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 [151,391,151,177] Reptilia Dactyloidae Anolis Animalia Squamata 47 48 Chordata species zapotecorum sp. nov.   Figs. 33–39     Holotype. SMF96425, an adult male from Pluma Hidalgo( 15.942410°N, 96.430440°W),  1350 m, Estado de Oaxaca, Mexico; collected  4 March 2013by Gunther Köhlerand Raúl Gómez Trejo Pérez. Field tag number GK- 4369.    Paratypes. IBH 26997–27004,  SMF 96418–24, 96426–27, same collecting data as holotype.  IBH 26998–27001, 27003–27004,  SMF96419, 96421–22, 96424, 96426–27are males,  IBH26997, 27002,  SMF96418, 96420, 96423are females.  Referred specimens ⎯  Mexico: Oaxaca: 7 miS of la Candelaria,  358 m: UIMNH 8906;  2.5 kmS Candelaria,  394 m: KU137684; road from San Jose Pacíficoto CandelariaLoxicha,  500–1469 m: UTA R-51711–16; road from Pochutlato Candelaria,  380 m: UTA R-53606; Finca de Juan García, near Santiago La Galera,  1010 m: IBH 26579, SMF 96246–47;  5.1 kmS Jalatengo,  1017 m: KU 137686–87;  Santiago la Galera,  1160 m: IBH 26513–14, 26521–22, 26578, 26582, 26612, SMF96240, 96248–51; near Tierra Blanca,  1287 m: SMF 96431–32; near La Soledad,  1350 m: IBH 27028–29, SMF 96429–30, 96558–59;  Tierra Blanca,  1369 m: SMF 96753–54;  0.2 kmN Jalatengo,  1458 m: KU137688;  Sierra Madre del Sur, Portillo de Rayo,  1519–1590 m: UTA R-51708, 51720–24; 2 miW La Soledad: UIMNH 8963;  La Soledad: UIMNH 8914, 8924, 8926, 8929, 8931, 8949, 8952, 8962, 8969-70; 1.0 km S Candelaria,  1621 m: KU137685;  Sierra Madre del Sur, along Hwy131, 15 miles ( 24.1 km) SW San Pedro Juchatengo,  1872 m: UTA R-51709; Mex Hwy175 and Chacalapilla River: UTA R-51720; Suchitepec,  2044 m: UTA R-51721; road from Portillo de Rayoto Pochutla,  677 m: UTAR-53603–05.   Diagnosis. A small species (our Species E of the  A. nebuloidescomplex, see above; SVL in largest male 50.0 mm, largest female 47.0 mm) of the genus  Anolis(sensu Poe 2004)that differs from all Mexican and Central American anoles except  A. nebuloides, A. nietoi, A. megapholidotus,and  A. stevepoeiby 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 pink dewlap in males and a very small pink dewlap in females ( Fig. 33).  Anolis zapotecorumdiffers from  A. nebuloides, A. nietoi,and  A. megapholidotusby having (1) a rather gradual transition over 3 to 4 scales between the enlarged dorsal scales and lateral body scales (vs. a more abrupt transition over only 1 or 2 scales); (2) the edges of the field of enlarged dorsal scale rows forming an undulating line due to single enlarged scales or groups thereof outside the main field (vs. the edges of this field forming well defined and in a more or less straight line) (3) and by having a pink male dewlap (vs. more reddish).  Anolis zapotecorumdiffers from  A. stevepoeiin hemipenial morphology (lobes well developed with two apical fields void of ornamentation—one on each lobe, and without a flap-like structure at tip of apex in  A. zapotecorumvs. lobes rudimentary with a single apical field void of ornamentation, and with a flap-like structure at tip of apex in  A. stevepoei) and in the average values of several morphometric and pholidotic characters, most obvious in (1) number of midventral scales in one head length; (2) dorsal scales between levels of axilla and groin; and (3) subdigital lamellae on Phalanges II–IV of Toe IV (see Table 8).   Description of the holotype. Adult male, as indicated by everted hemipenes and presence of large dewlap; SVL 47.5 mm; tail length 97.0 mm, tail complete; tail moderately compressed in cross section, tail height 2.6 mmand width 2.2 mm; axilla to groin distance 17.8 mm; head length 13.5 mm, head length/SVL ratio 0.28; snout length 5.9 mm; head width 8.1 mm; longest toe of adpressed hind limb reaching to mideye; shank length 13.2 mm, shank length/head length ratio 0.98; longest finger of extended forelimb reaching to a point 1 mmbeyond tip of snout; longest finger of adpressed forelimb reaching to anterior insertion of hind limbs. Scales on snout keeled, mostly unicarinate, some multicarinate; posterior head scales rugose or keeled; 6 postrostrals; 7 scales between nasals; 1 elongate prenasal scale on each side, partly fused with circumnasal, in contact with both rostral and first supralabial; circumnasal in contact with first supralabial; scales in deep prefrontal depression mostly keeled, keels mostly radiating from center of depression; supraorbital semicircles well developed, medially in contact with each other; supraorbital disc composed of an oval patch of 3 greatly enlarged, keeled scales; circumorbital row complete, therefore, enlarged supraorbital scales completely separated from supraorbital semicircles; 2 elongated, strongly overlapping superciliaries, anterior one larger than posterior one, followed posteriorly by 3 to 4 roundish to squarish scales of moderate size; 2 to 3 rows of small smooth scales extending between enlarged supraorbitals and superciliaries; parietal depression shallow; interparietal scale well developed, 2.4 x 2.0 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; 7 scales present between second canthals; 6 scales present between posterior canthals; 25 mostly keeled loreal scales in a maximum of 5 horizontal rows; 3 keeled subocular scales arranged in a single row; 6 supralabials to level below center of eye; 2 suboculars broadly in contact with 2 supralabials; ear opening 0.7 x 1.4 mm(length x height); mental distinctly wider than long, completely divided medially, bordered posteriorly by 4 postmentals, outer ones slightly larger than median ones; 6 infralabials to level below center of eye; sublabials not differentiated; keeled granular scales present on chin and throat; dewlap large, extending from level beyond anterior margin of eye to level of chest; 7–9 horizontal gorgetal-sternal rows with 10–21 scales per row; modal number of marginal pairs 2–4; a nuchal crest and a dorsal ridge present; dorsum of body with strongly keeled, imbricate, non-mucronate scales; about 10 to 14 medial rows greatly enlarged; largest dorsal scales about 0.95 x 0.95 mm(length x width); about 20 medial dorsal scales in one head length; about 34 medial dorsal scales between levels of axilla and groin; lateral scales smooth, granular and more or less homogeneous in size, average size 0.25 mmin diameter; ventrals at midbody strongly keeled, imbricate, mucronate, about 0.65 x 0.65 mm(length x width); about 36 medial ventral scales in one head length; about 46 medial ventral scales between levels of axilla and groin; 114 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; a pair of greatly enlarged postcloacal scales present, about 1.5 mmwide; 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; 26 (right)–25 (left) lamellae under phalanges II–IV of fourth toe; 8 scales under distal phalanx of fourth toe.   FIGURE 33.Dewlaps of  Anolis zapotecorumin life. (a) SMF 96559; (b) SMF 96426; (c) SMF 96247; (d) SMF 96418; (e) IBH 26514; (f) IBH 26513. (a–c) are males, (d–f) are females.   FIGURE 34.  Anolis zapotecorumin life. (a) SMF 96248; (b) SMF 96249; (c) SMF 96240; (d) IBH 27000. (a–c) are males, (d) is a female.   FIGURE 35.Holotype of  Anolis zapotecorum(SMF 96425): (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 36.  Anolis zapotecorum: head scalation in (a,c,e) SMF 96431; (b,d,f) SMF 96429. Scale bars equal 1.0 mm.   FIGURE 37.  Anolis zapotecorum: nasal region in (a) SMF 96426; (b) SMF 96429; superciliary region in (c) SMF 96429; (d) SMF 96431; chin region in (e) SMF 96426; (f) SMF 96431; cloacal region in (g) SMF 96248; (h) SMF 96424. Scale bars equal 1.0 mm.   FIGURE 38.  Anolis zapotecorum: dorsal scales in (a) SMF 96425; (b) SMF 96429; flank scales in (c) SMF 96431; (d) SMF 96429; ventral scales in (e) SMF 96426; (f) SMF 96429. Scale bars equal 1.0 mm. The completely everted hemipenis of SMF 96425 ( Fig. 39) 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. Coloration after five months preservation in 70% ethanol was recorded as follows: Dorsal surfaces of head and body Light Russet Vinaceous (246) grading into Light Neutral Gray (297) in parietal region and with an indistinct Dark Neutral Gray (299) lateral longitudinal stripe between levels of axilla and groin; dorsal surfaces of limbs Tawny (60) with indistinct Dark Drab (45) bands; dorsal surface of tail Tawny (60) with indistinct Fawn Color (258) bands; ventral surfaces of head, body, limbs, and tail Chamois (84) with a slight suffusion of Cinnamon-Drab (50); finger and toe pads Vandyke Brown (282); dewlap Vinaceous (247) grading into Rose Pink (243) on edges.   FIGURE 39.Hemipenis of  Anolis zapotecorum(SMF 96425): (left) sulcate view; (center) asulcate view; (right) lateral view. Scale bar equals 1.0 mm.  Variation. The paratypesagree well with the holotypein general appearance, morphometrics, and scalation (see Table 8). Variation is evident in some scalation characters as follows: The number of slightly to moderately enlarged dorsal scale rows varies from 11 to 14. All specimens have a single elongate prenasal. Suboculars in contact with supralabials (1–3 suboculars in contact with 1–3 supralabials) in all specimens examined. Of 30 individuals, the longest toe of adpressed hind leg reaching to a point between ear opening and eye in 3.3%, to posterior border of eye in 20.0%, to mideye in 66.7%, to anterior border of eye in 6.7%, and to beyond eye in 3.3%. All males have a pair of greatly enlarged postcloacal scales. The coloration in life of an adult male of (SMF 96248; Fig. 34a) was recorded as follows: Dorsal ground color Raw Umber (22) with Antique Brown (24) interorbital bar and Mars Brown (25) occipital blotch; lateral stripe Ground Cinnamon (270) edged above and below by Ferruginous (35) streaks; dorsum with Warm Sepia (40) chevrons; dorsal surface of limbs and tail with Warm Sepia (40) bars; ventral surfaces Light Buff (2), those on limbs and tail suffused with Drab (19); dewlap Spinel Pink (235) with Light Orange Yellow (7) marginals and Paris White (139) gorgetals; iris Warm Sepia (40). The coloration in life of another adult male (SMF 96240; Fig. 34c) was recorded as follows: Dorsal ground color Mikado Brown (42) with a suffusion of Olive Brown (278) dorsally and with a Burnt Siena (38) interorbital bar; lateral stripe Smoky Gray (267) edged above and below by series of Walnut Brown (27) splotches; dorsal surface of limbs and tail Antique Brown (24) with Warm Sepia (40) bands; ventral surface of head Pale Buff (1); ventral surface of body Pale Pinkish Buff (3); ventral surface of limbs and tail Ground Cinnamon (270) with Dark Drab (45) suffusions; dewlap Magenta (236) with Cream White (52) gorgetals and Chamois (84) marginals. The coloration in life of an adult female (SMF 96249; Fig. 34b) was recorded as follows: Dorsal ground color Raw Umber (22) with suffusions of Cinnamon (21) and with Mars Brown (25) stipples on flanks; ventral surface of head Light Buff (2); ventral surface of limbs and tail Tawny Olive (17); Venter Cream Color (12); dewlap Spinel Pink (235) with Cream White (52) gorgetals; iris Burnt Umber (48).   Etymology. The name  zapotecorumrefers to the Zapotecan people. The Zapotec civilization was an indigenous pre-Columbian civilization that flourished in the Valley of Oaxaca. Many people in the range of  Anolis zapotecorumstill speak the ancient Zapotec language.  Natural History Notes.  Anolis zapotecorumappears to be very abundant in the surroundings of La Galera, Buena Vista, and Pluma Hidalgo. During daytime surveys, they were encountered on leaf litter or perching low on tree trunks about 0.1 to 0.5 mabove the ground. At night, they were found sleeping on twigs, leaves, and ferns. Near Pluma Hidalgoat 10:30 hrs., an adult male of  A. zapotecorumwas observed using the same coffee tree to rest as a juvenile of  Anolis macrinii, with only a few centimeters in distance between them, although on a different branch.  Geographic Distribution and Conservation. As currently known,  Anolis zapotecorumis restricted to the Pacific versant of the eastern portion of the Sierra Madre del Sur in the south-central portion of the Mexican State of Oaxacaat elevations between 358 and 2044 masl( Fig. 40). Given its usual abundance wherever this species occurs, it seems justified to classify  A. zapotecorumas Least Concern based on the IUCN Red List Categories and Criteria ( IUCN 2012). 4013201517 SMF 96425 2013-03-04 SMF Mexico 1350 15.94241 Pluma 1 -96.43044 Gunther Kohler 47 48 GK- 4369 1 Hidalgo holotype 4013201426 IBH 26997 [199,574,366,391] IBH 47 48 IBH 26997-27004 1 paratype 4013201466 SMF 96418 [594,923,366,391] SMF 47 48 SMF 96418-24, 96426-27 1 paratype 4013201512 SMF 96425 [947,1365,366,391] 2013-03-04 SMF Mexico 1350 15.94241 Pluma 1 -96.43044 Gunther Kohler 47 48 1 Hidalgo paratype 4013201434 IBH 26998 IBH Mexico 47 48 IBH 26998-27001, 27003-27004 1 Hidalgo paratype 4013201306 SMF 96419 [499,993,402,427] SMF Mexico 47 48 96421-22, 96424, 96426-27 1 Hidalgo paratype 4013201490 IBH 26997 [1139,1367,402,427] IBH Mexico 47 48 27002 1 Hidalgo paratype 4013201359 SMF 96418 SMF Mexico 47 48 96420, 96423 1 Hidalgo paratype 4013201343 [434,1187,474,499] UIMNH la Candelaria Mexico 358 Oaxaca 47 48 UIMNH 8906 1 Oaxaca 4013201396 KU 137684 KU Mexico 394 Candelaria 47 48 1 Oaxaca 4013201325 [395,1219,510,535] UTA Mexico 985 Candelaria San Jose Pacifico 47 48 1 Oaxaca 4013201442 UTA Pochutla Mexico 380 Candelaria 47 48 1 Oaxaca 4013201488 IBH 26579, SMF 96246 Mexico 1010 Santiago La Galera Finca de Juan Garcia 47 48 1 Oaxaca 4013201516 KU 137686 [425,930,582,607] KU Mexico 1017 Jalatengo 47 48 KU 137686-87 1 Oaxaca 4013201387 IBH 26513, SMF 96240 SMF Mexico 1160 Santiago la Galera 47 48 IBH 26513-14, 26521-22, 26578, 26582, 26612, 96248-51 1 Oaxaca 4013201443 SMF 96431 [820,1335,618,643] SMF Mexico 1287 Tierra Blanca 47 48 SMF 96431-32 1 Oaxaca 4013201404 IBH 27028, SMF 96429 IBH, SMF Mexico 1350 La Soledad 47 48 IBH 27028-29, SMF 96429-30, 96558-59 1 Oaxaca 4013201427 SMF 96753 [864,1315,654,679] SMF Mexico 1369 Tierra Blanca 47 48 SMF 96753-54 1 Oaxaca 4013201307 KU 137688 KU Mexico 1458 Jalatengo 47 48 1 Oaxaca 4013201397 [514,1178,690,715] UTA Mexico 1555 Portillo de Rayo Sierra Madre del Sur 47 48 1 Oaxaca 4013201310 UIMNH Mexico La Soledad 47 48 UIMNH 8963 1 Oaxaca 4013201329 [534,1432,726,751] UIMNH Mexico La Soledad 47 48 UIMNH 8914, 8924, 8926, 8929, 8931, 8949, 8952, 8962, 8969-70 1 Oaxaca 4013201314 KU 137685 [151,647,762,787] KU Mexico 1621 Candelaria 47 48 1 Oaxaca 4013201462 UTA SW San Pedro Juchatengo Mexico 1872 Hwy Sierra Madre del Sur 47 48 1 Oaxaca 4013201319 [522,1125,798,823] UTA Mex Hwy Mexico Chacalapilla River 47 48 1 Oaxaca 4013201309 UTA Mexico 2044 Suchitepec 47 48 1 Oaxaca 4013201401 [250,952,834,859] UTA Pochutla Mexico 677 Portillo de Rayo 47 48 1 Oaxaca