articulation to tip of fourth finger), tail length (when complete), and total length with a common ruler to the nearest 1 mm or a dial caliper to the nearest 0.1 mm.

For the most part, we use the terminology of Echternacht (1971), Avila-Pires (1995), and Ugueto and Harvey (2010). As discussed by Ugueto and Harvey (2010), we consider the large scale abutting the anterior ocular angle and forming the lower edge of the orbit to be a subocular whether or not it contacts the supralabials. As in Cnemidophorus (Colli et al., 2003; Ugueto et al., 2010), the gular region in A. ameiva is divided into two areas. A line of granules uniting the posterior margins of the

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auditory meati separates the anterior gulars from the posterior gulars. Scales of the anterior region are small but not granular and arranged in transverse rows that tend to enlarge (sometimes more or less conspicuously) midventrally. Some specimens have a midventral patch of enlarged posterior gulars in front of the antegular fold. Scales between the supraoculars and supraciliaries may be entirely or mostly in a single series or in an entirely divided series. We recorded anterior extent of the circumorbital semicircles relative to adjacent supraoculars. We assigned shape of the central terminal scale of the preanal shield in males to one of the categories of Markezich et al. (1997).

We combined counts from both sides for supralabials (we consider the scale below the tip of third subocular to be the last supralabial), infralabials (last infralabial lies directly below last supralabial), supraoculars, supraciliaries, circumorbitals (counted as scales touching supraocular from posteriormost portion of last supraocular to last scale between anterior supraoculars and other cephalic scales), scales between supraoculars and supraciliaries, and femoral pores. We also counted the total number of parietals (including the interparietal), occipitals around the parietals, ventrals in a transverse row (at the 15th ventral row), transverse rows of ventrals (from gular fold to anterior margin of leg), transverse rows of dorsals (between interparietal and enlarged keeled scales at base of tail), scales around midbody (excluding ventrals), lamellae under the fourth finger, lamellae under the fourth toe (both palmar and pedal lamellae were routinely counted only on a specimen’s left side, except when damaged, in which case those on right side were counted), and number of preanal scales (from between femoral pores to terminal largest scale). In the accounts, means 6 standard deviation follow ranges when appropriate. We recorded measurements and scale counts for all specimens of the new species and for most of the material examined (Appendix 1; Table 1).

Following data collection, each of the characters was classified as nominal, ordinal, or interval (Kleinbaum et al., 1988). For univariate statistical analyses, we assigned specimens to one of seven geographically

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and taxonomically defined groups (Table 2).

For analysis of interval characters, we used the PAST software program (Hammer et al., 2001). For each of the interval characters, we tested for normality (Shapiro-Wilk test) and homogeneity of variance (Levene’s test). To test the null hypothesis that means are equal for the various characters, we used a one-way analysis of variance (ANOVA) when assumptions of normality and homoscedasticity were satisfied. When the data violated either or both assumptions, we used the Kruskal-Wallis nonparametric test. We used Tukey’s HSD test (when assumptions were satisfied) or a Bonferroni-corrected Mann-Whitney U-test (when assumptions were violated) for pairwise comparisons among the species.

In northern Venezuela, the ranges of two putative taxa come into contact. To look for possible cases of character displacement in the zone of contact, we regressed each of our meristic characters on geographic longitude.

Obvious divergence in character clines at the contact zone provided evidence of character displacement.

For mensural characters, we used analysis of covariance (ANCOVA), treating SVL as a covariate. An F-test for equality of the regression slopes was used to test the assumption of parallelism. As for other interval characters, we used Levene’s test and the Shapiro-Wilk test to test assumptions of homoscedasticity and normality before the ANCOVA. Sex was determined by subcaudal dissection except in those males that possessed everted hemipenis. Drawings were made with the aid of photographs from preserved specimens.

Data on habitat and natural history include our own observations and those from trusted sources. Museum abbreviations are those of Leviton et al. (1985). In the accounts, ranges are separated by a dash, whereas counts from opposite sides of specimens are separated by a backslash.

RESULTS Our analyses (see further below) found concordance (sensu Avise and Ball, 1990) of meristic, mensural, coloration, and ecological characters to define four taxa in Venezuela, and each of these taxa should be recognized as a distinct species. From here on we use

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specific names for the different taxa, which will be described in detail after the discussion of characters used in our analysis.

We found no evidence of morphological intergradation or hybridization among species of Ameiva recognized in this study. This is the case even when our samples approach each other geographically, suggesting absence of or very narrow hybrid zones. We follow the general lineage species concept as proposed by de Queiroz (1998) and do not recognize subspecies within the A. ameiva complex in Venezuela. Names are available for three of the four species, and we propose a new name for a new species from southern Monagas.

Several characters proved to be useful for diagnosing the four species and are here discussed in detail.

Meristic Characters and Scalation Six meristic characters differed among several pairs of populations (Tables 1 and 2) and proved particularly useful for inferring species boundaries. The most useful scale characters for identifying species of the A. ameiva complex are numbers of subdigital lamellae of the fourth toe, posterior gulars, midbody scale rows, occipitals, and granular scales between the supraoculars and supraciliaries (Table 1). In the following sections, we elaborate on three useful scale characters.

Occipital scales.—Ameiva atrigularis has fewer and relatively larger occipitals than other species, although this character is significant only in pairwise comparisons with A. praesignis or Ameiva sp. nov. Most (31 of 40 specimens, 80%) A. atrigularis have two or more enlarged occipitals that are conspicuously larger than the first row of dorsals. In A. ameiva, A. praesignis, and Ameiva sp. nov., the first row of occipitals is usually subequal in size to the first row of dorsals (Fig. 1).

Median anterior gulars.—Size of the median anterior gular scales distinguishes Ameiva atrigularis and Ameiva sp. nov. from the other two species. These scales are moderately to distinctly enlarged (grading to smaller scales laterally), quadrangular and juxtaposed in most A. ameiva (Fig. 2) and A. praesignis (Table 1). In contrast, most Ameiva sp. nov. and A. atrigularis have small or only barely enlarged median anterior gulars (Fig. 2;

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Table 1). Many Ameiva sp. nov. and A. atrigularis have rounded, juxtaposed median anterior gulars conspicuously separated by interstitial skin. This condition presumably could be an artifact of preservation, but it was not observed in A. ameiva or in most A. praesignis.

Enlarged posterior gular scales.—Ameiva ameiva has fewer transverse rows of posterior gular scales than the other three species (Tables 1 and 2). This difference is partially attributable to a median patch (sometimes not forming a distinct patch but scales grading to larger midventrally) of distinctly enlarged posterior gulars in most A. ameiva (77%). In contrast, these scales are small and do not form a differentiated patch in most (63%) A. atrigularis (Fig. 2). The median posterior gulars are intermediate in size in the remaining A. atrigularis, about one-half of the Ameiva sp. nov. specimens and most (38%) of A. praesignis (another 35% have these scales small as in A. atrigularis and the remaining 27% large like A. ameiva). We noted some variation in size of these scales between populations of A. atrigularis. Whereas 50% (n 5 16) of north-central A. atrigularis have small posterior gulars, 75% (n 5 24) of specimens from northeastern Venezuela and Trinidad have small posterior gulars.

Character Displacement and Parapatry We have not encountered two or more species of the A. ameiva complex in sympatry.

Nonetheless, ranges closely approach one another in several areas. For example, A. ameiva, Ameiva sp. nov., and A. praesignis all occur in close proximity to one another in the patchwork of forest and savannah of southern Anzoa´tegui and Monagas (Fig. 3). Where forests approach xeric coastal habitats, the ranges of A. atrigularis and A. praesignis come into contact in the vicinity of Caracas, Distrito Capital, and Vargas State.

Among the meristic characters, we identified possible character displacement in numbers of transverse dorsals and scales around midbody at this contact zone between A. atrigularis and northern A. praesignis. Transverse dorsal rows increase in number from east to west in A. atrigularis (Fig. 4). Individuals west of the Unare Depression (geographically

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closest to A. praesignis) have 301–361 dorsals, whereas those from northeastern Venezuela and Trinidad have 263–334 scales. The lowest counts occur in specimens from Trinidad, whereas specimens from Sucre and Isla de Margarita have intermediate values (320–327 dorsals). On the other hand, northern A. praesignis does not exhibit any apparent east– west cline in dorsal counts, except that two specimens (USNM 121194, 162763) from Vargas have the lowest counts of the entire sample. Scales around midbody show no apparent east–west cline in A. atrigularis, but increase from east to west in A. praesignis (Fig. 4). The two specimens of A. praesignis from Vargas (closest geographically to A. atrigularis) exhibit the lowest values.

Mensural Characters Most mensural characters considered by us exhibit sexual dimorphism. For this reason, we restricted our ANCOVAs between populations to males only. Moreover, since our sample from the highlands of Bolivar was relatively small, we did not consider it in our analyses of mensural characters. Our eastern and western samples of A. atrigularis do not differ in any mensural characters (i.e., ANCOVAs between these populations were nonsignificant). Southern specimens of A. praesignis have relatively shorter feet (F13,16 5 7.50, P 5 0.01) than northern A. praesignis, however no other characters differ between these populations.

Accordingly, we combined eastern with western A. atrigularis and northern with southern A. praesignis in all subsequent comparisons except for foot length where we treated the two populations of A. praesignis as separate operational taxonomic units (Table 3).

Populations of Ameiva differ in maximum snout–vent length and in relative head depth and tibia, foot and head (Table 3) lengths. After pooling the species, strict application of the Bonferroni correction for multiple tests requires that probabilities be less than 0.05/9 5 0.006 for foot length and less than 0.05/6 5 0.008 for the other characters. Applying this conservative criterion, Ameiva sp. nov. has a relatively shorter tibia than any of the other species (Fig. 5) and a shorter foot than A. ameiva or northern A. praesignis. Moreover, the very low probability that the foot lengths of

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Ameiva sp. nov. and A. atrigularis are the same (0.009 just barely falls short of the strict cutoff), strongly suggests that this character will distinguish these species as well (Fig. 5). Finally, inspection of Table 3 reveals that mensural characters likely distinguish A. ameiva and A. atrigularis from A. praesignis even though probabilities for these tests fall short of the strict Bonferroni-corrected cutoff values.

Ameiva praesignis is much larger than any of the other three species; some specimens reach lengths closer to other teiid genera such as Tupinambis. Males of this species reach a maximum SVL of 243 mm, whereas females attain 167 mm. Echternacht (1971) reported maximum SVL in Panama of 197 mm in males and 157 in females. Populations of southern A. praesignis from the Venezuelan Llanos appear to be somewhat smaller than those along the north-central coast and west of the Andes.

Nevertheless, Llanos specimens are still larger in size than any of the other three species.

Among a series of 36 specimens from this region, the largest male is 192 mm SVL. Ameiva atrigularis also attains a very large size, although smaller than A. praesignis.

Among the material studied, male A. atrigularis reach a maximum SVL of 186 mm. However, a specimen captured, and later released, in Fuentiduen˜o, Isla de Margarita, measured 190 mm SVL (Ugueto and Rivas, 2010). Ameiva ameiva is smaller than the two previous taxa (largest male examined was 170 mm in SVL).

Avila-Pires (1995) reported 174 mm as the maximum SVL for males in Brazilian Amazonia and this size exceeds any specimens that we examined. In contrast, Ameiva sp. nov. is considerably smaller than the other three species. The largest male among 32 specimens of this taxon was only 152 mm SVL, whereas the largest female was 142 mm.

Sexual Dimorphism Ameiva atrigularis exhibits the most extreme sexual size dimorphism. Males of this species have relatively longer, wider, and deeper heads, longer tibias, and longer hands (Fig. 6; Table 4). Moreover, allometry of each of the three head dimensions differs between the sexes as indicted by highly significant differences in slopes of the regression lines.

Different head width allometry characterizes

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TABLE 1.—Selected morphological and meristic characters of four species of Ameiva in Venezuela. Ranges (in parentheses) follow means 6 standard deviation.

Characters

Snout–vent length (SVL) Head length/SVL

Head width/SVL

Head depth/SVL

Tibia/SVL

Foot length/SVL

Hand length/SVL

Supralabials Infralabials Circumorbitals Supraorbital semicircles

Scales between supraoculars and supraciliaries Interparietals/ adjacent parietals

Occipitals Occipitals/first row of dorsals

Anterior gulars

Ameiva ameiva (n 5 22)

A. atrigularis (n 5 40)

A. pantherina (n 5 32)

A. praesignis (n 5 63)

Males: 170 mm Females: 142 mm Males: 0.25 6 0.00 (0.24–0.26) Females: 0.23 6 0.01 (0.23–0.26) Males: 0.14 6 0.01 (0.01–0.16) Females: 0.13 6 0.00 (0.11–0.14) Males: 0.15 6 0.00 (0.00–0.16) Females: 0.12 6 0.00 (0.12–0.13) Males: 0.24 6 0.01 (0.22–0.26) Females: 0.23 6 0.00 (0.22–0.24) Males: 0.42 6 0.03 (0.37–0.50) Females: 0.41 6 0.01 (0.38–0.43) Males: 0.16 6 0.01 (0.14–0.19) Females: 0.16 6 0.01 (0.14–0.26) 13.4 6 1.33 (12–17) 12.4 6 0.85 (12–15) 11.2 6 2.65 (6–17) Anterior third supraocular (4%) Mid-third supraocular (59%) Posterior third supraocular (37%) 28.3 6 7.18 (19–44)

Males: 186 mm Females: 146 mm Males: 0.25 6 0.00 (0.23–0.27) Females: 0.22 6 0.00 (0.22–0.24) Males: 0.15 6 0.01 (0.12–0.19) Females: 0.13 6 0.00 (0.12–0.15) Males: 0.15 6 0.01 (0.11–0.18) Females: 0.12 6 0.00 (0.11–0.14) Males: 0.23 6 0.01 (0.19–0.29) Females: 0.22 6 0.01 (0.21–0.25) Males: 0.39 6 0.03 (0.32–0.45) Females: 0.39 6 0.02 (0.34–0.42) Males: 0.16 6 0.00 (0.14–0.17) Females: 0.15 6 0.01 (0.14–0.19) 12.3 6 0.74 (12–15) 12.4 6 0.88 (10–15) 12.5 6 2.28 (6–17) Anterior third supraocular (10%) Mid-third supraocular (69%) Posterior third supraocular (21%) 27.3 6 6.38 (17–47)

Males: 152 mm Females: 138 mm Males: 0.24 6 0.01 (0.22–0.26) Females: 0.23 6 0.01 (0.21–0.26) Males: 0.14 6 0.01 (0.12–0.18) Females: 0.13 6 0.01 (0.12–0.17) Males: 0.14 6 0.01 (0.12–0.18) Females: 0.12 6 0.01 (0.11–0.15) Males: 0.21 6 0.01 (0.19–0.24) Females: 0.21 6 0.01 (0.18–0.24) Males: 0.35 6 0.03 (0.31–0.43) Females: 0.36 6 0.02 (0.32–0.41) Males: 0.16 6 0.02 (0.12–0.19) Females: 0.17 6 0.01 (0.14–0.23) 12.3 6 0.79 (11–14) 12.2 6 0.52 (12–14) 12.6 6 2.86 (9–24) Anterior third supraocular (16%) Mid-third supraocular (56%) Posterior third supraocular (28%) 29.7 6 4.71 (24–41)

Males: 243 mm Females: 167 mm Males: 0.24 6 0.01 (0.22–0.26) Females: 0.22 6 0.01 (0.21–0.26) Males: 0.15 6 0.02 (0.11–0.24) Females: 0.13 6 0.01 (0.11–0.19) Males: 0.14 6 0.01 (0.11–0.16) Females: 0.13 6 0.01 (0.11–0.2) Males: 0.21 6 0.01 (0.19–0.24) Females: 0.22 6 0.01 (0.20–0.24) Males: 0.35 6 0.03 (0.29–0.44) Females: 0.35 6 0.03 (0.30–0.41) Males: 0.16 6 0.01 (0.13–0.18) Females: 0.15 6 0.01 (0.12–0.18) 12.7 6 1.02 (12–16) 12.1 6 0.61 (10–14) 10.7 6 1.94 (6–17) Anterior third supraocular (9%) Mid-third supraocular (66%) Posterior third supraocular (25%) 22.6 6 4.98 (15–36)

Narrower (65%) Subequal (23%) Wider (12%) 14.6 6 1.49 (11–17) Smaller (4%) Subequal (63%) Larger (33%) 27.4 6 3.59 (18–32)

Narrower (51%) Subequal (36%) Wider (13%) 13.6 6 1.70 (10–16) Smaller (0) Subequal (20%) Larger (80%) 29.5 6 4.33 (22–40)

Narrower (31%) Subequal (47%) Wider (22%) 15.1 6 0.37 (14–18) Smaller (0) Subequal (56%) Larger (44%) 29.2 6 3.74 (20–36)

Narrower (53%) Subequal (19%) Wider (28%) 16.0 6 2.04 (12–21) Smaller (0) Subequal (75%) Larger (27%) 29.2 6 4.28 (22–40)

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TABLE 1.—Continued.

Characters

Size of anterior gulars

Posterior gulars Patch of enlarged posterior gulars

Dorsals Scales around body Scales around body/dorsals 3 100 Ventrals Longitudinal rows of ventrals Preanals

Femoral pores Enlarged brachials/ adjacent upper row Fourth finger lamellae Fourth toe lamellae

Ameiva ameiva (n 5 22)

A. atrigularis (n 5 40)

A. pantherina (n 5 32)

A. praesignis (n 5 63)

Small (10%) Medium (34%) Large (56%) 11.7 6 2.16 (7–17) Small (0) Medium (23%) Large (77%) 293.5 6 21.76 (259–339) 159.5 6 12.16 (129–187) 54.5 6 6.91 (43.2–72.2)

Small (45%) Medium (42%) Large (13%) 15.9 6 2.21 (13–23) Small (63%) Medium (37%) Large (0) 309.5 6 27.05 (263–361) 151.3 6 11.22 (134–179) 48.8 6 4.49 (38.4–60.8)

Small (84%) Medium (16%) Large (0) 16.5 6 1.64 (14–20) Small (44%) Medium (53%) Large (3%) 311.3 6 13.69 (291–343) 147.4 6 7.21 (137–163) 47.3 6 2.84 (43–53.9)

Small (27%) Medium (48%) Large (25%) 13.3 6 2.45 (9–18) Small (35%) Medium (38%) Large (27%) 289.3 6 23.19 (237–348) 133.0 6 11.26 (111–157) 45.8 6 3.36 (37.2–54.3)

30.5 6 0.89 (29–32) 10 6 0.00 (10–10) 4 (0) 5 (26%) 6 (44%) 7 (23%) 8 (7%) 37.5 6 4.39 (30–45) Subequal (15%) Larger (85%) 15.1 6 0.66 (14–16) 35.8 6 2.15 (32–40)

31.4 6 1.18 (29–34) 10 6 0.00 (10–10) 4 (10%) 5 (28%) 6 (41%) 7 (21%) 8 (0) 35.5 6 2.66 (31–42) Subequal (26%) Larger (74%) 15.5 6 0.88 (14–17) 36.3 6 2.10 (32–41)

all of the species, except perhaps Ameiva sp. nov. Ameiva ameiva exhibits similar dimorphism in the same traits except for hand length. Male A. praesignis are much longer than females and have relatively longer heads.

On the other hand, Ameiva sp. nov. is not sexually dimorphic for any of these mensural characters. Although we did not explicitly evaluate the power of our test, statistical power is very unlikely to be a problem in this analysis, because our sample size of Ameiva sp. nov. is comparable to A. atrigularis and greater than A. ameiva where similar tests were highly significant (Table 4).

31.8 6 1.22 (29–35) 10 6 0.00 (10–10) 4 (0) 5 (19%) 6 (66%) 7 (15%) 8 (0) 35.8 6 1.95 (32–40) Subequal (28%) Larger (72%) 15.4 6 0.79 (14–17) 32.7 6 1.76 (30–37)

31.3 6 1.11 (29–34) 10 6 0.00 (10–10) 4 (0) 5 (25%) 6 (34%) 7 (31%) 8 (10%) 33.9 6 3.24 (28–42) Subequal (0) Larger (100%) 15.8 6 0.87 (13–18) 34.0 6 2.45 (29–40)

Color Pattern Adult coloration.—Color pattern of adult Ameiva, especially that of adult males, is diagnostic and easily separates the four species. Adult male A. ameiva have a brown head, arms and anterior dorsum, cream or pale bluish throat, and green rump, legs and tail. Some specimens have the ventral aspect of the legs, sides of the tail and subcaudal region pale turquoise. Ameiva ameiva also has whitish, black-bordered ocelli on the flanks that do not extend onto the dorsum. Some Venezuelan specimens of this species have

NS 8.69, 0.000 7.09, 0.000 8.69, 0.000 NS NS

Posterior gulars (F 5 21.42, P 5 0.000) Ameiva sp. (Guianan Highlands) A. atrigularis (western) A. atrigularis (eastern) A. pantherina A. praesignis (northern) A. praesignis (southern)

NS NS NS 0.002 0.000 0.000

8.02, 0.000 6.16, 0.000 NS NS NS NS

Transverse dorsals (F 5 13.17, P 5 0.000) Ameiva sp. (Guianan Highlands) A. atrigularis (western) A. atrigularis (eastern) A. pantherina A. praesignis (northern) A. praesignis (southern) 0.000 0.000 0.000 0.000 0.000

A. atrigularis (eastern)

6.19, 0.000 NS 6.75, 0.000 7.01, 0.000

NS 6.74, 0.000 4.36, 0.034 4.561, 0.021

NS NS NS

5.92, 0.001 NS NS

Tukey’s pairwise comparisons: Q, p

A. atrigularis (western)

NS NS

NS NS

A. pantherina

NS NS NS NS NS

NS NS 0.014 0.000

NS 0.000 0.000

0.001 0.000

7.80, 0.000 6.19, 0.000 NS 7.80, 0.000 NS NS NS 4.95, 0.008 NS 4.95, 0.008 NS 7.18, 0.000 5.57, 0.002 7.18, 0.000 Bonferroni-corrected Mann-Whitney pairwise comparison probabilities

14.17, 7.98, 11.20, 7.42, 7.17,

10.08, 0.000 9.26, 0.000 NS 5.72, 0.001 5.52, 0.002

Ameiva sp. (Guianan Highlands)

NS

NS

NS

NS

A. praesignis (northern)

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Midbody scale rows (H 5 76.22, P 5 0.000) Ameiva sp. (Guianan Highlands) A. atrigularis (western) A. atrigularis (eastern) A. pantherina A. praesignis (northern) A. praesignis (southern)

8.60, 0.000 NS NS 5.26, 0.004 NS NS

Character

Subdigital lamellae under fourth toe (F 5 12.17, P , 0.000) Ameiva sp. (Guianan Highlands) A. atrigularis (western) A. atrigularis (eastern) A. pantherina A. praesignis (northern) A. praesignis (southern)

Ameiva ameiva

TABLE 2.—Summary of univariate comparisons for meristic characters among Ameiva ameiva (n 5 22), Ameiva sp. (n 5 5), eastern A. atrigularis (n 5 24), western A. atrigularis (n 5 16), A. pantherina (n 5 32), northern A. praesignis (n 5 28), southern A. praesignis (n 5 37). Characters satisfying assumptions of normality and homoscedasticity appear in the upper half of the table where statistics (F) and probabilities for one-way ANOVA precede results of Tukey’s pairwise comparisons. In the lower portion of the table, statistics (H) and probabilities for the Kruskal-Wallis test precede Bonferroni-corrected Mann-Whitney pairwise comparison probabilities. Nonsignificant pairwise comparisons are indicated with NS.

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NS NS NS NS NS NS

NS NS NS NS NS NS

Occipitals (H 5 33.05, P 5 0.000) Ameiva sp. (Guianan Highlands) A. atrigularis (western) A. atrigularis (eastern) A. pantherina A. praesignis (northern) A. praesignis (southern)

Granular scales (total of both sides) between supraoculars and supraciliaries (H 5 44.40, P 5 0.000) Ameiva sp. (Guianan Highlands) A. atrigularis (western) A. atrigularis (eastern) A. pantherina A. praesignis (northern) A. praesignis (southern)

Character

Ameiva ameiva

NS NS NS NS NS

NS NS NS NS NS

Ameiva sp. (Guianan Highlands)

TABLE 2.—Continued.

NS NS NS NS

NS NS NS NS

NS 0.011 0.025

0.005 0.002 0.001

A. atrigularis (eastern)

Tukey’s pairwise comparisons: Q, p

A. atrigularis (western)

0.000 0.000

NS NS

A. pantherina

NS

NS

A. praesignis (northern)

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FIG. 1.—Size of posterior occipitals versus first row of dorsal scales. (A) Ameiva atrigularis USNM 27788, showing occipitals conspicuously larger than first row of dorsals. (B) Ameiva pantherina LACM 31433, showing only slightly enlarged occipitals. (C) Ameiva praesignis USNM 162762 showing small occipitals subequal to first dorsal row.

several of the lateral ocelli fused forming short transverse bands on the flanks. For the most part, preserved specimens lose most of the coloration in life. Nevertheless, well-preserved adult A. ameiva often retain a blue or greenish tint on the rump, legs, and tail (Fig. 7). Among the material studied, most adult A. ameiva have larger and well-spaced black reticulations or small, somewhat round black spots on the head, neck or anterior dorsum that disappear or become very minute posteriorly (Fig. 7). The amount of black reticulations or spots on the

anterior dorsum and neck is variable, and a few specimens have only scattered minute black dots. Three specimens (USNM 80615, 80624, and 80655) are unique in that the black reticulations extend onto the rump; in USNM 80655 they even form a short and irregular black middorsal stripe. USNM 80621 also differs from other specimens in having reticulations more numerous on the rump than on the anterior dorsum.

Adult males of Ameiva atrigularis from Distrito Capital, Vargas, and adjacent northern

FIG. 2.—Condition of the anterior gulars (AG), posterior gulars (PG), and mesoptychial scales (MS) in Ameiva from Venezuela. (A) Ameiva atrigularis USNM 217033 with small median anterior and posterior gulars and small mesoptychial scales. (B) Ameiva ameiva USNM 80615 with moderately enlarged median anterior gulars, a patch of distinctly enlarged posterior gulars, and large mesoptychial scales. (C) Ameiva praesignis USNM 162763 with distinctly enlarged median anterior gulars, a distinct patch of enlarged posterior gulars and large mesoptychial scales.

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FIG. 3.—Map illustrating the distribution of species of the Ameiva ameiva complex in Venezuela. Solid symbols represent material studied and confirmed observations; open symbols represent data from literature: Barbour and Noble (1915), Ruthven (1922), Mu¨ller (1929), Schmidt (1932), Mu¨ller and Hellmich (1940), Marcuzzi (1950, 1954), DonosoBarros (1968), Donnelly and Myers (1991), Gonza´les and Prieto (1997), Murphy (1997), Esqueda and la Marca (1999), Gorzula and Sen˜aris (1999), Mijares-Urrutia and Arends (2000), Esqueda et al. (2001), Barrio-Amoro´s and BrewerCarı´as (2008), Ugueto and Rivas (in press). Records from the uplands of the Venezuelan Guiana tentatively referred to Ameiva sp.

Miranda are rather similar in coloration to A. ameiva. Lizards from these areas have a brown head, anterior dorsum, limbs, and anterior flanks, whereas their rump, posterior flanks, legs, and tail are bright green. The ventral aspect of the legs, subcaudal region and sides of the tail are cobalt blue. Flanks of this species also exhibit pale turquoise or whitish, blackbordered ocelli more or less aligned in vertical rows; the ocelli are never fused. However, in contrast to A. ameiva, adult male A. atrigularis from this area have a black throat, postocular region and sometimes sides of the head (Fig. 8). Adult male A. atrigularis from Sucre, Isla de Margarita, and Trinidad are never green but uniformly dark brown or dull olive-brown dorsally. These lizards have more extensive black coloring on the head where pigment

extends to the sides of the head, throat, chest and occasionally the ventral aspect of the arms.

The dorsum of adult A. atrigularis is covered with numerous minute black reticulations (Fig. 7); these reticulations are for the most part very small and remain the same size on the neck and dorsum.

Adult males of Ameiva sp. nov. have a unique dorsal pattern formed by large black blotches (sometimes round or squarish in shape), extending throughout all dorsal surfaces of the neck and dorsum (Fig. 7). This pattern is somewhat reminiscent of occasional specimens of A. ameiva, but Ameiva sp. nov. differ conspicuously from all A. ameiva in having much larger reticulations and very dark gray or black throats and abdomens (Fig. 8).

Readers are cautioned that throats and

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FIG. 4.—Character displacement between Ameiva atrigularis and A. praesignis in northern Venezuela and Trinidad. Upper graph: dorsals versus longitude. Lower graph: scales across body versus longitude.

abdomens of some preserved adult A. praesignis are dark gray or even blackish (Fig. 8) even though these surfaces are cream colored or blue in life. The dark color in preservative is likely to be an artifact of preservation. We have not seen live specimens of Ameiva sp. nov., and this species may have dark blue rather than black abdomens in life.

Most Ameiva praesignis have a distinct pale vertebral stripe or row of pale spots. Some individuals lack the middorsal stripe but have ocelli extending across the dorsum (Fig. 9); a few old adult males rarely lack or have faded ocelli on the dorsum. Presence or absence of a vertebral stripe or dorsal ocelli does not appear to be associated with a particular population nor is it sexually dimorphic. In our samples, the middorsal stripe is distinct in 70% (n 5 56) of the adults (.105 mm SVL).

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Eighteen percent have lateral pale ocelli extending onto the vertebral area, and only 12% (seven specimens) have no vertebral stripe or ocelli middorsally. Ameiva ameiva, A. atrigularis, and Ameiva sp. nov. always lack a pale vertebral stripe. Apart from a single specimen of A. atrigularis with ocelli on the posterior dorsum, only A. praesignis have rows of ocelli across the dorsum. The background coloration of adult A. praesignis is gray, bluish gray, or brownish. Northern populations often have each scale with minute black pigments producing a salt-and-pepper pattern. The chin and throat vary from cream colored to blue, often with profuse black speckling. We did not observe any substantial differences in coloration between northern and southern Venezuelan populations, although it is our general impression that northern populations tend to be more bluish than those from the Llanos.

Juvenile coloration.—All four species undergo conspicuous ontogenetic color change.

In the past, researchers largely neglected juvenile color pattern. Unfortunately, some characters are not always evident in preserved specimens and differences may be subtle.

Juvenile A. ameiva and A. atrigularis are in most regards identical. Photographs of juveniles from the Guianan uplands (Ameiva sp.) show lizards with a conspicuous light dorsolateral stripe bordering the upper margin of the broad lateral dark band. This stripe extends from the posterior border of the supraoculars to the anterior third of the tail (Fig. 10). In contrast, young A. ameiva, A. atrigularis, and Ameiva sp. nov. lack this stripe or have it faded and present only on the head and anterior dorsum. Juvenile Ameiva sp. and Ameiva sp. nov. have pairs of large dark spots on the dorsum that often cover the area between the vertebral region and the upper margin of the dorsolateral light stripe. In contrast, these spots are much smaller or faded in young A. ameiva and A. atrigularis (Fig. 10).

Young A. praesignis usually can be quickly identified by a faint to distinct pale vertebral stripe and numerous small white ocelli covering the dorsum (Fig. 11). Juvenile A. ameiva, Ameiva sp., Ameiva sp. nov., and A. atrigularis lack these characters. Nevertheless, these distinctive markings may be absent

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TABLE 3.—Summary of univariate comparisons of mensural characters among male A. ameiva (n 5 13), A. atrigularis (n 5 20), A. pantherina (n 5 16), and A. praesignis (n 5 22). (Note that strict application of the Bonferroni correction requires probabilities be less than 0.05/6 5 0.0083 for tibia length, head length, snout length, and head depth, and probabilities be less than 0.05/9 5 0.006 for foot length.)

ANCOVA F, P Character

Tibia length A. atrigularis A. pantherina A. praesignis Foot length A. atrigularis A. pantherina A. praesignis (Northern) A. praesignis (Southern)

Ameiva ameiva

A. atrigularis

A. pantherina

NS 21.98, 0.000 5.77, 0.022

10.97, 0.002 5.32, 0.027

8.238, 0.007

NS 18.09, 0.000 NS 22.95, 0.000

7.73, 0.009 NS 7.31, 0.011

9.078, 0.006 NS

Head length A. atrigularis A. pantherina A. praesignis

NS 5.361, 0.029 NS

5.13, 0.031 NS

NS

Snout length A. atrigularis A. pantherina A. praesignis

NS NS NS

NS 6.38, 0.016

NS

Head depth A. atrigularis A. pantherina A. praesignis

NS NS NS

NS 5.72, 0.022

NS

from hatchling A. praesignis, which otherwise look similar to juvenile A. atrigularis. At least in Venezuela and Colombia, most young A. praesignis have only a faint light dorsolateral stripe that does not extend to the tail. We did not examine recently collected juveniles from Central America, but photographs of young individuals from Panama and Fig. 4 of Echternacht (1971) show animals with narrow dorsolateral lines, extending to the rump and sometimes onto the anterior third of the tail.

Ecological Differences Each of the four species is associated with a particular habitat. Ameiva sp. nov. occurs in savannah and appears to be restricted to open formations of southern Monagas (E. Asens, personal communication), although we have little information on the preferred habitat of individuals of this species. Ameiva ameiva and A. atrigularis are most commonly encountered in deciduous, tropical, and premontane humid forests. Nevertheless, these lizards almost never occur inside primary forest.

Instead, they frequent natural or man-made forest clearings, tree falls or along roads that cut through forested areas (Avila-Pires, 1995; Hoogmoed, 1973; Ugueto and Rivas, 2010). In northern Venezuela and Isla de Margarita, A. atrigularis consistently avoids thornscrub, dry forests, savannahs, and other arid or open areas, and it is found up to (at least) 1635 m in humid forest on the central portion of Cordillera de La Costa (Gonza´les and Prieto, 1999). Ugueto and Rivas (in press) reported that in Isla de Margarita, A. atrigularis is replaced in dry arid areas by the smaller A. bifrontata, a species strongly associated with xeric or semiarid conditions. Ameiva ameiva and A. atrigularis are also common around perianthropic situations (Avila-Pires, 1995; Hoogmoed, 1973; Murphy, 1997) and are often ubiquitous in gardens and backyards.

Ameiva atrigularis is commonly found in parks of large cities such as Caracas. In Los Robles and Urbanizacio´n El Placer, Miranda, A. atrigularis is often observed in backyards and basking on the ground or in clearings

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FIG. 5.—Ameiva atrigularis and A. pantherina. Upper graph: tibia length versus snout–vent length (SVL); lower graph: foot length versus SVL.

along roads in areas of secondary growth previously occupied by premontane humid forest (900–1100 m). Murphy (1997) reported that A. atrigularis in Trinidad uses ‘‘areas of forest with open undergrowth where patches of light reach the forest floor, scrub lands, secondary growth, suburban yards and gardens, roadsides and beaches.’’

Ameiva ameiva appears to be more tolerant of open habitats than A. atrigularis. Gorzula and Sen˜aris (1999) reported that in the lowlands (below 400 m) of the Venezuelan Guayana, A. ameiva is a forest-edge species, whereas in the uplands (400–1400 m) it occurs in open savannahs and scrub. In Brazil, south of the Amazon River, this lizard is known to

FIG. 6.—Sexual dimorphism in Ameiva atrigularis.

Upper graph: hand length versus snout–vent length (SVL); middle graph: head length versus SVL; lower graph: head width versus SVL.

occur in thorn forests, close-canopy forest, and open fields (Vitt and Colli, 1994). In southern Brazil, A. ameiva has been reported to occur in sand dunes with scattered shrubs (Rocha, 2008).

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TABLE 4.—Sexual dimorphism in species of the Ameiva ameiva complex. Statistics and probabilities that sexes are the same and that slopes are the same are given for each species.

ANCOVA F, P A. ameiva (nine females, 13 males) Character

Dimorphism

Slope

A. atrigularis (19 females, 20 males) Dimorphism

Slope

Head length 9.06, 0.007 NS 24.12, 0.000 15.56, 0.000 Head width 10.46, 0.005 5.09, 0.038 8.50, 0.006 37.93, 0.000 Head depth 25.7, 0.000 17.8, 0.001 12.16,0.002 15.42, 0.001 Tibia length 6.35, 0.021 NS 4.20, 0.049 NS Foot length NS NS NS NS Hand length NS NS 12.19, 0.001 NS

In contrast, A. praesignis is predominantly associated with open, xeric communities and dry forests. This species has been reported from banana plantations, open and cleared areas, and savannahs and perianthropic situations in Central America (Echternacht, 1971; Savage, 2002); from flood-plain forests, deserts, and dry forests around Santa Marta, Colombia (Ruthven, 1922); from thorny xerophilous scrub, cultivated lands, tropophilous deciduous and semideciduous scrub, tropophilous semideciduous seasonal forests, coastal cloud forests, and cultivated lands in Falco´n State, northwestern Venezuela (Mijares-Urrutia and Arends, 2000); and from thornscrub in Maiquetı´a, Vargas, north-central Venezuela (Marcuzzi, 1954). Heatwole (1966) noted that A. praesignis is conspicuously absent from the heavily forested Darie´n Gap, between Panama

A. pantherina (17 females, 16 males) Dimorphism

Slope

NS NS NS NS NS NS

NS NS NS NS NS NS

A. praesignis (27 females, 22 males) Dimorphism

Slope

8.16, 0.006 NS NS 4.44, 0.040 NS NS NS NS – – NS NS

and northwestern Colombia, and that the species is expanding its range eastward in Panama by moving into areas cleared by man.

Test et al. (1966) reported that A. praesignis is common in ‘‘warmer, dryer, and more open country than cloud forest’’ in northern Aragua.

We observed A. praesignis around Cata and Ocumare de La Costa, northern Aragua, in thornscrub, beaches, and perianthropic situations (backyards, parks, roads). In this area, A. praesignis occurs in sympatry with a blue Cnemidophorus (probably C. lemniscatus splendidus). Both taxa only seem to penetrate deciduous forests of nearby Turiamo through roads and other man-made clearings. Ameiva praesignis is also sympatric with C. lemniscatus cf. splendidus along the arid coast of Vargas (personal observation), with A. bifrontata and C. l. splendidus in thornscrub and dry

FIG. 7.—Dorsal pattern in adult male Ameiva from Venezuela. (A) Ameiva atrigularis USNM 217051. (B) Ameiva ameiva USNM 80602. (C) Ameiva pantherina LACM 31424.

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FIG. 8.—Ventral coloration of four species of Ameiva from Venezuela. (A) Ameiva ameiva USNM 80623, from Bolivar.

(B) Ameiva atrigularis USNM 22526, from Miranda. (C) Ameiva praesignis USNM 121194, from Distrito Capital.

(D) Ameiva pantherina LACM 31431 (holotype), from Monagas. (E) Ameiva praesignis UF 12789, from Portuguesa.

(F) Ameiva praesignis UF 127286, from Cojedes.

forests of the Peninsula de Paraguana´, Falco´n (Markezich et al., 1997), and with A. bifrontata and C. lemniscatus in dry forests around Santa Marta, Colombia (Ruthven, 1922). East of the Andes, in the Llanos of Venezuela, Mu¨ller (1929) reported that this species occurs in gallery forests or forested areas around savannahs, whereas Staton and Dixon (1977) reported that A. praesignis occurs not only in ecotones between gallery forests and savannah, but also in open savannah.

Test et al. (1966) remarked, ‘‘a single green-tailed individual of either this species

[referring to Ameiva praesignis] or Cnemidophorus lemniscatus was seen a few times at Rancho Grande in the fall of 1951 and one similarly in January, 1960.’’ Neither A. praesignis nor C. lemniscatus are known to occur in the cloud forests of Rancho Grande, Henri Pittier National Park. Furthermore, A. praesignis never has a green tail at any age.

We suspect the green-tailed lizard observed

by Test et al. was A. atrigularis, which has been found in clearings within cloud forest in neighboring Miranda, Vargas, and Distrito Capital.

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FIG. 9.—Pattern variation in Ameiva praesignis from Venezuela. (A) USNM 217055 from Zulia. (B) UF 127341 from Portuguesa. (C) USNM 217029 from Carabobo. (D) UF 127337 from Portuguesa.

SPECIES ACCOUNTS Ameiva ameiva (Linnaeus, 1758) Figs. 12–13 Lacerta Surinamensis Seba (1734: 139).— Type unknown, based on Tabula 88, Figs. 1 and 2, from Suriname.

Lacerta Americana Seba (1734: 141).— Type unknown, based on Tabula 89, Fig. 3, from America.

Lacerta Tigrina Ceilonica Seba (1734: 143).— Type unknown, based on Tabula 90, Fig. 7, from ‘‘Ceylon’’ (in error).

Lacerta Ameiva Linnaeus (1758: 202).— Syntypes (NRM 120 and 124, and UUZM 12) from ‘‘America,’’ restricted by Hoogmoed (1973) to the confluence of the Cottica River and the Perica Creek, Suriname; Gmelin (1789: 1065); Daudin (1802: 98).

Seps surinamensis Laurenti (1768: 59).— Type unknown, based on Tabula 88, Figs. 1 and 2, of Seba (1734).

Seps zeylanicus Laurenti (1768: 59).—Type unknown, based on Tabula 89, Fig. 3 and Tabula 90, Fig. 7 (Lacerta Tigrina Ceilonica) of Seba (1734).

Ameiua (sic) americana Meyer (1795: 28; replacement name for Lacerta ameiva Gmelin [1789: 1065], which is based on Linnaeus, 1758: 202).

Lacerta litterata Daudin (1802: 112); after Seba (1734).

Lacerta graphica Daudin (1802: 112); after Seba (1734).

Am[eiva] lateristriga Cuvier (1817: 28).— Type unknown, based on Tabula 89, Fig. 3 of Seba (1734).

Ameiva vulgaris Lichtenstein (1823: 91; replacement name for Lacerta ameiva Linnaeus, 1758).

Tejus tritaeniatus Spix (1825: 22).—Lectotype (ZSM 750/0, designated by Hoogmoed and Gruber, 1993) from ‘‘Habitat in provincia Bahiae’’ (5Bahia, Brazil).

Ameiva maculata Gray (1838: 277).—Type unknown, based on Tabula 88, Fig. 1 of Seba (1734).

Ameiva guttata Gray (1845: 18).—Syntypes (BMNH III.8a–d, BMNH 1946.8.30.31–33) from ‘‘Demerara, Pernambuco and Brazil’’ (name first used by Gray, 1838, although not formally described until 1845).

Ameiva laeta Cope (1862: 65).—Cotypes (MCZ 10536–37) from Rio de Janeiro, Brazil (replacement name for Ameiva guttata Gray [1845] due to transference of North American Cnemidophorus guttatus Wiegmann [1834; now Aspidoscelis guttatus] to the genus Ameiva by Cope, 1862).

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FIG. 10.—Juveniles of four species of the Ameiva ameiva complex in Venezuela. (A) Uncatalogued Ameiva sp. from near Salto Aponwao, Bolivar; photo: A. Anker. (B) Uncatalogued Ameiva sp. from Chivato´n, Bolivar; photo: C. L. BarrioAmoro´s. (C) Uncatalogued A. atrigularis from Los Robles, Miranda; photo: G. N. Ugueto. (D) Uncatalogued A. atrigularis from Sector Sabas Nieves, Avila National Park, Distrito Capital; photo: A. Allais. (E) Uncatalogued A. ameiva from La Paragua, Bolivar; photo: J. C. Sen˜aris (courtesy of G. Rivas). (F) Uncatalogued subadult A. pantherina from Rio Amana, in the savannahs south of Maturı´n, Monagas; photo: E. Asens.

Ameiva petersii Cope (1868: 99).—Holotype number left blank in original description, probably deposited in the Academy of Natural Sciences, Philadelphia (ANSP), from ‘‘Napo or Maran˜on,’’ Ecuador.

Ameiva bifrontata Peters (1871: 398).— Syntypes (ZMB 7178–79) from ‘‘Pebas’’ and ‘‘Pozozu,’’ Pasco department, Peru.

Ameiva pleurotaenia Peters (1871: 652; replacement name for Ameiva bifrontata Peters [1871]).

Ameiva surinamensis (Laurenti, 1768): Boulenger (1885: 352; in part).

Cnemidophorus roeschemanni Werner (1909: 28).—Holotype juvenile (ZMH, lost in World War II, Hallermann, 1998, 2006) from ‘‘Provinz Beni,’’ Bolivia (Burt, 1931 included this

name as a synonym of A. ameiva ameiva, but his action was later questioned by Peters and Donoso-Barros, 1970, who remarked that A. ameiva was not known from Bolivia; A. ameiva occurs throughout the Bolivian lowlands [M. Harvey, personal observation; Dirksen and De la Riva, 1999; Langstroth, 2005], and we find no reason to refute the synonymy proposed by Burt, 1931).

Ameiva ameiva ameiva (Linnaeus, 1758): Barbour and Noble (1915: 462); Burt and Burt (1931: 304, 1933:51); Marcuzzi (1950: 101; in part); Peters and Donoso-Barros (1970: 18); Hoogmoed (1973: 225).

Ameiva ameiva bilineata Barbour and Noble (1915: 464).—Holotype UMMZ 46142 from ‘‘Dunoon, Demerara River, British Guiana’’

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FIG. 11.—Juveniles of Ameiva praesignis. (A) Uncatalogued juvenile from Hacienda Caracas, Rio Lajas Basin, Zulia; photo: T. Barros. (B) Uncatalogued juvenile from Caja Seca, Me´rida; photo: C. L. Barrio-Amoro´s. (C) Uncatalogued juvenile from Monte Cano, Peninsula de Paraguana´, Falco´n; photo: A. Miralles (courtesy of G. Rivas). (D) Uncatalogued juvenile from Barinas, Barinas; photo C. L. Barrio-Amoro´s. (E) Uncatalogued juvenile from Hato Masaguaral, Guarico; photo: N. D. Sly. (F) Uncatalogued juvenile from Hato Masaguaral, Guarico; photo N. D. Sly.

(description based on a juvenile male [106 mm SVL] with typical coloration of typical A. ameiva).

Ameiva ameiva laeta Cope (1862): Barbour and Noble (1915: 467); Peters and DonosoBarros (1970: 19).

Ameiva ameiva petersii Cope (1868): Barbour and Noble (1915: 466); Peters and Donoso-Barros (1970: 19).

Ameiva ameiva praesignis: Burt and Burt (1931: 305; in part; misidentification).

Ameiva ameiva petersi: Duellman (1978: 210; misspelling of Ameiva ameiva petersii Cope, 1862).

Ameiva ameiva (Linnaeus, 1758): Vanzolini (1986: 1); Hoogmoed and Gruber (1983: 394); Avila-Pires (1995: 491); Gorzula and Sen˜aris (1999: 148; in part); Duellman (2005: 346).

Diagnosis.—A medium-sized Ameiva distinguished from all congeners by the following combination of characters: (1) maximum SVL in males 174 mm (Avila-Pires, 1995); (2) dorsal head scales smooth; (3) frontal usually single; (4) frontoparietal and parietal plates in contact with interparietals; (5) 19–44 (both sides) scales, usually in single row between supraoculars and supraciliaries; (6) 11–17 occipitals, usually subequal to first dorsal row; (7) 18–32 anterior gulars; (8) middle anterior gulars polygonal and usually distinctly enlarged, less often only moderately enlarged; (9) patch of distinctly enlarged posterior gulars usually present, occasionally only moderately enlarged; (10) 7–16 posterior gulars between antegular and gular folds; (11) enlarged mesoptychial scales subequal or

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FIG. 12.—Cephalic morphology of Ameiva ameiva (USNM 561200, adult male, SVL 170 mm).

larger than largest gulars; (12) postbrachials moderately to distinctly enlarged; (13) 259– 339 scales between interparietals and base of tail; (14) 129–187 dorsal scales across midbody; (15) ventrals in 29–32 transverse rows, and 10 longitudinal rows; (16) in life, adult male bicolored: head, arms, and anterior

dorsum tan or light brown and posterior dorsum, legs and tail bright green, with pale lateral ocelli; (17) throat in adults cream colored; (18) no vertebral stripe or rows of ocelli across dorsum; (19) juveniles with small paired black spots on dorsum (often faded); pale dorsolateral line (often very indistinct)

FIG. 13.—Individuals of Ameiva ameiva in Venezuela. (A) Uncatalogued adult male from Sierra de Imataca, Bolivar; photo: C. L. Barrio-Amoro´s. (B) Uncatalogued adult female from Las Claritas, Bolivar; photo: D. Ascanio.

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bordering upper margin of broad black lateral stripe present only on head and anterior dorsum; (20) associated most commonly with forested habitats.

Description.—Maximum SVL in males 174 (Avila-Pires, 1995); largest Venezuelan male 170 mm (USNM 561200); maximum total length in males 535 mm (USNM 561200); maximum SVL in females 149 mm (AvilaPires, 1995); largest Venezuelan female 142 mm (USNM 162757); maximum total length in females 459 mm (USNM 80624).

Head pyramidal, 0.24–0.26 (0.25 6 0.00, n 5 13) times SVL in males, 0.23–0.26 (0.23 6 0.01, n 5 9) times SVL in females. Snout elongate, bluntly pointed; canthus rostralis distinct. Neck only slightly narrower than head and body. Body cylindrical. Limbs well developed; tibia 0.22–0.26 (0.24 6 0.01, n 5 13) times SVL in males, 0.22–0.24 (0.23 6 0.00, n 5 9) times SVL in females; foot 0.37– 0.50 (0.42 6 0.03, n 5 13) times SVL in males, 0.38–0.43 (0.41 6 0.01, n 5 9) times SVL in females. Tail round in cross section, tapering toward tip, 2.14–2.51 (2.34 6 0.13, n 5 8) times SVL in males, 2.50–2.60 (n 5 3) times SVL in females.

Rostral pentagonal, higher than wide, visible from above, bordered posteriorly by nasals, which form short or very short medial suture. Each nasal divided by oblique suture.

Nostril in lower part of suture and anterior to it, directed lateroposteriorly, taller than long.

Frontonasal subhexagonal or octagonal, in contact with nasals, loreal, and prefrontals.

Prefrontals paired, rectangular, or roughly pentagonal, with long medial suture about twice or three times as long as that between nasals (equal in USNM 80604), laterally in contact with loreal, first supraocular and first supraciliary. Frontal hexagonal or roughly pentagonal, longer than wide and wider anteriorly, its sutures with prefrontals angulate, slightly curved, those with frontoparietals almost forming straight line or in very wide angle; frontal laterally in contact with first, second, and, often, narrow part of third supraoculars. Frontoparietals paired, pentagonal, or trapezoidal, longer than wide and with long medial suture (in USNM 80656 and 162758 broken into two scales posteriorly), laterally in contact with third supraocular and

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small circumorbital scales bordering fourth up to middle (or rarely anterior) portion of third supraocular. Interparietal pentagonal, higher than wide, often narrower or subequal (but sometimes slightly wider) than adjacent parietals, its sutures with parietals straight or slightly oblique; interparietal bordered at each side by two (rarely one) large, irregular parietals divided by oblique suture; outermost parietal usually round in shape and larger or subequal to inner parietal (USNM 561202 lacks an interparietal scale and instead has two large median parietals that meet and form a broad suture). Parietal series composed of 4–6 (usually five) scales (including interparietal).

Occipitals 11–17 (14.6 6 1.49, n 5 22), irregular and subequal in size; occipitals usually subequal in size to first dorsal row, less often moderately larger or in USNM 80621 slightly smaller. Supraoculars four at each side; five on left side of USNM 80621, right side of USNM 80623, 80624, 80655; and six on the right side of USNM 80621 (small wedge-shaped scale between first and second supraocular counted as additional supraocular); supraoculars broken and divided into 8/9 scales at each side of USNM 80656; second largest, fourth smallest. Circumorbital semicircles formed by 3–9 scales at each side, 6–17 (11.2 6 2.65, n 5 22) combining both sides, reaching middle or posterior portion of third supraocular (to anterior portion in USNM 162759); only one scale row present between third supraocular and prefrontals; fourth supraocular separated from parietals by up to four rows of circumorbital scales. Laterally, all supraoculars except first, usually separated from supraciliaries by single row of small rectangular scales (occasionally doubled in the middle), 19–44 (28.3 6 7.18, n 5 22) combining both sides. Supraciliaries six or seven, first highest, second and/or third longest, remaining short and subequal. Loreal very large, trapezoidal, and single; loreal in contact with nasal, frontonasal, first supraciliary, first and second suboculars, and third and fourth supralabials (occasionally also very narrowly with second supralabial). Suboculars four, large; first subocular in preocular position, irregularly trapezoidal, longer than wide and as wide as second subocular; first subocular in contact with loreal, first supra-

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ciliary, small scales of eyelid, and second subocular; margin with loreal slightly curved or straight (separated from first supraciliary by small scale on right side of USNM 561200).

Third subocular longest; all, except first, in contact with supralabials; second subocular of both sides of USNM 80623 transversally divided. Continuous keel from first subocular through entire length of third subocular.

Postoculars small, approximately in two rows of four or five irregularly trapezoidal scales.

Palpebral disc semiopaque, with enlarged scales on lower eyelid. Supralabials usually six or seven, rarely eight or nine at each side, fifth below center of eye; followed to commissure of mouth by smaller scales. Row of enlarged supratemporals decreasing in size posteriad. Temporal region with rectangular and round scales, slightly smaller centrally than peripherally. External auditory meatus large, vertically oval, bordered by granular scales, its anterior margin semicircular, its posterior margin straight. Tympanum recessed. All dorsal and lateral head scales juxtaposed and smooth.

Symphysal anteriorly ellipsoid, its posterior sutures forming wide angles with infralabials and postsymphysal. Postsymphysal single, pentagonal, in contact with first infralabials, followed by three pairs of enlarged chinshields. First pair of chinshields in broad medial contact; first and occasionally part of second pair in contact with infralabials.

Remaining chinshields separated from infralabials by one row of sublabials. Medial chin scales moderately small, elongate, convex, smooth, juxtaposed, hexagonal or subehexagonal, in slightly oblique rows, all subequal in size. Usually six or seven infralabials (rarely eight), fifth below center of eye followed to commissure of mouth by smaller scales.

Gular region divided in two areas: anterior region with polygonal and flat scales in slightly oblique rows that grade to larger scales at center of throat, delimited posteriorly by line uniting lower margin of ear openings; middle anterior gular scales moderately to distinctly enlarged, 18–32 (27.4 6 3.59, n 5 22) scales along midline of chin from anteriormost to posteriormost anterior gular. Posterior gular region covered by smaller polygonal scales, in transverse rows;

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posteriormost median posterior gulars usually forming medial patch of distinctly enlarged scales, occasionally only moderately enlarged in some specimens; 7–17 (11.7 6 2.16, n 5 21) scales midventrally from anteriormost posterior gular to antegular fold. Mesoptychial scales moderately enlarged (larger or, less often, subequal to largest gulars), in about one or two rows, hexagonal, flat, smooth, and juxtaposed.

Scales on nape and sides of neck similar to dorsals. Dorsals and scales on flanks granular (slightly larger on dorsum than laterally), round, smooth, subimbricate or juxtaposed, in approximately transverse rows; 259–339 (293.5 6 21.76, n 5 22) scales between occipitals and base of tail. Scales around midbody (excluding ventrals) 129–187 (159.5 6 12.61, n 5 22). Ventrals large, smooth, rectangular (wider than long), imbricate, in 10 longitudinal (at midbody) and 29–32 (30.5 6 0.91, n 5 22) transverse rows; transition between ventral scales and those on flanks sharp. Preanal shield with 5–7 rows of enlarged scales (eight in USNM 162756); pattern of terminal scales of the preanal plate variable: often one large or two median scales followed posteriorly by two subequal scales, or followed by three scales (middle one smallest), or preanals divided and irregularly disposed in series of four or five enlarged and subequal scales; preanal shield rarely formed by two hexagonal scales followed by three similar and subequal scales. Preanal plate surrounded anteriorly and laterally by smaller scales; posteriorly by much smaller scales. Femoral pores in continuous row along each thigh; medially with short gap; 15–24 pores on each leg, 30–45 (37.5 6 4.39, n 5 22) pores combining both legs. Each pore between four or five scales.

Scales on base of tail rectangular or pentagonal, smaller than ventrals, longer than wide, mostly keeled (smooth on base of tail on ventral surface). Caudal scales imbricate, in transverse rows continuous around tail, except first few rows incomplete ventrally. Distally caudal scales become longer and narrower, distinctly keeled, in transverse and approximate longitudinal rows.

Arms with row of very large, smooth, slightly imbricate, trapezoid antebrachial

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scales on anterodorsal aspect of forearms and similar but smaller brachial scales on upper arms that extend almost to shoulder. One or occasionally two moderately to distinctly enlarged brachial rows with smaller accessory row bordering upper margin and grading into granules posteriorly; antebrachial and brachials separated by one or two rows of small scales at elbow. Scales of dorsoposterior, posterior, and ventral aspect of arms similar to, but slightly larger than dorsals, except for group (approximately two rows) of postbrachial scales on posterior aspect of upper arms, which are slightly larger and irregular. Legs with large, smooth, imbricate scales on anterior and ventral aspects of thighs and ventral aspect of shanks. Row of large, trapezoid scales anteriorly on thighs, gradually become smaller and irregular toward row of femoral pores. On ventral aspect of shank three rows of very large scales, anterior two trapezoidal, posterior one rhomboidal; enlarged scales of shank decreasing in size from anterior toward posterior row. Elsewhere on legs scales similar to dorsals.

Subdigital lamellae transversely enlarged and single, moderately to distinctly tuberculate towards base. Tubercles most prominent under base of third toe; lamellae of innermost toe continued to tibio-tarsal articulation. On palms, lamellae of inner finger continuing and increasing in size towards wrist; single large scale present on postaxial side of hand, at a short distance from wrist. Lamellae 14–16 (15.1 6 0.66, n 5 22) under fourth finger, 32–40 (35.8 6 2.15, n 5 22) under fourth toe.

Coloration in life.—Ameiva ameiva exhibits conspicuous ontogenetic color changes (Figs. 10 and 13). Young specimens have a broad black or dark reddish brown (black-bordered) band extending from the postocular region through the sides of the head (above ear opening) and along the flanks to the tail where it breaks into irregular dark blotches. As individuals grow, light spots appear within the black band and on the rest of the flanks. This band often has a faint upper and a distinct lower light margin. The upper margin often fades at midbody, and the lower one extends to the inguinal region. A similar lower pale stripe is also present on sides of the tail. The flanks below the pale stripe are marked with dark brown or black spots, forming

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a checkered pattern, or are brownish with light spots in subadults. Dorsally, the head and body are uniformly tan or green anteriorly (head, neck, and anterior dorsum) and brown posteriorly (posterior dorsum) with small, paired black spots on each side of the vertebral region (usually inconspicuous, sometimes faded); subadults sometimes have uniformly green heads and dorsa. Laterally, the head is whitish with few black spots, and the canthus rostralis is often darker (as a continuation of the dorsolateral band). The limbs and tail are brown with dark brown spots and dots. The throat is uniformly whitish or with a few blackish dots; the abdomen is whitish with some dark markings, especially laterally.

In adult males, the head, anterior dorsum, anterior flanks and arms are light brown or tan, whereas posterior surfaces are grass green (Fig. 13). The tail of some specimens is graybrown. Dorsally, the head, neck, and body usually are marked with black reticulations or dots (absent in some specimens). When present, these markings continue onto the green portion of the body or, more frequently, are limited to the brown area. Laterally, the head of adult males is slightly lighter and often marked by small black spots. Numerous distinct whitish ocelli with narrow to moderate black margins are distinct on the flanks (sometimes ocelli have a green cast on the posterior flanks). The ocelli are usually aligned in vertical rows and do not extend onto the dorsum; a few ocelli sometimes merge forming short vertical bands. Pale turquoise spots cover the paraventral region, and the chin, throat, chest and anterior abdomen are whitish, occasionally with some black mottling on the throat. Posteriorly, the abdomen, ventral aspect of the legs and subcaudals are usually pale blue in adult males. The subcaudals and sides of the tail are whitish in some specimens and pale turquoise in others (becoming more intense towards the tip); the subcaudals often have black edges or dark speckling.

Adult female A. ameiva are similar to males but retain some of the juvenile pattern (Fig. 13). Females often have remnants of the lateral dark band; they also have a black bordered, whitish, ventrolateral stripe extending from midflank to the inguinal region.

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Additionally, females have fewer ocelli on the flanks and are usually whitish on all ventral surfaces instead of bluish. At all stages of development, A. ameiva has a reddish brown iris.

Coloration in preservative.—The green coloration present in adults and juveniles fades to olive, dark green, or bluish in preserved specimens. Brown areas among adults often turn light olive–brown, whereas the grass green tones turn bluish or dark green. Small black reticulations are usually present on the head and anterior dorsum and fade towards the rump. USNM 80615, 80621, and 80624 (all from southern Anzoa´tegui) have distinct black reticulations, becoming more conspicuous and broader posteriorly.

USNM 80655 has scattered black spots on the dorsum and an irregular black vertebral stripe extending from midbody to the sacrum.

USNM 80653 has only a few black spots on the head, neck, and anterior dorsum; this specimen also preserves some of the dark lateral band present in juveniles. The throat and other ventral surfaces of preserved juveniles are uniformly whitish; the throat of USNM 80656 has scattered black spots; and most ventral surfaces of USNM 162758–59 (from Amazonas) are pale gray or bluish. The throat of most adult specimens is uniformly whitish, occasionally with a few scattered black dots. The abdomen is uniformly whitish, suffused with blue laterally and with a few black spots toward the sides. Ventral aspects of legs are whitish suffused with blue anteriorly, with black blotches in this same area.

Ventral surfaces (except posteriormost portion of abdomen, anal region, ventral aspect of arms, subcaudal region, and blotches on ventral aspect legs) of USNM 162756–57 (from Amazonas) are blue.

Distribution and natural history.—This species is widely distributed throughout Amazonia in southern Venezuela, the Guianas, Brazil, southern Colombia, eastern Ecuador, Peru and Bolivia. In Venezuela, A. ameiva is found throughout Amazonas and Bolivar south of the Orinoco River, up to 380 m (Gorzula and Sen˜aris, 1999), although it probably occurs at slightly higher elevations.

Populations from Paulo, Mount Roraima, the confluence of the Rı´o Arabopo´ and Rı´o

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Kukena´n and remaining uplands (above 400 m) in southeastern Bolivar (Burt and Burt, 1931; Donnelly and Myers, 1991; Gorzula and Sen˜aris, 1999) may be referable to a different taxon (see remarks). Ameiva ameiva has also been reported from Delta Amacuro (Donoso-Barros, 1968), and it is here reported from La Soledad, southern Anzoa´tegui, north of the Orinoco River (Fig. 3).

Gorzula and Sen˜aris (1999) reported that A. ameiva is a forest-edge species in the lowlands and an open savannah or scrub species in the uplands of the Venezuelan Guayana. Throughout its entire distribution, A. ameiva has been found in open, sunny areas or clearings (natural or manmade) in areas with at least some vegetation cover in inundated or terra-firma tropical forests, Amazonian savannahs, restinga, caatingas, and cerrados (Avila-Pires, 1995; Duellman, 2005, personal observation; Hoogmoed, 1973; Vitt and Colli, 1994). The natural history of this species has been relatively well documented, especially regarding its basic ecology, reproduction, and diet (e.g., Colli, 1991; Cruz-Neto and Gordo, 1996; Duellman, 2005; Magnusson, 1987; Magnusson et al., 1985; Rocha, 2008; Silva et al., 2003; Simmons, 1975; Vitt, 1982; Vitt and Colli, 1994; Vitt et al., 2000; Zaluar and Rocha, 2000). However, virtually nothing is known about the natural history of A. ameiva in Venezuela.

Remarks.—Vitt and Colli (1994) found A. ameiva to be morphologically conservative throughout Brazil and reported no conclusive geographic variation within the populations they examined. Three subspecies have been reported from south of Venezuela, Colombia, and the Guianas: A. a. ameiva, A. a. laeta, and A. a. petersii. We examined few specimens from Peru and Bolivia (Appendix). Nonetheless, these specimens and photographs of specimens in life from southern Brazil, Peru, and Bolivia show animals basically identical to those from southern Venezuela, Guyana, Amazonian Brazil, and southern Colombia.

The color characteristics indicated by Peters and Donoso-Barros (1970) to separate the nominal taxon from A. a. laeta and A. a. petersii appear to be based on ontogenetic variation and we consider these names to be

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synonyms of A. a. ameiva until the validity of these taxa is tested.

There is geographic variation in squamation between specimens from Amazonas and those from Bolivar and Anzoa´tegui. In most scale characters examined, specimens from Amazonas are similar to the other populations studied. However, individuals from Amazonas have 40–45 (42.3 6 1.79, n 5 8) lamellae under the fourth toe, whereas those from Bolivar and Anzoa´tegui have 30–38 (34.8 6 2.58, n 5 14). We do not know if this difference represent clinal variation or variation between isolated demes.

Burt and Burt (1931) reported A. a. praesignis from Paulo, Mt. Roraima, based on a series of specimens housed in the American Museum of Natural History (AMNH). According to Burt and Burt (1931), some young and middle-sized specimens of A. a. praesignis have a series of paired, undulate or branching, light areas on the dorsum. The same authors reported that specimens from Roraima have ‘‘paired, undulate, dorsal light areas’’ and thus considered them conspecific with A. a. praesignis. We examined five of the Roraima specimens (AMNH 36328, 36330–33, all juveniles). These specimens have the paired dark spots typical of juveniles of the A. ameiva complex and no trace of light undulate areas.

Furthermore, paired, branching, light areas were also never observed in A. praesignis.

Nevertheless, the Roraima sample is noteworthy as we found them to differ conspicuously in some morphological characters from A. praesignis and A. ameiva (Tables 2 and 5). These specimens were collected in the uplands surrounding Mount Roraima, in southeastern Bolivar, and may represent a distinct species.

Comments by Gorzula and Sen˜aris (1999) suggest that these highland Ameiva may inhabit a different habitat than their counterparts in the lowlands. Furthermore, S. Gorzula in Gorzula and Sen˜aris (1999) commented: ‘‘SG also had the impression that in southern Venezuela the upland specimens were smaller than those from the lowlands. In the uplands there was an apparent absence of large green males.’’ We found color differences between A. ameiva and among specimens of all ages and sexes from the Guayana uplands. In particular the amount of green color in adult males, large

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dorsal spots present in juveniles and adults and a distinct pale line bordering the upper margin of the lateral dark stripe of juveniles (Figs. 10 and 14) seem to differ between upland and lowland samples.

Donnelly and Myers (1991) reported that the largest male among their sample from the summits of Cerro Guaiquinima (1030–1150 m) was 112 mm in SVL, whereas the largest female was 91 mm. These specimens were sexually mature and the female had two small eggs in each oviduct. They also indicated all the specimens had broad dark lateral bands. Although some of the scale counts differ from those from the Roraima specimens we examined (femoral pores and lamellae under fourth toe), we believe the Guaiquinima Ameiva is possibly conspecific with the Roraima population.

Status of the Roraima Ameiva requires further study, and we refrain from assigning AMNH 36328, 36330–33 and the Cerro Guaiquinima specimens to any of the four species of the A. ameiva complex present in Venezuela.

Ameiva atrigularis Garman Figs. 15–17 Ameiva surinamensis var. atrigularis Garman (1887: 2).—Lectotype male (MCZ 186018; designated herein) from Trinidad.

Ameiva atrigularis Garman (1887): Barbour and Noble (1915: 460); Burt and Burt (1931: 307).

Ameiva ameiva melanocephala Barbour and Noble (1915: 465).—Holotype male (MCZ 9993, examined by authors) from Cumanacoa, Venezuela; Peters and Donoso-Barros (1970: 19).

Ameiva ameiva ameiva Linnaeus (1758): Marcuzzi (1950: 101; in part).

Ameiva ameiva Linnaeus (1758): Roze (1964: 237); Murphy (1997: 152); Gorzula and Sen˜aris (1999: 148; in part); Rivas et al.

(2005: 351); Ugueto and Rivas (2010: 187).

Ameiva ameiva melahocephala: DonosoBarros (1968: 115; in part; misspelling).

Ameiva ameiva tobagana Cope (1879): Peters and Donoso-Barros (1970: 20; misidentification).

Ameiva ameiva atrigularis Garman (1887): Tuck and Hardy (1973: 241).

Designation of lectotype.—Garman based his description on five specimens all from Trinidad: MCZ 186018–19 (previously MCZ 6079–80), MCZ 6081, ZMB 29431, NMW

Bicolored as body, with pale blue or white ocelli with narrow black borders in males; ocelli can merge to form short transversal stripes; a distinct black-bordered white lateral stripe in females

Pale brown with or without black spots

Cream-colored, with or without scattered black dots

Bicolored, anterior green, posterior brown, or completely brown with (usually) large paired black spots usually distinct

With broad black or dark brown lateral band with conspicuous pale stripes bordering upper and lower margins of dark band, upper pale stripe occasionally faded; pale ocelli appear only in larger subadults

Flanks in adults

Sides of head in adults

Throat in adults

Dorsum in juveniles

Flanks in juveniles

A. ameiva

Anterior tan or light brown, posterior green; small black spots or reticulations often only on brown area

Dorsum in adults

Characters

With broad black lateral band with conspicuous pale stripe bordering lower margin of dark band, upper stripe faded; pale ocelli appear only in larger subadults

Bicolored, anterior green, posterior brown, or completely brown with small paired black spots usually faded

Black or dark gray

Brown with dark postocular band or completely black

Anterior tan, posterior green in north-central Venezuela, completely dark brown or olive–brown in northeastern Venezuela, Trinidad and Tobago; usually with minute but numerous black reticulations Bicolored as body or brown, with pale blue or white ocelli with distinct black borders in males; ocelli usually do not merge to form short transversal stripes; a distinct black-bordered white lateral stripe in females

A. atrigularis

A. praesignis

Not known in life, in preservative with broad black lateral band with conspicuous pale stripe bordering lower margin of dark band, upper stripe faded; pale ocelli appear only in larger subadults

Not known in life, probably brown, with pale ocelli with distinct black borders in males; ocelli not merging to form short transversal stripes; a distinct black-bordered white lateral stripe in females Not known in life, uniformly purplish in preservative Not known in life, dark gray in preservative, sometimes with bluish cast Not known in life, in preservative purplish with large paired black spots distinct, sometimes with satellite smaller spots around them

Bicolored, anterior green, posterior brown, or completely brown with (usually) small paired black spots distinct or faded; sometimes with pale vertebral stripe or with white spots on sides of dorsum With broad black lateral band with conspicuous pale stripe bordering lower margin of dark band, upper stripe often faded; pale ocelli appear sometimes even in smallest of juveniles

Pale gray, pale brown, or bluish with or without minute black dots Cream colored or blue, sometimes profusely speckled with black

Gray, brown, or bluish; with white, yellow, or pale blue ocelli with narrow black borders in males; ocelli in rows that usually do not merge to form short transversal stripes; a distinct black-bordered white lateral stripe in females

Purplish, probably brown or Gray–brown or bluish gray, each olive in life, with very scale often with black pigments large, leopard-like reticulations producing salt-and-pepper on dorsum, smaller in females appearance; dorsum with a yellow or whitish vertebral pale stripe, or with transverse rows of ocelli

A. pantherina

TABLE 5.—Selected coloration characters useful in identifying species of the Ameiva ameiva complex in Venezuela.

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FIG. 14.—Individuals of Ameiva sp. in Venezuela. (A), (B): Uncatalogued adult male from Chivato´n, Bolivar.

(C) Uncatalogued adult female from same locality. (D) Uncatalogued juvenile from same locality. All photos: C. L. BarrioAmoro´s.

23408 (Tiedemann et al., 1994). We examined the first two of these specimens and here designate MCZ 186018 (Fig. 16), an adult male in good condition, as lectotype of A. atrigularis.

The lectotype has supralabials 6/6; infralabials 6/6; supraoculars 5/5; circumorbitals 6/7 extending to mid third supraocular; 16/13 scales between supraorbital and supraciliary scales; parietals 5; occipitals 14; occipitals larger than first dorsal row; 34 anterior gulars; mid anterior gulars small; 15 posterior gulars, midposterior gulars moderately enlarged; mesoptychial scales subequal to largest anterior gulars; dorsals between occipitals and base of tail 311; 166 scales around midbody (excluding ventrals); 10 longitudinal ventral rows; 32 transverse ventral rows; 20/20 femoral pores; 6 scales forming preanal plate; 17 lamellae under fourth finger on right hand; 37 lamellae under fourth toe on right foot.

Diagnosis.—A medium-sized Ameiva distinguished from all congeners by the following combination of characters: (1) maximum SVL in males 186 mm; (2) dorsal head scales smooth; (3) frontal single; (4) frontoparietal and parietal plates in contact with interparietals; (5) 17–47 (both sides) scales, usually in single row, between supraoculars and supraciliaries; (6) 10–18 occipitals, usually larger than first dorsal

row; (7) 20–40 anterior gulars; (8) middle anterior gulars polygonal or rounded and usually small or moderately enlarged; (9) posterior gulars usually small, less often with medial patch of moderately enlarged posterior gulars; (10) 13–23 posterior gulars between antegular and gular fold; (11) enlarged mesoptychial scales subequal or larger than largest gulars; (12) postbrachials moderately to distinctly enlarged; (13) 263–361 scales between occiput and rump; (14) 134–179 dorsal scales across midbody; (15) ventrals in 29–34 transverse and 10 longitudinal rows; (16) adult male coloration in life uniformly brown in northeastern Venezuela, Isla de Margarita, and Trinidad or bicolored (anteriorly brown and posteriorly green) in north-central Venezuela, usually with minute black dorsal reticulations and pale blue or whitish lateral ocelli; (17) throat in adults black or dark gray; (18) no vertebral light stripe and only rarely ocelli present on dorsum; (19) juveniles often with faded dorsal paired black spots, indistinct pale dorsolateral line bordering upper margin of broad black lateral stripe distinct in front of arm; (20) associated with forests or forest clearings.

Description.—Maximum SVL in males 186 mm (USNM 217052), maximum total length in males of 526 mm (USNM 27788);

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FIG. 15.—Cephalic morphology of Ameiva atrigularis (USNM 217052, adult male, SVL 186 mm).

maximum SVL in females 146 mm (USNM 217053), maximum total length in females of 420 mm (USNM 121197). Head pyramidal, 0.23–0.27 (0.25 6 0.00, n 5 19) times SVL in males; 0.22–0.24 (0.22 6 0.00, n 5 11) times SVL in females. Snout elongate, bluntly pointed; canthus rostralis distinct. Neck only slightly narrower than head and body. Body cylindrical. Limbs well developed; tibia 0.19– 0.29 (0.23 6 0.01, n 5 18) times SVL in males and 0.21–0.25 (0.22 6 0.01, n 5 11) times SVL in females; foot 0.32–0.45 (0.39 6 0.03, n 5 19) times SVL in males and 0.34–0.42 (0.39 6 0.02, n 5 11) times SVL in females.

Tail round in cross section, tapering toward tip; 2.25–2.58 (2.37 6 0.08, n 5 9) times SVL in males; 2.04–2.24 (n 5 3) times SVL in females.

Rostral pentagonal, higher than wide, clearly visible from above, bordered posteriorly by

nasals, which form short or very short medial suture. Each nasal divided by oblique suture.

Nostril in lower part of suture and anterior or slightly anterior to it, directed lateroposteriorly, wider than tall. Frontonasal octagonal or subhexagonal, in contact with nasals, loreal, and prefrontals. Prefrontals paired, pentagonal or trapezoidal, with medial suture equal or about twice (rarely three times) as long as that between nasals; in USNM 27788 prefrontals separated medially by frontal and frontonasal; prefrontal laterally in contact with loreal, first supraocular, and first supraciliary. Frontal hexagonal, longer than wide and wider anteriorly; posterior portion of frontal in USNM 217961 horizontally divided into small accessory scale; sutures with prefrontals angulate and straight or wavy, those with frontoparietals forming very wide angle or almost straight line; frontal laterally in contact with first, second,

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FIG. 16.—Lectotype of Ameiva atrigularis (MCZ 186018).

and narrow part of third supraoculars, or just with first and second. Pair of pentagonal or trapezoidal frontoparietals, longer than wide and with long medial suture and sometimes

broken into one or more scales posteriorly, laterally in contact with third supraocular and small circumorbital scales bordering fourth up to posterior or middle portion of third

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FIG. 17.—Individuals of Ameiva atrigularis in Venezuela. (A) Uncatalogued adult male from Jardı´n Bota´nico de Caracas, Distrito Capital; photo: J. M. Saez Sierra. (B), (D) Uncatalogued adult male from Fuentiduen˜o, Isla de Margarita; photo: G. N. Ugueto. (C) Uncatalogued juvenile from Sector Los Robles, La Unio´n, Miranda; photo: G. N. Ugueto. (E) Uncatalogued adult female from Sector Los Robles, La Unio´n, Miranda; photo: G. N. Ugueto.

(F) Uncatalogued adult female from Jardı´n Bota´nico de Caracas, Distrito Capital; photo: A. Afonso.

supraocular (rarely to anterior third supraocular or to suture with frontal). Interparietal hexagonal, higher than wide (longitudinally divided in CM S 4952); often narrower, less frequently subequal and rarely slightly wider than adjacent parietals; sutures with parietals straight or less often curved; interparietal bordered at each side by two (rarely one) large, irregular parietals divided by oblique suture; outermost parietal usually rounder in shape and smaller or subequal to inner parietal.

Occipitals irregular, 10–16 (13.6 6 1.70, n 5 40), subequal in size or central pair larger than rest; occipitals usually much larger than first

row of dorsals posterior to them, occasionally subequal in size. Supraoculars four at each side, three on left side of USNM 217050 and both sides of CM S 7448, five on both sides of USNM 217047 and 217052, CM S 6606, CM S 7446, and right side of USNM 217061, six and five on left and right side, respectively, of USNM 217051 (smaller wedge-shaped scales between supraoculars counted as additional supraocular); second supraocular largest (third on right side of USNM 27788), fourth smallest.

Circumorbital formed by 3–9 scales at each side, 6–17 (12.5 6 2.28, n 5 40) combining both sides, reaching posterior or middle

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portion of third supraocular, reaching anterior third in USNM 217052, to suture with frontal in CM S 4952, CM S 6507, and to anterior third supraocular on right side and second supraocular on left side of USNM 217051; only one scale row between third supraocular and prefrontals; fourth supraocular separated from parietals by 2–4 rows of circumorbital scales.

Laterally, all supraoculars except first, usually separated from supraciliaries by single row (sometimes doubled in middle portion) of small rectangular scales, 17–47 (27.3 6 6.38, n 5 40) combining both sides. Supraciliaries six or seven (rarely five, eight in USNM 217051, CM S 6505, and right side of CM S 6507 and 6509), first highest, second and/or third longest, remaining short and subequal. Loreal very large, trapezoidal and single, in contact with nasal, frontonasal, prefrontal, first supraciliary, first and second suboculars, and third and fourth supralabials (rarely also narrowly with second supralabial). Suboculars four; first subocular in preocular position, irregularly trapezoidal, longer than wide and slightly to distinctly narrower than second subocular (rarely subequal), in contact with loreal, first supraciliary, small scales of eyelid, and second subocular; suture between first subocular and loreal slightly curved or straight. Third subocular longest; all suboculars except first in contact with supralabials; second subocular broken into two scales on right side of CM S 6509. Continuous keel from first through entire length of third subocular. Postoculars small to moderate in size, approximately in two rows of 3–5 irregularly trapezoidal scales. Palpebral disc semiopaque with enlarged scales on lower eyelid. Supralabials usually six, less often seven at each side (eight on right side of USNM 217053), fifth below center of eye, followed to commissure of mouth by smaller scales. Row of enlarged supratemporals decreasing in size posteriad. Temporal region with irregularly shaped and round scales, distinctly smaller centrally than peripherally. External auditory meatus large, vertically oval, bordered by granular scales, its anterior margin semicircular, its posterior one straight. Tympanum recessed. All dorsal and lateral head scales juxtaposed and smooth.

Symphysal anteriorly ellipsoid, its posterior sutures forming wide angles with infralabials

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and postsymphysal. Postsymphysal single and pentagonal, in contact with first and second infralabials, followed by three pairs of enlarged chinshields. First pair of chinshields in ample medial contact; first pair, or sometimes first and second pairs, in contact with infralabials. Second chinshield usually separated from infralabials by row of small scales; remaining chinshields separated from infralabials by one row of sublabials. Medial chin scales moderately small, elongate, convex, smooth, juxtaposed, irregular or somewhat round, in slightly oblique rows, all subequal but increasing in size posteriorly. Six or seven infralabials, fifth below center of eye followed to commissure of mouth by smaller scales.

Gular region divided into two areas: anterior region with round or polygonal and flat scales in slightly oblique rows that usually grade to moderately larger scales medially or that remain subequal; anterior region delimited by line uniting lower margin of ear openings; middle anterior gular scales usually moderately enlarged, or often subequal (rarely distinctly enlarged), 22–40 (29.5 6 4.33, n 5 40) scales along midline of chin from anteriormost to posteriormost anterior gular. Posterior gular region covered by smaller polygonal or rounded scales, in transverse rows; posteriormost median gulars usually small and subequal to lateral posterior gulars, or occasionally forming patch of moderately enlarged scales; 13–23 (15.9 6 2.21, n 5 40) scales midventrally from anteriormost posterior gular to antegular fold. Mesoptychial scales moderately enlarged, larger or subequal to largest gulars (smaller in CM S 4948, 4950, 6500, and 6501), in about two rows, hexagonal, flat, smooth, and juxtaposed.

Scales on nape and sides of neck similar to dorsals. Dorsals and scales on flanks granular (all of roughly same size), round, smooth, subimbricate or juxtaposed, in approximately transverse rows; 263–361 (309.5 6 27.05, n 5 33) scales between occipitals and base of tail.

Scales around midbody (excluding ventrals) 134–179 (151.3 6 11.22, n 5 32). Ventrals large, smooth, rectangular (wider than long), imbricate, in 10 longitudinal (at midbody) and 29–34 (31.4 6 1.18, n 5 40) transverse rows.

Transition between ventrals and scales on flanks sharp. Preanal shield with 5–7, rarely

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four rows of enlarged scales; pattern of terminal scales of the preanal plate variable; most often formed by two hexagonal scales followed by three similar and subequal scales or a large median scale followed posteriorly by two subequal scales, or much less often followed by three scales (middle one smallest), or preanals divided and irregularly disposed in a series of four or five enlarged and subequal scales. Preanal plate surrounded anteriorly and laterally by smaller scales; posteriorly by much smaller scales. Femoral pores in a continuous row along each thigh; medially with a short gap; 15–21 pores on each leg, 31– 42 (35.5 6 2.66, n 5 40) pores combining both legs; each pore between four scales.

Scales on base of tail rectangular or pentagonal, smaller than ventrals, longer than wide, mostly keeled (smooth on base of tail on ventral surface). Caudal scales imbricate, in transverse rows continuous around tail, except first few rows incomplete ventrally. Distally scales become longer and narrower, distinctly keeled, in transverse and approximately longitudinal rows.

Arms with row of very large, smooth, slightly imbricate, trapezoidal antebrachial scales on anterodorsal aspect of forearms and similar but smaller brachial scales on upper arms that extend almost to shoulder. Brachial row moderately to distinctly enlarged with smaller accessory row bordering upper margin and grading into granules posteriorly. Antebrachials and brachials in contact by continuous row of slightly enlarged scales at elbow or separated by 1–2 small scales. Dorsoposterior, posterior, and ventral aspect of arms covered in scales similar to, but slightly larger than dorsals, except for group (approximately one to two rows) of enlarged and imbricate postbrachial scales on posterior aspect of upper arms. Legs with large, smooth, imbricate scales on anterior and ventral aspects of thighs and ventral aspect of shanks. Row of large, trapezoidal scales anteriorly on thigh, gradually becoming smaller and irregular toward femoral pores. On ventral aspect of shanks three rows of very large scales, anterior two trapezoidal, posterior one rhomboidal, decreasing in size posteriorly. Elsewhere on legs scales similar to dorsals.

Subdigital lamellae transversely enlarged and single, moderately to distinctly tuberculate

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toward base. Tubercles most prominent under base of third toe; lamellae of outer toe continuing to heel. On palms, lamellae of first finger continuing to wrist, increasing in size toward it; single large scale (or up to three) present on postaxial side, following line of fifth finger, short distance from wrist. Lamellae under fourth finger 14–17 (15.5 6 0.88, n 5 40), under fourth toe 32–41 (36.3 6 2.10, n 5 40).

Coloration in life.—Ameiva atrigularis exhibits conspicuous geographic and ontogenetic color variation. Color pattern remains constant among juveniles and hatchlings throughout its distribution. In young specimens a broad black lateral band is always present, extending from the tip of the snout or postocular region through the sides of the head (above ear opening) and flanks, to the sides of the tail where it breaks into irregular dark blotches. As individuals grow light spots appear within the black band and on the rest of the flanks. The upper margin of this band is bordered by an inconspicuous pale dorsolateral stripe (starting from posterior supraocular) that fades on posterior dorsum; the lower margin of the black lateral band is bordered by a distinct white lower lateral stripe (sometimes with green wash on head) that extends from the posterior subocular area to the inguinal region. A similar pale stripe is also present on sides of tail (Fig. 10). Dorsally, the head and body are uniformly tan or green anteriorly and brown posteriorly with small, paired black spots on each side of the vertebral region (in most specimens very inconspicuous or almost completely absent). Laterally, the head is whitish with conspicuous black spots. The flanks are brownish with dark spots (sometimes very inconspicuous), forming an irregular checkered pattern or longitudinal dark stripes. The limbs and tail are brown with dark brown spots and dots. The throat is uniformly whitish or with large dark spots; the abdomen is whitish with some dark markings, especially laterally.

Adult males from localities in Distrito Capital, Vargas, and Miranda State are bicolored and resemble A. ameiva (Fig. 17).

These individuals have the top of the head and anterior portion of dorsum and arms tan or light brown, whereas the posterior dorsum, legs, and tail are green. The sides of the head vary from

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slightly darker than top of head to dark gray or blackish. Some specimens have a broad black postocular stripe extending from posterior corner of eye to above ear opening and sometimes reaching the sides of the neck. The labial region varies from black to brownish, but the cheeks are often paler, grayish, or whitish.

In contrast, the throat and chin are often dark gray or black. The green dorsum, legs, and tail have numerous small black reticulations; these reticulations are more apparent on the dorsum (including the brown portion). The flanks are brown or tan, with conspicuous light turquoise or whitish, black-bordered ocelli more or less aligned in vertical stripes but not usually merging with each other. Some specimens may have a bluish or green cast on the flanks, which also have numerous minute black reticulations. Light turquoise ocelli, similar but smaller than those on flanks, are also present on legs and on lateral aspect of tail.

The tail is green on top with blackish reticulations, the sides and underside often become bright cobalt blue toward posterior half. The abdomen is pale bluish white with some black spotting (particularly evident laterally); the anal region is often somewhat yellowish.

Adult males from Sucre, Isla de Margarita and Trinidad are similar to those described above, but all dorsal surfaces are dark brown (rarely tan or olive brown) with numerous minute black reticulations. In these males the black coloration of the head and throat is more conspicuous; these specimens often have the sides of the head, chin, throat, chest, and even sometimes the anterior aspect of forearms uniformly jet black; the degree of black on the sides of the head varies individually and some specimens have light brown or grayish cheeks (Fig. 17). The sides of the tail are blue as in Miranda specimens.

One specimen observed in Guarenas, Miranda resembled more those from Sucre, Margarita and Trinidad in being completely brown but had less black on the sides of the head.

Coloration of adult females also varies geographically, albeit less obviously than in males. Throughout its distribution, females usually preserve remnants of the juvenile color pattern. An adult specimen observed in Los Robles, Miranda (Fig. 17) had a light brown head with slightly darker brown sides

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and a very indistinct broad dark brown postocular stripe that extended posteriorly.

Most of the dorsum (except the rump), was grass green and unspotted. The rump, flanks, limbs, and tail are tan. The flanks had whitish spots merging into a longitudinal white stripe from midflanks to the inguinal region; this stripe had conspicuous black margins; the rest of the flanks with black reticulations. The limbs and tail are brown (tail never blue as in male), also with black reticulations and small spots; a whitish lateral stripe is also present on the base of the tail. Several adult females observed in the Jardı´n Bota´nico of Caracas, Distrito Capital, were similar to the coloration described above, but the green dorsal coloration extended to rump and there were several dark brown dorsal spots disposed in pairs.

Additionally, these specimens had a black postocular band and the head was gray laterally. Adult females observed at several localities on Isla de Margarita and in the State of Sucre were similar to males (with conspicuous black sides of head and throat), but often had far less conspicuous ocelli but had a black-bordered lateral white stripe. These specimens also had less dorsal black reticulations than males. Adult females lacked the blue tail coloration. The iris is reddish brown at all stages of development.

Coloration in preservative.—Most of the dorsal color fades in preserved specimens, but diagnostic elements of the pattern remain obvious in most specimens. In preservative, the adult background color varies from purplish to olive brown. Many of the pale ocelli on flanks of CM S 4950 merge, forming short transveral bars. As in life, the dorsal paired spots at each side of the vertebral region are usually faint or absent in the juveniles. Females retain most of the juvenile pattern up to a SVL of 120 mm. The smallest male exhibiting an almost fully developed adult pattern is 101 mm SVL (CM S 6500). USNM 217052 is unique among the series in that the lateral ocelli invade rump and small ones are even apparent on the vertebral region. The ventral coloration tends to darken considerably in preservative. Most adults and subadults examined have gray, dark gray, or blackish chin and throat but also have the chest and most of the abdomen gray or dark gray; posterior margin of the ventrals pale; pale

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area becoming broader on more posteriad ventrals; in some specimens most of the abdomen has a blue cast, other individuals (as in life) have the black coloration restricted to the chin, throat, sides of head, chest, and ventral aspect of arms, whereas the remaining ventral surfaces are pale; USNM 217051 has a dark gray chin and throat, pale gray chest, and dark gray abdomen (with bluish cast posteriorly) and white spots laterally; USNM 217034 has all anterior ventral surfaces bluish with black spots on the throat and sides of abdomen. Some juveniles have pale gray or dark bluish-gray throats with numerous black dots; the smallest specimens have uniformly pale or bluish throats and abdomens. The posterior portions of the abdomen, anal, and subcaudal regions are pale (these areas profusely spotted with dark gray in USNM 217053); the ventral aspects of the thighs have dark gray and whitish spots, whereas that of the shanks is mostly immaculate.

Distribution and natural history.—Ameiva atrigularis occurs in north-central and northeastern Venezuela (including Isla de Margarita) and on Trinidad In Venezuela, A. atrigularis occurs throughout forested regions of Distrito Capital and the states of Vargas and Miranda. It is apparently absent from the arid lowlands of northern Anzoa´tegui and reappears again in the state of Sucre, where it extends eastward to the Peninsula de Paria.

On Isla de Margarita it is associated with forested areas (Ugueto and Rivas, 2010) on both peninsulas of the island (Fig. 3). Esqueda et al. (2001) reported a specimen of A. a. melanocephala (5A. atrigularis) from Sierra de Perija´, Zulia and mentioned this was the westernmost locality for the taxon. However, this record was based on a misidentified specimen of A. praesignis (which is found in the area), judging by the presence of a distinct pale vertebral stripe (J. E. Garcı´a-Pe´rez, personal comment). Esqueda et al. (2001) also reported A. a. melanocephala from Puerto Cabello, Carabobo State. If the identification of this specimen is correct then it would represent the westernmost locality for this species. Unfortunately, we were unable to examine this specimen and this record is not included in the map of Fig. 3.

Ameiva atrigularis is locally common.

Ugueto and Rivas (in press) reported this

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species from perianthropic situations, semideciduous and premontane evergreen forests up to 600 m on Isla de Margarita. Gonza´les and Prieto (1999) reported A. atrigularis from the area of Quebrada Onda and Quebrada Paujı´ (northern Miranda) at 1639 m, but the species probably extends to even higher elevations in the central coastal range.

Like other teiids, A. atrigularis is an alert, fast-moving, diurnal, and heliothermic lizard. All of the specimens we observed were active on warm sunny days. Active individuals were never encountered on overcast or rainy days. Ugueto and Rivas (in press) reported lizards were most active between 0900 and 1330 h during the month of October on Isla de Margarita.

Gonza´les and Prieto (1999) reported active specimens have body temperatures ranging from 36.28 to 41uC. When first encountered, most individuals run for short distances and either stop or continue moving along slowly. In Los Robles, Miranda, we observed green adults often remained motionless amidst vegetation on roadsides allowing close approach, but once feeling discovered they fled rapidly. Many individuals were observed moving alertly between patches of vegetation, constantly flicking their tongues. Ugueto and Rivas (in press) reported that this species is capable of bipedal movement for short distances.

Ameiva atrigularis is known to consume a wide variety of arthropods, small vertebrates, eggs, and even fruit (Boos and Quesnel, 1968; Murphy, 1997; Roze, 1964). Ugueto and Rivas (in press) observed two subadult specimens eating a dead land crab on the main road of La Sierra at the Cerro Copey National Park, Isla de Margarita. Murphy (1997) reported individuals of this species chasing C. lemniscatus in a coconut plantation on Trinidad. The author could not confirm if A. atrigularis preyed on the smaller Cnemidophorus or were just keeping them out of their territory.

An adult Mastigodryas boddaerti (Sentzen) was observed preying on a subadult male in Los Robles, Miranda. Gonzales and Prieto (1999) reported males are more numerous than females during August in the state of Miranda, Venezuela. These authors suggested females may stay inside nest chambers laying eggs or avoiding wasting energy unnecessarily.

Gonza´les and Prieto (1999) also suggested

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that hatchlings are probably found during November and December, when arthropods are more abundant in the area. Quesnel (1979) reported that courtship and mating occur during May, June, October, and November on Tobago. Quesnel also reported multiple copulation events (each averaging 2 min and 3 s) were observed during a few hours. Murphy (1997) reported mating occurs in June and July on Trinidad.

Remarks.—Color differences between western bicolored populations (brown anteriorly, green posteriorly) and eastern brown populations do not appear to be correlated with additional characters that might warrant recognition of two taxa. Green A. atrigularis from north-central Venezuela have 301–361 transverse dorsal scales, whereas brown lizards have 263–334 scales. However, there appears to be a west-to-east cline in the number of dorsals (Fig. 4); highest values are found among specimens from Vargas and Distrito Capital, whereas lowest values were found on Trinidad. Specimens from Sucre and Isla de Margarita have somewhat intermediate values (320–327 dorsals). We also found slight variation in size of the posterior gulars between green (western) and brown (eastern) A. atrigularis. Brown populations had a high frequency of small posterior gulars, whereas only roughly half of the specimens of green populations had small posterior gulars. Although both morphs in Venezuela are separated by a stretch of arid lowlands in the northern portion of Anzoa´tegui State (Unare Depression), we believe these differences are slight and do not merit recognizing the green lizards as a separate subspecies.

Cope (1879) described Ameiva surinamensis tobaganus based on a single specimen purportedly collected on the island of Tobago. According to Cope, A. s. tobaganus differed from the continental form in color pattern, disposition, and greater subdivision of scales on the limbs and abdomen, and in greater length of the foot.

Almost a decade later, Garman (1887) named two insular ‘‘variants’’ of A. surinamensis: A. s. var. atrigularis from the neighboring island of Trinidad and A. s. var. aquilina from Grenada and St. Vincent. The type series of A. s. var. aquilina included 8 specimens from Grenada and 39 from Saint Vincent. According to the

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original description, this taxon differed from A. s. var. atrigularis in having more femoral pores and lacking a black chin or throat. Apparently, Garman was unaware of Cope’s description of A. s. tobaganus and did not compare his specimens of either form with the taxon described from neighboring Tobago.

Barbour and Noble (1915) described a new subspecies (A. a. melanocephala) from two localities in northern and eastern Venezuela (La Guayra [5La Guaira] and Cumanacoa).

According to the authors, this taxon differed from the nominal subspecies in having larger median gulars and a black throat. Barbour and Noble also considered the populations of Trinidad, Tobago, and the Lesser Antilles (Grenada and St. Vincent) to be full species: A. atrigularis, A. tobagana, and A. aquiline, respectively. Inexplicably, they did not compare A. a. melanocephala with A. atrigularis even though adults of both forms have black chins and throats. Burt and Burt (1931) synonymized A. a. melanocephala with the nominal form but continued to recognize A. atrigularis. Nevertheless, these authors cautioned that A. atrigularis and melanic Venezuelan specimens formerly referred to A. a. melanocephala might be the same species. Donoso-Barros (1968) did not accept the taxonomic arrangement proposed by Burt and Burt (1931) and reported A. ameiva melanocephala from Sucre and Vargas. Additionally, he reported that A. a. melanocephala intergrades with A. a. vogli in southern Monagas, but gave no further details. On the other hand, Mu¨ller (1929) claimed that no intermediate specimens between A. a. melanocephala and A. a. praesignis were known.

Finally, Peters and Donoso-Barros (1970) recognized A. a. melanocephala on Margarita Island and on the mainland and A. a. tobagana on Trinidad and Tobago, thus sinking A. atrigularis in the synonymy of A. a. tobagana.

Tuck and Hardy (1973) concluded that the type of A. ameiva tobagana was likely collected in Grenada, the Grenadines, or St. Vincent (populations then allocated to A. a. aquilina) rather than on Tobago. These authors allocated the Trinidad and Tobago populations to A. a. atrigularis but did not compare this subspecies to mainland taxa.

Although we did not examine specimens from Tobago, we obtained several color

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FIG. 18.—Cephalic morphology of Ameiva pantherina (holotype LACM 31431, adult male, snout–vent length 147 mm).

photographs of lizards from this island in life.

Lizards on Tobago have an identical color pattern to those from Trinidad and northeastern Venezuela. Thus, we consider Tobago Ameiva to probably represent A. atigularis.

However, because we did not examine specimens from that island, we refrain to formally assigning them to the latter taxon.

We did not examine Ameiva from the Lesser Antilles. Nevertheless, we had the opportunity to observe several photographs of lizards from Grenada. Ameiva on Grenada exhibit coloration substantially different from those of Tobago, Trinidad, or northeastern Venezuela.

The head of males is pale green or grayish with black reticulations and the chin and throat are pale blue. These lizards are not A. atrigularis.

The name currently associated with these Lesser Antillean populations is A. ameiva aquilina Garman (1887). Considering the allopatric distribution and their distinctiveness

in morphology and coloration, A. ameiva aquilina likely represents a different taxon from those on mainland South America, Isla de Margarita, and Trinidad. Their taxonomic status requires further study Ameiva pantherina sp. nov.

Figs. 18–20 Ameiva ameiva melanocephala Donoso-Barros (1968: 115; in part, misidentification).

Holotype.—An adult male (LACM 31431, Figs. 18, 19) from 60 km southeast of Maturı´n, Monagas, Venezuela, collected in 1958 by B. B. Butterworth.

Paratopotypes (20).—Nine males and 11 females (LACM 31422–41) with the same data as the holotype.

Paratypes (11).—Three males and four females (LACM 31414–21) from 42 km southeast of Maturı´n, Monagas, Venezuela, collected

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149

FIG. 19.—Dorsal and ventral view of holotype of Ameiva pantherina (LACM 31431).

in 1958 by B. B. Butterworth. Three males and two females (USNM 216982, 217048–50) from Hato Mata de Bejuco, 54 km south–southeast Maturı´n, Monagas, Venezuela, collected in 1968 by an unknown collector, donated by C. O. Handley Jr. An adult male (USNM 217047) from Hato Mata de Bejuco, 55 km southsoutheast of Maturı´n, Monagas, Venezuela, collected in 1968 by an unknown collector, donated by C. O. Handley, Jr. Diagnosis.—A medium-sized Ameiva distinguished from all congeners by the following combination of characters: (1) maximum SVL

in males 152 mm; (2) dorsal head scales smooth; (3) frontal single; (4) frontoparietal and parietal plates in contact with interparietals; (5) 24–41 (both sides) scales, usually in single row, between supraoculars and supraciliaries; (6) 14–18 occipitals, usually subequal to first dorsal row; (7) 20–36 anterior gulars; (8) middle anterior gulars polygonal or rounded and usually small, rarely moderately enlarged; (9) patch of moderately enlarged posterior gulars frequently present, less often all posterior gulars small; (10) 17–20 posterior gulars between antegular and gular folds; (11) en-

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FIG. 20.—Pattern variation in Ameiva pantherina. (A) LACM 31418 (snout–vent length [SVL] 145 mm). (B) LACM 31421 (SVL 114 mm). (C) LACM 31428 (SVL 127 mm). (D) LACM 31430 (SVL 146 mm). (E) USNM 217047 (SVL 127 mm). (F) USNM 217048 (SVL 141 mm).

larged mesoptychial scales subequal or larger than largest gulars; (12) postbrachials moderately to distinctly enlarged; (13) 291–343 scales between occiput and rump; (14) 137–163 dorsal scales across midbody; (15) ventrals in 29–35 transverse and 10 longitudinal rows; (16) adult male coloration in life unknown, in preservative purplish with large and conspicuous black with dorsal reticulations and whitish lateral ocelli; (17) throat in adults dark gray; (18) no vertebral light stripe and ocelli never present on dorsum; (19) juveniles with distinct paired black spots on dorsum and with indistinct whitish dorsolateral line bordering upper margin of broad black lateral stripe; (20) associated with savannahs.

Description.—Maximum SVL in males 152 mm (LACM 31432), maximum total length in males 487 mm (LACM 31430); maximum SVL in females 138 mm (LACM USMN

217050), maximum total length in females 413 mm (LACM 31428). Head pyramidal, 0.22–0.26 (0.24 6 0.01, n 5 15) times SVL in males; 0.21–0.26 (0.23 6 0.01, n 5 17) times SVL in females. Snout elongate, bluntly pointed; canthus rostralis distinct. Neck only slightly narrower than head and body. Body cylindrical. Limbs well developed; tibia 0.19– 0.24 (0.21 6 0.01, n 5 15) times SVL in males; 0.18–0.24 (0.21 6 0.01, n 5 17) times SVL in females; foot 0.31–0.43 (0.35 6 0.03, n 5 15) times SVL in males; 0.32–0.41 (0.36 6 0.02, n 5 16) times SVL in females. Tail round in cross section, tapering toward tip, 2.20–2.33 (2.27 6 0.04, n 5 4) times SVL in males; 2.05–2.57 (2.28 6 0.14, n 5 11) times SVL in females.

Rostral pentagonal, higher than wide, visible from above, bordered posteriorly by nasals, which form very short medial suture.

Each nasal divided by oblique suture. Nostril

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in lower part of suture and anterior to it, directed lateroposteriorly, taller than long.

Frontonasal subhexagonal or octagonal, in contact with nasals, loreal and prefrontals.

Prefrontals paired, rectangular or roughly pentagonal, with very long medial suture about three times (sometimes twice) as long as that between nasals; laterally in contact with loreal, first supraocular and first supraciliary. Frontal hexagonal, longer than wide and wider anteriorly, its sutures with prefrontals forming wide angle (forming an almost straight line in holotype), slightly curved or straight, its sutures with frontoparietals almost forming a straight line or a very wide angle; frontal laterally in contact with first, second and (narrowly) third supraoculars (often just with first and second or with first, second, and circumorbirtals). Pair of pentagonal or trapezoidal frontoparietals (often broken into two, rarely three scales posteriorly), longer than wide and with long medial suture; laterally in contact with third supraocular (narrowly with second in some specimens), and small circumorbital scales bordering fourth up to middle or anterior portion of third supraocular, occasionally extending to frontal suture.

Interparietal irregularly pentagonal or hexagonal, higher than wide, often subequal or narrower (but sometimes slightly wider) than adjacent parietals; interparietal longitudinally divided in LACM 31421 and 31425; sutures with parietals slightly oblique and curved or straight; interparietal bordered at each side by two (rarely one) large, irregular parietals divided by oblique suture; outermost parietal usually round or somewhat oval in shape and larger or subequal (rarely smaller) to inner parietal. Parietal series composed of 4–5 (almost always five) scales including interparietal. Occipitals 14–18 (15.1 6 0.37, n 5 32), irregular and heterogeneous or subequal in size; occipitals usually subequal in size to first dorsal row, less often moderately to distinctly larger. Supraoculars four or five at each side, six on left side of LACM 31436, right side of LACM 31432 and 31424 (small wedge-shaped scale between first and second supraocular counted as additional supraocular in most specimens with five supraoculars). Circumorbital semicircles formed by 4–12 scales at each side, 9–24 (12.6 6 2.84, n 5 32) combining

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both sides (usually 9–16), reaching middle, posterior, or occasionally anterior portion of third supraocular (extending to or slightly beyond frontal suture in LACM 31420, 31424, 31430, and 31432; extending completely around supraoculars except first supraocular in LACM 31423); only one scale row between third supraocular and prefrontals; fourth supraocular separated from parietals by up to three rows of circumorbital scales. Laterally, all supraoculars except first, usually separated from supraciliaries by single or partially doubled row of small rectangular scales; 24– 41 (29.7 6 4.71, n 5 32) combining both sides. Supraciliaries six or seven (on one side), first highest, second longest, remaining short and subequal. Loreal very large, trapezoidal and single, in contact with nasal, frontonasal, first supraciliary, first and second suboculars, and third and fourth (and occasionally narrowly with second) supralabials. Suboculars four; first subocular in preocular position, irregularly trapezoidal, longer than wide and slightly to distinctly narrower than second subocular; in contact with loreal, first supraciliary, small scales on ocular region, and second subocular (first subocular on right side of LACM 31415 in very narrow contact with supralabial); a smaller accessory scale is situated below first subocular on both sides of LACM 31432, separating loreal from second subocular; margin with loreal slightly curved or straight. Third subocular longest; all, except first, in contact with supralabials; second suborbital divided on right side of LACM 31426. Continuous keel from first subocular through entire length of third subocular. Postoculars small, approximately in two rows of four or five irregularly trapezoidal scales. Palpebral disc semi-opaque with enlarged scales on lower eyelid. Supralabials usually six or seven, rarely five at each side, fifth below center of eye; followed to commissure of mouth by smaller scales. Row of enlarged supratemporals decreasing in size posteriad. Temporal region with polygonal or rounded scales, slightly smaller centrally than peripherally. External auditory meatus large, vertically oval, bordered by granular scales; anterior margin semicircular, posterior one straight. Tympanum recessed. All dorsal and lateral head scales juxtaposed and smooth.

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Symphysal anteriorly ellipsoid, its posterior sutures forming wide angles with infralabials and postsymphysal. Postsymphysal single and pentagonal; in contact with first and second infralabials, followed by three pairs of enlarged chinshields. First pair in ample medial contact; only first, or first and part of second pairs, in contact with infralabials. Remaining chinshields separated from infralabials by one row of sublabials. Medial chin scales moderately small, elongate, convex, smooth, juxtaposed, hexagonal or subhexagonal, in slightly oblique rows, all subequal in size. Infralabials six or seven, fifth below center of eye followed to commissure of mouth by smaller scales.

Gular region divided into two areas: anterior region with round or polygonal and flat scales in slightly oblique rows that usually remain subequal or rarely grade to larger scales medially, delimited posteriorly by line uniting lower margin of ear openings; middle anterior gular scales usually small, rarely moderately enlarged, 20–36 (29.2 6 3.74, n 5 32) scales along midline of chin from anteriormost to posteriormost anterior gular. Posterior gular region covered by smaller polygonal scales in transverse rows; posteriormost median gulars usually forming medial patch of moderately enlarged scales, occasionally small in some specimens (distinctly enlarged in USNM 217050); 14–20 (16.5 6 1.64, n 5 32) scales midventrally from anteriormost posterior gular to antegular fold. Mesoptychial scales moderately enlarged (larger or, less often, subequal to largest anterior gulars), in about one or two rows, hexagonal, flat, smooth, and juxtaposed.

Scales on nape and sides of neck similar to dorsals. Dorsals and scales on flanks granular (slightly larger on dorsum than laterally), round, smooth, subimbricate or juxtaposed, in approximately transverse rows; 291–343 (311.3 6 13.69, n 5 32) scales between occipitals and base of tail. Scales around midbody (excluding ventrals) 137–163 (147.4 6 7.21, n 5 32). Ventrals large, smooth, rectangular (wider than long), imbricate, in 10 longitudinal (at midbody) and 29–35 (31.8 6 1.22, n 5 32) transverse rows; transition between ventrals and scales on flanks sharp.

Preanal shield with 5–7 rows (usually six) of enlarged scales; pattern of terminal scales of the preanal plate somewhat variable; often

[No. 25

large median scale followed posteriorly by two subequal scales, or by three scales (middle one smallest). Preanal plate surrounded anteriorly and laterally by smaller scales; posteriorly by much smaller scales. Femoral pores in continuous row along each thigh, with short gap medially; 15–20 pores on each leg, 32–40 (35.8 6 1.95, n 5 32) pores combining both legs. Each pore between four scales.

Scales on base of tail rectangular or pentagonal, smaller than ventrals, longer than wide, mostly keeled (smooth on base of tail on ventral surface). Caudal scales imbricate, in transverse rows, continuous around tail, except first few rows incomplete ventrally.

Distally caudals longer and narrower, distinctly keeled, in transverse and approximately longitudinal rows.

Arms with rows of very large, smooth, slightly imbricate, trapezoidal antebrachial scales on anterodorsal aspect of forearms and similar but smaller brachial scales on upper arms that extend almost to shoulder. Brachial row distinctly enlarged with smaller accessory row bordering upper margin and grading into granules posteriorly. Antebrachials and brachials in contact or separated by smaller scales at elbow. Dorsoposterior, posterior, and ventral aspect of arms with scales similar to dorsal, but slightly larger, except for group (approximately one or two rows) of postbrachial scales on posterior aspect of upper arms, which are slightly to moderately enlarged and irregular.

Legs with large, smooth, imbricate scales on anterior and ventral aspects of thighs, and ventral aspect of shanks. Row of large, trapezoidal scales anteriorly on thigh, gradually becoming smaller and irregular toward pores.

On ventral aspect of shanks, three rows of very large scales, anterior two trapezoidal, posterior one rhomboidal, decreasing in size from anterior toward posterior row. Elsewhere on legs scales similar to dorsals.

Subdigital lamellae transversely enlarged and single, moderately to distinctly tuberculate towards base. Tubercles most prominent under base of third toe, lamellae of outer toe continuing to heel. On palms, lamellae of inner finger continuing to wrist as large scales, increasing in size towards it; single large scale on postaxial side, following line of fifth finger, at short distance from wrist. Lamellae under

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fourth finger 14–17 (15.4 6 0.79, n 5 32), under fourth toe 30–37 (32.7 6 1.76, n 5 32).

Data of holotype.—SVL 147 mm; head length 38.7 mm; tibia length 28.8 mm; foot length 48 mm; hand length 22 mm; tail regenerated; supralabials 6/6; infralabials 7/6; supraoculars 4/4; circumorbitals 6/6 extending to mid third supraocular; 14/14 scales between supraorbital and supraciliary scales; parietals 5; occipitals 14; occipitals larger than first dorsal row; 29 anterior gulars; mid anterior gulars small; 17 posterior gulars, midposterior gulars moderately enlarged; mesoptychial scales larger than largest anterior gulars; dorsals between occipitals and base of tail 293; 158 scales around midbody (excluding ventrals); 10 longitudinal ventral rows; 33 transverse ventral rows; 18/19 femoral pores; six scales forming preanal plate; 15 lamellae under fourth finger on right hand; 34 lamellae under fourth toe on right foot.

Coloration in life.—The only information we have on the coloration in life of this species is based on a photograph of a juvenile (Fig. 10) taken on the savannahs south of Maturı´n, on the margins of the Amana River, Monagas. This specimen appears to have a coloration similar to other juvenile Ameiva.

The top of the head and anterior dorsum are green and there is a broad black (anteriorly) to reddish brown (posteriorly) lateral band extending from the postocular region to the flanks; there are no conspicuous light lines bordering this stripe. A series of white ocelli are distinct on the flanks and extend to the dorsolateral region. The throat of this specimen seems to be suffused with black, which is consistent with the color pattern exhibited by the preserved material.

Coloration in preservative.—Like other species of the A. ameiva complex, this species exhibits conspicuous ontogenetic color changes. Young specimens have a broad black longitudinal lateral band that extends from the postocular region through the sides of the head (above ear opening) and flanks, fading on the sides of the tail. As individuals grow, light spots appear within the black band and rest of flanks. This band often has a conspicuous whitish line bordering its lower margin that extends from the postocular to inguinal regions; a similar pale stripe is also presents

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on the sides of the tail. The upper margin of the dorsolateral band is also bordered by a much less conspicuous whitish stripe that extends from the posterior supraoculars and fades midway along the dorsum. The flanks below the pale stripe have an inconspicuous black stripe; the rest of the flanks are purplish.

Dorsum of the head and body uniformly brownish with medium-sized, paired black spots at each side of the vertebral region (most conspicuous towards sacrum). The sides of the head are uniformly grayish. Limbs and tail are brown with dark spots and dots. The throat, chest, ventral aspect of arms and most of the abdomen are gray or dark gray (sometimes with a greenish or bluish tint); the posterior margin of each ventral scale is pale; most of abdomen pale in LACM 31437. The ventral aspect of the legs, posteriormost portion of the abdomen, anal and subcaudal regions are whitish with a few scattered blackish spots on the tail base.

Adult males have a purplish or gray-brown dorsum with conspicuous large, broad, black reticulations or blotches; these markings remain conspicuous along dorsum from nape to the base of the tail. Flanks have small, scattered black reticulations and pale ocelli with distinct black margins more or less arranged in vertical rows; ocelli do not extend onto dorsum in most specimens, but in LACM 31430 ocelli extend onto dorsolateral region.

Adult males LACM 31414, 31418, and USNM 217047 preserve the whitish lower lateral stripe on lower flanks; LACM 31414 has very few lateral ocelli and LACM 31418 and USNM 217047 preserve the dorsolateral dark band. The top and sides of the head are uniformly purplish brown without spots. The limbs are purplish as the body with black reticulations (smaller and less conspicuous than those on dorsum). The tail is slightly lighter than the body with numerous but scattered light and dark scales, tending to form irregular rings. Chin, throat, chest, ventral aspect of arms, and most of the abdomen are gray or dark gray; the posterior margin of each ventral scale is pale, becoming wider posteriorly (abdomen mostly blue with some black on posterior margin of ventrals in USNM 217048–217050); the posteriormost portion of the abdomen, anal area, and

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subcaudal region are pale (a few dark scales on the subcaudal region present). The ventral aspect of thighs is pale with few black spots or dark gray with pale spots. The ventral aspect of shanks is uniformly pale or with some dark markings. USNM 217047 differs from all the other specimens in having a pale gray throat with black dots, whereas the chest, abdomen, subcaudal region, and ventral aspect of limbs are whitish with a few black dots scattered on the chest and abdomen.

Adult females preserve much of the juvenile pattern, including the black dorsolateral band and the lower light stripe. Unlike juveniles, females have very few whitish ocelli on flanks (including a series of ocelli that often borders the upper margin of the black dorsolateral stripe), and a series of black reticulations along dorsum (distinct from nape to the base of tail), or with a series of paired black or dark gray spots like those in juveniles. Ocelli are basically absent in some specimens. The dorsal reticulations are similar to those present in males but much smaller; particularly minute in LACM 31426 and 31438. The throat is gray to dark gray, often with scattered blackish dots. The chest, ventral aspect of arms, and most of the abdomen are gray; posterior margin of the ventral scales pale. The ventral aspect of legs, posteriormost portion of abdomen, anal, and subcaudal regions are pale with a few scattered blackish spots on the base of the tail.

Distribution and natural history.—Ameiva pantherina is endemic to the savannahs of central and southern Monagas (Fig. 3). Nothing is known about the natural habits of this species. The savannahs of Monagas represent the easternmost portion of the eastern Llanos and have well-drained, sandy soils of Pleistocene origin. These grasslands are the highest elevations of the Venezuelan Llanos at approximately 200 m (Lo´pez et al., 2003). The rattlesnake Crotalus vegrandis is also endemic to this general area. Interestingly, like A. pantherina, this smaller relative of the larger and widespread C. durissus is apparently restricted to these savannahs.

Etymology.—The specific epithet pantherina is a feminine adjective derived from the Greek word panther, meaning ‘‘cat,’’ in reference to the leopard-like dorsal pattern exhibited by adults of this species.

[No. 25

Ameiva praesignis (Baird and Girard, 1852) Figs. 21–23 Cnemidophorus praesignis Baird and Girard (1852: 129) Syntypes (USNM 5519 and UMMZ 3823) from Chagres, Panama.

Ameiva praesignis (Baird and Girard, 1852): Cope (1862: 67).

Ameiva praesigna (Baird and Girard, 1852): Bocourt (1874: 265; unjustified emendation).

Cnemidophorus maculatus Fischer (1878: 95; dated 1879).—Holotype possibly deposited in the Zoologischen Museum of Hamburg (ZMH) but does not appear cited in Hallermann (1988), current whereabouts unknown; from Sabana Larga, Colombia.

Ameiva surinamensis (Laurenti, 1768): Boulenger (1885: 22; in part).

Ameiva ameiva maculata (Fischer, 1878): Barbour and Noble (1915: 467).

Ameiva ameiva praesignis (Baird and Girard, 1852): Barbour and Noble (1915: 468); Burt and Burt (1931: 305; in part); Marcuzzi (1950: 102); Test et al. (1966: 19); DonosoBarros (1968: 115); Peters and Donoso-Barros (1970: 20).

Ameiva ameiva vogli Mu¨ller (1929: 100).— Holotype (ZSM 1/1929, holotype and paratypes apparently lost during Second World War, fide Franzen and Glaw, 2007) from ‘‘Barinas, Zamora, Venezuela.’’ Donoso-Barros (1968: 116); Peters and Donoso-Barros (1970: 20).

Ameiva ameiva ornata Mu¨ller and Hellmich (1940: 1790).—Holotype (ZSM 118/1937; not mentioned in Franzen and Glaw, 2007) from La Puerta, Fusagasuga´, Colombia, 1200 m; Peters and Donoso-Barros (1970: 19).

Ameiva ameiva fischeri Peters and DonosoBarros (1970: 19; replacement name for Cnemidophorus maculatus Fischer because the name A. maculata Gray [1838] was already in existence and a synonym of A. a. ameiva).

Ameiva ameiva (Linnaeus, 1758): Echternacht (1971: 14).

Diagnosis.—A medium-sized Ameiva distinguished from all congeners by the following combination of characters: (1) maximum SVL in males 243 mm; (2) smooth dorsal head scales; (3) frontal single; (4) frontoparietal and parietal plates in contact with interparietals; (5) 15–36 (both sides) scales, usually in a single row, between supraoculars and supra-

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155

FIG. 21.—Cephalic morphology of Ameiva praesignis (USNM 217028, adult male, snout–vent length [SVL] 180 mm).

ciliaries; (6) 12–21 occipitals, usually subequal to first dorsal row; (7) 22–40 anterior gulars; (8) middle anterior gulars polygonal or rounded and usually moderately enlarged, less often small, and rarely distinctly enlarged; (9) patch of moderately enlarged posterior gulars present, or all posterior gulars small, rarely posterior gulars in a distinctly enlarged patch; (10) 22–40 posterior gular scales between antegular and gular folds; (11) enlarged mesoptychial scales subequal or larger than largest gulars; (12) postbrachials moderately to distinctly enlarged; (13) 237–348 scales between occipitals and base of tail; (14) 111–157 dorsal scales across midbody; (15) ventrals in 29–34 transverse rows, and 10 longitudinal rows; (16) adult male coloration in life gray, bluish gray or gray–brown, each scale often with black pigments, with pale lateral ocelli; (17) throat in adults cream colored or blue; (18) a distinct whitish or yellow vertebral light stripe often present or ocelli present across dorsum; (19) juveniles with paired black spots and with

indistinct pale dorsolateral line bordering upper margin of broad black lateral stripe, vertebral pale stripe or spots often on dorsum; (20) associated with open localities, dry forests, and savannahs.

Description.—Largest species of Ameiva, with a maximum SVL in males 243 mm (USNM 162764), maximum total length in males 737 mm (USNM 162764); maximum SVL in females 167 mm (USNM 162761), maximum total length in females 511 mm (USNM 142392). Head pyramidal, 0.22–0.26 (0.24 6 0.01, n 5 29) times SVL in males; 0.21–0.26 (0.22 6 0.01, n 5 23) times SVL in females. Snout elongate, bluntly pointed; canthus rostralis distinct. Gular region and cheeks usually swollen in large males. Neck as wide or only slightly narrower than head and body. Body cylindrical. Limbs well developed; tibia 0.19–0.24 (0.21 6 0.01, n 5 28) times SVL in males; 0.20–0.24 (0.22 6 0.01, n 5 23) times SVL in females; foot 0.29–0.44 (0.35 6 0.03, n 5 28) times SVL in males; 0.30–0.41

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[No. 25

FIG. 22.—Individuals of Ameiva praesignis in Venezuela. (A) Uncatalogued adult male from Monte Cano, Peninsula de Paraguana´, Falco´n; photo: C. L. Barrio-Amoro´s. (B) Uncatalogued adult male from Hato Masaguaral, Guarico; photo: N. D. Sly. (C) Uncatalogued adult male from Caja seca, Me´rida; photo: C. L. Barrio-Amoro´s. (D) Uncatalogued adult female from Caja Seca, Me´rida; photo: C. L. Barrio-Amoro´s. (E) Uncatalogued adult female male from Masaguaral, Guarico; photo: N. D. Sly.

(0.35 6 0.03, n 5 23) times SVL in females.

Tail slightly depressed proximally, round in cross section distally, tapering toward tip, 2.03–2.48 (2.28 6 0.12, n 5 13) times SVL in males; 2.01–2.56 (2.29 6 0.15, n 5 15) times SVL in females.

Rostral pentagonal, higher than wide, well visible from above, bordered posteriorly by nasals, which form a short or very short medial suture (nasals medially separated and rostral in distinct contact with frontonasal in USNM 121194). Each nasal divided by oblique

suture. Nostril in lower part of suture and anterior to it, directed lateroposteriorly, taller than long. Frontonasal hexagonal or octagonal, in contact with nasals, loreal, and prefrontals.

Prefrontals paired, roughly pentagonal or trapezoidal, with long medial suture about two or three times as long as that between nasals (equal to that between nasals in CMS 76014a), laterally in contact with loreal, first supraocular, and first supraciliary; well separated from nasal. Frontal hexagonal or roughly pentagonal, longer than wide and wider

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FIG. 23.—Top: copulating pair of Ameiva praesignis from Hato Masaguaral, Guarico, Venezuela; photo: N. D. Sly. Bottom: female A. praesignis from San Diego de Cabrutica, Anzoa´tegui; photo: E. Wefer.

anteriorly (posterior portion horizontally divided in USNM 127285, 162764, and CMS 76014a), its sutures with prefrontals angulate or semicircular, slightly curved or straight, those with frontoparietals almost forming a wide angle or occasionally almost forming straight line; frontal laterally in contact with first, second, and, often, narrow part of third

supraoculars (in USNM 217057 in contact with first and second supraocular and with anteriormost scale of circumorbital series).

Frontoparietals paired, pentagonal or trapezoid, longer than wide and with long medial suture; laterally in contact with third supraocular and small circumorbital scales bordering fourth up to midportion of third supraocular

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(only in contact with circumorbitals in USNM 217057). Interparietal hexagonal or rarely pentagonal, higher than wide; usually narrower or subequal (rarely slightly wider) than adjacent parietals; sutures with parietals straight or rarely slightly oblique; interparietal bordered at each side by two large (very rarely one), irregular parietals divided by oblique suture; outermost parietal usually round in shape and larger, subequal, or less often smaller to inner parietal. Parietal series composed of 3–6 (usually five) scales (including interparietal). Occipitals 12–21 (16 6 2.04, n 5 63), irregularly shaped, mostly subequal in size; occipitals usually subequal in size to first dorsal row, rarely larger. Supraoculars four at each side, three on both sides of UF 127336 (a narrow scale wedged between second and third supraoculars on both sides of UF 127287 counted as an additional supraocular, a similar scale between third and fourth supraoculars on both sides of USNM 121196 also counted as additional supraocular), fourth supraocular absent in USNM 127336; second or third supraocular largest, fourth smallest. Circumorbital semicircles formed by 3–10 scales at each side, 6–17 (10.7 6 1.94, n 5 61) combining both sides, reaching middle or posterior portion of third supraocular (only rarely reaching suture with frontal); only one scale row between third supraocular and prefrontals; fourth supraocular separated from parietals by up to four rows of circumorbital scales (usually three rows).

Laterally, all supraoculars except first, usually separated from supraciliaries by single row (rarely completely or partially double row) of small rectangular scales, 15–36 (22.6 6 4.98, n 5 61) combining both sides. Supraciliaries six or seven at each side (eight or nine in FLMNH 155874), first highest, second, or third longest, remaining scales short and subequal. Loreal very large, trapezoidal and single in contact with nasal, frontonasal, prefrontal, first supraciliary, preocular, first subocular (very rarely separated by an elongated preocular), and third and fourth supralabials (occasionally also in narrow contact with second supralabial). Suboculars four, large; first subocular in preocular position, rarely divided into two scales, irregularly trapezoidal, longer than wide and narrower to as wide as second

[No. 25

subocular; in contact with loreal, first supraciliary, small scales on ocular region, and second subocular; margin with loreal slightly curved.

Very rarely second and third suboculars fused.

Third subocular longest; all in contact with supralabials. Continuous keel from first through entire length of third subocular. Postoculars small, approximately in two rows of 3–5 irregularly trapezoidal scales. Palpebral disc semiopaque with enlarged scales on lower eyelid. Supralabials usually six or seven (eight in USNM 142391) at each side, fifth below center of eye; followed to commissure of mouth by smaller scales. Row of distinctly enlarged supratemporals decreasing in size posteriad.

Temporal region with rectangular and round scales, slightly smaller centrally than peripherally. External auditory meatus large, vertically oval, bordered by granular scales, its anterior margin semicircular, its posterior margin straight. Tympanum recessed. All dorsal and lateral head scales juxtaposed and smooth or slightly rugose.

Symphysal anteriorly ellipsoid, its posterior sutures forming wide angles with infralabials and postsymphysal. Postsymphysal single and pentagonal; in contact with first and second infralabials, followed by three pairs of enlarged chinshields. First pair of chinshields in ample medial contact; first, and occasionally second pairs in contact with infralabials (rarely third pair may be in narrow contact). Remaining chinshields separated from infralabials by one row of sublabials. Medial chin scales moderately small, elongate, convex, smooth, juxtaposed, hexagonal or subhexagonal, in slightly oblique rows, all subequal in size. Six (rarely seven) infralabials, fifth below center of eye followed to commissure of mouth by smaller scales. Gular region divided in two areas: anterior region with enlarged polygonal and flat scales in slightly oblique rows that grade to larger scales at center of throat, delimited posteriorly by line uniting lower margin of ear openings; middle anterior gular scales moderately enlarged or subequal to lateral scales (very rarely scales distinctly enlarged), 22–40 (29.2 6 4.28, n 5 62) scales along midline of chin from anteriormost gular to antegular fold. Posterior gular region covered by smaller polygonal (or rarely slightly rounded) scales, in transverse rows; posteriormost median posterior gulars often moderately enlarged

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(rarely forming a patch of distinctly enlarged scales) or small scales margining the antegular fold, 9–18 (13.3 6 2.45, n 5 62) scales midventrally from anteriormost posterior gular to antegular fold. Mesoptychial scales moderately enlarged (larger or subequal to largest gulars; rarely slightly smaller), in about one or two rows, hexagonal, flat, smooth, and juxtaposed.

Scales on nape and sides of neck similar to dorsals. Dorsals and scales on flanks granular (slightly larger on dorsum than laterally in some specimens), round, smooth, subimbricate or juxtaposed, in approximately transverse rows; 237–348 (289.3 6 23.19, n 5 60) scales from occiput to base of the tail. Scales around midbody (excluding ventrals) 111–157 (133.0 6 11.26, n 5 60). 37.2–54.3 (45.8 6 3.91, n 5 60). Ventrals large, smooth, rectangular (wider than long), imbricate, in 10 longitudinal (at midbody) and 29–34 (31.4 6 1.16, n 5 63) transverse rows. Transition between ventrals and scales on flanks sharp.

Preanal shield with 4–8 rows of enlarged scales; pattern of terminal scales of the preanal plate variable; often a large or two subequal median scales followed posteriorly by two subequal or slightly larger scales, or followed by three scales (middle one smallest and wedged between the other two), or preanals divided and irregularly disposed in series of four or five enlarged and subequal scales. Preanal plate surrounded anteriorly and laterally by smaller scales; posteriorly by much smaller scales. Femoral pores in continuous row along each thigh; medially with a short gap; 14–21 pores on each leg, 28–42 (33.9 6 3.24, n 5 62) pores combining both legs. Each pore between three or four scales.

Scales on base of tail rectangular or pentagonal, smaller than ventrals, longer than wide, mostly keeled (smooth on base of tail on ventral surface). Caudal scales imbricate, in transverse rows continuous around tail, except first few rows incomplete ventrally. Distally scales become longer and narrower, distinctly keeled, in transverse and approximate longitudinal rows.

Arms with a row of very large, smooth, slightly imbricate, trapezoid antebrachial scales on anterodorsal aspect of forearms, and similar but smaller brachial scales on

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upper arms that extend almost to shoulder.

Brachial row in one moderately to distinctly enlarged row with a smaller accessory row bordering upper margin and grading into granules dorsally. Antebrachial and brachials connected or separated by up to four rows of small scales at elbow level. Dorsoposterior, posterior, and ventral aspect of arms with scales similar to, but slightly larger than, dorsals, except for a group (approximately two rows) of postbrachial scales on posterior aspect of upper arms (near elbow), which are slightly larger and irregular. Legs with large, smooth, imbricate scales on anterior and ventral aspects of thighs, and ventral aspect of shanks. On thighs there is a row of large, trapezoid scales anteriorly, and from this row toward row of pores the scales gradually become smaller and irregular. On ventral aspect of shanks three rows of very large scales, anterior two trapezoid, posterior one rhomboid; they decrease in size from anterior toward posterior row. Elsewhere on legs scales similar to dorsals.

Subdigital lamellae transversely enlarged and single, moderately to distinctly tuberculate towards base. Tubercles most prominent under base of third toe. On palms, lamellae of inner finger continue up to wrist, increasing in size toward it; single large scale present on postaxial side, following line of fifth finger, short distance from wrist. Lamellae under inner four toe continue up to tibio-tarsal articulation. Lamellae under fourth finger 13–18 (15.8 6 0.87, n 5 60), under fourth toe 29–40 (34.0 6 2.45, n 5 58).

Coloration in life.—This species exhibits conspicuous ontogenetic and moderate geographic color variation (Figs. 11, 22, 23).

Juvenile and hatchling coloration remains constant throughout its distribution. Young specimens have a broad black or dark brown lateral stripe extending at each side from postocular region through the sides of the head (above ear opening) to the sides of tail, where it breaks into irregular blotches. As individuals grow, light spots appear within the black band, rest of the flanks and dorsolateral area. This band often is accompanied by an inconspicuous light dorsolateral stripe along its upper border and fades on the posterior dorsum (this stripe is occasionally distinct and

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TABLE 6.—Characters of Ameiva from the Guyanan uplands contrasted with two species confused with it. Ranges (in parentheses) follow means 6 standard deviation.

Characters

Scales between supraoculars and supraciliaries Occipitals Occipitals/first row of dorsals Anterior gulars Size of anterior gulars Posterior gulars Patch of enlarged posterior gulars Dorsals Scales around body Scales around body/dorsals 3 100 Femoral pores Fourth finger lamellae Fourth toe lamellae

Ameiva ameiva (n 5 22)

Ameiva sp. (n 5 5)

A. praesignis (n 5 63)

27.3 6 7.18 (19–44) 14.6 6 1.49 (11–17) Smaller (4%) Subequal (63%) Larger (33%) 27.4 6 3.59 (18–32) Small (10%) Medium (35%) Large (56%) 11.7 6 2.16 (7–17) Small (0) Medium (23%) Large (77%) 293.5 6 21.76 (259–339) 159.5 6 12.61 (129–187)

20.6 6 1.15 (20–22) 13.6 6 2.30 (11–17) Smaller (0) Subequal (60%) Larger (40%) 25.4 6 3.20 (21–28) Small (40%) Medium (40%) Large (20%) 12.2 6 2.06 (10–15) Small (20%) Medium (40%) Large (40%) 248.7 6 16.99 (225–262) 133.7 6 23.68 (114–163)

22.6 6 4.98 (15–35) 16.0 6 2.04 (12–21) Smaller (0) Subequal (73%) Larger (27%) 30.4 6 4.62 (23–40) Small (13%) Medium (48%) Large (25%) 13.3 6 2.45 (9–18) Small (35%) Medium (38%) Large (27%) 289.3 6 23.19 (237–348) 133.0 6 11.36 (111–157)

54.5 37.5 15.1 35.8

6 6 6 6

5.96 4.39 0.66 2.15

(43.2–72.2) (30–45) (14–16) (32–40)

may have narrow black margins; in most young specimens from Panama, Colombia and a few from Venezuela, this stripe extends to tail). A narrower whitish line margins the lower border of the dark lateral band and extends to the inguinal region (sometimes breaking into spots); a similar pale stripe is also present on sides of tail. Dorsum of head and body are uniformly tan or green anteriorly and brown posteriorly, with small to large but narrow, paired black spots at each side of the vertebral region. A pale vertebral stripe or numerous white ocelli on flanks that extend across dorsum (or at least dorsolateral region) are present in most specimens (these light markings tend to be absent in hatchlings). The sides of the head are whitish with conspicuous black spots and a blackish stripe along canthus. Flanks are brownish with dark brown or dark gray spots (sometimes very inconspicuous) that form an irregular checkered pattern or longitudinal stripes. The limbs and tail are brown with dark brown spots and dots. The throat is uniformly whitish or with large dark gray spots. The abdomen is whitish with numerous to very few dark markings, especially laterally.

Adult male A. praesignis are gray–brown or bluish gray, often with black pigments on each

54.6 34.0 14.4 30.7

6 6 6 6

10.90 (43.8–65.7) 2.21 (29–34) 0.89 (14–16) 2.21 (29–34)

45.8 33.9 15.8 34.0

6 6 6 6

3.36 3.24 0.87 2.45

(37.2–54.3) (28–42) (13–18) (29–40)

scale, producing a salt-and-pepper effect. A pale lemon yellow or whitish vertebral stripe (often with irregular borders) is often present along dorsum (usually extending from shoulder level to sacrum), in contact or separated from whitish, pale yellow, or blue ocelli (ocelli often pale turquoise on paraventral area) with narrow black borders. Ocelli extend toward flanks, often aligned in vertical lines; in some specimens a few of the lateral ocelli occasionally fuse, forming short, vertical bands. Some specimens lack the light vertebral stripe but have transverse rows of ocelli extending across the dorsum (very rarely faded or absent on the dorsum of old adults); in these specimens the ocelli are often most conspicuous on the posterior half of the body. Individuals with vertebral stripes are usually present in all populations, but the incidence of striped individuals appears to vary slightly geographically (see remarks below). The flanks are usually grayish like the dorsum, but in some specimens they are brownish or have a reddish cast toward the vertebral stripe. The top of the head is gray, like the body, and has black reticulations that are less conspicuous on the slightly lighter sides. The arms are gray with black reticulations; the legs have ocelli like those on the body but smaller. The tail is

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grayish with minute black pigments and whitish spots irregularly disposed that become turquoise toward tail tip; the sides of the tail are bluish, becoming more intensely blue toward the tip. The chin and throat are whitish or pale blue with grayish spotting or in some specimens with numerous black minute dots (amount individually variable).

The abdomen and ventral aspects of limbs (darker on legs) and tail are turquoise blue suffused with black spots (most evident on ventral aspects of limbs).

Slight geographic color variation was also observed in females, although in less degree than in males. In all populations observed, females retained some of the juvenile color pattern. However, female A. praesignis sometimes have remnants of a pale, black-margined dorsolateral stripe that sometimes extends to the tail (Figs. 22 and 23) that was never observed in female A. ameiva or A. atrigularis.

The flanks have a longitudinal lower lateral, white or bluish stripe from midflank to inguinal region; this stripe may be margined by black stripes or not. Females often have whitish ventral surfaces, but in some populations blue-throated females were observed.

At all stages of development, A. praesignis has a reddish-brown iris.

Coloration in preservative.—Preserved specimens retain much of the pattern observed in life, but the background color fades to muddy brown, purplish, or dark gray. The smallest adult male in our sample with a completely adult coloration is 101 mm SVL. The adult male UF 127333 has a conspicuous dorsal pattern formed by black reticulations that merge, forming a marbled pattern on posterior dorsum and sacrum. This same specimen and males UF 127335 and 155874 have lost most of their flank ocelli except posteriorly where they also invade the sacrum. The chin, throat, chest, and most of abdomen fade to blue, bluish gray or dark blackish gray (some specimens can appear almost black ventrally). Some specimens have black speckling or spotting on the throat. Pale spots and black blotches are often distinct laterally on abdomen. The posterior portion of the abdomen, ventral aspect of legs, anal region, and subcaudal area are whitish with blue or gray blotches. Some specimens have all ventral surfaces immaculate whitish or pale gray; UF

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155874 has all ventral surfaces blackish (edges of scales pale); UF 217038 and 217041 have black speckling throughout all ventral surfaces.

The abdomen of some juveniles may be uniformly whitish, pale gray, or profusely checkered with dark blue and black.

Distribution and natural history.—In Venezuela, A. praesignis occurs throughout dry, xeric habitats, grasslands, and savannahs in the northern and western portions of the country west of the Andes and throughout the savannahs and Llanos of central and western Venezuela (Fig. 3). Its range extends throughout Colombia (Barbour and Noble, 1915; Mu¨ller and Hellmich, 1940; Ruthven, 1922), central and western Panama (Echternacht, 1971) and extreme southwestern Costa Rica (Savage, 2002). Although the distribution of Ameiva praesignis may seem odd at first, other reptiles of northern Venezuela associated with open or dry habitats have similar ranges (e.g., the gecko Phyllodactylus ventralis and snake Coluber [Masticophis] mentovarius).

In Ocumare de La Costa and Cata, northern Aragua, this species is very common in peranthropic situations. At these localities, A. praesignis was commonly observed in backyards and parks, where it is sympatric with Cnemidophorus lemniscatus cf. splendidus. This species has also been recorded from the thornscrub in Maiquetia, Vargas (Marcuzzi, 1954). Ruthven (1922) reported it from flood-plain forests, deserts, dry forests, and clearing in Santa Marta, Colombia. Staton and Dixon (1977) reported A. praesignis inhabits both open grasslands and ecotones between savannah and gallery forest in the Llanos of Apure and Guarico, Venezuela.

Additionally, Staton and Dixon (1977) reported A. praesignis becomes active 1– 1.5 h after sunrise, when individuals bask for approximately 30 min before foraging. The same authors reported that at midmorning, specimens travel in ‘‘loose packs.’’ However, they indicated it was unclear if the lizards formed these groups as part of their foraging behavior. Staton and Dixon (1977) also observed intraspecific antagonistic behavior.

Adult specimens were seen chasing subadults and young away. Ameiva praesignis appears to be well adapted to life in the seasonal Venezuelan central and western Llanos. Sta-

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ton and Dixon (1977) reported that in Apure and Gua´rico, adult A. praesignis aestivated during the dry season and emerged shortly after rains. According to the same authors, young specimens occasionally remain active during the dry season. Staton and Dixon (1977) reported newborns during November in the Venezuelan Llanos.

Remarks.—We examined specimens associated with the names praesignis, vogli, and fischeri from Panama, northern Colombia, and Venezuela. We found the diagnostic characters of these purported taxa to represent nothing more than intraspecific variation and consider all these populations to be conspecific. Although we did not examine material associated with A. a. ornata, we believe the diagnostic characters described in the original description are not sufficient for recognizing this taxon and tentatively consider it a synonym of praesignis.

Echternacht (1971) reported that specimens from eastern Panama differ from those of western Panama in having a distinct pale vertebral stripe as adults. A vertebral stripe is also absent in the only specimen known from southwestern Costa Rica (Savage, 2002).

According to Bennet (1968), a Pleistocene eruption of El Valle Volcano could have separated eastern and western Panamanian A. praesignis for sufficient time to explain divergence in color pattern between both populations.

Ameiva a. vogli from the Venezuelan Llanos was reported to lack the vertebral stripe (Donoso-Barros, 1968; Mu¨ller, 1929; Peters and Donoso-Barros, 1970). We did not examine the material reported by Mu¨ller (1929); nonetheless, we observed a pale vertebral stripe in most specimens from the Venezuelan Llanos. We found no sexual dimorphism in the presence of this stripe. Among adult northern A. praesignis (above 105 mm SVL) from localities west of the Andes in northern Venezuela, Colombia, and eastern Panama, 76% (n 5 27) have a pale vertebral stripe.

Similarly, 62% (n 5 29) of adult southern lizards from the Venezuelan Llanos have this stripe. Additionally, 24% of the southern and 12% of the northern population have pale ocelli extending across the dorsum or along the vertebral line. Only 12% of the northern and

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14% of the southern adult A. praesignis lack both vertebral ocelli and a stripe.

As indicated in the results section, the only difference we found between northern and southern samples of A. praesignis is foot length relative to SVL. This character is particularly divergent among males. Male southern A. praesignis from the slightly wetter grasslands and Llanos have shorter feet than northern lizards from arid thornscrub, dry forests, and grasslands. Possibly, the shorter foot of southern lizards is an adaptation for a slightly wetter, grassy habitat. Southern A. praesignis has the shortest foot among all Ameiva populations examined (Tables 1 and 6). However, we do not believe that this single character warrants recognizing southern lizards (referable to A. a. vogli) as a different taxon.

KEY TO THE SPECIES OF AMEIVA IN VENEZUELA 1.

(A) Frontal scale horizontally divided in two or three scales; one row of distinct, transversely enlarged postbrachial scales that extend midway along arm; dorsum in adults usually with five longitudinal pale stripes; juveniles with pattern similar to adults; maximum SVL A. bifrontata complex below 135 mm (B) Frontal scale almost always entire (at least in Venezuela, Colombia, Panama and Trinidad); nasals broadly separated from prefrontals; patch of slightly to moderately enlarged postbrachial scales near elbow; dorsum of adults without longitudinal stripes or with single vertebral stripe; juveniles with pattern different from that of adults; maxi2 mum SVL above 150 mm (A) Maximum SVL below 190 mm; 129–187 (excluding ventrals) scales around midbody; dorsum of adults often green posteriorly, pale vertebral stripe never present, ocelli almost never extending across dorsum; juveniles never with white ocelli across dorsum or 3 only rarely on dorsolateral area (B) Maximum SVL 243 mm; 111–157 (usually below 140) scales around midbody; dorsum in adults never green posteriorly, with pale vertebral stripe or with ocelli across dorsum; juveniles often with white ocelli A. praesignis across dorsum or dorsolateral area (A) Medial anterior gulars usually small or moderately enlarged; 13 or more posterior gulars (usually 15 or more); posterior gulars small or moderately enlarged, not or rarely forming enlarged medial patch; throat black 4 in adults (north of Orinoco River) (B) Medial anterior gulars usually distinctly enlarged; 7–16 posterior gulars (usually 12 or less); posterior gulars usually forming _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _

2.

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _

_ _ _ _ _

3.

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _

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HERPETOLOGICAL MONOGRAPHS distinctly enlarged patch; throat whitish in adults (mostly south of the Orinoco River, except in southern Anzoa´tegui and Delta A. ameiva Amacuro) (A) Maximum SVL 186 mm; occipitals usually larger than first dorsal row; medial anterior gulars often moderately enlarged; dorsum uniformly brown or green posteriorly with minute black reticulations (forest edges of north-central and northeastern Venezuela, Isla de Margarita, Trinidad, and A. atrigularis possibly Tobago) (B) Maximum SVL 152 mm; occipitals usually subequal to first dorsal row; medial anterior gulars usually small; dorsum brown with large, black, broad reticulations (saA. pantherina vannahs of southern Monagas; _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _

4.

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _

_ _ _ _ _ _ _ _

Included in the A. bifrontata complex are the nominal subspecies found throughout northern Venezuela, A. b. insulana Ruthven from Los Testigos Archipelago and A. provitaae Garcı´a-Pe´rez from Lagunillas, Me´rida.

The taxon from Los Testigos Archipelago apparently differs from A. b. bifrontata in the size of the postbrachial scales, and having brachials and antebrachials separated by small scales (Ruthven, 1924). According to the original description, A. provitaae differs from A. bifrontata in a few details of squamation, coloration and a larger SVL (Garcı´a-Pe´rez, 1995). Nevertheless, our current understanding of the taxa associated with A. bifrontata is poor and several of the diagnostic characters used to distinguish among the subspecies of A. bifrontata and A. provitaae appear to be far more variable than previously thought (G. Ugueto and M. Harvey, personal observations). For this reason we prefer to treat A. bifrontata as a complex of closely related taxa. The authors are currently working to clarify the status of these taxa.

DISCUSSION Numerous Neotropical species previously considered to be widespread have proven to be composed of several closely related taxa (e.g., Bergmann and Russell, 2007; Henderson et al., 2009; Passos and Fernandes, 2008; Rodrigues, 1987), and the A. ameiva complex is a perfect example of this history of underestimated taxonomic diversity. Prior to this work, this teiid lizard was most commonly treated as a widespread and morphologically variable taxon. Although as many as 11 subspecies were

163

described, most recent authors preferred to consider this A. ameiva as monotypic (e.g., Avila-Pires, 1995). Revalidation of two names previously relegated to the synonymy of A. ameiva and discovery of A. pantherina corrects some of the confusing taxonomic problems associated with this genus of large, ubiquitous lizard.

With the taxa recognized and described in this study, the genus Ameiva now contains 37 species. Our study leaves several important questions unanswered. Status of some insular populations currently considered subspecies of A. ameiva should be reconsidered. It is likely that A. a. tobagana and A. a. fulginosa represent full species. Additionally, the Ameiva from the Guayana uplands in southeastern Venezuela require further evaluation.

We were unable to allocate five juvenile specimens from ‘‘Paulo,’’ Mount Roraima to any of the species examined in this study.

These specimens differ from all other species of the A. ameiva complex in several characters (Tables 1 and 5). Currently, the authors are working to clarify the status of this population.

As in other reptile genera, such as Drymarchon (Wu¨ster et al., 2001), Gonatodes (Rivero-Blanco, 1979), Mabuya (Miralles et al., 2009), and Cnemidophorus (Markezich et al., 1997; Ugueto et al., 2010; Ugueto and Harvey, in press), Venezuela appears to be an important center of diversity in the A. ameiva complex. High local diversity in these genera is likely due to Venezuela’s intermediate geographic position between the Amazonian region, Central America, and the Antilles, and because of the remarkable variety of ecosystems found within this country (Miralles et al., 2005).

A COMMENT ON THE IDENTITY OF THE AMEIVA LIZARDS INTRODUCED IN THE STATE OF FLORIDA, SOUTHEASTERN UNITED STATES Neill (1957) first reported A. ameiva from an unspecified locality in Florida. A year later, Duellman and Schwartz (1958) reported the species had been introduced to vacant lots at two localities in Miami–Dade County, Florida, since (at least) 1954. King and Krakauer (1966) reported A. ameiva was introduced to three localities in Miami, and Wilson and Porras (1983) reported this species for the first time from Key Biscayne, Miami–Dade

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County. Currently, introduced Ameiva sp. are known from Deerfield Beach, Broward County (Krysko et al., 2005), South Miami, Hialeah, and Key Biscayne in Miami–Dade County (Smith and Krysko, 2007). Individuals have also been collected in Monroe and Palm Beach Counties (Smith and Krysko, 2007), but the status of these populations is still unknown. Furthermore, we are aware of two previously unreported localities where A. ameiva is abundant and likely well established: north of Biscayne Boulevard and 125th Street, North Miami, Dade County, and near the intersection of Stirling and Davie Roads in Davie, Broward County, Florida.

Replicating the taxonomic history of this species complex in its native distribution, the identity of the A. ameiva populations introduced in southeastern Florida has been confusing (Conant, 1975; Meshaka et al., 2004). King and Krakauer (1966) referred populations from Suniland (South Miami) and Hialeah to A. a. ameiva and populations from approximately 25 city blocks from NW 79th to 76th streets and from 36th Avenue to Le Jeune Road to A. a. petersii. Based on information from Echternacht, Conant (1975) suggested Miami Ameiva sp. may represent intergrades between the two South American subspecies. Wilson and Porras (1983) did not assign the population from Key Biscayne to any subspecies and questioned Conant’s suggestion, arguing ‘‘it is clear that the adult males of the Suniland population are predominantly green, whereas those of all other populations in Dade County are predominantly blue and larger in size. All of the existing populations are in disparate areas of the city, unconnected to one another, and all, except the one on Key Biscayne, appear to be on decline.’’ Wilson and Porras (1983) considered both subspecies in question to be the nominal taxon and A. a. petersii, although based on a comment by A. Echternacht, they considered the identity of Floridian lizards as ‘‘unclear.’’ Meshaka et al. (2004) once again resurrected the concept of the Florida populations being the result of intergradation between two introduced subspecies. Our observations contradict those of Meshaka et al. (2004), as it appears that all populations of Ameiva sp. in south Florida

[No. 25

are still isolated and have not expanded much from their original places of introduction. It appears that new localities probably represent new introductions.

We observed Ameiva lizards in Hollywood and Davie, Broward County, in North Miami, Miami–Dade County, and in the locality of Suniland, Miami–Dade County. At all these localities adult males were bicolored (brown anteriorly and bright green posteriorly) and did not exceed 160 mm in SVL (Fig. 24).

Based on these characteristics and examination of a few specimens from two of these localities, it is clear these populations are referable to A. ameiva (sensu stricto). The population in Hollywood was extant but very limited in distribution at the time of our last visit (April 2006), and only a few specimens were observed. The population of North Miami was also limited in distribution, but at the time of our last visit (April 2009) appeared to be very healthy. During a very short survey of this area, 26 specimens were observed in less than 1 h; 18 of these were juveniles.

Meshaka et al. (2004) reported they were unable to find this species at Suniland; we found this population was limited in distribution (extending throughout a park and backyards of adjacent homes) but was very healthy during the time of our last visit in May 2009.

In 1.5 h, 21 specimens of all ages and sexes were observed. All these populations appear to be isolated from each other and their origin can probably be traced to releases associated with pet dealers or pet owners (Wilson and Porras, 1983).

Another population is present in Key Biscayne and it was probably derived from releases of lizards from a zoological park exhibit (Meshaka et al., 2004). This population was still extant at the time of our last visit to the area (May 2009), and it is formed by very large lizards (Meshaka et al., 2004, reported a maximum SVL of 200 mm for males and 159 mm for females collected at Cape Florida State Recreation Area in Key Biscayne), mostly gray-blue in coloration when adults (Fig. 23). Several adults specimens observed had a distinct vertebral light stripe on the posterior part of the dorsum or ocelli across the back. All of the Ameiva sp. on Key Biscayne are A. praesignis. Meshaka et al.

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165

FIG. 24.—Populations of Ameiva introduced in southeastern Florida, United States of America. Top: A. ameiva, adult male from North Miami, Miami–Dade County. Bottom: A. praesignis, adult male from Key Biscayne, Miami–Dade County. Photos: G. N. Ugueto

(2004) incorrectly thought that the bluish color and large size of this population are traits associated with A. a. ameiva.

In summary, two species of Ameiva occur in south Florida: A. ameiva restricted to a few

localities in Miami–Dade and southern Broward counties, and A. praesignis restricted to Key Biscayne, Miami–Dade County.

Acknowledgments.—For loan of specimens under their care and kindly checking the status of several type

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HERPETOLOGICAL MONOGRAPHS

specimens in their institutions we thank J. A. Campbell (UTA), R. F. Feeney (LACM), J. Hallermann (ZMH), B. Hollingsworth (SDNHM), D. Kizirian (AMNH), C. J. McCarthy (BMNH), R. A. Nussbaum (UMMZ), A. Resetar (FMNH), R. P. Reynolds (USNM), S. P. Rogers (CM), J. Rosado (MCZ), G. Schneider (UMMZ), and H. Voris (FMNH). We are very grateful to A. Afonso, A. Allais, A. Anker, D. Ascanio, E. Asens, C. L. BarrioAmoro´s, T. Barros, J. E. Garcı´a Pe´rez, D. Kjerlkerud, A. Miralles, G. Rivas, J. M. Saez Sierra, J. C. Sen˜aris, N. D. Sly, and E. Wefer who generously allowed us to use their photographs and/or observations in this study.

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APPENDIX I Specimens Examined Ameiva ameiva (27).—BOLIVIA: Santa Cruz: Finca Dos Milanos (UTA R-34735–39), El Refugio (UTA R38077–79). PERU: Loreto: Yarinacocha, Rı´o Ucayali (FMNH 56026, 56030–34). VENEZUELA: Amazonas: Tamatama, Rı´o Orinoco, NMNH 80653–56; Boca Mavaca, 84 km SSE of La Esmeralda, NMNH 162756–59.

Anzoa´tegui: Corrientoso, near Carapa NMNH 80615–16; Soledad NMNH 80621–24. Bolı´var: Ciudad Bolı´var NMNH 80602,04; Santa Marı´a de Erebato, NMNH 561200–06.

Ameiva atrigularis (42).—VENEZUELA: Distrito Capital: El Valle (USNM 128914). Gua´rico: Parque Nacional Guatopo, 15 km NW of Altagracia de Orituco (USNM 217032). Miranda: Quebrada Siquire, 11 km ENE of Santa Lucia (USNM 121197); 1 km S of Rı´o Chico (USNM 216981); Curupao, 5 km NNW of Guarenas (USNM 217033–34); Hacienda Bejuquero, 1 km S of Rı´o Chico (USNM 217060–62); Santa Lucia, 11 km ENE on Quebrada Siquire (CM S 7445–48). Nueva Esparta: Isla de Margarita, 3 km NNE of La Asuncio´n (USNM 217051); Isla de Margarita, near Salamanca, 2 km N and 1 km E of La Asuncio´n (USNM 217052). Sucre: Manacal, 26 km ESE of Caru´pano (USNM 217053); Cumanacoa (MCZ 9993, holotype of A. a. melanocephala). Vargas: La Guaira (USNM 22526). TRINIDAD: Unspecified locality: MCZ 186018–19 (syntypes of A. atrigularis). Tunapuna/ Piarco: St. George, Mount Saint Benedict (CM S 4845, 4947–53, 6498–501, 6503, 6505–509).

Ameiva bifrontata (27).—VENEZUELA: Nueva Esparta: Isla Cubagua (SDMNH 34906, 34941–46); Isla de Margarita, Los Robles (USNM 79217–21). Sucre: Caru´pano (UMMZ 107086). Trujillo: Sabana Mendoza (UMMZ 57436–42). Yaracuy: San Felipe (UMMZ 55532). COLOMBIA: Cesar: Valle Dupar, Santa Marta Mountains (UMMZ 55024–30).

Ameiva pantherina (32).—VENEZUELA: Monagas: Hato Mata de Bejuco, 54 km SSE of Maturı´n, USNM 216982, 217048–50; Hato Mata de Bejuco, 55 km SSE of Maturı´n, USNM 217047; 42 km SE of Maturı´n, LACM 31414–21; 60 km SE of Maturı´n, LACM 31422–41.

Ameiva praesignis (65).—VENEZUELA: Apure: Hato Cariben, Rı´o Cinaruco, 60 km NW of Puerto Paez (USNM 162760). Aragua: Pie del Cerro La Victoria (USNM 121195–96); El Limo´n (NMHN 142391–93).

Carabobo: Sabana Aguirre, 5.5 km SE of Montalban (USNM 217028–29). Cojedes: Finca La Coromoto, Camoruco, Municipio Cojedes (FMNH 127284, FMNH 155874). Gua´rico: San Juan de Los Morros (USNM 72752); Palenque (USNM 128915); Estacio´n Biolo´gica de los Llanos, 9 km SE of Calabozo (USNM 217035–45); Calabozo Airport, Distrito Miranda (LACM 130661).

Me´rida: Coromoto, 4 km S and 6.5 km E of Tabay (USNM 162761–62). Portuguesa: San Jorge, Municipio

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Piritu (FMNH 127284–89). Trujillo: El Dividive, 28 km NW of Valera (USNM 162764); El Dividive, 30 km NW of Valera (USNM 162790); Valle Verde near Santa Apolonia, 52 km WNW of Valera (USNM 162791–94). Vargas: El Limo´n (USNM 121194, 162763). Zulia: Can˜o Azul, El Rosario, 65 km WNW of Encontrados (USNM 217054); Hacienda Platanal, near Cerro Azul, 33 km NW of La Paz (USNM 217055–56); near Cerro Azul, 39 km NW of La

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Paz (USNM 217057). COLOMBIA: Cesar: Valle Dupar, Santa Marta Mountains (UMMZ 54915–16); Vallencia, Santa Marta Mountains (UMMZ 54917–21). PANAMA: Canal Zone: El Vigı´a (UMMZ 76012); Gatuncillo (UMMZ 76013); Alhajuela (UMMZ 76014, four specimens); Madden Dam (UMMZ 76015, two specimens).

Ameiva cf. ameiva (5).—VENEZUELA: Bolı´var: Paulo, Mt. Roraima, AMNH 36328, 30–33.