Anolis liogaster Boulenger 1905

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 : 158-167

Anolis liogaster Boulenger 1905

Figs. 131–136

Anolis liogaster Boulenger 1905: 245 ; type locality: “Omilteme, Guerrero, 7600 ft.,” Mexico. Syntypes: BMNH 1946.8.8.53–54 (examined by the authors). Gadow 1905, Barbour 1934, Smith &Taylor 1950 a,b, Davis 1954, Stuart 1955, Etheridge 1959, Davis & Dixon 1961, Duellman 1961, Shelford 1963, Duellman 1965, Smith 1972, Smith et al. 1972, Fitch & Henderson 1973, 1976, Smith & Smith 1976, Lieb 1981, Fitch & Hillis 1984, Gorman et al. 1984, Buth et al. 1985, Saldaña de la Riva & Pérez-Ramos 1987, Muñoz-Alonso 1988, Flores-Villela & Muñoz-Alonso 1990, MuñozAlonso & Flores-Villela 1990, Flores-Villela 1993, Flores-Villela & Muñoz-Alonso 1993, Flores-Villela & Gerez 1994, Lieb 1995, Pérez-Ramos et al. 2000, Lieb 2001, Poe 2004, Liner 2007, Fläschendräger & Wijffels 2009, Flores-Villela et al. 2010, Urbina-Cardona & Flores-Villela 2010, Wilson & Townsend 2010, Köhler et al. 2013 a, Nieto Montes de Oca et al. 2013, Wilson et al. 2013, Köhler 2014, Köhler et al. 2014

Anolis adleri Smith 1972: 179 ; type locality: “ 2.2 km (by road) W Patio de Aviación, Guerrero, Mexico, 2160 m.” Holotype: UMMZ 131685 (examined by the authors). Smith et al. 1972, Fitch & Henderson 1976, Lieb 1981, Flores-Villela 1993, Flores-Villela & Gerez 1994, Lieb 2001, Fläschendräger & Wijffels 2009, Flores-Villela et al. 2010, Köhler 2012 a, Nieto Montes de Oca et al. 2013

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

Norops adleri: Nicholson et al. 2012

Diagnosis. A small to moderate-sized species (SVL in largest male 50.0 mm, largest female 54.0 mm) of the genus Anolis (sensu Poe 2004) that differs from all other Mexican and Central American congeners except A. dunni , A. gadovii , A. omiltemanus , and A. peucephilus by having (1) smooth ventral scales; (2) an oval patch of usually three greatly enlarged supraorbital scales; (3) a pair of greatly enlarged postcloacal scales in males. Anolis liogaster differs from A. dunni and A. gadovii by having 10–15 rows of dorsal scales moderately enlarged (vs. the middorsal scales not or only 2-4 rows slightly enlarged in A. dunni and A. gadovii ). It further differs from A. dunni by having a uniform purple to pink male dewlap (vs. pinkish to orange red male dewlap with semicircular pale streaks and blotches in A. dunni ). It also differs from A. gadovii by the absence of a bold reticulated body pattern (vs. such a pattern present in A. gadovii ). Anolis liogaster differs from A. omiltemanus and A. peucephilus by having longer hind legs with the longest toe of adpressed hind leg reaching to a point between posterior and anterior margin of eye or occasionally to a point between ear opening and eye (vs. to level of ear opening or to a point between shoulder and ear opening in A. omiltemanus and A. peucephilus ), usually only a single pair of greatly enlarged sublabial scales in contact with infralabial scales (vs. usually two pairs in A. omiltemanus and A. peucephilus ), and a pink to purple male dewlap (vs. orange yellow in A. omiltemanus and A. peucephilus ).

Description. Anolis liogaster is a small to moderate-sized anole (maximum recorded SVL 50.0 mm in males, 54.0 mm in females); dorsal head scales in internasal region weakly keeled, some smooth or rugose, other dorsal head scales smooth or rugose; moderately deep frontal depression present, shallow parietal depression; 5–8 postrostrals; anterior nasal usually single, occasionally divided, the lower scale in contact with rostral and first supralabial or, exceptionally, only with rostral scale; 6–8 internasals; canthal ridge sharply defined; scales comprising supraorbital semicircles well defined, posterior ones smooth, anterior ones weakly keeled, largest scale in semicircles larger than largest supraocular scale; supraorbital semicircles usually broadly in contact, commonly separated by a complete row of scales; 1–3 scales separating supraorbital semicircles and interparietal at narrowest point; interparietal well defined, greatly enlarged relative to adjacent scales, surrounded by scales of small to moderate size, longer than wide, usually larger than ear opening; enlarged supraoculars usually faintly keeled, separated from supraorbital semicircles by a complete row of small scales, or these scales narrowly in contact; 2–5 rows of granular scales between enlarged supraoculars and superciliaries; 2–3 elongate superciliaries, anterior one longest, followed posteriorly by a series of 4–5 rounded or squarish scales of moderate size; usually 3 enlarged canthals, the second canthal largest; 4–9 scales between second canthals; 5–11 scales present between posterior canthals; loreal region slightly concave, 16–39 mostly keeled (some smooth or rugose) loreal scales in a maximum of 4–6 (commonly 4 or 5) horizontal rows; 5–7 supralabials to level below center of eye; suboculars keeled, in broad contact with supralabials (1–3 suboculars in contact with 1–4 supralabials); ear opening vertically oval, oriented slightly obliquely; scales anterior to ear opening granular, about four times as large than those posterior to ear opening; 5–7 infralabials to level below center of eye; 2–5 (usually 4) postmentals, outer pair usually at least four times larger than adjacent median postmental scales; usually one, occasionally zero or two, enlarged sublabials in contact with infralabials on each side; smooth or faintly keeled granular scales present on chin and throat; male dewlap large, extending from level below anterior margin of eye to level of chest; 8–9 horizontal gorgetal-sternal rows with 7–13 scales per row; modal number of marginal pairs 2–4; female dewlap very small to small; a nuchal crest and a dorsal ridge present in males; scales on middorsum subimbricate, weakly keeled, with rounded posterior margins, grading into smaller granular lateral scales; 10–17 middorsal scale rows slightly to moderately enlarged, two vertebral rows occasionally larger than adjacent rows and usually a few smaller scales interspersed in enlarged rows; lateral scales usually homogeneous, occasionally slightly heterogeneous; 37–58 dorsal scales along vertebral midline between levels of axilla and groin in males, 44–65 in females; 22–36 dorsal scales along vertebral midline contained in one head length in males, 22–34 in females; ventral scales on midsection about twice the size of largest dorsal scales; scales on midventer smooth, subimbricate to imbricate with rounded posterior margins, usually somewhat heterogeneous in size; 35–47 ventral scales along midventral line between levels of axilla and groin in males, 33–45 in females; 24–38 ventral scales contained in one head length in males, 18–28 in females; 102–128 scales around midbody in males, 92–132 in females; tube-like axillary pocket absent; precloacal scales smooth or weakly keeled; males with a pair of greatly enlarged postcloacal scales; tail moderately compressed in cross section, tail height/tail width 1.16–1.50 in males, 1.05–1.59 in females; basal subcaudal scales smooth or faintly keeled, all other caudal scales strongly keeled; lateral caudal scales homogeneous, although an indistinct division in segments is discernible; dorsomedial caudal scale row slightly enlarged, keeled, not forming a crest; scales on anterodorsal surface of brachium and on dorsal surface of antebrachium imbricate, weakly keeled, unicarinate; 20–28 subdigital lamellae on Phalanges II–IV of Toe IV of hind limbs; 6–9 subdigital lamellae on distal phalanx of Toe IV of hind limbs; digital pads dilated, about 3 times the width 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 12 .

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The coloration in life of an adult male from Omiltemi (SMF 96199; Fig. 132a ) was recorded as follows: Dorsal ground color Ground Cinnamon (270) with Brussels Brown (33) chevrons that end in lateral ocelli; center of ocellus Smoky White (261) surrounded by Burnt Umber (48); mid-dorsal area between chevrons Cinnamon Drab (50); lateral stripe Smoky White (261) edged by Brussels Brown (33); postorbital stripe Verona Brown (37) with Kingfisher Rufous (28) bands; dorsal surface of tail Mikado Brown (42) with Prout´s Brown (47) bands; ventral surface of head Cinnamon Drab (50) with Dark Neutral Gray (299) speckles; venter Chamois (84) with suffusions of Cinnamon Drab (259); ventral surface of limbs Fawn Color (258); dewlap Spinel Pink (235) with Cream White (52) gorgetals; iris Burnt Umber (48). The coloration in life of another adult male from Omiltemi (SMF 96201; Fig. 132b ) was recorded as follows: Dorsal ground color of body limbs and tail Dark Drab (45) with Raw Umber (280) vertebral spots and with irregular Cinnamon-Drab (50) zigzag band and Cream White (52) vertical line; venter Chamois (84); ventral surface of limbs Cinnamon Drab (259); ventral surface of tail Chamois (84); dewlap Spinel Pink (239) with Cream White (52) gorgetals.

The coloration in life of an adult female from Omiltemi (SMF 96200; Fig. 132e ) was recorded as follows: Dorsum with a Salmon Color (83) vertebral line edged by Raw Umber (280); dorsal surface of head Drab (19) with a Russet (44) interorbital bar; dorsal surfaces of limbs Army Brown (46) with Cinnamon Rufous (31) blotches and bands; dorsal surface of tail Clay Color (18); lateral sides of body Clay Color (18) with Yellow Ocher (14) lateral stripe and with Pale Horn Color (11) semicircular markings; ventral surface of head, body and limbs Cream White (52) with Dark Pearl Gray (290) suffusions.

The completely everted hemipenis of SMF 96199 ( Fig. 136 ) is a small, unilobate organ; sulcus spermaticus bordered by well developed sulcal lips and opening into a single apical field void of ornamentation; a low asulcate processus near tip of apex but no asulcate ridge present; no surface ornamentation discernible.

Natural History Notes: In and around the village of Omiltemi, Anolis liogaster is a common species that can be found on wooden fences, on the lower portions of pine tree trunks, and on the ground.

Geographic Distribution and Conservation. As currently known, Anolis liogaster is restricted to the hills in the region of Omiltemi in the south-central portion of the Mexican State of Guerrero at elevations between 1995 and 2493 masl ( Fig. 111 ). Given its usual abundance wherever this species occurs, we consider the conservation status of A. liogaster to be Least Concern based on the IUCN Red List Categories and Criteria ( IUCN 2012).

Specimens examined ⎯ Mexico: Guerrero: 1.0 mi E Omiltemi: UTA 4409 ; 17.2 mi W Asoleadero , 190 m: UMMZ 130982 ; 2.2 km W Patio de Aviación , 2160 m: UMMZ 130981 , 131685 ; 3 mi W Omiltemi , 2493 m: MVZ 57178 ; Omiltemi , 2175 m: FMNH 108507-08 , 125620-21 , 125624 , IBH 26599-601 , 26605-07 , MCZ R-85021–22, 78718, SMF 96200-02 ; Omiltemi , 2060 m: SMF 96199 ; Omiltemi , 1995 m: BMNH 1946.8.8.53-54, SMF 96203- 06 ; Omiltemi , 2135 m: USNM 47748-52 , 148865 ; W Chilpancingo on the slope that leads to a canyon between the villages of Filo de Caballo and Carrizal de Bravo : UMMZ 229869 ; 37.7 km SW Filo de Caballo , 2256 m: KU 182540–43 ; 0.5 km E Filo de Caballo , 2294 m: IBH 6429 ; village of Los Morros, W Chilpancingo, 62.8 km from Zumpango del Río via Casa Verde , 2180 m: UMMZ 229861-62 , 229864 , 229866-67 ; Morro Viejo of the village Los Morros, W Chilpancingo: UMMZ 229870 .

Figures

FIGURE 111a. Map indicating collecting localities of species related to Anolis liogaster. Each symbol can represent one or more adjacent localities. Black circles: Anolis dunni; white circles: A. liogaster; black squares: A. taylori; white squares: A. gadovii.

FIGURE 131. Dewlaps of Anolis liogaster in life. (a) SMF 96199; (b) IBH 26600; (c) IBH 26602; (d) IBH 26605. (a–b) are males, (c–d) are females.

FIGURE 132. Anolis liogaster in life. (a) SMF 96199; (b) SMF 96201; (c) IBH 26600; (d) SMF 96202; (e) SMF 96200; (f) IBH 26599. (a–d) are males, (e–f) are females.

FIGURE 133. Anolis liogaster: head scalation in (a,c,e) SMF 96199; (b,d,f) SMF 96200. Scale bars equal 1.0 mm.

FIGURE 134. Anolis liogaster: nasal region in (a) IBH 26606; (b) SMF 96203; superciliary region in (c) IBH 26600; (d) SMF 96204; chin region in (e) IBH 26599; (f) SMF 96206; cloacal region in (g) SMF 96199; (h) SMF 96201. Scale bars equal 1.0 mm.

FIGURE 135. Anolis liogaster: dorsal scales in (a) SMF 96202; (b) SMF 96203; flank scales in (c) IBH 26605; (d) SMF 96205; ventral scales in (e) IBH 26605; (f) SMF 96203. Scale bars equal 1.0 mm..

FIGURE 136. Hemipenis of Anolis liogaster (SMF 96199): (left) sulcate view; (right) asulcate view. Scale bar equals 1.0 mm

Tables

TABLE 12. Selected measurements, proportions and scale characters of the species related to Anolis liogaster. Range is followed by mean value and standard deviation in parentheses. For abbreviations see text.

		Anolis dunni ♂ 25 ♀ 14	Anolis taylori ♂ 30 ♀ 19	Anolis gadovii ♂ 23 ♀ 17	Anolis liogaster ♂ 19 ♀ 22	Anolis omiltemanus ♂ 24 ♀ 21	Anolis peucephilus ♂ 3 ♀ 4
maximum SVL	males	58.5	73.0	76.0	50.0	47.0	46.0
	females	51.0	58.0	63.0	54.0	48.0	45.0
TL / SVL	males	1.75–2.09 (1.92±0.10)	1.95–2.11 (2.02±0.06)	1.82–1.94 (1.87±0.06)	1.90–2.27 (2.08±0.14)	1.52–1.91 (1.65±0.09)
	females	1.86–1.97 (1.90±0.04)	1.94–2.16 (2.03±0.08)	1.61–1.91 (1.79±0.13)	1.75–2.07 (1.93±0.14)	1.42–1.71 (1.56±0.08)
VDT / HDT	males	1.13–1.48 (1.31±0.10)	1.17–1.70 (1.52±0.13)	1.14–1.55 (1.32±0.12)	1.16–1.50 (1.28±0.09)	1.05–1.50 (1.33±0.14)	1.21–1.31 (1.28±0.05)
	females	1.19–1.60 (1.35±0.14)	1.17–1.44 (1.33±0.08)	1.11–1.25 (1.18±0.06)	1.05–1.59 (1.26±0.15)	1.00–1.50 (1.18±0.15)	1.24–1.50 (1.36±0.11)
AGD / SVL	males	0.37–0.44 (0.40±0.02)	0.37–0.44 (0.40±0.02)	0.37–0.48 (0.40±0.03)	0.34–0.45 (0.40±0.03)	0.37–0.48 (0.42±0.03)	0.39–0.40 (0.39±0.00)
	females	0.38–0.46 (0.42±0.02)	0.41–0.46 (0.43±0.01)	0.36–0.45 (0.42±0.02)	0.38–0.47 (0.42±0.02)	0.39–0.48 (0.44±0.03)	0.39–0.50 (0.43±0.05)
HL / SVL	males	0.24–0.28 (0.27±0.01)	0.23–0.29 (0.26±0.01)	0.24–0.27 (0.26±0.01)	0.27–0.33 (0.29±0.01)	0.26–0.31 (0.28±0.01)	0.26–0.28 (0.27±0.01)
	females	0.25–0.28 (0.26±0.01)	0.24–0.28 (0.26±0.01)	0.24–0.27 (0.26±0.01)	0.25–0.29 (0.28±0.01)	0.22–0.31 (0.27±0.02)	0.24–0.28 (0.26±0.01)
HL / HW	males	1.53–1.76 (1.67±0.06)	1.60–1.84 (1.72±0.07)	1.52–1.82 (1.68±0.07)	1.59–1.75 (1.66±0.05)	1.56–1.79 (1.64±0.06)	1.71–1.79 (1.76±0.04)
	females	1.58–1.80 (1.69±0.05)	1.61–1.86 (1.73±0.06)	1.57–1.80 (1.69±0.07)	1.46–1.69 (1.60±0.06)	1.52–1.70 (1.59±0.06)	1.59–1.75 (1.69±0.07)
SL / SVL	males	0.11–0.13 (0.11±0.01)	0.11–0.13 (0.11±0.01)	0.11–0.12 (0.11±0.00)	0.12–0.15 (0.13±0.01)	0.13–0.14 (0.13±0.01)	0.12–0.13 (0.13±0.00)
	females	0.10–0.13 (0.12±0.01)	0.11–0.13 (0.12±0.01)	0.11–0.13 (0.12±0.00)	0.11–0.13 (0.12±0.01)	0.12–0.15 (0.13±0.01)	0.11–0.12 (0.12±0.01)

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