Anolis anatoloros, Ugueto & Barros & Sánchez- Pacheco & García-Pérez, 2007

Ugueto, Gabriel N., Barros, Gilson Rivas Fuenmayor Tito, Sánchez- Pacheco, Santiago J. & García-Pérez, Juan E., 2007, A revision of the Venezuelan Anoles I: A new Anolis species from the Andes of Venezuela with the redescription of Anolis jacare Boulenger 1903 (Reptilia: Polychrotidae) and the clarification of the status of Anolis nigropunctatus Williams 1974, Zootaxa 1501 (1), pp. 1-30 : 17-28

Anolis anatoloros sp. nov.

( Figs. 4–6 )

Holotype.— MHNLS 17872 ; an adult male ( Fig. 5 ), collected 10 June 2006 by J. M. Guayasamín and C. L. Barrio-Amorós on the flat leaf of a bush, 1.5 m aboveground on San Isidro, Barinas (8°50’83”N; 70°34’23”W), 1480 m, Venezuela.

Paratypes.— MHNLS 17871 ; an adult female , MBLUZ 896 , MHNLS 17874 ; both adult males; collected with same data as holotype . EBRG 4587 ; a subadult female, collected 14 June 2005 on the same locality as holotype by E. Camargo , R. Rivero and C. L. Barrio-Amorós . MCNG 2500 ; an adult female, collected 16 September 1999 on road between Boconó and La Vega de Guaramacal, Guaramacal National Park, Trujillo, 1900 m, Venezuela . MCNG 2501 ; an adult male, collected 20 March 2005 on the same locality as the previous paratype but at 2000 m . MCNG 2502 ; an adult male, collected 13 November 1995 on road between Chabasquén and Córdova , Portuguesa, 1400 m, Venezuela .

Diagnosis.—A medium-sized anole with males reaching maximum SVL of 68 mm and females 66 mm. Head with moderately large, smooth or weakly keeled scales; supraocular semicircles usually distinct and separated by 0–1 scales; a series of 1–3 bordering the upper margin of the supraciliar series (first one or occasionally second one conspicuously large); 0–1 scales between interparietal and supraorbital semicircles; scales around interparietal large, including those posterior to it. Dorsal scales small, weakly keeled; an enlarged double row is present in males; a tall nuchal fold is present in males, which continues along the vertebral region of dorsum; ventrals larger than dorsals, smooth, squarish or roundish. Tibia 0.20–0.22 times SVL; digital expansions well developed and distinct. Males yellow green with some dark reticulations; females with a light ver- tebral stripe bordered by very dark stripes or with conspicuous vertebral dark bands that extend considerably onto the flanks. Dewlap in males turquoise blue becoming green peripherally with greenish scales; in female dark brown with black spots and pale scales. Known from the eastern slopes of the Venezuelan Andes in San Isidro, Barinas, Guaramacal, Trujillo and Chabasquén, Portuguesa.

Comparisons.— Anolis anatoloros differs from other anoles in the same ways A. jacare does and thus, it has been carefully compared with it. Anolis species are conservative in squamation and morphology and these two are no exception. However, both differ in important details of lepidosis and coloration. Major differences among the two species are summarized in table 2. There is a series of 1–3 scales bordering the inner margin of the supraciliar scales in Anolis anatoloros . The first scale of this series is conspicuously larger than the rest in all specimens except EBRG 4587, in which the second scale of the series is noticeably larger (first scale minute). In contrast, A. jacare has three or more small, somewhat squarish scales above the supraciliar series that are sometimes almost as small as those immediately next to them on the supraocular region. In some specimens of A. jacare , some of the scales above the supraciliar series may be visibly larger than the rest, but these are never the first or second scales and are never as large as those shown by A. anatoloros .

In Anolis anatoloros , the size of the scales around the interparietal varies from slightly smaller to subequal or larger than that scale and are always larger than those between nostrils. The scales bordering the posterior margin of the interparietal are subequal or only slightly smaller than those located laterally and anteriorly. Typically, in A. jacare , the scales around the interparietal are much smaller, especially those bordering its pos- terior margin. The latter are smaller than the scales located laterally and anterior to the interparietal. In a few specimens the interparietal is very small and thus, the scales around it look sometimes subequal but are still proportionally much smaller than those present in A. anatoloros . The scales around the interparietal are usually smaller or subequal to those between nostrils (only larger in three specimens). One specimen (EBRG 163) has a very large interparietal and large scales around it, however, this condition is probably aberrant; in other scale characters this specimen agrees with typical A. jacare .

In seven out of eight Anolis anatoloros , four suboculars where in direct contact with the supralabials and in the right side of MHNLS 17872 and the left of EBRG 4587 five scales where in distinct contact. Nevertheless only three scales where in contact on the left side of MBLUZ 896 and on the right side of EBRG 4587. In A. jacare there are usually two or three subocular scales in contact with the supralabials, only four specimens out of 34 had four suboculars in contact with supralabials in at least one of the sides of the head. In neither species are the tail verticils conspicuous, but they are usually visible on the second third of the tail. In the middle portion, A. anatoloros shows six dorsal and four ventrals scales whereas A. jacare normally shows five or four dorsals and three or four ventrals. The anterior verticils of A. anatoloros are composed by 7–9 dorsal scales whereas A. jacare shows 5–7. Also, the nuchal fold of males is conspicuously taller in males A. anatoloros . Unfortunately, this character is best observed in males in life as nuchal and dorsal folds are noticeably reduced or lost during preservation in many specimens.

Both species exhibit marked sexual dimorphism in coloration. Although males show a somewhat similar dorsal pattern (dark reticulations, sometimes forming crossbars), the ground color in life of male Anolis jacare tends to always present a distinctive brown or yellow cast whereas A. anatoloros appears to be consistently greener (compare fig 3 with fig 6). The supralabials below the eye are whitish, sometimes with a yellow tint in A. jacare and yellow green in A. anatoloros ; this characteristic seems to remain constant even during metachrosis. In all male A. jacare observed in life from Mérida, there was a conspicuous whitish or light, oblique stripe on the shoulder right below the scapular spot. This stripe was not visible in the five male A. anatoloros observed in life. As is common in closely related anole species, one of the most conspicuous differences is the dewlap coloration of males. While male A. jacare consistently presented a yellow-ochre or orange-yellow dewlap with white, pale green or dark brown scales, A. anatoloros showed a turquoise green dewlap (bluer towards the base and anteriorly) with greenish scales. This difference is also visible in preservative, all male A. jacare examined showed a uniformly pale dewlap with either pale or dark scales. In contrast, the three A. anatoloros males had a gray or blue-gray dewlap with similarly colored or paler scales that was decidedly darker than the dewlap of male jacare .

Contrary to males, females are much more similar in coloration and are more difficult to differentiate. Females of both species exhibit two main color morphs, one striped and one banded. In female A. anatoloros MHNLS 17871, the banded pattern in life appeared much bolder than in any of the female A. jacare observed. In the latter species, the dorsal banding extended only tenuously laterally (frequently breaking into dots) while in the female A. anatoloros the banding extended conspicuously laterally forming a distinct barred pattern (fig 3, fig 6). The striped pattern was observed in the other two female anatoloros . Although this morph is very similar to the striped morph of female A. jacare , here also the pattern appears to be slightly bolder in A. anatoloros , especially the dorsolateral dark stripes and spots on flanks. As a general rule, the dewlaps of A. anatoloros females were darker. The three females known showed a dark brown dewlap with inconspicuous darker spots and pale scales. Female A. jacare observed showed a yellowish, greenish or brownish dewlap with conspicuous dark spots and pale scales. However, difference in dewlap coloration is not as evident in preserved females.

Description.—The holotype (MHNLS 17872) is the largest male with SVL 68 mm; maximum total length in males 210 mm (MBLUZ 896); maximum SVL in females 66 mm (MCNG 2500); maximum total length in females 213 mm (MCNG 2500). Head 0.26–0.29 (0.27 ± 0.012, n = 8) times SVL, 1.70–1.79 (1.76 ± 0.023, n = 8) times as long as wide, and 1.03–1.19 (1.10 ± 0.066, n = 8) times as wide as high. Snout 0.55–0.61 (0.57 ± 0.017, n = 8) times the head length, moderately long, moderately blunt or rounded and not swollen. Frontal region barely to moderately depressed; frontal ridges barely to moderately distinct posteriorly, only barely anteriorly; supraorbital ridges from inconspicuous to moderately distinct; occipital ridges from barely distinct to noticeable; parietal region slightly to moderately depressed anterior to the occiput, (the latter may be conspicuously swollen in males, not swollen in females). Occipital region somewhat swollen or not. Intertemporal region slightly protuberant. Neck slightly narrower than head and body. Body and tail slightly compressed, distance between axilla and groin 0.37–0.42 (0.39 ± 0.018, n = 7) times SVL. Limbs well developed, hind limbs 0.68–0.72 (0.69 ± 0.016, n = 6) times SVL, tibia 0.20–0.22 (0.21 ± 0.006, n = 8) times SVL. Tail 2.17– 2.26 times SVL.

Rostral wider than high, higher medially with moderately to distinctly indented posterior margin; just barely visible from above and not projecting forward. Postrostrals 6–8, irregularly polygonal and smooth or very weakly keeled. Circumnasal with its antero-inferior or inferior portion in distinct contact with rostral or completely separated by one scale. Scales on snout, between nostrils, not or only slightly swollen, distinctly longer than wide, some slightly narrow, smooth or with a few scales weakly keeled, and juxtaposed; middorsal scales between nostrils larger than those lateral to them; all smaller to those on and posterior to the frontal region, where scales are larger, polygonal, flat, smooth or slightly rugose, and juxtaposed. Scales on the center and posterior to the frontal depression smaller. 6–9 scales between circumnasals; 7–10 scales across snout at level of second canthal. Canthus rostralis well defined, mostly straight or slightly curved anteriorly. Five or six enlarged canthals slightly to distinctly keeled, increasing in size posteriorly; first canthal very small on both sides of MHNLS 17874, MCNG 2500, MCNG 2502 and on the left side of MBLUZ 896; 2 nd or 3 rd canthal longer than the rest (on the left side of MBLUZ 896, 3 rd and 4 th subequal). 17–22 scales between the rostral and the interparietal scales. Supraorbital semicircles moderately distinct or barely so, separated by one scale or in contact. Second scale the largest of the supraorbital series or 5 th in the left side and 4 th on the right side of MBLUZ 896 an left side of MCNG 2500 or 3 rd on the left side and 4 th on the right side of MHNLS 17871; first one in distinct contact or separated from the first canthal by one scale. Supraocular region with a group of large, smooth or weakly keeled, juxtaposed, irregularly polygonal and flat or slightly swollen scales, larger near the supraocular semicircles; elsewhere with granules (particularly infero-posteriorly); the group of large supraocular scales is completely or almost completely separated from the supraorbital semicircles by smaller scales (though one or two of the larger scales may be in narrow contact). There is a series of 1–3 conspicuously large, scales bordering the upper margin of the supracilar scale; in almost all specimens the first scale is considerably large and elongate, followed by one or two much smaller and square ones; only in EBRG 2587 is the first scale small and squarish and then followed by a large, elongate one. Supraciliar series formed by a very long scale that reaches almost the half of the orbit then followed by granules. Interparietal irregularly shaped, roughly hexagonal or somewhat elliptical; 0.09–0.12 (0.10 ± 0.012, n = 6) times the head length, 1.69–2.34 (2.02 ± 0.249, n= 6) times as long as wide; slightly larger, roughly equal or slightly smaller than the ear-opening in size. Interparietal separated from supraorbital semicircles by 0–1 scales, only in MBLUZ 896 in very narrow contact; moderately larger than the surrounding scales and with a noticeable but small “pineal organ” (somewhat obscured in 17871). Parietal region covered with flat, smooth or slightly rugose, irregularly polygonal, more or less homogeneously-sized scales, which decrease in size noticeably only on the parietal depression; always larger than those between nostrils and slightly smaller or subequal to those on frontal region (which are in turn frequently larger than those on the center of the frontal depression). Scales immediately next to interparietal, large, only slightly smaller than that scale (those posterior to it slightly smaller or subequal to those located laterally and anterorly); in MCNG 2502 those anterior to the interparietal larger than that scale. 7–10 (most specimens with seven) scales around interparietal. Occipital and supratemporal region covered by small, juxtaposed scales, slightly larger anteriorly. Loreal scales irregularly polygonal, longer than wide, and more or less in longitudinal rows; all mostly subequal or lowest one (bordering supralabials) only slightly larger; flat, smooth or slightly rugose with no median keel distinct. The uppermost row (below can- thals) of MHNLS 17871, MCNG 2500, MCNG 2502 and EBRG 4587 is narrower than the rest, slightly swollen and has a distinct median keel. 3–7 scales in a vertical row at level of second canthal. Suboculars 5–6, moderately distinct, flat smooth or weakly keeled; anterior ones usually wider than high, posterior ones the opposite, but most subequal in size; four in contact with supralabials (five on the left side of EBRG 2587 and on the right side of MHNLS 17872, three on the left side of MBLUZ 896 and on the right side of EBRG 2587). Eyelids covered with minute granules, with two rows of larger scales bordering the rim. Supralabials 6–8 to below center of eye. Temporal region with small, granular scales, separated from eyelids by a single or double row of slightly larger scales, and from parietal as well as supratemporal regions by a double intertemporal row of moderately enlarged scales. Ear-opening small, vertically oval, its lower margin roughly at the same level of the commissure of mouth; with smooth margins (slightly denticulate anterior one in MHNLS 17872) and short auditory meatus.

Mental large, roughly crescent-shaped or somewhat trapezoidal, indented by adjacent scales; semidivided by an usually weak median cleft, which does not continue as a short midventral sulcus; medial line bordered on each side by first infralabial, one sublabial, and one or two, small, median scales. Infralabials 5–8 to below the center of the eye. Chin with a row of large, weakly keeled sublabials, 2–4 in contact with infralabials; medially, scales are distinctly smaller, decreasing in size towards the midventral line and posteriad. Scales on throat slightly larger, especially towards the dewlap. Dewlap in males large, reaching to a point between axilla and midbelly; laterally with longitudinal, widely separated rows of single, elongate, laterally flattened and narrow scales which are larger than those on the sides of the neck; along rim scales wider, smooth, round and distinctly imbricate; lateral dewlap scales longer but narrower than ventrals, those along rim subequal or larger than ventrals. In females, dewlap small, just reaching the insertion of the arms; laterally with longitudinal, narrowly separated rows of single, somewhat elongate, compressed and smooth scales that are larger than those on the sides of the neck and smaller than ventrals, the scales along the rim similar but imbricate, also smaller than ventrals.

Scales on nape small, slightly swollen or not, subimbricate or juxtaposed. A double middorsal rows barely to distinctly enlarged in males, starting on nape and extending posteriorly. A vertebral fold is present on the males from occiput to shoulders then continuing as a low, very inconspicuous fold along the vertebral region on dorsum and disappearing on sacrum. The vertebral crest was considerably more distinct in life. Dorsals somewhat swollen or not (those on vertebral region flatter), vertebral rows usually subimbricate, other rows frequently juxtaposed, all weakly keeled or smooth. A double row of enlarged vertebral scales are distinct on dorsum (barely on EBRG 4587 and MCNG 2500). 75–92 (82.1 ± 7.0, n = 6) dorsal scales in a vertebral line between axilla and groin. Scales on flanks similar, mostly juxtaposed and in males usually somewhat conical and swollen (flatter in females), changing gradually toward ventrals, which are larger, roughly quadrangular or somewhat oval, subimbricate or imbricate and smooth. 62–77 (70.3 ± 5.0, n = 6) ventral scales in a midventral line between axilla and groin. 116–143 (127.6 ± 7.8, n = 6) scales around midbody. Preanal plate with small, juxtaposed to subimbricate, relatively uniform scales. Two very distinct, very large postanal scales are present in males MBLUZ 896 and 17874. Male MHNLS 17872 with a conspicuous, horizontally elongate and large scale that covers most of the postanal area followed by a similar but smaller one. Enlarged postanal scales absent in females.

Base of tail with numerous keeled, subimbricate scales dorsally and laterally; ventrally, scales smooth (occasionally some slightly keeled), increasing in size slightly towards midventral line, flat and subimbricate. There is a double middorsal enlarged row (it is still a singular, although a double row) (hardly visible in EBRG 4587). Tail distally with large, polygonal, imbricate scales, in a few longitudinal rows; the pair of ventral, and the adjacent ventrolateral rows larger; all scales with distinct keels, which form longitudinal ridges. Verticils mostly distinct on the mid portion of the tail; when visible anteriorly with 7–9 dorsals and five or four ventrals. Middle verticils with six dorsals and four ventral scales. Tip of the tail with rather indistinct verticils, when visible with five or four dorsals and three ventrals.

Larger scales on forelimbs slightly larger than dorsals, keeled, imbricate or subimbricate, larger on anterior aspect of forearm, elbow and around wrist; ventral aspect of forelimbs with smooth, smaller or subequal scales. Hind limbs with large (subequal to ventrals), weakly keeled, imbricate scales on their anterior aspect, grading to smaller dorsally and almost granular on the posterior aspect of lower legs; ventral aspect of thighs with small and smooth scales, slightly larger on the lower legs, particularly anteriorly and posteriorly. Digital expansions very well developed; 18–22 (20.4 ± 1.5, n = 12, 6 specimens) lamellae under the fourth finger to end of digital expansion; 30–36 (33.8 ± 1.7, n = 11, 6 specimens) lamellae under fourth toe to end of digital expansion, 20–24 (21.7 ± 1.3, n = 10, 5 specimens) between the II and III phalanges.

Meristic data of holotype.–SVL 68 mm, tail length 116.5 mm (broken tip), head length 19.88 mm, head width 11.11 mm. head height 9.55 mm, snout length 11.23 mm, axilla–groin length 28.59 mm, hind limb length 46.56 mm, tibia length 14.36 mm, interparietal length 2.32 mm, interparietal height 1.37 mm, ear opening length 0.88 mm, ear opening height 1.66 mm. Dorsal scales in a longitudinal row between axilla and groin 79, ventral scales in a longitudinal row between axilla and groin 74, scales around midbody 131, lamellae under the end of digital expansion of fourth finger 19 (left) 18 (right), lamellae under the end of digital expansion of fourth toe 30 (left) 31 (right), lamellae under the second and third phalanges of fourth toe 20 on both sides. Scutellation as depicted in fig.5.

Coloration in life.—This species is similar in coloration to Anolis jacare and, as that species, shows marked sexual dichromatism. Males: Yellowish green, pea green or red-brown above, sometimes slightly greener on shoulders and neck. There is a conspicuous dorsal pattern formed by dark reticulations, similar to those of male A. jacare . The reticulations are mostly dark gray or brown on body but on the sides of the neck and anterior flanks they change their color to dark or turquoise green. In MCNG 2501 the reticulations are grass green. The vertebral region is pale brown and interrupting this area are six or ten dark tan or gray-brown spots with narrow blackish margins. These spots may extend laterally as barely visible or distinct (intensity variable depending on metachrosis), tan or dark tan, oblique bars. These bars tend to be more or are only apparent on the flanks and have black and light (less distinct and not always visible) dots along their length. The bars are not apparent in MCNG 2501. Vertebrally, the tan spots may have a light colored spot in their center. The head is brownish-green or red-brown with numerous dark or green reticulations most apparent on the interocular and parietal regions as well as along the canthus rostralis and in between nostrils. A moderately distinct or ill-defined postocular stripe (tan with a few light dots within it and having narrow black margins) extends from the postero-inferior corner of the eye to above the ear opening where it fades into the dorsal pattern. Occasionally the postocular stripe is only indicated by its upper margin. Labials anteriorly greenish with dark spots or brownish with green spots, lighter yellow-green below the eye and continuing as an irregular stripe of the same color to the upper margin of the ear opening and sides of the neck on several specimens. A blackish or greenish stripe is present below the eye occasionally extending but very tenuously on the supralabials directly below it. Below the lateral yellow-green stripe on the sides of the neck there is a large, dark tan and oblique spot with conspicuous black margins and two light spots within it of the background color. The visibility of this spot varies during metachrosis, in lighter phases the spot almost disappears being just indicated by reticulations that become wider on the area but do not form a conspicuous spot. In one of the males, this spot was just indicated by two irregularly shaped dark tan and black spots (anterior one the largest). When the spot is visible, the reticulations around it are of a turquoise green color. Bright yellow spots are also visible on the shoulders and on the adjacent portions of the neck. Eyelids yellow-green. Limbs of the same background color of the body with dark reticulations that may form narrow but well-defined crossbars. Those on thighs, wider and usually better defined; those on forelimbs sometimes just reduced to numerous dark dots. Digits with dark rings. Tail yellow-green, with a yellow-brown tint on its base dorsally and on sacrum. Vertebral region of the tail slightly lighter than the rest anteriorly. Conspicuous dark brown to blackish rings present becoming wider posteriorly. Some of these rings occasionally meet laterally. Chin pale greenish, yellow green or pale turquoise just with a few indistinct dark or green dots. Chest and abdomen mostly immaculate. Dewlap pale turquoise in the center and anteriorly fading into greenish peripherally or completely pale turquoise (darker anteriorly). Scales on dewlap always greenish sometimes with slightly darker spots.

Females: As in Anolis jacare , females of this species presented two distinct color morphs. Striped morph:

MCNG 2500 was yellowish-green above with a pale greenish-white longitudinal, vertebral stripe from the occipital area to the dorsal surface on the base of the tail, where it faded. This stripe was margined by one, sinuous dark reddish-brown stripe that started on the beginning of the neck and extended onto the base of the tail. This stripe had a sinuous lower margin and presented three short lateral projections, one on the scapular region, one on mid-body and one on the posterior dorsum. Below the two posteriormost lateral projections there was a large, vertically oval, dark reddish-brown spot. Both lateral spots were formed by dark, connecting reticulations and were very irregular in shape. Most dorsal surfaces, except on the pale, vertebral stripe, covered with dark vermiculations and reticulations; in other parts of the flanks and dorsum, these dark markings connected to form irregular markings and some aligned rather longitudinally, forming narrow stripes right along the pale vertebral stripe. Some scattered, pale but very inconspicuous spots were also present on the sides of the body, between the lateral dark spots and were also aligned vertically (except one that was present right beyond the scapular region, posterior to the arm). A reddish-brown, slightly obliquely elongated spot was distinct on the sides of the neck; it was margined with black and this color invaded the spot anteriorly. Head yellowish-green with some inconspicuous blackish vermiculations on the snout. The supraocular region as well as the tip of the snout (but more lightly) had a reddish-brown tint and there were two parallel, narrow, black interocular lines; a similar occipital one radiated from the upper-posterior corner of the eye. Sides of head slightly lighter with blackish dots on loreal region and labials. A postocular stripe (formed by dark reticulations) was distinct; it extended to the sides of the neck and then broke into reticulations. A pale, irregularlymargined, unspotted stripe extended from below the eye, passing through the upper margin of the ear opening extending laterally to the large spot on each side of the neck. Some blackish reticulations were present below this pale stripe, particularly evident below the ear opening.

Arms yellowish-green with a few scattered, pale yellowish spots between narrow crossbars formed by black dots and reticulations. Legs yellowish-green with light yellowish, round spots and some distinct but moderately narrow, blackish crossbars. All digits with blackish crossbars or spots. Tail continuing the dorsal pattern on the base then changing to brown in background color with a few irregular, lateral blackish areas, more evident antero-laterally.

The subadult female EBRG 4587 was reddish brown on the neck and most of the dorsum. There was a slight suffusion of green on the scapular region and the posterior portion of the body as well as the sacrum was bright green. There was a light grayish-white vertebral stripe that extended from occiput to the dorsal aspect of the tail; it was slightly narrower on occiput and neck and its widest point was on sacrum and anterior tail. At each side of this stripe was a slightly wider, dark brown stripe that extended from the insertion of the neck to the tail on its anterior third, vanishing posteriorly. Black dots and dark smudges were present on the flanks, more or less disposed in transversal rings. The head was dark brown from the tip of the snout to the right posterior to the interocular region, changing to reddish brown posterior to that. A dark brown postocular stripe was distinct from the postero-inferior corner of the eye to the insertion of the neck where it merged with the dark dorsal stripe. Less well defined parietal and interocular stripes were also present. Eyelids yellowish. Sides of the head lighter, with dark spots anteriorly, mostly immaculate pale green below the eye (except for an ill-defined stripe that extended from below the eye to supralabials).

Forelimbs with a mixture of reddish brown and green and dark reticulations forming ill-defined crossbars on lower arms. Legs green with dark brown dots forming more or less well-defined rings. Digits with dark rings. Tail continued the body pattern anteriorly becoming more uniformly brown towards its tip. Chin grayish with dark brown dots, becoming slightly yellowish on throat. Abdomen whitish with some indistinct dark dots. Dewlap grayish with blackish spots, appearing completely dark with pale scales.

Banded morph: The female MHNLS 17871 was very boldly patterned. It was bright green with six dark tan, broad, oblique and very distinct crossbars (first one reduced to a vertebral spot). These bands were slightly wider vertebrally and had moderately distinct blackish margins that broke into spots laterally. A few inconspicuous dark reticulations and yellowish smudges were also present between the bands. These bands interrupted a pale gray-brown vertebral stripe that extended from occiput to the dorsal aspect of the tail. The crossbars constricted vertebrally. An oval dark tan spot on the side of the neck was extended dorsally where it connected with the second dorsal crossbar. Another dark tan spot was present on the posterior part of the scapular region and a similar but much smaller one was present on the insertion of the arm. The head was green as the body with an inconspicuous dark reticulation. There was a more or less defined dark tan postocular band that extended from the postero-inferior corner of the eye to above the ear opening and from there, dorsally where it became almost invisible and it merged with the first dorsal band on neck. Less defined parietal, interocular and canthal dark stripes were also present. Eyelids yellow-green. Labials greenish with inconspicuous dark spots anteriorly, becoming mostly immaculate yellow-green below the eye and temporal region. A few reddish areas around the ear opening.

Limbs green with well-defined dark tan bands. These bands showed narrow black margins. The digits had dark rings. Tail continuing the body pattern anteriorly changing posteriorly to gray-brown and with wider dark rings. The rings faded towards the tail tip. Chin greenish with inconspicuous dark smudges. Dewlap dark brownish with black spots, appearing completely dark with pale green scales.

Coloration in preservative.–The marked sexual dichromatism observed in life is clearly visible in the type series after preservation. Males: The holotype and the other two males show a very similar coloration. Pale brown or pale gray-brown with five (holotype) or six (male paratypes) dark brown, slightly oblique crossbars on dorsum that extend well onto flanks, reaching the paraventral area. These bands have conspicuous dark reticulations within them. A small similar band is restricted to the vertebral region on the neck. These bands are just indicated vertebrally and much less distinct on the flanks of the male paratypes (particularly indistinct in MHNLS 17874). The three males show a conspicuous dark reticulated pattern on flanks and dorsum. The reticulations and vermiculations also break into dots, mostly towards the vertebral region. On the two male paratypes the areas between the transversal bands on the flanks are lighter (pale gray MBLUZ 896 or pale greenish gray 17874). These light areas are not evident on the holotype. Light areas are also distinct as squarish spots between the dark bands on vertebral region of MBLUZ 896. A large, oval, slightly obliquely dark brown spot is present on the sides of the neck and scapular region of the three males. This spot has a few round spots of the background color inside it. It is more conspicuous in the holotype and in MBLUZ 896. Head of the same color as body or slightly browner, with inconspicuous dark reticulations. An indistinct postocular band formed by dark reticulations is most visible in MBLUZ 896, but very indistinct in the other two males. Supralabials whitish (slightly yellowish below the eye in holotype), extending as a narrow band to above the upper margin of ear opening and in MHNLS 17874 (and to less extent on MBLUZ 896) extending obliquely on scapular region above the lateral oval spot. Small dark dots and spots are present on the anterior and mid supralabials, rest of the area immaculate.

Forelimbs with dark, narrow rings (holotype), conspicuous and moderately broad bands with darker margins ( MHNLS 17874 ) or inconspicuous, narrow rings ( MBLUZ 896 ). Hindlimbs with conspicuous dark brown bands (again more conspicuous in MHNLS 17874 ), two on thighs and three on tibia. A few conspicuous whitish dots are present on the posterior end of tibia on the holotype and MHNLS 17874 . Digits with dark rings. Tail with conspicuous dark rings that become wider and less distinct towards the tip. Chin whitish (more distinct in holotype and MBLUZ 896 ) changing into brownish on throat and with numerous dark gray (holotype) or brown and gray ( MHNLS 17874 ) or dark brown ( MBLUZ 896 ) dots. Chest and abdomen pale brownish, whitish on paraventral region on the holotype and MBLUZ 896 , or chest whitish and abdomen yellowish brown (darker midventrally) as on MHNLS 17874 ; with some indistinct dots centrally that become distinct laterally (dark in holotype; light in MHNLS 17874 ; dark on chest, light on abdomen of MBLUZ 896 ). Ventral aspect of MBLUZ 896 and holotype’s limbs, whitish and mostly immaculate (forelimbs) or with dark reticulations (hindlimbs). On MHNLS 17874 , forelimbs mostly immaculate, hindlimbs with very minute gray-brown and whitish dots, forming an inconspicuous salt-and-pepper appearance. Ventral aspect of MBLUZ 896 and holotype’s tail brownish midventrally, whitish paraventrally, rings more apparent towards mid portion. MHNLS 17874 with a light brown tail, becoming darker posteriorly, rings becoming apparent towards mid portion. Dewlap bluish-gray with brownish scales in the holotype; or gray (darker anteriorly) with brownish lateral scales and whitish along the rim in MHNLS 17874 ; or gray with dark gray scales laterally and brown scales along the rim in MBLUZ 896 .

Females: The two females show two different color patterns, similar to the two morphs present in Anolis jacare . MHNLS 17872: Gray brown, slightly lighter vertebrally, with five, conspicuous dark brown transversal bands that extend conspicuously on the flanks. These bands distinctly constrict on the vertebral region, forming distinct “butterfly-wings” figures when seen from above. Black margins of these bands are more conspicuous along the vertebral region. The anterior band, splits laterally into two oblique stripes; the anterior one extends above the shoulder to the sides of the neck; the posterior one extends to the post-axillar region. The vertebral region next to the dorsal bands is much lighter. Pairs of dark spots (most round, a few irregular) are present vertebrally between the dark transversal bands. The last two dorsal bands are closer together than the others. A dark brown spot is present right after the insertion of the arm on the upper axillar area. Head brown with indistinct dark spots; a more or less conspicuous dark interocular band is visible as well as an irregular, narrow band across anterior snout. A dark brown, irregular postocular band extends posteriorly and merges along the vertebral area of neck, forming a band very similar to those on body. Supralabials anteriorly pale brownish with dark crossbars, changing to whitish below the eye with dark spots; the whitish color extends only briefly and irregularly to the upper margin of the ear opening. Limbs with conspicuous dark bands and a few, inconspicuous light dots (more apparent on tibiae). Tail continuing the dorsal color anteriorly, rings becoming wider and less conspicuous towards its tip. Chin grayish, becoming darker (and browner posteriorly) and dark dots. Chest brownish, becoming slightly lighter on abdomen (but still darker midventrally) and with dark dots laterally.

Limbs with dark reticulations; thighs with light and dark spots. Tail brownish with dark, irregular dots and spots, becoming like dorsal aspect of tail towards the mid portion. Dewlap very dark brown with very indistinct darker dots, appearing very dark altogether with brownish scales (much paler than the skin color).

MCNG 2500: Pale gray above with a pale gray longitudinal and vertebral stripe that starts on the occipital area and extends on the dorsal surface of the base of the tail, fading afterwards. This stripe is of the same width throughout its length, about 6 scales wide. Bordering this stripe there is one dark brown area that starts on the beginning of the neck and extends onto the base of the tail. These brown areas are rather diffuse and are wider (extending more laterally) on the sides of the neck. Very inconspicuous and tenuous brownish bands are only barely apparent on the flanks; these connect with the dark brown areas that border the pale vertebral stripe. All body and neck, except on the pale gray stripe but still visible on the dark brown areas, showing blackish reticulations and reticulations; some of these connect and form rather irregular (sometimes oblique) markings and some may tend to form narrow longitudinal stripes right along the pale vertebral stripe. A brown, horizontally or obliquely elongated, brown spot is present on the sides of the neck. This spot is narrowly margined with black and shows some black reticulations inside the brown area. Head olivaceous-gray with some inconspicuous blackish reticulations (but far less defined and noticeable than the ones present on the flanks). The supraocular region yellowish-brown with a dark narrow line in the middle of the area on each side. Sides of the head slightly lighter only with barely noticeable dark dots. The temporal region is brown; there is a postocular stripe (formed by dark reticulations) that merges with the dark areas bordering the pale vertebral stripes and the body reticulations. A moderately distinct, unspotted area extends from the upper margin of the ear opening extending laterally to above the large spot on each side of the neck.

Arms with a few scattered, pale yellowish spots and black dots, some of them merging and forming narrow crossbars (most notorious one near wrist). Legs gray with some conspicuous, light buff-colored spots (more conspicuous one on thighs near the insertion of the limb) and some blackish reticulations, usually forming crossbars. All digits with blackish crossbars. Tail continuing the dorsal pattern on the base then changing to brown in background color with a few irregular blackish areas, but tip uniformly brown. Chin pale gray laterally, becoming slightly brownish centrally and on throat, with scattered blackish dots (more apparent towards the sides of the neck). Abdomen and ventral aspect of limbs and tail, grayish-white with only a few blackish dots towards the sides of the body and a few indistinct on the ventral aspect of the thighs. Dewlap blackish, very dark with pale gray scales (most with a slight brownish tint).

EBRG 4587: Gray brown with a conspicuous whitish vertebral stripe that starts on the occiput and extends onto the dorsal aspect of the tail, where it fades into the background color. The stripe stays mostly the same width along the dorsum (7–9 scales wide). Very conspicuous and broad, dark brown stripes margin the vertebral one at each side. These stripes are wider and have a more or less irregular lower margin; they extend from the middle neck to the middle portion of the tail where they fade; they are wider along the middle flanks. Narrow, light longitudinal, short lines are present along the vertebral light stripe on the dark areas. Some dark, irregular dark smudges and spots are present on the flanks, aligned in transversal bands; these bands merge with the longitudinal dark dorsolateral stripes. Numerous light round spots and dots are present along the flanks. Head gray brown with indistinct dark dots and reticulations; there is a dark postocular stripe that extends somewhat irregularly towards the dorsal aspect of the neck where it merges with the dark dorsolateral area. Supralabials light with dark bands anteriorly, becoming whitish with dark spots below the eye; the whitish color extends irregularly to the upper margin of the ear opening and obliquely on the sides of the neck. A dark brown irregular area is present below this light region.

Forelimbs with dark and light dots, the dark ones tending to form crossbars on lower arms. Hindlimbs with alternate light and dark rings formed by small spots on a gray brown background. Digits with dark rings. Tail continuing the dorsal coloration anteriorly and on its mid portion where it fades into a pale gray color. There are a few indications of brown rings on the posterior third of the tail. Chin whitish becoming brownish (especially posteriorly) with dark gray (anteriorly) and dark brown (posteriorly) dots. Chest brownish, becoming paler on abdomen (slightly darker midventrally); mostly immaculate except for a few dark dots laterally. Ventral aspect of limbs with whitish and a few dark brown dots. Ventral aspect of the tail brownish with some dark dots on the base; dark rings become apparent towards the mid portion; ventral coloration between the rings white, contrasting with the gray color of dorsal surfaces. Dewlap as in MHNLS 17872.

Habitat and natural history.–All but one of the eight specimens were collected in moderately intervened premontane and montane wet forests between 1480 and 2000 m. MCNG 2502 was collected at midday perched on a fence post along cattle pasture at 1400 m. Four specimens (three males and one female, MHNLS 17871 - 17872,17874 , MBLUZ 896 ) were captured the same night (between the 19:00 and 20:00 h) while each was sleeping on the flat leaves of medium-sized bushes, approximately 1.5 m aboveground. MCNG 2500 was collected while it jumped on the ground of a forest clearing on direct sunlight at 13:00 h. No other anoles species have been collected at any of the localities were Anolis anatoloros has been found. The only other reptile observed at the type locality was the colubrid snake Liophis epinephelus (C.L. Barrio-Amorós, pers. comm.) .

Distribution and Material Examined.– Anolis anatoloros is known only from the eastern slopes of the Venezuelan Andes in the states of Barinas, Portuguesa and southeastern Trujillo ( Fig. 4 ). It likely also occurs on eastern Mérida.

Etymology.—The name of the new species is a noun in apposition derived from the Greek noun anatoli meaning east and the Greek noun oros meaning mountain. It alludes to the presence of this species on the eastern slopes of the Venezuelan Andes.

Figures

FIGURE 4. Distribution of Anolis jacare (closed circles) and Anolis anatoloros (open circles) in Venezuela and Colombia. 1) Estoraques, 2) Sisavita; 3) El Diamante; 4) Toledo; 5) Villa Páez, Betania; 6) Paramo La Negra; 7) Doradas; 8) La Macana; 9) La Pedregosa; 10) Road between Mérida and El Morro; 11) Localities in Mérida City; 12) La Mucuy; 13) Quebrada Honda; 14) Chabasquén; 15) Guaramacal; 16) San Isidro.

FIGURE 5. Anolis anatoloros, adult male, holotype MHNLS 17872. a, b, c: dorsal, lateral and ventral views of the head; d: ventral view of left foot; e: dorsum; f: venter. All scale bars shown in the drawing represent 1 mm.

FIGURE 6. Anolis anatoloros, coloration in life. a: Adult male MHNLS 17872, San Isidro, Barinas; b: Adult male MCNG 2501, Guaramacal, Trujillo; c: Adult male MHNLS 17872 dewlap; d: Adult female MHNLS 17871 dewlap, San Isidro, Barinas; e: Adult female MCNG 2500 striped morph, Guaramacal, Trujillo; f: Adult female MHNLS 17871 banded morph; g: Subadult female EBRG 4587 striped morph, San Isidro, Barinas.