Anolis quercorum Fitch 1978

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 : 85-93

Anolis quercorum Fitch 1978

Anolis quercorum Fitch 1978: 6 ; type locality: “ 26 km SE Nochixtlán (2.5 km NW Cuesta Blanca, Highway 190), Oaxaca, Mexico ”. Holotype: KU 176050. Lieb 1981, Fitch & Hillis 1984, Gorman et al. 1984, Flores-Villela 1993, Flores-Villela & Gerez 1994, Lieb 1995, Liner 2007, Fläschendräger & Wijffels 2009, Flores-Villela et al. 2010, Urbina-Cardona & Flores-Villela 2010, Wilson & Townsend 2010, Wilson et al. 2013, Köhler et al. 2014, Köhler 2014

Anolis nebuloides: Gehlbach & Colette 1957

Norops quercorum: Savage & Guyer 1989 , Nicholson 2002, Castiglia et al. 2010, Nicholson et al. 2012

Diagnosis. A small species (SVL in largest male 46.0 mm, largest female 43.5 mm) of the genus Anolis (sensu Poe 2004) that differs from all Mexican and Central American anoles except A. carlliebi and A. sacamecatensis by having a combination of (1) strongly keeled ventral scales; (2) usually a patch of three greatly enlarged supraocular scales; (3) 12–20 rows of slightly to moderately enlarged dorsal scales that are smaller than ventral scales; (4) short hind legs, longest toe of adpressed hind leg usually reaching to ear opening or to a point between ear opening and eye, ratio shank length/SVL 0.22–0.26; (5) circumnasal usually in contact with first supralabial; and (6) a large pink dewlap in males and a very small pinkish orange-red dewlap in females ( Fig. 65 ). Anolis quercorum differs from A. carlliebi and A. sacamecatensis in the ranges and average values of several morphometric and pholidotic characters (see Table 9 ), most obvious in (1) number of middorsal scales in one head length; (2) ratio shank length/ HL; and (3) subdigital lamellae on Phalanges II–IV of Toe IV. Also, Anolis quercorum differs from A. carlliebi and A. sacamecatensis in hemipenis morphology (hemipenis distinctly bilobed in A. quercorum vs. unilobed in A. carlliebi and slightly bilobed in A. sacamecatensis ) and by substantial genetic distances (see above).

Description. Anolis quercorum is a small anole (maximum recorded SVL 46.0 mm in males, 43.5 mm in females); dorsal head scales in internasal region smooth to weakly keeled, occasionally strongly keeled, unicarinate, other dorsal head scales smooth to weakly keeled in prefrontal region, in parietal region smooth to rugose; shallow to deep prefrontal depression present, shallow parietal depression; 4–7 postrostrals; anterior nasal usually single, exceptionally divided, the lower scale in contact with rostral and first supralabial; 4–8 internasals; canthal ridge sharply defined; scales comprising supraorbital semicircles well defined, smooth to strongly keeled, largest scale in semicircles usually subequal or larger than largest supraocular scale; supraorbital semicircles usually broadly in contact, occasionally separated by a complete row of scales; 1–4 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 subequal to ear opening; enlarged supraoculars usually a patch of 3 greatly enlarged scales in a single row (occasionally a patch of 4–7 moderately to greatly enlarged, somewhat irregular arranged scales in two rows) rugose to weakly keeled, separated from supraorbital semicircles by a complete row of small scales, or these scales narrowly in contact (broadly in contact in SMF 96456); 2–3 scales between enlarged supraoculars and superciliaries; usually 2 occasionally 3 elongate superciliaries, anterior one longest, followed posteriorly by a series of 3–4 rounded or squarish keeled scales of moderate size; usually 3 enlarged canthals, the second canthal largest; 4–9 scales between second canthals; 6–10 scales between posterior canthals; loreal region slightly concave, 11–29 mostly keeled (some smooth to rugose) loreal scales in a maximum of 3–5 horizontal rows; 5–7 supralabials to level below center of eye; suboculars keeled, broadly in contact with supralabials (2–3 suboculars in contact with 2–4 supralabials); ear opening vertically oval; scales anterior to ear opening keeled, juxtaposed, about four times larger than granulars posterior to ear opening; 5–8 infralabials to level below center of eye; 3–5 postmentals (usually 4), outer pair slightly to distinctly larger than adjacent median postmental scales; 0–3(commonly 0 or 1) enlarged sublabials in contact with infralabials on each side; keeled granular scales present on chin and throat; male dewlap of large size extending onto chest; 7–8 horizontal gorgetal-sternal rows with 14–18 scales per row; modal number of marginal pairs 2–3;female dewlap very small or small; a nuchal crest and a dorsal ridge present in males; scales on middorsum weakly to strongly keeled, subimbricate with rounded posterior margins; 12–20 middorsal scale rows slightly to moderately enlarged, with a gradual transition to lateral scales; lateral scales keeled, more or less homogeneous; 48–67 dorsal scales along vertebral midline between levels of axilla and groin in males, 49–71 in females; 22–36 dorsal scales along vertebral midline contained in one head length in males, 22–38 in females; ventral scales on midsection larger than largest dorsal scales; scales on midventer usually strongly keeled, imbricate, usually mucronate, occasionally weakly keeled, subimbricate, non-mucronate; 33–52 ventral scales along midventral line between levels of axilla and groin in males, 33–49 in females; 22–40 ventral scales contained in one head length in males, 18–28 in females; 106–140 scales around midbody in males, 100–126 in females; tube-like axillary pocket absent; precloacal scales smooth to weakly keeled; males with a pair of greatly enlarged postcloacal scales; tail moderately compressed in cross section, tail height/tail width 1.15–1.79 in males, 1.11–1.54 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 subimbricate to imbricate, strongly keeled, unicarinate, non-mucronate; scales on dorsal surface of antebrachium strongly keeled, unicarinate, subimbricate to imbricate, some slightly mucronate; 19–26 subdigital lamellae on Phalanges II–IV of Toe IV of hind limbs; 5–7 subdigital lamellae on distal phalanx of Toe IV of hind limbs; digital pads dilated, slightly more than 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. Of 14 individuals, the longest toe of adpressed hind leg reaching almost to ear opening in 3.3%, to ear opening in 60.0%, and to a point slightly beyond ear opening in 10.0%. For variation in selected scalation and morphometric characters see Table 9 .

The coloration in life of an adult male from 2 km SE Cuesta Blanca (SMF 96457) was recorded as follows: Dorsal ground color of body, head, limbs and tail Drab (19); limbs with Tawny Olive (17) bands; tail with Olive Brown (278) bands; ventral surfaces of body, limbs and tail Light Buff (2); dewlap Pale Purple (223) with Pale Pinkish Buff (3) gorgetals; iris Dark Drab (45).

The coloration in life of an adult female from 2 km SE Cuesta Blanca (IBH 26982) was recorded as follows: Dorsal ground color Mikado Brown (42) with a suffusion of Cinnamon Brown (43) on head; vertebral line Dark Salmon Color (59) grading into Orange-Rufous (56) on tail and edged by Cinnamon Brown (43) on dorsum; ventral surface of head Light Buff (2) ventral surface of limbs Drab (19); ventral surface of tail Salmon Color (58); dewlap Spinel Pink (235) with Pale Pinkish Buff (3) gorgetals; iris Sprout´s Brown (47). The coloration in life of another adult female from 2 km SE Cuesta Blanca (IBH 26983; Fig. 66d ) was recorded as follows: Dorsal ground color Drab (19) with a weakly defined Medium Fawn Color (257) zigzag band; dorsal surface of limbs True Cinnamon (260) with Cinnamon Drab (259) bands; ventral surface of head and body Pale Buff (1); ventral surface of limbs Pale Pinkish Buff (3); ventral surface of tail Beige (254); iris Sprout´s Brown (47).

The almost completely everted hemipenis of SMF 96439 ( Fig. 70 ) is a bilobate organ; sulcus spermaticus bordered by well developed sulcal lips and opening into two apical fields void of ornamentation, one on each lobe; a large finger-like asulcate processus and a weak asulcate ridge present; apex strongly calyculate, truncus with transverse folds.

Natural History Notes: The habitat of Anolis quercorum is leaf litter in oak forest. Here this typical ground anole can be found in amazing population densities. In the vicinity of the type locality of this species (2 km SE Cuesta Blanca, 17.33065°N, 97.158420°W, 2250 masl), we saw at least 50 individuals within one hour of searching in a relatively small area (<1 ha). When disturbed, these lizards darted a meter or so across the leaf litter and then usually froze and were well camouflaged against the vivid background colors.

Geographic Distribution and Conservation. As currently known, Anolis quercorum is distributed on the Pacific versant of the western central and northwestern portions of the Mexican State of Oaxaca at elevations between 1616 and 2250 masl ( Fig. 71 ). Given its usual abundance wherever this species occurs, it seems justified to classify A. quercorum as Least Concern based on the IUCN Red List Categories and Criteria ( IUCN 2012).

Specimens examined— Mexico: Oaxaca: 0.9 mi N (by road) El Cerezal, Sierra Aloapaneca (N of city of Oaxaca) Mex Hwy 175: UTA R-8421; 2 km SE Cuesta Blanca, 2250 m: IBH 26973–76 , 26982–83 , SMF 96441–55 , 96457–58 , 96747–51 , 98021 , 98735 ; 1.7 mi SE Nochixtlán : MCZ R-133097–98; 26 km SE Nochistlán , 2.5 km NW Cuesta Blanca : KU 176051–57 ; 3.5 WSW Tlaxiaco : UTA R-3568–71; 32 mi NW Oaxaca, 2164 m: UMMZ 115057 ; 32.5 mi NW Oaxaca, 2134 m: UMMZ 112636 (12 specimens); 3.1 km NW Llano Verde : UTA R-11502–03; 5.6 km E Llano Verde : KU 193181, 193188–89 ; 12.8 km W Llano Verde : MCZ R-171142–44; 52 mi N Oaxaca on MEX 190 2.4 mi N El Cumbre: UMMZ 131485 (nine specimens); Hotel Victoria , 1616 m: UTA R-25789; Oaxaca: FMNH 108789-92 ; Oaxaca Motor Court: Oaxaca City : MCZ R-86374–75; grounds of Oaxaca Port Motel : MCZ R-79562–64; 3 mi N La Heradura : MCZ R-177946–49, 54954–58; Tamazulapán del Progreso , 1990 m: IBH 26977, SMF 96438–40, 96456 ; Cerro San Felipe : MCZ R-93671–72; 15 km NW (on road) San Juan del Estado, distrito Etla : MVZ 144181 ; El Tejocote, 2078 m: UTA R-25788, 52818–23.

Taxonomy of the Mexican anoles related to Anolis boulengerianus Thominot 1887

Thominot (1887) described Anolis boulengerianus based on three specimens (now MNHN 6554, 1994.1670–71; Fig. 72 and 73 ), examined by GK, from “Téhuantépec ( Mexique).” For most of the time since its original description, this nominal species had remained in the synonymy of A. nebulosus or A. nebuloides (e.g., Barbour 1934, Smith and Taylor 1950 a, Guibé 1954, Brygoo 1989). Davis (1954) described A. subocularis from “one mile southwest of Tierra Colorado, 900 ft., Guerrero ” (holotype now TCWC 8675, Fig. 74 ), examined by GK. In 1978, Fitch described A. isthmicus based on specimens from “ 19.5 km WNW Tehuantepec, Oaxaca, Mexico ” and nearby localities. In his comparisons, he differentiated it from a selection of anole species from southern Mexico, but did not include A. boulengerianus in his considerations. Recently, Nieto Montes de Oca et al. (2014) demonstrated that the type material of A. boulengerianus is conspecific with that of A. isthmicus and therefore synonymized the latter with A. boulengerianus . Because of priority, A. boulengerianus is the valid name for the species previously known as A. isthmicus .

For our analysis, we defined three OTUs in this complex: OTU1 is formed by specimens from south-central and southeastern Guerrero; OTU2 by specimens from the coastal plain between Huatulco and Puerto Escondido in south-central Oaxaca; and OTU 3 by specimens from the hills west and northwest of Tehuantepec in southeastern Oaxaca. The analysis of the CO1 and 16s gene fragments revealed the existence of three genetic clusters ( Fig. 75 ). Genetic distances between our OTUs 2 and 3 are low in both genes (5.7–7.7% in CO1; 1.5–2.8 in 16s). In contrast, our OTU1 has a genetic distance to OTU 2 and 3, respectively, of 13.9–17.6 (CO1) and 6.9–7.8 (16s), respectively. We observed differences in male dewlap coloration between OTUs 2 and 3: in OTU 2, the male dewlap is pinkish red without paler areas around gorgetals whereas in OTU 3 the male dewlap is orange yellow with paler areas around gorgetals. Also, the eastern populations (our OTU 3) occur in a much more arid habitat than the western populations (our OTU 2). These findings are evidence for the existence of three species in this complex. Our OTU 1 contains the type locality of A. subocularis and therefore this name has to be applied to this species. The type localities of A. boulengerianus and A. isthmicus , respectively, are both contained in our OTU3; thus the older name (i.e., A. boulengerianus ) is the valid name for this species and the nominal species A. isthmicus remains in the synonymy of A. boulengerianus . No scientific name is available for our OTU 2, and we therefore describe it as a new species below.

Figures

FIGURE 65. Dewlaps of Anolis quercorum in life. (a) SMF 96451; (b) IBH 26974; (c) IBH 26977; (d) SMF 96458; (e) IBH 26983; (f) IBH 26975. (a–d) are males, (e–f) are females.

FIGURE 66. Anolis quercorum in life. (a) SMF 96448; (b) SMF 96438; (c) SMF 96447; (d) IBH 26983; (e) IBH 26976; (f) SMF 96456. (a–c) are males, (d–f) are females.

FIGURE 70. Hemipenis of Anolis quercorum (SMF 96439): (left) sulcate view; (center) asulcate view; (right) lateral view. Scale bar equals 1.0 mm.

FIGURE 71. Map indicating collecting localities of species related to Anolis quercorum. Each symbol can represent one or more adjacent localities. Black circles: Anolis carlliebi; white circles: A. quercorum; white squares: A. sacamecatensis.

FIGURE 72. Syntypes of Anolis boulengerianus: (a) MNHN 6554 dorsal view; (b) MNHN 6554 ventral view; (c) MNHN 1994.1671 ventral view; (d) MNHN 1994.1671 lateral view; (e) MNHN 1994.1670 dorsal view; (f) MNHN 1994.1670 ventral view.

FIGURE 73. Syntypes of Anolis boulengerianus: (a) MNHN 6554 dorsal view of head; (b) MNHN 6554 lateral view of head; (c) MNHN 6554 ventral view of head; (d) MNHN 6554 details of dorsum.

FIGURE 74. Holotype of Anolis subocularis (TCWC 8675): (a) dorsal view; (b) lateral view; (c) ventral view; (d) lateral view of head; (e) dorsal view of head; (f) ventral view of head. Scale bars equal 5.0 mm in (a–c) and 1.0 mm in (d–f), respectively.

FIGURE 75. Phylogenetic tree of specimens of the species related to Anolis boulengerianus from a maximum-likelihood analysis of DNA sequences of two mitochondrial genes: (a) 16S rRNA and (b) CO1. A scale bar is indicated. The numbers at nodes are bootstrap values. The tree is rooted with the species Anolis macrinii. For designation of OTUs see text.