Anolis gaigei Ruthven, 1916

Köhler, Gunther, Batista, Abel, Vesely, Milan, Ponce, Marcos, Carrizo, Arcadio & Lotzkat, Sebastian, 2012, Evidence for the recognition of two species of Anolis formerly referred to as A. tropidogaster (Squamata: Dactyloidae), Zootaxa 3348, pp. 1-23 : 14-18

Anolis gaigei Ruthven, 1916

Figures 1 b; 3c, d; 4 a, c, e, g; 6 d, e, f; 7 c, d; 11

Anolis gaigei Ruthven 1916:6 ; holotype (UMMZ 48304) from “San Lorenzo, Santa Marta Mountains, Colombia, elevation of 2, 700 ft.” Ruthven (1922).

Anolis limifrons: Martínez Cortés & Rodriguez (2005 ; in part.: Fig. 22B)

Anolis polylepis: Martínez Cortés and Rodriguez (2003, 2005 ), Rodríguez et al. (2004), Ibáñez (2006; in part.).

Anolis tropidogaster: Swanson (1945) , Evans (1947), Donoso-Barros (1968), Myers and Rand (1969), Peters and Donoso-Barros (1970; in part.), Campbell (1971), Sexton et al. (1964, 1971), Kiester (1979), Kourany & Telford (1981), Rand and Myers (1990), Quintero and Cambra (1993), Auth (1994; in part.), Williams et al. (1995; in part.), Telford (1996), Ibáñez et al. (1996), Ibáñez et al. (1997 "1995"), Young et al. (1999; in part.), Ibáñez et al. (2001; in part.), Poe (2004, in part.), Pinto et al. (2008), Jaramillo et al. (2010; in part.).

Norops tropidogaster: Villa et al. (1988) , Köhler (2000; in part.), Nicholson (2002), Köhler (2003; in part.), Nicholson et. al. (2005), Köhler (2008; in part.), Steffen (2009).

Diagnosis. A medium-sized species (SVL in largest specimen examined 52.5 mm) of the genus Anolis (sensu Poe 2004) that differs from all other Lower Central American beta anoles (sensu Etheridge 1967) in that it is longlegged (longest toe of adpressed hind leg reaches to at least center of eye, usually to a point between anterior border of eye and nostril), has strongly keeled mucronate imbricate ventral scales, a large almost orange red (in life) colored dewlap with a yellowish margin in males, a pair of greatly enlarged postcloacal scales in males, a small unilobed hemipenis in males, and no tube-like axillary pocket. Anole species from Lower Central America that are somewhat similar in appearance to A. gaigei are A. tropidogaster , A. polylepis , and A. cupreus . The males of A. tropidogaster have a mostly uniform purplish red dewlap, a bilobed hemipenis, and lack a pair of greatly enlarged postcloacal scales. For a more detailed comparison of A. gaigei and A. tropidogaster see the Diagnosis section for the latter species. Anolis polylepis and A. osa have smooth ventral scales at midbody and a larger, mostly uniform orange male dewlap (in some parts of its range in Costa Rica, considerable variation was observed in male dewlap coloration of A. polylepis , see Köhler et al. 2010). Also, male A. polylepis have a bilobate hemipenis. The males of Anolis cupreus lack a pair of greatly enlarged postcloacal scales, have a brown to pink dewlap with an orange margin, and a bilobate hemipenis.

Description. Anolis gaigei is a medium-sized anole (maximum recorded SVL 52.5 mm in males, 52.0 mm in females); dorsal head scales ( Fig. 11) in internasal region keeled, in prefrontal, parietal, and frontal areas rugose to tuberculate; scales in distinct prefrontal depression slightly wrinkled, parietal depression absent; 5–7 (6.06 ± 0.54) postrostrals; anterior nasal usually single, occasionally divided, usually in contact with rostral and first supralabial ( Fig. 9 , Tab. 2); 5–9 (7.03 ± 1.01) internasals; canthal ridge sharply defined; scales comprising supraorbital semicircles weakly keeled, largest scale in semicircles about same size as largest supraocular scale; supraorbital semicircles well defined; 1–4 (2.10 ± 0.62) scales separating supraorbital semicircles at narrowest point; 2–5 (3.03 ± 0.65) scales separating supraorbital semicircles and interparietal at narrowest point; interparietal well defined, greatly enlarged relative to adjacent scales, surrounded by scales of moderate size, longer than wide, usually larger than ear opening; supraorbital disc composed of 6–12 distinctly enlarged keeled scales; enlarged supraoculars not in contact with supraorbital semicircles; usually a single elongated superciliary, or, if 2 elongate superciliaries, posterior one much shorter than anterior one; 2–3 enlarged canthals; 8–14 (10.93 ± 1.14) scales between second canthals; 10–16 (13.10 ± 1.32) scales present between posterior canthals; loreal region slightly concave, 22–59 (40.25 ± 5.79) mostly keeled (some smooth or rugose) loreal scales in a maximum of 5–8 (6.31 ± 0.60) horizontal rows; 6–9 (7.31 ± 0.65) supralabials to level below center of eye; suboculars keeled, suboculars separated from supralabials by 0–1 (0.69 ± 0.47) scale row; ear opening vertically oval; scales anterior to ear opening granular, similar in size to those posterior to ear opening; 5–9 (6.35 ± 0.80) postmentals, outer pair largest; keeled granular scales present on chin and throat; male dewlap extending well onto chest, anterior insertion at level of center of eye, posterior insertion about 3.0 mm beyond level of axilla; 8–9 horizontal gorgetal-sternal rows with 11–15 scales per row, rows somewhat irregular; female dewlap small or absent; no nuchal crest or dorsal ridge; 2 middorsal scale rows slightly enlarged, weakly keeled, dorsal scales lateral to middorsal series gradually larger than granular lateral scales; no enlarged scales scattered among granular laterals; 53–75 (62.7 ± 5.57) dorsal scales along vertebral midline between levels of axilla and groin in males, 55–83 (62.7 ± 6.45) in females; 31–56 (38.5 ± 5.35) dorsal scales along vertebral midline contained in one head length in males, 28–52 (37.1 ± 7.10) in females; ventral scales on midsection about the same size as largest dorsal scales; ventral body scales strongly keeled, imbricate; 43–58 (50.0 ± 3.94) ventral scales along midventral line between levels of axilla and groin in males, 40–54 (47.6 ± 3.79) in females; 29–48 (35.0 ± 4.31) ventral scales contained in one head length in males, 25–38 (30.5 ± 4.44) in females; 94–138 (119.0 ± 9.68) scales around midbody in males, 101–128 (116.1 ± 7.5) in females; tubelike axillary pocket absent; preanal scales not keeled; males with a pair of greatly enlarged postcloacal scales; tail laterally compressed in cross section, tail height/tail width 1.05–1.47 (1.20 ± 0.09); basal subcaudal scales smooth; lateral caudal scales keeled, homogeneous; dorsal medial caudal scale row slightly enlarged, keeled, not forming a crest; most scales on lateral surface of antebrachium weakly keeled, unicarinate; 20–28 (25.45 ± 1.65) subdigital lamellae on Phalanges II–IV of Toe IV of hind limbs; SVL 36.0–52.5 (45.6 ± 3.40) mm in males, 37.0–52.0 (44.7 ± 4.75) mm in females; HL 10.5–13.2 (12.1 ± 0.65) mm in males, 10.6–12.1 (11.5 ± 0.48) mm in females; tail length 59.0–115.0 (95.0 ± 11.63) mm in males, 82.0–100.7 (92.3 ± 8.33) mm in females; shank length 11.2–15.6 (13.6 ± 0.85) mm in males, 10.5–14.0 (12.4 ± 0.82) mm in females; tail length/SVL 1.37–2.34 (2.09 ± 0.21) in males, 1.75–2.27 (2.10 ± 0.13) in females; HL/SVL 0.26–0.30 (0.27 ± 0.01) in males, 0.24–0.29 (0.26 ± 0.02) in females; shank length/SVL 0.28–0.32 (0.30 ± 0.01) in males, 0.23–0.33 (0.29 ± 0.02) in females; shank length/HL 1.04–1.24 (1.12 ± 1.04) in males, 0.94–1.18 (1.08 ± 0.05) in females. Of 20 specimens examined, the longest toe of the adpressed hind leg reached to mid-eye in 2 individuals (10%), to anterior margin of eye in 4 individuals (20%), and to a point between eye and nostril in 14 individuals (70%).

Coloration in life of an adult male (SMF 91918) was recorded as follows: Dorsal ground color Sayal Brown (223C) with a Beige (219D) vertebral band, edged by Raw Umber (223) pigment; Raw Umber (223) line continuing anteriorly through eye to tip of snout; dorsum of head Clay Color (26) with a medially interrupted Dark Brownish Olive (129) interorbital bar and a Sepia (219) nuchal spot, followed posteriorly by a Beige (219D) longitudinal line; forelegs Sayal Brown (223C); hind legs Cinnamon-Brown (33) with Raw Umber (23) crossbars; dorsal surface of tail Raw Umber (23) grading into Buff (24) distally and with indistinct Cinnamon-Rufous (40) bands; ventral surfaces of head, body, and limbs Pale Horn Color (92); ventral surface of tail suffused with Orange-Rufous (132C); dewlap Chrome Orange (16), grading into Orange Yellow (18) on anterior and distal margins; gorgetals dirty white; iris Kingfisher Rufous (240). Coloration in life of another adult male (SMF 91529) was recorded as follows: Dorsal and lateral surfaces of body and forelimbs Tawny Olive (223D); two broad Natal Brown (219A) longitudinal stripes extending from eye paravertebrally to base of tail, suffused with Walnut Brown (221B); a series of Sepia (119) blotches between occipital region and base of tail; dorsal surface of head Raw Umber (123), laterally grading into Tawny Olive (223D); ventral ground color Pale Horn Color (92), suffused with Orange-Rufous (132C) beneath tail; dorsal and lateral surfaces of tail and hind limbs Sayal Brown (223C) with the suggestion of diffuse Orange-Rufous (132C) crossbars; iris Robin Rufous (340); dewlap Burnt Orange (116), especially anterior portions suffused with Grayish Olive (43); anterior base of dewlap Warm Buff (118), posterior base Pale Horn Color (92); dewlap scales dirty white. Coloration in life of an adult female (SMF 91917) was recorded as follows: Dorsal ground color Tawny Olive (223D) with a Clay Color (123B) vertebral stripe, edged by Verona Brown (223B) pigment; dorsum of head Cinnamon Brown (33); forelegs Tawny Olive (223D); hind legs True Cinnamon (139) with Orange-Rufous (132C) spots and crossbars; dorsal surface of tail Tawny Olive (223D) with faint Orange Rufous (132C) crossbars; chin dirty white; venter Pale Pinkish Buff (121D); ventral surface of tail suffused with Orange- Rufous (132C); gular region Spectrum Orange (17) grading into Orange Yellow (18) on anterior margin; gorgetals Pale Pinkish Buff (121D); iris Robin Rufous (340). The coloration of an adult male from the Canal Zone (SMF 85304) was recorded as follows: Middorsum Army Brown (219B) bordered by a Burnt Umber (22) dorsolateral stripe; flanks Dark Drab (119B) with Drab-Gray (119D) punctuations; venter Drab-Gray (119D) suffused with Dark Drab (119B); dewlap Chrome Orange (16) with Orange Yellow (18) anterior border.

Natural history notes. Wherever we found it in central and western Panama, Anolis gaigei is an extremely common anole that reaches high population densities, being the most obvious lizard at many sites where it occurs. It is usually encountered in low vegetation between 0.5 and 2.0 m above the ground. Occasionally, individuals can be observed on the ground. At night, these animals sleep in the usual anole-like fashion on twigs or the upper surface of leaves. Although A. gaigei inhabits a wide range of habitats, it seems to be a typical species of the semideciduous forest and of bushy savannahs ( Fig. 12 ). In cattle pasture they depend on the trees along the fences (“living fences,” where living trees make up the actual fence posts). Aspects of the ecology of this species were studied by Sexton et al. (1971), Campbell (1971), and Quintero and Cambra (1993).

Geographic distribution. As currently known, Anolis gaigei is distributed from near the eastern city limit of Davíd, Chiriquí, along the Pacific versant of western and central Panama, including the Azuero Peninsula, to at least the Canal Zone, and widely distributed in the Santa Marta Mountains of Colombia as well as possibly northwestern Venezuela ( Figs. 2 , 13 ). The documented vertical range of the species is from near sea level to about 900 m. The elevation stated to be “8, 0 0 0 ft.” (= 2438 m) for a supposed locality of this species (Pueblo Viejo, Santa Marta Mountains, Colombia) by Ruthven (1916: 8) seems to be exceedingly high and therefore unlikely for this species. However, the associated specimens (UMMZ 48322–23) are clearly referable to A. gaigei .

Figures

FIGURE 1. (a) Type A hemipenis (SMF 91956); (b) Type B hemipenis (SMF 91902). See text for details. Scale bar = 1.0 mm.

FIGURE 2. Map indicating known collecting sites mentioned in text of anoles formerly referred to as Anolis tropidogaster. Each symbol can represent one or more nearby localities. Circles: Type A hemipenes; squares: Type B hemipenes; triangles: localities of Anolis tropidogaster - like specimens not verified by authors (sources: MHUA catalogue, Donoso-Barros 1968, Carvajal-Cogollo and Urbina-Cardona 2008). See text for details.

FIGURE 9. Designation of nasal scalation types. (a) Nasal type 1 (MHCH 1636); (b) Nasal type 2 (SMF 91901). Scale bars = 1.0 mm. See Text for details.

FIGURE 12. Habitat of Anolis gaigei (a) near Santo Domingo, Los Santos, Panama, 40 masl; (b) at Finca La Providencia, near Ponuga, Veraguas, Panama, 20 masl.

FIGURE 13. Map indicating collecting sites of selected lowland anoles inhabiting the Pacific versant of Panama and Costa Rica. Each symbol can represent one or more nearby localities. Circles: Anolis tropidogaster; squares: A. gaigei; white triangles: localities of A. tropidogaster - like specimens not verified by authors; black inverted triangles: A. polylepis; stars: A. osa; hexagons: A. cupreus. See text for details.