Anolis boulengerianus Thominot 1887

Köhler, Gunther, Pérez, Raúl Gómez Trejo, Petersen, Claus Bo P., Méndez, Fausto R. & Cruz, De La, 2014, A revision of the Mexican Anolis (Reptilia, Squamata, Dactyloidae) from the Pacific versant west of the Isthmus de Tehuantepec in the states of Oaxaca, Guerrero, and Puebla, with the description of six new species, Zootaxa 3862 (1), pp. 1-210 : 103-111

Anolis boulengerianus Thominot 1887

Figs. 84–89

Anolis boulengerianus Thominot, 1887:182 ; type locality: “Téhuantépec ( Mexique)”. Syntypes: MNHN 6554 and 1994.1670–71. Smith and Taylor 1950b, Nieto Montes de Oca et al. 2014

Anolis isthmicus Fitch 1978: 2 ; type locality: “ 19.5 km WNW Tehuantepec, Oaxaca, Mexico ”. Holotype: KU 176034. Lieb 1981, Fitch & Hillis 1984, Gorman et al. 1984, Liner 1994, Flores-Villela 1993, Flores-Villela & Gerez 1994, Lieb 1995, 2001, Liner 2007, Fläschendräger & Wijffels 2009, Flores-Villela et al. 2010, Johnson et al. 2010, Urbina-Cardona & Flores-Villela 2010, Wilson et al. 2013, Wilson & Townsend 2010, Nieto Montes de Oca et al. 2013, Köhler et al. 2014

Anolis nebuloides: Hartweg & Oliver 1940 , Werler & Smith 1952, Guibé 1954, Pianka & Smith 1959, Chrapliwy & Fugler 1955, Brygoo 1989

Anolis nebulosus: Barbour 1934 (in part), Smith and Taylor 1950 (in part).

Norops isthmicus: Savage & Guyer 1989 , Nicholson 2002, Nicholson et al. 2012

Diagnosis. A small to moderate-sized species (SVL in largest male 56.0 mm, largest female 51.0 mm) of the genus Anolis (sensu Poe 2004) that differs from all Mexican and Central American anoles except A. carlliebi , A. immaculogularis , A. quercorum , A. subocularis , and A. sacamecatensis by having a combination of (1) strongly keeled ventral scales; (2) usually a patch of three greatly enlarged supraocular scales; (3) 13–20 rows of slightly to moderately enlarged dorsal scales that are smaller than ventral scales; (4) moderately long hind legs, longest toe of adpressed hind leg usually reaching to level of posterior border of eye or beyond, ratio shank length/SVL 0.24–0.32; and (5) a large orange yellow dewlap in males and a small orange dewlap in females. Anolis boulengerianus differs from A. carlliebi , A. quercorum , and A. sacamecatensis by having longer legs with the longest toe of adpressed hind leg usually reaching to level of posterior border of eye or beyond, ratio shank length/ SVL 0.24–0.31 (vs. to ear opening or to a point between ear opening and eye, ratio shank length/SVL 0.20–0.26). Anolis boulengerianus differs from A. subocularis in male dewlap coloration in life (orange yellow with paler areas around gorgetals in A. boulengerianus vs. pinkish red with paler areas around gorgetals in A. subocularis ) and in hemipenial morphology (hemipenis slightly bilobate with two apical fields void of ornamentation, one on each lobe and with a finger-like asulcate processus and an indistinct asulcate ridge in A. boulengerianus vs. hemipenis unilobate with a single apical field void of ornamentation and without an asulcate processus or asulcate ridge in A. subocularis ). Also, in A. boulengerianus , the subocular scales are usually broadly in contact with the supralabials (vs. subocular scales separated from supralabials by one scale row or these scales in contact in A. subocularis ). Anolis boulengerianus differs from A. immaculogularis in male dewlap coloration in life (orange yellow with paler areas around gorgetals in A. boulengerianus vs. pinkish red without paler areas around gorgetals in A. immaculogularis ).

Description. Anolis boulengerianus is a small to moderate-sized anole (maximum recorded SVL 56.0 mm in males, 51.0 mm in females); dorsal head scales in internasal region keeled, uni- to tricarinate, other dorsal head scales mostly keeled in prefrontal and frontal region, smooth or keeled in parietal region; moderately deep prefrontal depression present, shallow parietal depression; 5–7 postrostrals; anterior nasal single or divided, the lower scale in contact with rostral and first supralabial; 6–8 internasals; canthal ridge sharply defined; scales comprising supraorbital semicircles well defined, weakly keeled, largest scale in semicircles subequal or larger than largest supraocular scale; supraorbital semicircles usually in contact, commonly separated by a complete row of scales; 0–2 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, larger than ear opening; enlarged supraoculars usually a patch of 3 greatly enlarged scales in a single row, usually separated from supraorbital semicircles by a complete row of small scales, or these scales occasionally narrowly in contact; 1–3 scales between enlarged supraoculars and superciliaries; 2 or 3 elongate superciliaries, anterior one longest, followed posteriorly by a series of 4–5 rounded or squarish keeled scales of moderate size; usually 3 enlarged canthals, the second canthal largest; 6–9 scales between second canthals; 7–11 scales between posterior canthals; loreal region slightly concave, 20–40 smooth to strongly keeled loreal scales in a maximum of 5–6 horizontal rows; 6–8 supralabials to level below center of eye; suboculars keeled, broadly in contact with supralabials (1–3 suboculars in contact with 2–4 supralabials); ear opening vertically oval, oriented slightly obliquely; scales anterior to ear opening keeled, juxtaposed to subimbricate, about four times larger than granulars posterior to ear opening; 5–8 infralabials to level below center of eye; 3–6 postmentals (usually 4), outer pair slightly to distinctly larger than adjacent median postmental scales; 0–2 (usually 1) enlarged sublabials in contact with infralabials on each side; keeled granular scales present on chin and throat; male dewlap moderate-sized to large, extending from level below anterior margin of eye to level of chest; 7–8 horizontal gorgetal-sternal rows with 8–15 scales per row; modal number of marginal pairs 2–4; female dewlap very small; a nuchal crest and a dorsal ridge present in males; scales on middorsum strongly keeled, subimbricate with rounded posterior margins; 12–18 middorsal scale rows slightly to moderately enlarged, with a gradual transition to lateral scales; lateral scales keeled, granular, usually homogeneous; 45–60 dorsal scales along vertebral midline between levels of axilla and groin in males, 48–73 in females; 24–34 dorsal scales along vertebral midline contained in one head length in males, 24–40 in females; ventral scales on midsection slightly to distinctly larger than largest dorsal scales; scales on midventer strongly keeled, imbricate, mucronate; 43–57 ventral scales along midventral line between levels of axilla and groin in males, 38–54 in females; 30–42 ventral scales contained in one head length in males, 22–34 in females; 110–140 scales around midbody in males, 116–126 in females; tube-like axillary pocket absent; precloacal scales weakly to strongly keeled; males almost always with a pair of greatly enlarged postcloacal scales; tail moderately compressed in cross section, tail height/tail width 1.18–1.56 in males, 1.19–1. 46 in females; all caudal scales strongly keeled, homogeneous, although an indistinct division in segments is discernible; dorsal medial caudal scale row hardly enlarged, strongly keeled, not forming a crest; scales on anterodorsal surface of brachium imbricate, weakly to strongly keeled, unicarinate, slightly mucronate; scales on dorsal surface of antebrachium subimbricate to imbricate, weakly to strongly keeled, uni- to tricarinate, slightly mucronate; 21–27 subdigital lamellae on Phalanges II–IV of Toe IV of hind limbs; 6–8 subdigital lamellae on distal phalanx of Toe IV of hind limbs; digital pads dilated, about twice the size of distal phalanx. In all specimens examined, the longest toe of the adpressed hind leg reaches to level of tympanum or to a point between shoulder and tympanum. For variation in selected scalation and morphometric characters see Table 10 .

The coloration in life of an adult male from Santa Cruz Bamba (SMF 96385; Fig. 85c ) was recorded as follows: Dorsal ground color Drab-Gray (256) suffused with Ground Cinnamon (270) on mid-dorsal and Sayal Brown (49) interorbital bar; dorsal surface of limbs Drab (19) with Tawny Olive (17) bars; tail Drab (19) with Cream Color (12) blotches on base of tail and with Grayish Horn Color (268) bands; ventral surfaces of limbs and body Pale Pinkish Buff (3) suffused with Beige (254); ventral surface of tail Drab (19); dewlap Chrome Orange (74) with Light Buff gorgetals; iris Warm Sepia (40).

The coloration in life of an adult female from Santa Cruz Bamba (SMF 96719) was recorded as follows: Dorsal ground color Drab (19) with Raw Umber (23) in vertebral region and with Marsh Brown chevrons; dorsal surface of tail Dark Drab (45) with Verona Brown (37) chevrons and an indistinct Pale Pinkish Buff (3) lateral line; dorsal surface of head Drab (19) with Clay Color (20) interorbital bar; dorsal surface of limbs Warm Sepia (40) with Verona Brown bars; ventral surface of body Light Buff (2) with Drab (19) suffusions; ventral surface of limbs Drab (19) with Cream Color (12) suffusions; chin Pale Buff (1) with Drab (19) speckles; dewlap Flame Scarlet (73) grading into Chrome Orange (74) towards posterior margin and Cream White (52) gorgetals; iris Olive (126).

The completely everted hemipenis of SMF 96279 ( Fig. 89 ) is a small, slightly bilobate organ; sulcus spermaticus bordered by well developed sulcal lips and opening into two apical fields void of ornamentation, one on each lobe; a finger-like asulcate processus and an indistinct asulcate ridge present; no surface ornamentation discernible.

Natural History Notes: Anolis boulengerianus inhabits the rocky slopes in dry forest. All individuals of A. boulengerianus were found clinging to trunks and branches of bushes and small trees during daytime, the lizard’s body often oriented horizontally.

Geographic Distribution and Conservation. As currently known, Anolis boulengerianus is restricted to the hills west and northwest of Tehuantepec in southeastern Oaxaca at elevations between 55 and 1315 masl ( Fig. 76 ). Given its presumably small geographic range, degraded habitat, and threat from deforestation, it seems justified to classify A. boulengerianus as Critically Endangered based on criterion B1ab(iii) of the IUCN Red List Categories and Criteria ( IUCN 2012).

Specimens examined ⎯ Mexico: Oaxaca: Cerro Sacamecates, 1315 m: SMF 96719; Tehuantepec: FMNH 108663, MNHN 1994.1670–71, 6554; near Tehuantepec, 427 m: FMNH 108664, USNM 46983–86; Portillo de la Conception: FMNH 108662, UIMNH 20054; Santa Cruz Bamba, 55 m: IBH 26575, 26990–91, SMF 81592, 96197, 96279, 96386, 96752; Mex Hwy 185, 20 km N 7 km E Juchitán de Zaragoza, 200 m: MCZ R-146493; north of Juchitán: SMF 80763; near Chivela: MCZ R-25030–31; Gingola, 10 km from Mixtequilla: SMF 77184; La Sierra Madre: UIMNH 56172; Pie de la Montana de la Sierra Madre, Zanatepec: UIMNH 56166; 5.3 km (airline) NNW San Pedro Totolapa, 1140 m: SMF 96385.

Figures

FIGURE 76. Map indicating collecting localities of species related to Anolis boulengerianus. Each symbol can represent one or more adjacent localities. Black circles: Anolis boulengerianus; white circles: A. immaculogularis; black squares: A. subocularis.

FIGURE 84. Dewlaps of Anolis boulengerianus in life. (a) IBH 26691; (b) SMF 96385; (c) SMF 96297; (d) IBH 26575; (e) SMF 96197; (f) SMF 96386. (a–d) are males, (e–f) are females.

FIGURE 85. Anolis boulengerianus in life. (a) SMF 96279; (b) IBH 26990; (c) SMF 96385; (d) IBH 26575; (e) SMF 96386; (f) IBH 26991. (a–c) are males, (d–f) are females.

FIGURE 86. Anolis boulengerianus: head scalation in (a,c,e) SMF 96197; (b,d,f) SMF 96297. Scale bars equal 1.0 mm.

FIGURE 87. Anolis boulengerianus: nasal region in (a) SMF 96197; (b) IBH 26575; superciliary region in (c) SMF 96297; (d) IBH 26575; chin region in (e) SMF 96297; (f) IBH 26575; cloacal region in (g) SMF 96297; (h) IBH 26575. Scale bars equal 1.0 mm.

FIGURE 88. Anolis boulengerianus: dorsal scales in (a) IBH 26575; (b) SMF 96197; flank scales in (c) IBH 26575; (d) SMF 96297; ventral scales in (e) IBH 26575; (f) SMF 96197. Scale bars equal 1.0 mm.

FIGURE 89. Hemipenis of Anolis boulengerianus (SMF 96279): (left) sulcate view; (right) asulcate view. Scale bar equals 1.0 mm.