Anolis microlepidotus Davis 1954

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: 16-24

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Anolis microlepidotus Davis 1954


Anolis microlepidotus Davis 1954  

Figs. 9–14 View FIGURE 9 View FIGURE 10 View FIGURE 11 View FIGURE 12 View FIGURE 13 View FIGURE 14

Anolis microlepidotus Davis 1954: 4   ; type locality: “four miles west of Chilpancingo, 5800 ft., Guerrero ”, Mexico. Holotype: TCWC 10276. Etheridge 1959, Davis & Dixon 1961, Smith et al. 1964, Duellman 1965, Smith & Taylor 1966, Fitch et al. 1976, Smith & Smith 1976, Lieb 1981, Flores-Villela 1993, Flores-Villela & Gerez 1994, Lieb 1995, Lieb 2000, Pérez-Ramos et al. 2000, 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, Wilson et al. 2013, Köhler et al. 2014

Anolis forbesi Smith & Van Gelder 1955: 147   ; type locality: “ 5 miles east of Izúcar de Matamoros, Puebla,” Mexico. Holotype: UIMNH 35553. Etheridge 1959, Smith et al. 1964, Smith & Taylor 1966, Fitch & Henderson 1973, Smith & Smith 1976, Lieb 1981, Flores-Villela 1993, Flores-Villela & Gerez 1994, Lieb 1995, Liner 2000, Urbina- Cardona & Flores-Villela 2010, Lieb 2001, Phillips 2003, Liner 2007, Fläschendräger & Wijffels 2009, Wilson & Townsend 2010, Köhler 2012 a, Wilson et al. 2013, Köhler et al. 2014

Anolis forbesorum: Michels & Bauer (2005   “2004“)

Norops forbesi: Savage & Guyer 1989   , Liner 2000, Nicholson 2002, Nicholson et al. 2012

Norops microlepidotus: Nicholson 2002   , Nicholson et al. 2012

Norops micropholidotus   [sic]: Savage & Guyer 1989

Diagnosis: A small species of Anolis   that differs from all other Mexican and Central American congeners by the combination of having (1) keeled ventral scales; (2) small middorsal scales (22–40 dorsal scales in one head length; 65–84 dorsal scales between levels of axilla and groin); (3) hind legs relatively short (fourth toe of adpressed hind leg reaching to ear opening or to a point between ear opening and eye; ratio of shank length/SVL 0.22–0.25); (4) flank scales usually somewhat heterogeneous with enlarged scales scattered among small flank scales; (5) postcloacal scales in males moderately to greatly enlarged or not enlarged at all; (6) male dewlap dull orange yellow with paler semicircular streaks and blotches ( Fig. 9 View FIGURE 9 ). Among the Anolis species   treated in this contribution, A. microlepidotus   is most similar to A. nebulosus   from which it differs in male dewlap coloration (dull orange yellow with paler semicircular streaks and blotches in A. microlepidotus   vs. uniform orange or orange with a white anterior margin in A. nebulosus   ) and smaller dorsal scales (number of dorsal scales in one head length 32–48, mean 41.2, in A. microlepidotus   vs. 26–38, mean 30.0, in A. nebulosus   ).

Description. Anolis microlepidotus   is a small anole (maximum recorded SVL 46.5 mm in males, 44.0 mm in females); dorsal head scales in internasal region rugose or keeled, in prefrontal, parietal, and frontal areas smooth or rugose; moderately deep prefrontal depression present, no parietal depression; 5–7 postrostrals; anterior nasal single, in contact with rostral and first supralabial; 5–8 internasals; canthal ridge sharply defined; scales comprising supraorbital semicircles well defined, posterior ones smooth, anterior ones with a rounded keel, largest scale in semicircles subequal or larger than largest supraocular scale; supraorbital semicircles usually broadly in contact, occasionally separated by one scale row; 0–3 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, much larger than ear opening; enlarged supraoculars separated from supraorbital semicircles by a complete row of small scales, or these scales narrowly in contact; 1–2 scales between enlarged supraoculars and superciliaries; 2–3 elongate superciliaries, anterior one longest, followed posteriorly by a series of 4–7 small rounded or squarish scales; usually 3 enlarged canthals, the second canthal largest; 4–7 scales between second canthals; 6–10 scales between posterior canthals; loreal region slightly concave, 15–40 mostly keeled (some smooth or rugose) loreal scales in a maximum of 3–6 horizontal rows; 6–7 supralabials to level below center of eye; suboculars keeled, in broad contact with supralabials (2–3 suboculars in contact with 2–5 supralabials); ear opening vertically oval, oriented slightly obliquely; scales anterior to ear opening granular, about twice as large as those posterior to ear opening; 6–8 infralabials to level below center of eye; 4–7 (usually 4) postmentals, outer pair slightly to distinctly larger than adjacent median postmental scales; 0–2 (commonly 1 or 2) enlarged sublabials in contact with infralabials on each side; faintly keeled granular scales present on chin and throat; male dewlap of moderate to large size extending onto chest; 8–10 horizontal gorgetal-sternal rows with 8–14 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 weakly to moderately keeled, subimbricate; 15–23 middorsal scale rows not enlarged, although slightly larger than laterals; usually a few enlarged scales scattered among granular laterals, or lateral scales more or less homogeneous; 65–84 dorsal scales along vertebral midline between levels of axilla and groin in males, 67–81 in females; 36–46 dorsal scales along vertebral midline contained in one head length in males, 32–48 in females; ventral scales on midsection about twice the size of largest dorsal scales; scales on midventer distinctly keeled, subimbricate, usually with rounded posterior margins, occasionally slightly mucronate; 45–64 ventral scales along midventral line between levels of axilla and groin in males, 47–53 in females; 28–38 ventral scales contained in one head length in males, 24–36 in females; 112–134 scales around midbody in males, 112–130 in females; tube-like axillary pocket absent; precloacal scales smooth or weakly keeled; males with or without two moderately to greatly enlarged postcloacal scales; tail moderately compressed in cross section, tail height/tail width 1.10–1.73 in males, 1.20–1.50 in females; all caudal scales strongly keeled and mucronate; lateral caudal scales, homogeneous although an indistinct division in segments is discernible; dorsal medial caudal scale row not enlarged; scales on dorsal surface of antebrachium strongly keeled, unicarinate, subimbricate to imbricate, mucronate; 17–20 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, 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 7 View TABLE 7 .

The coloration in life of an adult male from near Chilpancingo ( SMF 96213 View Materials ) was recorded as follows: Dorsal ground color Cinnamon Drab (50) with an Army Brown (46) interorbital bar and with Burnt Umber (48) spots in vertebral region of anterior body; hind limbs with Olive Brown (278) bands; tail with Tawny (60) bands; subocular region Pale Buff (1); ventral surface of body Light Buff (2); ventral surface of limbs and tail Drab (19); dewlap Pratt´s Rufous (72) with light Orange Yellow (77) bands and blotches; iris Vandyke Brown (282).

The coloration in life of another adult male from near Chilpancingo ( SMF 96214 View Materials ) was recorded as follows: Dorsal ground color Drab-Gray (256) grading mid-dorsally into Light Neutral Gray (297) and with Dark Grayish Brown (284) spots on anterior middorsum; interorbital bar and lateral stripe Glaucous (289); ventral surface of head Pale Buff (1); ventral surface of body, limbs, and tail Light Buff (2) with Drab (19) mottling; dewlap Mars Brown (25) with Clay Color (20) bars and blotches and with Cream White (52) gorgetals.

The coloration in life of an adult male from 8 km west Izúcar de Matamoros ( SMF 96383 View Materials ) was recorded as follows: Dorsal ground color Tawny Olive (17) with a Pale Pinkish Buff (3) vertebral region that contains Grayish Olive (274) chevrons; ventral surfaces of body, limbs and tail Pale Pinkish Buff (3); dewlap Burnt Orange (10) with Orange Yellow (8) streaks and blotches; iris Warm Sepia (40). The coloration in life of another adult male from 8 km west Izúcar de Matamoros ( SMF 96716 View Materials ) was recorded as follows: Dorsal surface of body Drab (19) with Cream Color (12) vertebral line grading into Chestnut (30) on tail and edged on dorsum by an inner line of Light Yellow Ocher (13) and by an outer line of Raw Umber (23); dorsal surfaces of head Tawny Olive (17) with Clay Color (18) interorbital bar; ventral surfaces of body, limbs, and tail Pale Horn Color (11); chin Pale Buff (1); dewlap Burnt Orange (10) with Light Buff (2) gorgetals; iris Sayal Brown (41).

The coloration in life of an adult female from 8 km west Izúcar de Matamoros ( SMF 96715 View Materials ) was recorded as follows: Dorsal ground color Chamois (84) with Pale Buff (1) vertebral and lateral line and with Grayish Horn Color (268) dots in vertebral region; dorsal surface of head Medium Fawn Color (257) with Amber (51) interorbital bar; dorsal surface of limbs Pale Cinnamon (55) with Olive-Brown (278) bars; ventral surfaces of body, limbs and tail Cream White (52) grading into Straw Yellow (53) towards tip of tail; dewlap Flame Scarlet (73) with suffusions of Medium Chrome Orange (75); iris Cinnamon (21).

The incompletely everted hemipenis of SMF 96384 View Materials ( Fig 14 View FIGURE 14 ) is a moderate-sized, bilobate organ; sulcus spermaticus bordered by well developed sulcal lips; an asulcate ridge present; apex strongly calyculate, truncus and asulcate ridge with transverse folds.

Geographic Distribution and Conservation. As currently known, Anolis microlepidotus   occurs at intermediate elevations (1245–1900 masl) from the Chilpancingo region northwest to about the city of Izúcar de Matamoros ( Fig. 15 View FIGURE 15 ). Given its small geographic range, degraded habitat, and threat from deforestation, we consider the conservation status of Anolis microlepidotus   as Critically Endangered based on criterion B1ab(iii) of the IUCN Red List Categories and Criteria ( IUCN 2012).

Natural History Notes: Anolis microlepidotus   seems to be quite adaptable with regard to the tolerated habitats. We found this species both in very dry situations such as in a open dry forest between the cities of Chilpancingo and El Zumpango del Río, in more mesic forest edge situations in Petaquillas near Chilpancingo, and within the sparse vegetation on a massive wall at the edge of a small village just a few kilometers west of Chilpancingo on the road to Omiltemi. All specimens collected by us were found at night while the lizards were sleeping on low vegetation 0.5 to 3.0 m above the ground.

Specimens examined ¾ Mexico: Guerrero: Amojileca, near Chilpancingo , 1550 m: SMF 96213 View Materials   ; between Chilpancingo and Zumpango del Río , 1385 m: SMF 93844 View Materials   ; Chilpancingo , 1260 m: MCZ   R-33696–97; 4 mi W Chilpancingo : TCWC 10009 View Materials   ; Petaquillas, near Chilpancingo , 1245 m: IBH 26572 View Materials –73 View Materials , SMF 96214 View Materials   ; Omiltemi : 1900 m, UTA   R-4513; 2.5 mi S Almolonga : MCZ R   78727, TCWC 10276 View Materials , 11395 View Materials , 11397 View Materials   ; 2 km E Tixtla , 1360 m: KU 105703 View Materials   ; 1–3 km al E de la Escalera , 1558 m: IBH 6438 View Materials   ; 0.5–1.0 km al W de Ixcateopan , 1390 m: IBH 6451 View Materials   . Puebla: 8 km E Izúcar de Matamoros , 1350 m: UIMNH 35553 View Materials , SMF 96715 View Materials –17 View Materials   ; San Juan Epatlan , 1373 m: SMF 96382 View Materials –84 View Materials   ; 4.8 km W Teotlalco , 1128 m: IBH 23613 View Materials   .

Taxonomy of the Mexican anoles related to Anolis nebuloides Bocourt 1873  

The species of anoles related to Anolis nebuloides   are readily differentiated from all other Mexican and Central American congeners by having a combination of (1) several moderately to greatly enlarged dorsal scales, often larger than ventral scales; (2) moderately to strongly keeled ventral scales; (3) an oval patch of usually three greatly enlarged supraorbital scales; and (4) usually a pair of greatly enlarged postcloacals in males. Anolis nebuloides   was described by Bocourt (1873) based on four specimens (now MNHN 2494, 1994.0984–86, examined by GK; Fig. 16 View FIGURE 16 ) from “Putla, province d’Oaxaca ( Mexique).” This syntype series consists of three adult females (MNHN 2494, 1994.0984, 1994.0986) in favorable condition and an adult male (MNHN 1994.0985) in poor condition (very soft); all four specimens lack most of the tail. In 1933, Smith described his new species Anolis megapholidotus   based on specimens collected “between Rincon and Cajones (about 40–45 kilometers south of Chilpancingo), Guerrero, Mexico ” (holotype: FMNH 100105, examined by GK; Fig. 17 View FIGURE 17 ). The type locality was later stated to correspond to what is now known as Agua de Obispo ( Smith & Taylor 1950a: 61). Holman (1964) described Anolis simmonsi   based on a single adult male (UIMNH 52899, examined by GK; Fig. 18 View FIGURE 18 ) from “Rio Canoa, 16.25 miles west of Pinotepa National, Oaxaca, Mexico.” Interestingly, Holman (1964) compared his new species only with A. pygmaeus Alvarez   del Toro & Smith 1956 and did not recognize the similarity of UIMNH 52899 with A. megapholidotus   and A. nebuloides   . Recently, Nieto Montes de Oca et al. (2013) synonymized Anolis simmonsi Holman 1964   with Anolis nebuloides Bocourt 1873   .

In order to shed some light on the genetic and morphological variation in the populations of nebuloides   -like anoles in the Mexican states of Oaxaca and Guerrero, two of us (GK and RGT) visited numerous localities where these lizards occur including the areas of the type localities of the nominal species nebuloides   , megapholidotus   , and simmonsi   . For our analysis, we defined seven OTUs in this complex: OTU1 is formed by specimens from Agua de Obispo and surroundings, southcentral Guerrero; OTU2 by specimens from Santa Cruz el Rincón and surroundings, eastern Guerrero; OTU3 by specimens from Iliatenco and surroundings, central eastern Guerrero; OTU4 by specimens from Putla, western Oaxaca; OTU5 by specimens from San Juan Colorado, Villanueva and surroundings, southwestern Oaxaca; OTU6 by specimens from along the road from San Gabriel Mixtepec to El Vidrio, southern central Oaxaca; OTU7 by specimens from the foothills of the Sierra Madre del Sur, north of Pochutla, southern central Oaxaca.

The results of a molecular genetic analysis based on CO1 and 16s gene fragments indicated a high degree of genetic differentiation among several populations ( Fig. 19 View FIGURE 19 ). Our molecular results reveal several distinct genetic clusters. The analysis of the CO1 sequences show five genetic clusters that are separated by mean genetic distances of 10.1–14.0%. Geographic distance by itself is not responsible for these genetic distances, as evidenced by the low genetic differentiation between our OTUs 2 and 3 (6.0%) and between OTUs 4 and 5 (5.0%), respectively. The analysis of the 16s sequences indicate the same genetic clusters as in the CO1 tree but with smaller average genetic distances among our OTUs 1, 2, and 3 and also between OTUs 2 and 6. In the analysis of combined CO1 and 16s sequences, these five genetic clusters are recovered and have mean genetic distances of 7.9–11.8%. We take this high level of genetic differentiation among these five genetic clusters as evidence for lack of gene flow and in conclusion recognize these five clusters as species level units. In external morphology, these five genetic clusters are more conservative and not easily differentiated ( Table 8 View Table 8 ). However, subtle differences among most of these clusters are evident, supporting the recognition of each of these as a distinct species. Also, we observed differences in hemipenial morphology among these clusters. Thus, the five species we recognize are Species A (OTU 1), Species B (OTUs 2 and 3), Species C (OTUs 4 and 5), Species D (OTU 6), and Species E (OTU 7). Our Species A is from the type locality of A. megapholidotus   ; thus, this is the valid name for this species. Our OTU 4 (species C) is from the type locality of A. nebuloides   and therefore this name has to be applied to this species. The type locality of the nominal species A. simmonsi   is near our OTU 5 ( A. nebuloides   sensu stricto); thus, A. simmonsi   remains in the synonymy of A. nebuloides   . Therefore, no names are available for our Species B, D, and E, and therefore we describe each of them as a new species below.


Forschungsinstitut und Natur-Museum Senckenberg


Museum of Comparative Zoology


University of Texas at Arlington


Departamento de Geologia, Universidad de Chile


Biodiversity Institute, University of Kansas


Universidad Nacional Autonoma de Mexico, Instituto de Biologia














Anolis microlepidotus Davis 1954

Köhler, Gunther, Pérez, Raúl Gómez Trejo, Petersen, Claus Bo P., Méndez, Fausto R. & Cruz, De La 2014

Anolis forbesi

Smith, H. M. & van Gelder, R. G. 1955: 147

Anolis microlepidotus

Davis, W. B. 1954: 4