Anolis stevepoei, Köhler & Pérez & Petersen & Méndez & Cruz, 2014

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 : 40-47

Anolis stevepoei sp. nov.

Figs. 26–31

Holotype. SMF 96232, an adult male from about 15.8 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.2205°N, 97.14883°W), 1900 m, Estado de Oaxaca, Mexico; collected 16 November 2012 by Gunther Köhler and Raúl Gómez Trejo Pérez. Field tag number GK-4130 .

Paratypes. All from Estado de Oaxaca, Mexico: IBH 26523, SMF 96244–45 from Rancho El Sagrado , near San Gabriel Mixtepec (16.08914°N, 97.06263°W), 630 m; IBH 26581, 26584 , SMF 96243 from near San Gabriel Mixtepec (16.08108°N, 97.08115°W), 755 m; IBH 26517, SMF 96242 from about 6.5 airline km N San Gabriel Mixtepec on road to El Vidrio (16.15547°N, 97.08112°W), 1030 m; KU 87316–24 from 12 km NNW San Gabriel Mixtepec, 1040 m; SMF 96745–46 from about 10.9 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.19286°N, 97.09767°W), 1326 m; SMF 96739–41 from about 11.1 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.19212°N, 97.10509°W), 1382 m; IBH 26511, SMF 96236, 96428 from about 11.3 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.19280°N, 97.10821°W), 1400 m; KU 137714 from 31.2 km N San Gabriel Mixtepec, 1480 m; SMF 96742–44 from about 11.6 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.19483°N, 97.11397°W), 1481 m; KU 101115, 101119 , MCZ R-92965–76 from 30 km N San Gabriel Mixtepec, 1530 m; SMF 96736–38 from about 11.3 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.18964°N, 97.11994°W), 1587 m; IBH 26509, SMF 96235 from about 11.5 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.19083°N, 97.12045°W), 1600 m; KU 137715 from 34.3 km N San Gabriel Mixtepec, 1600 m; KU 101124 from 33 km N San Gabriel Mixtepec, 1675 m; KU 137716 from 35.9 km N San Gabriel Mixtepec, 1680 m; SMF 96731–33 from about 12.8 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.20120°N, 97.12538°W), 1698 m; IBH 26512, SMF 96237 from about 12.9 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.20245°N, 97.12621°W), 1700 m; IBH 26608 from about 13.9 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.20843°N, 97.13646°W), 1750 m; IBH 26518 from about 14.2 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.20946°N, 97.13947°W), 1800 m; SMF 96728–30 from about 14.3 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.21085°N, 97.13647°W), 1803 m; KU 101125 from 37 km N San Gabriel Mixtepec, 1860 m; SMF 96234 from about 14.8 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.21621°N, 97.13947°W), 1870 m; SMF 96233 from about 15.8 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.21736°N, 97.14757°W), 1920 m; SMF 96726–27 , 96734–35 from about 15.9 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.21813°N, 97.14731°W), 1925 m; SMF 96231 from about 16.7 airline km NNW San Gabriel Mixtepec on road to El Vidrio (16.22822°N, 97.15132°W), 1980 m; IBH 26511, SMF 96231, 96233–35 are males, IBH 26509, 26512 , 26518 , 26608 , SMF 96236–37 are females, IBH 26517, 26581 , 26584 , SMF 96242–43 are juveniles. All IBH and SMF paratypes collected by Gunther Köhler and Raúl Gómez Trejo Pérez. All MCZ paratypes collected by William E. Duellman and John D. Lynch.

Referred specimens ⎯ Mexico: Oaxaca: Soledad: KU 38260; 29 km SSE San Pedro Juchatengo, 1980 m: KU 87338-40; 17.2 km N San Pedro Mixtepec, 710 m: KU 137692; 18.1 km N San Pedro Mixtepec, 720 m: KU 137693–94; 20 mi N (on road) Puerto Escondido, 525 m: UMMZ 130954; 4.8 mi SW Sola de Vega, 2040 m: UTA R-4290–91; 4.8 miles (7.7 km) S Sola de Vega, 1774 m: UTA R-51719; 23.0 km SW San Pedro Juchatengo, Sierra Madre del Sur, along Hwy 131, 1575 m: UTA R-51710; 20 mi (on Hwy 131) SW San Pedro Juchatengo: MCZ R- 100379; 4.7 km NE Sola de Vega, 1534 m: UTA R-51717–18; 18 km S Sola de Vega on MEX Hwy. 131, 1937 m: MVZ 144180.

Diagnosis. A small species (our Species D of the A. nebuloides complex, see above; SVL in largest male 50.0 mm, largest female 50.5 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 species we are describing below (our Species E of the A. nebuloides complex) 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 0.90–1.07; (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. 26 ). In external morphology, A. stevepoei is most similar to A. nebuloides , A. nietoi , A. megapholidotus and a species we are describing below (our Species E of the A. nebuloides complex). Anolis stevepoei differs from A. nebuloides , A. nietoi , and A. megapholidotus by having (1) a rather gradual transition over 3 to 4 scales between the enlarged dorsal scales and the 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 well defined and in a more or less straight line) (3) and by having a pink to purple male dewlap (vs. more reddish). Anolis stevepoei differs from Species E in hemipenial morphology (lobes rudimentary with a single apical field void of ornamentation, and with a flap-like structure at tip of apex in A. stevepoei vs. 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 Species E) 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 50.0 mm; tail incomplete; tail moderately compressed in cross section, tail height 2.5 mm and width 1.8 mm; axilla to groin distance 20.1 mm; head length 13.6 mm, head length/SVL ratio 0.27; snout length 5.9 mm; head width 8.4 mm; longest toe of adpressed hind limb reaching to mideye; shank length 13.9 mm, shank length/head length ratio 1.02; longest finger of extended forelimb reaching to nostril; longest finger of adpressed forelimb reaching to anterior insertion of hind limbs. Scales on snout keeled, unicarinate; posterior head scales mostly rugose or keeled, some smooth; 5 postrostrals; 6 scales between nasals; 1 elongate prenasal scale on each side, partly fused with circumnasal, in contact with both rostral and first supralabial; circumnasal separated from supralabial by one scale row; scales in deep prefrontal depression mostly keeled, keels mostly oriented longitudinally; supraorbital semicircles well developed, separated medially by one scale row at narrowest point; 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; 3 elongated, strongly overlapping superciliaries, anteriormost one larger than posteriormost one, followed posteriorly by 3 to 4 roundish to squarish scales of moderate size; 1 to 3 rows of small smooth scales extending between enlarged supraorbitals and superciliaries; parietal depression shallow; interparietal scale well developed, 2.2 x 1.9 mm (length x width), surrounded by scales of moderate size; 2 scales present between interparietal and supraorbital semicircles; canthal ridge distinct, composed of 3 large (second largest) and 3 small anterior canthal scales; 6 scales present between second canthals; 6 scales present between posterior canthals; 27 (right)– 27 (left) mostly keeled loreal scales in a maximum of 6 (right)– 5 (left) horizontal rows; 2 very elongate keeled subocular scales arranged in a single row; 6 supralabials to level below center of eye; 3 suboculars broadly in contact with 3 supralabials; ear opening 0.8 x 1.6 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; 8–10 horizontal gorgetal-sternal rows with 9–16 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; 10 to 14 medial rows greatly enlarged; largest dorsal scales about 0.75 x 0.65 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 0.20 mm in diameter; ventrals at midbody strongly keeled, imbricate, mucronate, about 0.55 x 0.55 mm (length x width); about 32 medial ventral scales in one head length; about 45 medial ventral scales between levels of axilla and groin; 112 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 0.7 mm wide; 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; 23 (right)–22 (left) lamellae under phalanges II–IV of fourth toe; 7 scales under distal phalanx of fourth toe.

Coloration after nine months preservation in 70% ethanol was recorded as follows: Dorsal surfaces of head, body, limbs, and tail Cinnamon-Drab (50) with suffusions of Sepia (286) in vertebral region of body and anterior tail, with paravertebral transverse Sepia (286) streaks and with an indistinct Sepia (286) lateral longitudinal stripe between levels of ear and groin; lateral and ventral surfaces of head Cream White (52); ventral surfaces of body and limbs Pale Buff (1); ventral surface of tail Cinnamon-Drab (50) suffused on edges with Sepia (286); finger and toe pads Vandyke Brown (282); dewlap Deep Vinaceous (248) grading into Light Lavender (201) on posterior portion.

Variation. The paratypes agree well with the holotype in general appearance, morphometrics, and scalation (see Table 8 ). Variation was evident in some scalation characters as follows: The number of slightly to moderately enlarged dorsal scale rows varies from 10 to 14. All but one specimen have a single elongate prenasal; only SMF 96233 has a divided prenasal. Suboculars in contact with supralabials (1–3 suboculars in contact with 1–4 supralabials) in all but one specimen (SMF 96233), which has the suboculars separated from supralabials by one complete scale row. Of 35 individuals, the longest toe of adpressed hind leg reaching to level of ear opening in 5.7%, to a point between ear opening and eye in 11.4%, to posterior border of eye in 22.9%, to mideye in 40.0%, to anterior border of eye in 14.3%, and to beyond eye in 5.7%. All males have a pair of greatly enlarged postcloacal scales.

The coloration in life of an adult male (SMF 96234) was recorded as follows: Dorsal surface of head Brussels Brown (33) anteriorly and grading posteriorly into Mikado Brown (42) with a Burnt Umber (48) interorbital bar; dorsal ground color Cinnamon-Drab (50); lateral stripe indicated by two series of Mahogany Red (34) streaks; dorsal surface of Limbs Cinnamon Brown (43) with Raw Umber (280) bands; ventral surface of head Light Buff (2) with indistinct Clay Color (18) transverse streaks; ventral surface of body Chamois (84); ventral surfaces of limbs and tail Ground Cinnamon (270) with suffusions of Pale Pinkish Buff (3); dewlap Spinel Pink (235) with Light Buff (2) gorgetals; iris Raw Umber (280) with a Pale Emerald Green (141) inner ring.

The coloration in life of an adult female (IBH 26518) was recorded as follows: Dorsal ground color Dark Drab (45) grading into Mikado Brown (42) on flanks and with Cinnamon Brown (43) chevrons, interorbital bar, and bars on limbs and tail; indistinct Cinnamon Drab (50) lateral stripe; ventral surface of body Pale Pinkish Buff (3); ventral surface of limbs and tail Drab (19); dewlap with suffusions of Vinaceous (247) and with Pale Pinkish Buff (3) gorgetals; iris Sepia (279) with a Greenish Glaucous (271) inner ring.

The completely everted hemipenis of SMF 96742 ( Fig. 32 ) is a small organ with rudimentary lobes and a flaplike structure at tip of apex; sulcus spermaticus bordered by well developed sulcal lips and opening at base of apex into a single large apical field void of ornamentation; 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 finely calyculate, truncus and asulcate ridge with transverse folds.

Etymology. The name stevepoei is a patronym honoring Steven Poe who has contributed substantially to our knowledge of the phylogeny and taxonomy of anoles.

Natural History Notes. We found Anolis stevepoei to be an abundant species along the road from San Miguel Mixtepec to El Vidrio. At night, we spotted numerous individuals sleeping on low vegetation (ferns, bushes, and branches of small trees) at 0.3 to 2.0 m above ground level. At Rancho Sagrado near San Miguel Mixtepec, we encountered several individuals that were active on the ground. On 23 June 2013 at 1:09 hrs. RGT observed a Leptodeira maculata on the hunt for an individual of A. stevepoei . The snake was climbing a parallel twig to a branch where an adult male of Anolis stevepoei was sleeping. The colubrid took around 35 minutes to slowly climb the 50 cm shrub twig but the Anolis was able to jump away before the snake could strike.

Geographic Distribution and Conservation. As currently known, Anolis stevepoei is restricted to the Pacific versant of the Sierra Madre del Sur in the mountainous region north of Puerto Escondido in the south-central and southwestern portions of the Mexican State of Oaxaca at elevations between 350 and 2040 masl ( Fig. 40 ). Given its usual abundance wherever this species occurs, it seems justified to classify A. stevepoei as Least Concern based on the IUCN Red List Categories and Criteria ( IUCN 2012).

Figures

FIGURE 26. Dewlaps of Anolis stevepoei in life. (a) SMF 96231; (b) SMF 96232; (c) SMF 96233; (d) SMF 96234; (e) IBH 26608; (f) IBH 26518. (a–d) are males, (e–f) are females.

FIGURE 27. Anolis stevepoei in life. (a) SMF 96231; (b) SMF 96232; (c) SMF 96233; (d) IBH 26518; (e) IBH 26608; (f) IBH 26512. (a–c) are males, (d–f) are females.

FIGURE 28. Holotype of Anolis stevepoei (SMF 96232): (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 29. Anolis stevepoei: head scalation in (a,c,e) SMF 96237; (b,d,f) SMF 96743. Scale bars equal 1.0 mm.

FIGURE 30. Anolis stevepoei: nasal region in (a) SMF 96233; (b) SMF 96232; superciliary region in (c) SMF 96233; (d) SMF 96727; chin region in (e) SMF 96730; (f) SMF 96727; cloacal region in (g) SMF 96230; (h) SMF 96231. Scale bars equal 1.0 mm.

FIGURE 31. Anolis stevepoei: dorsal scales in (a) SMF 96237; (b) SMF 96232; flank scales in (c) SMF 96237; (d) SMF 96232; ventral scales in (e) SMF 96237; (f) SMF 96232. Scale bars equal 1.0 mm.

FIGURE 32. Hemipenis of Anolis stevepoei (SMF 96742): (left) sulcate view; (center) asulcate view; (right) lateral view. Scale bar equals 1.0 mm.

FIGURE 40. Map indicating collecting localities of species related to Anolis nebuloides. Each symbol can represent one or more adjacent localities. Black squares: Anolis megapholidotus; white squares: A. nietoi; black circles: A. nebuloides; black triangles: A. stevepoei; white circles: A. zapotecorum.