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 Author Köhler, Gunther gkoehler@senckenberg.de Author Pérez, Raúl Gómez Trejo Author Petersen, Claus Bo P. Author Méndez, Fausto R. Author Cruz, De La text Zootaxa 2014 2014-09-19 3862 1 1 210 journal article http://dx.doi.org/10.11646/zootaxa.3862.1.1 1175-5326 4930959 3FA375FE-E4E0-4509-BE02-EE5E786B07C6 Anolis nebuloides Bocourt 1873 Figs. 45–50 Anolis nebuloides Bocourt 1873: 74 ; type locality: “Putla, province d’Oaxaca ( Mexique )” Syntypes : MNHN 2494, 1994.0984- 86. Troschel 1874 , O’Shaughnessy 1875 , Boulenger 1885 , Günther 1885 –1902, Cope 1887 , Barbour 1934 , Smith 1939 , Burt & Myers 1942 , Smith 1944 , Stuart 1948 , Smith & Taylor 1950a , b, Beltrán 1953 , Davis 1954 , Etheridge 1959 , Holman 1964 , Etheridge 1965 , Hardy & McDiarmid 1969 , Smith & Smith 1973 , Smith et al. 1972, Fitch & Henderson 1973 , Smith & Smith 1976 , Lieb 1981 , Fitch & Hillis 1984 , Lister & García Aguayo 1992 , Flores-Villela 1993, Flores- Villela & Gerez 1994, Lieb 1995 , 2001 , Poe 2004 , Liner 2007 , Fläschendräger & Wijffels 2009 , Flores-Villela et al. 2010 , Urbina-Cardona & Flores-Villela 2010 , Wilson & Townsend 2010 , Wilson et al. 2013 , Nieto Montes de Oca et al. 2013 , Köhler et al. 2014 , Köhler 2014 Anolis simmonsi Holman 1964: 52 ; type locality: “Rio Canoa, 16.25 miles west of Pinotepa National, Oaxaca , Mexico ”. Holotype : UIMNH 52899. Smith et al. 1964 , Smith & Taylor 1966 , Smith 1968 , Webb & Baker 1969 , Fitch & Henderson 1973 , Smith & Smith 1976 , Flores-Villela 1993, Flores-Villela & Gerez 1994 , Phillips 2003 , Fläschendräger & Wijffels 2009 , Urbina-Cardona & Flores-Villela 2010 , Wilson & Townsend 2010 , Wilson et al. 2013 , Köhler 2012 a , Nieto Montes de Oca et al. 2013 Norops nebuloides : Savage & Guyer 1989 , Nicholson 2002 , Castiglia et al. 2010, Nicholson et al. 2012 Norops simmonsi : Savage & Guyer 1989 , Nicholson 2002 , Nicholson et al. 2012 Diagnosis . A small species (SVL in largest male 46.0 mm, largest female 43.0 mm) of the genus Anolis (sensu Poe 2004 ) that differs from all Mexican and Central American anoles except A. megapholidotus , A. nietoi , A. stevepoei , and A. zapotecorum by having a combination of (1) strongly keeled ventral scales; (2) usually a patch of three greatly enlarged supraocular scales; (3) 10–12 rows of greatly enlarged dorsal scales that are larger than ventral scales; (4) short hind legs, longest toe of adpressed hind leg usually reaching to posterior margin of eye, occasionally to a point between levels of ear opening and eye or to mideye (to ear opening in single female), ratio shank length/SVL 0.24–0.29; (5) circumnasal usually in contact with first supralabial; and (6) a large pinkish red dewlap in males and a very small pink dewlap in females ( Fig. 45 ). Anolis nebuloides differs from A. stevepoei , and A. zapotecorum by having (1) a rather abrupt transition between the enlarged dorsal scales and lateral body scales (vs. a more gradual transition over 3 to 4 scales); (2) the edges of the field of enlarged dorsal scale rows well defined and in a more or less straight line (vs. the edges of this field forming an undulating line due to single enlarged scales or groups thereof outside the main field) (3) and by having a more reddish male dewlap (vs. pink). Anolis nebuloides differs from A. megapholidotus and A. nietoi in the ranges and average values of several morphometric and pholidotic characters, most obvious in (1) number of middorsal scales in one head length; and (2) number of subdigital lamellae on Phalanges II–IV of Toe IV (see Table 8 ). FIGURE 46. Anolis nebuloides in life. (a) SMF 96215; (b) SMF 96215; (c) IBH 26510; (d) SMF 96217; (e) SMF 96216; (f) SMF 96218. (a–c) are males, (d–f) are females. FIGURE 47. Anolis nebuloides : head scalation in (a,c,e) SMF 96215; (b,d,f) SMF 96217. Scale bars equal 1.0 mm. Description. Anolis nebuloides is a small anole (maximum recorded SVL 46.0 mm in males, 43.0 mm in females); dorsal head scales in internasal region strongly keeled, mostly unicarinate, other dorsal head scales mostly keeled in prefrontal region, in parietal region mostly weakly to strongly keeled, some rugose; shallow to deep prefrontal depression present, shallow parietal depression; 5–6 postrostrals; anterior nasal single, 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 usually smaller than largest supraocular scale; supraorbital semicircles usually separated from each other by one complete row of scales, commonly these scales in narrow or broad contact; 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 larger than ear opening, occasionally subequal to ear opening; enlarged supraoculars usually a patch of 3 greatly enlarged scales in a single row, rugose to weakly keeled, separated from supraorbital semicircles by a complete row of small scales, or these scales narrowly in contact; 2–3 scales between enlarged supraoculars and superciliaries; 3 elongate superciliaries, anterior one longest, followed posteriorly by a series of 3–4 rounded or squarish scales of moderate size; usually 3 enlarged canthals, the second canthal largest; 6–9 scales between second canthals; 7–10 scales between posterior canthals; loreal region slightly concave, 20–33 mostly keeled (some smooth to rugose) loreal scales in a maximum of 5–7 (commonly 5 or 6) horizontal rows; 5–7 supralabials to level below center of eye; suboculars keeled, in contact with supralabials (1–3 suboculars in contact with 1–4 supralabials); ear opening vertically oval; scales anterior to ear opening keeled, juxtaposed, about four times larger than granulars posterior to ear opening; 5–7 infralabials to level below center of eye; 4–6 postmentals (usually 4), outer pair slightly to distinctly larger than adjacent median postmental scales; 0–2 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; 8–10 horizontal gorgetal-sternal rows with 9–14 scales per row; modal number of marginal pairs 2–4; female dewlap very small or small; a nuchal crest and a dorsal ridge present in males; 10–14 middorsal scale rows moderately to greatly enlarged, abruptly larger than lateral scales; scales on middorsum strongly keeled, subimbricate with rounded posterior margins, some slightly mucronate; enlarged dorsal scales abruptly larger than keeled granular lateral scales; lateral scales more or less homogeneous; 34–44 dorsal scales along vertebral midline between levels of axilla and groin in males, 36–44 in females; 20–24 dorsal scales along vertebral midline contained in one head length in males, 18–24 in females; ventral scales on midsection smaller than largest dorsal scales; scales on midventer strongly keeled, subimbricate to imbricate, mucronate; 31–45 ventral scales along midventral line between levels of axilla and groin in males, 33–38 in females; 20–33 ventral scales contained in one head length in males, 18–24 in females; 96–132 scales around midbody in males, 90–116 in females; tube-like axillary pocket absent; precloacal scales weakly to strongly keeled; males with or without a pair of moderately to greatly enlarged postcloacal scales; tail moderately compressed in cross section, tail height/tail width 1.14–1.64 in males, 1.24–1.27 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, slightly mucronate; scales on dorsal surface of antebrachium strongly keeled, unicarinate, subimbricate to imbricate, slightly mucronate; 20–26 subdigital lamellae on Phalanges II–IV of Toe IV of hind limbs; 6–7 subdigital lamellae on distal phalanx o 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 8 . FIGURE 48. Anolis nebuloides : nasal region in (a) SMF 96219; (b) SMF 96217; superciliary region in (c) SMF 96215; (d) SMF 96217; chin region in (e) SMF 96219; (f) SMF 96217; cloacal region in (g) SMF 96215; (h) SMF 96411. Scale bars equal 1.0 mm. FIGURE 49. Anolis nebuloides : dorsal scales in (a) SMF 96215; (b) SMF 96217; flank scales in (c) SMF 96215; (d) SMF 96217; ventral scales in (e) SMF 96219; (f) SMF 96217. Scale bars equal 1.0 mm. The coloration in life of an adult male ( SMF 96215 ; Fig. 46a ) from near Putla was recorded as follows: Dorsal ground color Cinnamon (21) with Drab (19) lateral stripe edged above and below by Ferruginous (35); ventral surface of head Light Buff (2); venter Pale Pinkish Buff (3); ventral surfaces of limbs and tail Drab (19); dewlap Spinal Pink (235) with Chamois (84) marginals and Cream White (52) gorgetals; iris Warm Sepia (40). The color in life of an adult female ( SMF 96216 ; Fig. 46e ) from near Putla was recorded as follows: Dorsal ground color Verona Brown (37) with Burnt Umber (48) chevrons; dorsal surfaces of limbs Burnt Umber (48) with Burnt Siena (38) bars on shank; dorsal surface of tail Verona Brown (37) with Dark Yellow Buff (54) dashes and chevrons; ventral surface of head Light Buff (2) with a suffusion of Buff (5); venter Pale Pinkish Buff (3) with a suffusion of Light Orange Yellow (7); ventral surface of limbs and tail Drab (19); dewlap Spinal Pink (235) with Pale Buff (1) gorgetals; iris Maroon (39). The color in life of another adult female ( SMF 96218 ; Fig. 46f ) from near Putla was recorded as follows: Mid-dorsal stripe Tawny (60); edged by Russet (44); dewlap Dark Rose (238); with Cream White (53) gorgetals. The coloration in life of an adult male from the road between San Juan Colorado and Villanueva ( SMF 96414 ) was recorded as follows: Dorsal ground color Tawny Olive (17) with a Vandyke Brown (282) vertebral line, Cream White (52) spots and a lateral line that is edged by Salmon Color (59); dorsal surface of head Cinnamon (255); ventral surface of body Pale Sulphur Yellow (92) with suffusions of Light Flesh Color (250); chin with Medium Neutral Gray (298) bars; dewlap Spinal pink (253); iris Buff (115) . The coloration in life of an adult female from the road between San Juan Colorado and Villanueva ( SMF 96409 ) was recorded as follows: Dorsal surface of head Medium Chrome Orange (75); dorsal ground color of body Buff (15) with a Salmon color (83) vertebral line edged by Grayish Horn Color (268) and some Smoke Gray (267) suffusions; ventral surface of body Chamois (84) with Pink (242) suffusions; ventral surface of legs Raw Sienna (32); dewlap Rose (234); iris Cinnamon (255). The coloration in life of another adult female from the road between San Juan Colorado and Villanueva ( SMF 96415 ) was recorded as follows: Dorsal ground color Cinnamon (21) with a Warm Sepia (40) vertebral line that divides into three portions in neck area; dorsal surface of limbs head and tail Brussels Brown (33); ventral surface of body Cream Color (12); ventral surface of limbs and tail Tawny Olive (17); chin with Drab (19) bars; dewlap Medium Rose (233); iris Clay Color (20). The completely everted hemipenis of SMF 96219 ( Fig. 50 ) is a bilobate organ; sulcus spermaticus bordered by well developed sulcal lips and opening at base of apex into two apical fields void of ornamentation, one on each lobe; distal portion of sulcal lips with several transverse folds at level of bifurcation; a distinct asulcate ridge and a knob-like asulcate processus present; apex strongly calyculate, truncus and asulcate ridge with transverse folds. Natural History Notes: Anolis nebuloides was observed during the day in leaf litter and on the base of tree trunks in a pine-oak forest some kilometers north of the town of Putla where it appeared to be very abundant; only two specimens where caught at night sleeping in tall grass on the edge of the highway. FIGURE 50. Hemipenis of Anolis nebuloides (SMF 96219): (left) sulcate view; (center) asulcate view; (right) lateral view. Scale bar equals 1.0 mm. Geographic Distribution and Conservation . As currently known, Anolis nebuloides is restricted to the Pacific versant of Sierra Madre del Sur in the southwestern portion of the Mexican State of Oaxaca at elevations between 980 and 1575 masl ( Fig. 40 ). Given its usual abundance wherever this species occurs, it seems justified to classify A. nebuloides as Least Concern based on the IUCN Red List Categories and Criteria ( IUCN 2012 ). Specimens examined ⎯ Mexico : Guerrero : road from San Juan Colorado to Villanueva , 1170 m : SMF 96408 - 17 ; Oaxaca : Putla de Guerrero : MNHN 2494 , 1994.0984 – 86 ; 10 km N Putla de Guerrero , on Highway 125, 1005 m: USNM 304887 ; near Putla de Guerrero , 980 m : IBH 26508 , 26510 , 26515–16 , 26519–20 , 26524 , SMF 96215 –20 , 97011–12 ; road from Putla de Guerrero to San Andres Chicahuaxtla , Municipio de Putla de Guerrero de Guerrero , 1375 m : UTA R-52814. Taxonomy of the Mexican anoles related to Anolis quercorum Fitch 1978 In 1978, Fitch described his new species Anolis quercorum based on a series of specimens collected “ 26 km SE Nochixtlán ( 2.5 km NW Cuesta Blanca , Highway 190), Oaxaca , Mexico ” ( holotype KU 176050 ). Since the original description, this species showed up on several checklists but little remains known about its morphological variation and natural history . For our analysis, we defined five OTUs in this complex: OTU1 is formed by specimens from Tamazulapan , northwestern Oaxaca ; OTU2 by specimens from Cuesta Blanca , western central Oaxaca ; OTU3 by specimens from the valleys of Ixtlan de Juárez and Tehuacán-Cuicatlán , northwestern Oaxaca and adjacent Puebla ; OTU4 by specimens from Matatlán , eastern central Oaxaca ; OTU5 by specimens from Cerro Sacamecates , southeastern Oaxaca . The analysis of the 16s sequences revealed three well-supported clades, i.e., (1) our OTUs 1 and 2, (2) our OTU 3, and (3) our OTUs 4 and 5 ( Fig. 52a ). Mean genetic distances among these clades are 4.8–7.1%. Geographic distance by itself is not responsible for these genetic distances as evidenced by the low genetic differentiation between our OTUs 1 and 2 (1.5%) and between OTUs 4 and 5 (3.4%), respectively. The analysis of the CO1 sequences ( Fig. 52b ) indicates the same genetic clusters as in the 16s tree. We take this high level of genetic differentiation among these three genetic clusters as evidence for lack of gene flow and in conclusion recognize these three clusters as species level units. In external morphology, these three genetic clusters are more conservative and not easily differentiated ( Table 9 ). 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 three species we recognize are Species A (OTUs 1 and 2), Species B (OTU 3), Species C (OTUs 4 and 5). Our Species A includes the type locality of A. quercorum ; thus, this is the valid name for this species. No names are available for our Species B and C, and therefore we describe each of them as a new species below.