Anolis nebuloides : Smith 1939 Anolis quercorum : Ramirez-Bautista et al. 2002a 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 Köhler, Gunther Pérez, Raúl Gómez Trejo Petersen, Claus Bo P. Méndez, Fausto R. Cruz, De La Zootaxa 2014 2014-09-19 3862 1 1 210 Köhler & Pérez & Petersen & Méndez & Cruz, 2014 Köhler & Pérez & Petersen & Méndez & Cruz 2014 [151,335,604,630] Reptilia Dactyloidae Anolis Animalia Squamata 69 70 Chordata species carlliebi sp. nov.   Figs. 52–57    Anolis nebuloides: Smith 1939   Anolis quercorum: Ramirez-Bautista et al.2002a,b, Ramirez-Bautista 2003, Canseco-Márquez & Gutiérrrez-Mayén 2006, 2010     Holotype. SMF96714, an adult female from Ixtlán de Juárez, university campus ( 17.316350°N, 96.483470°W, WGS84),  1945 m, Estado de Oaxaca, Mexico; collected  13 June 2013by Raúl Gómez Trejo Pérez. Field tag number GK-4501.    Paratypes. All from the Mexican state of Oaxaca: GK-4502( IBHuncatalogued), same collecting data as holotype;  UMMZ 99820( two specimens), from Ixtlán de Juárez, collected  11 May 1947by Hellmuth Wagner;  UTAR-8421, from 0.9 mi N (by road) El Cerezal on Mexican Hwy175, Sierra Aloapaneca, by Jonathan A. Campbell;  UTAR-52816–17, from Tutepetongo, Municipio Cuicatlán, 17.732°N, 96.866°W, 1619 m, by Eric N. Smith;  MVZ 164275–76, from Ixtlán de Juárez, Vivero Rancho Teja, 17°19’50’’N, 96°29’14’’W.  GK-4502 ( IBHuncatalogued), MVZ144181, 164275–76, and UMMZ99820 are adult females, UTAR-52816 is an adult male, and UTAR-8421 is a subadult male.  Referred specimens ⎯  Mexico: Oaxaca: Cuicatlán: USNM 47614;  2miE Ixtepeji: KU320768.   Diagnosis. A small species (our Species B of the  Anolis quercorumcomplex, see above; SVL in largest male 46.0 mm, largest female 41.0 mm) of the genus  Anolis(sensu Poe 2004)that differs from all Mexican and Central American anoles except  A. quercorumand a species described below (our Species C of the  A. quercorumcomplex) 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 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 0.23–0.25; (5) circumnasal usually in contact with first supralabial; and (6) a large pink dewlap in males and a very small red or pink dewlap in females.  Anolis carlliebidiffers from  A. quercorumand the species described below (our Species C of the  A. quercorumcomplex) 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; and (2) ratio shank length/HL. Also,  Anolis carlliebidiffers from  A. quercorumand the species described below (our Species C of the  A. quercorumcomplex) in hemipenis morphology (hemipenis unilobed in  A. carlliebivs. slightly to distinctly bilobed in the other two species) and by substantial genetic distances (see above).   Description of the holotype. Adult female, as indicated by the shape of the base of tail and the absence of a large dewlap; SVL 45.0 mm; tail incomplete; tail moderately compressed in cross section, tail height 2.2 mmand width 1.8 mm; axilla to groin distance 20.2 mm; head length 11.1 mm, head length/SVL ratio 0.25; snout length 4.9 mm; head width 6.6 mm; longest toe of adpressed hind limb reaching to ear opening; shank length 10.8 mm, shank length/head length ratio 0.97; longest finger of extended forelimb reaching to tip of snout; longest finger of adpressed forelimb reaching to a point 5.3 mmin front of anterior insertion of hind limbs. Dorsal head scales in internasal region keeled, unicarinate; other dorsal head scales rugose or keeled; 6 postrostrals; 6 scales between nasals; 1 elongate prenasal scale on each side, in contact with both rostral and first supralabial; circumnasal separated from supralabial by one scale row; prefrontal depression absent; supraorbital semicircles well developed, narrowly in contact medially; supraorbital disc composed of an oval patch of 3 greatly enlarged, weakly keeled scales; circumorbital row complete on right side, incomplete on left side with on enlarged supraorbital scale in contact with supraorbital semicircles; 3 elongated, strongly overlapping superciliaries, the two anterior ones much larger than posterior one, followed posteriorly by 5 to 6 roundish to squarish scales of moderate size; 1 to 2 rows of small keeled scales extending between enlarged supraorbitals and superciliaries; parietal depression absent; interparietal scale well developed, 1.8 x 1.3 mm(length x width), surrounded by scales of moderate size; 1 scale present between interparietal and supraorbital semicircles; canthal ridge distinct, composed of 3 large (second largest) and 3 small anterior canthal scales; 5 scales present between second canthals; 8 scales present between posterior canthals; 19 (right)– 20 (left) mostly keeled loreal scales in a maximum of 5 horizontal rows; 4 keeled subocular scales arranged in a single row; 6 (right)– 5 (left) supralabials to level below center of eye; 3 suboculars broadly in contact with 4 supralabials; ear opening 0.5 x 1.0 mm (length x height); mental distinctly wider than long, completely divided medially, bordered posteriorly by 4 postmentals, outer ones much larger than median ones, a single small scale interspersed on each side between postmental, infralabial, and mental; 6 infralabials to level below center of eye; one greatly enlarged sublabial on each side; keeled granular scales present on chin and throat; dewlap very small; dorsum of body with weakly keeled, subimbricate, non-mucronate scales; about 18 medial rows slightly to moderately enlarged; largest dorsal scales about 0.39 x 0.34 mm(length x width); about 32 medial dorsal scales in one head length; about 62 medial dorsal scales between levels of axilla and groin; lateral scales keeled, granular and more or less homogeneous in size, average size 0.20 mmin diameter; ventrals at midbody strongly keeled, imbricate, mucronate, about 0.47 x 0.49 mm(length x width); about 22 medial ventral scales in one head length; about 51 medial ventral scales between levels of axilla and groin; 124 scales around midbody; all caudal scales keeled; middorsal caudal scales moderately enlarged, not forming a crest; lateral caudal scales without whorls of enlarged scales, although an indistinct division in segments is discernible; no tube-like axillary pocket present; scales on dorsal surface of forelimb weakly keeled; digital pads dilated, dilated pad about 3 times width of non-dilated distal phalanx; distal phalanx narrower than and raised from dilated pad; 24 (right)–22 (left) lamellae under phalanges II–IV of Toe IV of hind limbs; 7 (right) scales under distal phalanx of Toe IV of hind limbs.   FIGURE 52.  Anolis carlliebiin life: (a) two males from Ixtlán de Juárez, Oaxaca; (b) a male from San Juan Tepeuxila, Oaxaca. Photos by Christopher Ehecatl Vega Flores (a) and Luis Canseco Marquéz (b).   FIGURE 53.Holotype of  Anolis carlliebi(SMF 96714): (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 54.  Anolis carlliebi: head scalation in (a,c,e) MVZ 144181; (b,d,f) UTA R-52816. Scale bars equal 1.0 mm.   FIGURE 55.  Anolis carlliebi: nasal region in (a) UTA R-52818; (b) SMF 96714; superciliary region in (c) UTA R-52818; (d) MVZ 144181; chin region in (e) MVZ 164275; (f) SMF 96714; cloacal region in (g) UTA R-52816. Scale bars equal 1.0 mm.   FIGURE 56.  Anolis carlliebi: dorsal scales in (a) UTA R-52818; (b) SMF 96714; flank scales in (c) UTA R-52818; (d) SMF 96714; ventral scales in (e) UTA R-52816; (f) SMF 96714. Scale bars equal 1.0 mm. Coloration in life was recorded as follows: Dorsal ground color Drab (19); dorsal surface of head Clay Color (18); with Raw Umber (23) interorbital bar and Cream Color (12) lines from eyes to neck; vertebral region Light Buff (2) with Mars Brown (25) chevrons; dorsal surface of limbs and tail Antique Brown (24); ventral surface of body, limbs and tail Pale Horn Color (11) with Pale Pinkish Buff (3) speckles; dewlap Spectrum Orange (9) with Pale Buff (1) gorgetals; iris Cinnamon-Rufous (31). Coloration after four months preservation in 70% ethanol was recorded as follows: Dorsal surfaces of head and body Medium Neutral Gray (298) with large confluent Dusky Brown (285) blotches on middorsum and with a Grayish Horn Color (268) postocular stripe, edged with Burnt Umber (48); limb Medium Neutral Gray (298) suffused with Dusky Brown (285); tail Drab (19) with faint Hair Brown (277) vertical bands; ventral surfaces of head, body, limbs, and tail Pale Buff (1) with a suffusion of Medium Neutral Gray (298) on venter; finger and toe pads Vandyke Brown (282).  Variation. The paratypesagree well with the holotypein general appearance, morphometrics, and scalation ( Table 9). They differ in some scalation characters as follows: The number of slightly to moderately enlarged dorsal scale rows varies from 14 to 20. All specimens have a single elongate prenasal. Suboculars in contact with supralabials (2–4 suboculars in contact with 1–4 supralabials) in all specimens examined. All males have a pair of greatly enlarged postcloacal scales. The coloration in life of an adult female paratypefrom Ixtlán de Juárez (GK-4502, IBH) was recorded as follows: Dorsal ground color Drab (19) with Drab-Gray (256) vertebral line, Olive Brown (278) dots and a Light Buff (2) lateral line; dorsal surface of head Verona Brown (37) with Brussels Brown (33) interorbital bar; dorsal surface of limbs Cinnamon-Rufous (31) with Yellow Ocher (14) bars; dorsal surface of tail Drab (19) with suffusion of Robin Rufous (29); ventral surface of body, limbs and tail Light Buff (2) with Pale Pinkish Buff (3) suffusions; dewlap Orange Yellow (8) grading into Burnt Orange (19) towards front and Pale Buff (1) gorgetals; iris Natal Brown (49).   FIGURE 57.Hemipenis of  Anolis carlliebi(UTA R-52816): (left) sulcate view; (right) asulcate view. Scale bar equals 1.0 mm. The completely everted hemipenis of UTA-R 52816 ( Fig. 57) is a small unilobate organ; sulcus spermaticus bordered by weakly developed sulcal lips and opening into a single apical field; no asulcate processus present; no surface ornamentation discernable.   Etymology. The name  carlliebiis a patronym honoring Carl S. Lieb who has contributed substantially to our knowledge of Mexican anoles.  Natural History Notes. The two SMF typespecimens of  Anolis carlliebiwere collected during a 4.5 hours night search in a pine oak forest at the campus of the Ixtlán University. The specimens were found sleeping on twigs between 0.3 and 1.0 m above the ground. Most of the following is summarized from the information provided in Ramirez-Bautista et al.(2002a,b). The habitat of  Anolis carlliebiincludes xerophilic brush land, thorn forest, and deciduous tropical forest, heavily altered in most of the species’ range by human activities. During daytime, this lizard is mostly found on the leaf litter covered floor, but also on rocks and low vegetation. It is also found in disturbed areas, dump-yards, and along house fences, and is easy to spot at night sleeping on shrub twigs. It feeds on small beetles ( Coleoptera, family Curculionidae), true bugs (Homoptera, family Cicadellidae), and Hymenoptera(families Ichneumonidaeand Braconidae) ( Canseco-Márquez & Gutiérrrez-Mayén 2010). The limited available data suggest seasonal reproduction in  A. carlliebi( Ramirez-Bautista et al.2002b). CansecoMárquez & Gutiérrrez-Mayén (2010) noted that one femalecollected in May in Santiago Dominguillo laid an egg and that a male collected in October in San Lucas Teteletitlán had enlarged testicles. Ramirez-Bautista et al.(2002a)reported a male courting a female on a  Prosopistree in September. Canseco-Márquez & Gutiérrrez-Mayén (2010)reported observations of juveniles in November in Santa María Texcatitlán.  Geographic Distribution and Conservation. As currently known,  Anolis carlliebiis restricted to the Tehuacán-Cuicatlán Valley and the adjacent Ixtlán Valley in the northern central portion of the Mexican State of Oaxacaat elevations of 650 to 2,515 masl( Fig. 71). Given its usual abundance wherever this species occurs, it seems justified to classify  A. carlliebias Least Concern based on the IUCN Red List Categories and Criteria ( IUCN 2012). 4013201413 SMF 96714 2013-06-13 SMF de Juarez Mexico 1945 17.31635 Ixtlan de Juarez 1 -96.48347 69 70 WGS84, GK-4501 1 Oaxaca holotype 4013201513 SMF 96714 1947-05-11 IBH de Juarez & Jonathan A. Campbell Mexico Ixtlan de Juarez 1619 17.732 Oaxaca 76 -96.866 Hellmuth Wagner 69 70 GK-4502 1 Oaxaca paratype 4013201327 SMF 96714 [267,1436,987,1012] 1947-05-11 IBH de Juarez & Jonathan A. Campbell Mexico Ixtlan de Juarez 1619 17.732 Oaxaca 76 -96.866 Hellmuth Wagner 69 70 GK-4502 1 Oaxaca paratype 4013201445 SMF 96714 1947-05-11 IBH de Juarez & Jonathan A. Campbell Mexico Ixtlan de Juarez 1619 17.732 Oaxaca 76 -96.866 Hellmuth Wagner 69 70 GK-4502 1 Oaxaca paratype 4013201317 SMF 96714 1947-05-11 IBH de Juarez & Jonathan A. Campbell Mexico Ixtlan de Juarez 1619 17.732 Oaxaca 76 -96.866 Hellmuth Wagner 69 70 GK-4502 1 Oaxaca paratype 4013201336 SMF 96714 [232,1235,1099,1124] 1947-05-11 IBH de Juarez & Jonathan A. Campbell Mexico Ixtlan de Juarez 1619 17.732 Oaxaca 76 -96.866 Hellmuth Wagner 69 70 GK-4502 1 Oaxaca paratype 4013201456 [434,948,1210,1236] USNM Mexico Cuicatlan 69 70 USNM 47614 1 Oaxaca 4013201481 KU 320768 [959,1269,1211,1236] KU Mexico Ixtepeji 69 70 1 Oaxaca