Anolis unilobatus Köhler & Vesely 2010

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 : 190-200

Anolis unilobatus Köhler & Vesely 2010

Figs. 158–164

Anolis unilobatus Köhler & Vesely 2010: 217 ; type locality: “ Awasbila , a village along Río Coco, 14°47’N, 84°45’W, 60 m , Departamento Gracias a Dios, Honduras.” Holotype: SMF 87133. Nieto Montes de Oca et al. 2013, Köhler 2014, Köhler et al. 2014

Anolis kidderi: Werler & Smith 1952 ,

Anolis sallaei: Boulenger 1885 (in part.), Barbour & Loveridge 1929

Anolis sallaei wellbornae: Smith & Kerster (1955) , Lynch & Smith 1966

Anolis sericeus: Hartweg & Oliver 1940 , Smith & Taylor 1950a (in part.), Etheridge 1959, (in part.), Lee 1980 (in part.), Lieb 1995 (in part.), Poe 2004 (in part.), Liner 2007 (in part.), Wilson et al. 2013 (in part.), Wilson & Townsend 2010 (in part.)

Norops sericeus: Savage & Guyer 1989 (in part.), Nicholson 2002 (in part.)

Norops unilobatus: Nicholson et al. 2012

Diagnosis. A small species (SVL in largest male 48.5 mm, largest female 48.5 mm) of the genus Anolis (sensu Poe 2004) that differs from all Mexican and Central American anoles except A. sericeus and A. wellbornae by the following combination of characters: (1) male dewlap yellowish orange with large blue to purple blotch ( Fig. 159 ); (2) short legged (longest toe of adpressed hind limb reaching to a point between shoulder and ear opening, rarely beyond ear opening); (3) ear opening very small (less than ¼ the size of interparietal plate); (4) anterior superciliary conspicuously large and elongate; (5) ventral scales strongly keeled and mucronate; (6) males without a pair of enlarged postcloacal scales. Anolis unilobatus differs from A. sericeus and A. wellbornae by having a unilobate hemipenis (bilobate in A. sericeus and A. wellbornae ). Furthermore, A. unilobatus can be distinguished from A. sericeus in dewlap size ( A. unilobatus : male dewlap larger than 100 mm 2, no female dewlap; A. sericeus : male dewlap smaller than 50 mm 2, females with a dewlap of about the size of the male dewlap).

Description. Anolis unilobatus is a small anole (maximum recorded SVL 48.5 mm in males, 48.5 mm in females); dorsal head scales in internasal region weakly to strongly keeled, in prefrontal, parietal, and frontal areas usually flat, occasionally rugose or weakly keeled; weak to moderately deep prefrontal depression present, parietal depression usually absent, if present very shallow; 5–8 postrostrals; a single anterior nasal in contact with rostral and first supralabial, occasionally only with rostral scale; 7–9 internasals; canthal ridge sharply defined; scales comprising supraorbital semicircles well defined, posterior ones smooth, anterior ones weakly keeled, largest scale in semicircles subequal or larger than largest supraocular scale; supraorbital semicircles usually broadly in contact, occasionally separated by a complete row of scales; 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 anteriorly and by small to moderate size scales posteriorly, usually longer than wide, larger than ear opening; enlarged supraoculars usually separated from supraorbital semicircles by a complete row of small scales, occasionally these scales narrowly in contact; 2–3 scales between enlarged supraoculars and superciliaries; a single large elongated superciliary, followed posteriorly by a series of 7–8 rounded to squarish scales of moderate size; 4 enlarged canthals, the second canthal usually largest; 6–9 scales between second canthals; 8–12 scales present between posterior canthals; loreal region slightly concave, 23–47 mostly keeled (some smooth or rugose) loreal scales in a maximum of 5–6 horizontal rows; 6–9 supralabials to level below center of eye; suboculars keeled, in broad contact with supralabials (1–3 suboculars in contact with 2–4 supralabials); ear opening vertically oval, oriented slightly obliquely; scales anterior to ear opening granulars, about two to three times as large than those posterior to ear opening; 5–8 infralabials to level below center of eye; 4–6 (usually 6) postmentals, outer pair not enlarged relative to adjacent median postmental scales; no sublabials in contact with infralabials; small keeled scales present on chin and throat; male dewlap moderate-sized, extending from level below center of eye to level of chest; 8–9 horizontal gorgetal-sternal rows with 6–12 scales per row; modal number of marginal pairs 1–2; female dewlap very small; a nuchal crest and a dorsal ridge present in males; scales on middorsum weakly keeled, subimbricate, with rounded posterior margins; 12–22 middorsal scale rows slightly to moderately enlarged, with a gradual transition to lateral scales; lateral scales keeled, granular, usually homogeneous, occasionally slightly heterogeneous; 54–74 dorsal scales along vertebral midline between levels of axilla and groin in males, 56–74 in females; 30–38 dorsal scales along vertebral midline contained in one head length in males, 28–44 in females; ventral scales on midsection larger than largest dorsal scales; scales on midventer strongly keeled, mucronate, subimbricate to imbricate; 41–55 ventral scales along midventral line between levels of axilla and groin in males, 37–51 in females; 22–32 ventral scales contained in one head length in males, 22–32 in females; 112–130 scales around midbody in males, 110–132 in females; tube-like axillary pocket absent; precloacal scales smooth or weakly keeled; postcloacal scales not enlarged; tail slightly to moderately compressed in cross section, tail height/ tail width 1.18–1.54 in males, 1.06–1.40 in females; caudal scales strongly keeled, homogeneous, without whorls of enlarged scales, although an indistinct division in segments is discernible; dorsal medial caudal scale row slightly enlarged, not forming a crest; scales on anterodorsal surface of brachium and on dorsal surface of antebrachium usually imbricate, strongly keeled, unicarinate, mucronate; 20–26 subdigital lamellae on Phalanges II–IV of Toe IV of hind limbs; 5–8 subdigital lamellae on distal phalanx of Toe IV of hind limbs; digital pads dilated, two to three times the width 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, rarely beyond tympanum. For variation in selected scalation and morphometric characters see Table 14 .

The coloration in life of an adult male (SMF 96277) was recorded as follows: Dorsal surface of body Light Buff (2); dorsal surface of limbs Cinnamon (21); ventral surface of body, limbs and tail Pale Pinkish Buff (3); dewlap Orange Yellow (8) with Light Pratt´s Rufous (71) stripes and a Campanula (200) blotch. The coloration in life of another adult male (IBH 26571) was recorded as follows: Dorsal surface of body Light Buff (2); dorsal surface of limbs Cinnamon (255); Snout Light Orange Yellow (7); interorbital bar Umber (51); ventral surfaces of body Chamois (84); dewlap Orange Yellow (8) with light Pratt´s Rufous (71) Stripes and a Campanula (200) blotch; iris Tawny Olive (17).

The coloration in life of an adult female (SMF 96470) was recorded as follows: Dorsal color of body and tail Cream White (52) with a Cinnamon Drab (50) vertebral line edged by two Sulphur Yellow (91) lines; dorsal surface of legs Cinnamon (21); ventral surface of body, legs, and tail Pale Pinkish Buff (3); chin Smokey White (261) with Warm Sepia (40) dots; dewlap Medium Chrome Orange (75) with Spectrum Blue blotch; iris Brussels Brown (33). The coloration in life of another adult female (SMF 96278; Fig. 159d ) was recorded as follows: Dorsal ground color Pale Pinkish Buff (3) with suffusions of Tawny Olive (17) on head and with a medially interrupted Clay Color (20) interorbital bar; dorsal surface of limbs Dark Drab (45); ventral surfaces of body, limbs and tail Buff (2); dewlap Burnt Orange (10) with a Campanula (20) blotch and with Pale Buff (1) gorgetals; iris Raw Umber (22).

The completely everted hemipenis of SMF 84457 ( Fig. 164 ) is a small, unilobate organ; sulcus spermaticus bifurcating at base of apex into two branches that open into two apical fields, one on each side of apex; a large ridge-like double fold across sagittal plane of apex present; no surface ornamentation discernable.

Natural History Notes: In most areas where this species occurs, it is commonly encountered. In the vicinity of Zanatepec, Oaxaca, we collected six individuals of this species within about one hour of night searching. In the more western portions of its range, A. unilobatus seems to become more and more uncommon, and we encountered only few specimens of this species around the cities of Puerto Escondido and Huatulco, respectively. West of the Isthmus of Tehuantepec, A. unilobatus occurs syntopically with A. boulengerianus , which usually is far more abundant than A. unilobatus . Thus, aside from climatic factors, ecological competition might be one reason for the relative scarcity of A. unilobatus in the western portion of its range.

Geographic Distribution and Conservation. Anolis unilobatus is distributed from Costa Rica across most of Nicaragua (except northwestern corner of the country), Honduras, (except southern portion of the country), along the Caribbean versant of Guatemala and Chiapas, Mexico, to the Isthmus of Tehuantepec where it crosses to the Pacific versant and continues along the Pacific versant of Chiapas and Guatemala to about Mazatenango and northward to eastern Guerrero; from sea level to about 1200 masl (see Fig. 165 for localities in study area). Given its large geographic range and abundance at most localities where this species occurs, it seems justified to classify A. unilobatus 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, 750 m: SMF 96470 ; Oaxaca: 23 km WNW (on road) Tehuantepec , 175 m: IBH 26989, SMF 96471 ; 23.8 mi NW (by road) Tehuantepec: UTA R-4326; Huatulco , 90 m: SMF 96276 ; Parque National Huatulco , 10 m: IBH 26992 ; Puerto Escondido, Punta Colorada , 40 m: SMF 96280 ; Puerto Escondido, Universidad del Mar , 80 m: IBH 26569 ; Zanatepec , 50 m: IBH 26570–71 , 26574 , 26609 , SMF 96277–78 ; Chimalapa , 650 m: SMF 96275 ; north of Juchitán : SMF 78463, 78466–68 ; 5 km W Mixtequilla : SMF 84066, 84595 .

Key to species of Mexican Anolis from the Pacific versant west of the Isthmus de Tehuantepec in the states of Oaxaca, Guerrero, and Puebla.

1a Ventral scales smooth................................................................................. 2

1b Ventral scales faintly to strongly keeled................................................................... 8

2a Supraocular scales arranged in 2–3 rows of moderately enlarged scales; prenasal scale divided; SVL of adults 85–105 mm .......................................................................................... Anolis macrinii

2b A patch of 3–4 greatly enlarged, ovoid supraocular scales in a single row; a single elongate prenasal scale; SVL of adults <76 mm ........... ........................... .. ......................................... .. ............. 3

3a Middorsal scales not or only scarcely enlarged, usually only 2–4 median dorsal scale rows slightly enlarged............. 4

3b 10–15 rows of median dorsal scales moderately enlarged..................................................... 6

4a Dorsum usually with a distinct dark brown vermiculation; suboculars separated from supralabials by one scale or these scales narrowly in contact (then usually a single subocular scale in contact with one supralabial); male dewlap uniformly pink to purple in life................................................................................ Anolis gadovii

4b Dorsal pattern not as above; suboculars broadly in contact with supralabials; male dewlap pinkish to orange red with semicircular pale streaks and blotches in life..................................................................... 5

5a Dorsum with a bold color pattern consisting of contrasting white longitudinal body stripes in large males; a red dewlap with bold bluish purple semicircular markings in adult males in life....................................... Anolis taylori

5b Dorsum without such contrasting pattern; male dewlap red or orange red with yellow or whitish markings in life..................................................................................................... Anolis dunni

6a Fourth toe of adpressed hind leg usually reaching to a point between posterior and anterior margin of eye or occasionally to a point between ear opening and eye; ratio shank length / SVL 0.24–0.30; usually a single pair of greatly enlarged sublabial scales in contact with infralabial scales; male dewlap pink to purple in life, females with a very small to small pink dewlap.......................................................................................... Anolis liogaster

6b Fourth toe of adpressed hind leg reaching to level of ear opening or to a point between shoulder and ear opening; ratio shank length / SVL 0.18–0.23; usually two pairs of greatly enlarged sublabial scales in contact with infralabial scales; male dewlap orange yellow in life, females with a very small dirty white dewlap............................................. 7

7a Fourth toe of adpressed hind leg reaching usually to ear opening, occasionally to slightly beyond ear opening or to a point between shoulder and ear opening; circumnasal separated from first supralabial by the presence of a subnasal; a slightly smaller dewlap in females, in largest female about 41 mm 2 ..................................... Anolis omiltemanus

7b Fourth toe of adpressed hind leg reaching to a point between levels of axilla and ear opening; circumnasal usually in contact with first supralabial; a slightly larger dewlap in females, in largest female about 64 mm 2 .............. Anolis peucephilus

8a Ear opening very small (less than ¼ the size of interparietal plate); a single conspicuously large and elongate superciliary; short legged (longest toe of adpressed hind limb reaching to a point between shoulder and ear opening, rarely beyond ear opening); males without a pair of enlarged postcloacal scales; male dewlap in life yellowish orange with large blue to purple blotch......................................................................................... Anolis unilobatus

8b Ear opening not very small (more than ¼ the size of interparietal plate); usually 2–3 elongate superciliaries; short legged or not; males usually with a pair of moderately to greatly enlarged postcloacal scales; male dewlap in life not as above...... 9

9a SVL to 95 mm in males, to 96 mm in females; supraocular scales arranged in 2–3 rows of moderately enlarged scales; prenasal scale divided; ventral scales faintly to weakly keeled; females with a dewlap of moderate size............ Anolis macrinii

9b SVL of adults <60 mm; a patch of 3–4 greatly enlarged, ovoid supraocular scales in a single row; a single elongate prenasal scale; ventral scales weakly to strongly keeled; female dewlap very small to small................................ 10

10a 8–12 medial rows greatly and abruptly enlarged, largest dorsal scales about the same size as or larger than ventral scales; usually <23 middorsal scales in one head length; male dewlap pinkish-red or purple in life............................ 11

10b 0–20 medial rows slightly and gradually enlarged, largest dorsal scales smaller than ventral scales; usually>20 middorsal scales in one head length; male dewlap color variable....................................................... 15

11a A rather gradual transition over 3 to 4 scales between the enlarged dorsal scales and the lateral body scales with 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; legs longer, ratio shank length / HL 0.90–1.11, the majority of individuals in the range of 0.95–1.09; subdigital lamellae on Phalanges II–IV of Toe IV 21–29, the majority of individuals in the range of 23–26; a pink to purple male dewlap in life; distributed in south-central Oaxaca ................................................................ 12

11b A rather abrupt transition over only 1 or 2 scales between the enlarged dorsal scales and the lateral body scales with the edges of the field of enlarged dorsal scale rows well defined and in a more or less straight line; legs shorter, ratio shank length / HL 0.81–1.04, the majority of individuals in the range of 0.90–0.97; subdigital lamellae on Phalanges II–IV of Toe IV 17–26, the majority of individuals in the range of 20–23; a reddish orange pink male dewlap in life; distributed in south-central Guerrero and southwestern Oaxaca ............................................................................. 13

12a Hemipenial lobes rudimentary with a single apical field void of ornamentation, and with a flap-like structure at tip of apex (sulcate side); distributed along the Pacific versant of the Sierra Madre del Sur north of Puerto Escondido in the southern centralwestern portion of the Mexican State of Oaxaca ................................................ Anolis stevepoei

12b Hemipenial lobes well developed with two apical fields void of ornamentation, one on each lobe, and without a flap-like structure at tip of apex (sulcate side); distributed along the Pacific versant of the eastern portion of the Sierra Madre del Sur in the southern central portion of the Mexican State of Oaxaca ....................................... Anolis zapotecorum

13a Subdigital lamellae on Phalanges II–IV of Toe IV 17–23, the majority of individuals in the range of 18–21; 14–20, mean 15.8, middorsal scales in one head length; distributed along the Pacific versant of Sierra Madre del Sur in the southern central portion of the Mexican State of Guerrero ................................................... Anolis megapholidotus

13b Subdigital lamellae on Phalanges II–IV of Toe IV 20–26, the majority of individuals in the range of 21–24; 16–28, mean 20.5, middorsal scales in one head length; distributed along the Pacific versant of Sierra Madre del Sur in the southern eastern portion of the Mexican State of Guerrero or in the southwestern portion of the Mexican State of Oaxaca ................. 14

14a Number of medial dorsal scales between levels of axilla and groin 27–38; hemipenial lobes rudimentary with a single apical field void of ornamentation; distributed along the Pacific versant of Sierra Madre del Sur in the southern eastern portion of the Mexican State of Guerrero .................................................................... Anolis nietoi

14b Number of medial dorsal scales between levels of axilla and groin 34–44; hemipenial lobes well developed with two apical fields void of ornamentation, one on each lobe; distributed along the Pacific versant of Sierra Madre del Sur in the southwest-ern portion of the Mexican State of Oaxaca ................................................... Anolis nebuloides

15a Hind legs longer, fourth toe of adpressed hind leg usually reaching to level of posterior border of eye or beyond, ratio shank length / SVL usually>0.25, ratio shank length / HL usually>0.97; subdigital lamellae on Phalanges II–IV of Toe IV 21–28.................................................................................................... 16

15b Hind legs shorter, fourth toe of adpressed hind leg reaching to ear opening or to a point between ear opening and eye; ratio shank length / SVL usually <0.25, ratio shank length / HL usually <0.96; subdigital lamellae on Phalanges II–IV of Toe IV 17–26............................................................................................. 18

16a Male dewlap in life pinkish red with paler areas around gorgetals; subocular scales separated from supralabials by one scale row or these scales in contact; hemipenis unilobate with a single apical field void of ornamentation and without an asulcate processus or asulcate ridge; distributed in south-central and southeastern Guerrero ................... Anolis subocularis

16b Male dewlap in life pinkish red without paler areas around gorgetals or orange yellow with paler areas around gorgetals; subocular scales usually broadly in contact with supralabials; hemipenis slightly bilobate with two apical fields void of ornamentation, one on each lobe and with a finger-like asulcate processus and an indistinct asulcate ridge; distributed in southern Oaxaca ............................................................................................ 17

17a Male dewlap in life pinkish red without paler areas around gorgetals; distributed in south-central Oaxaca ................................................................................................. Anolis immaculogularis

17b Male dewlap in life orange yellow with paler areas around gorgetals; distributed west and north of the city of Tehuantepec..................................................................................... Anolis boulengerianus

18a Number of middorsal scales in one head length 36–46 in males, 32–48 in females; number of midventral scales in one head length 24–36 in females; subdigital lamellae on Phalanges II–IV of Toe IV 17–20; male dewlap in life dull orange yellow with paler semicircular streaks and blotches................................................... Anolis microlepidotus

18b Number of middorsal scales in one head length 22–40, usually <36; number of midventral scales in one head length 18–28 in females; subdigital lamellae on Phalanges II–IV of Toe IV 19–26; male dewlap in life not colored as above............ 19

19a Flank scales usually heterogeneous, enlarged conical or keeled scales scattered among granular scales; midventral scales between levels of axilla and groin 46–62 in males, 52–54 in females; head longer, ratio HL/HW usually>1.72; in life, dorsum of head of similar color as that of lateral head, not demarcated by a pale line; male dewlap in life orange with or without a white anterior margin.................................................................... Anolis nebulosus

19b Flank scales usually homogeneous, no enlarged conical or keeled scales scattered among granular scales; midventral scales between levels of axilla and groin 33–52 in males, 33–51 in females; head shorter, ratio HL/HW usually <1.70; in life, dorsum of head of different color, mostly darker, than that of lateral head, demarcation usually emphasized by a pale line; male dewlap in life pink or purple................................................................................. 20

20a Hemipenis unilobate and without an asulcate processus and without an asulcate ridge; number of middorsal scales in one head length usually <29 in males; legs longer, ratio shank length/HL usually>0.93.......................... Anolis carlliebi

20b Hemipenis slightly to distinctly bilobate and with an asulcate processus and an asulcate ridge; number of middorsal scales in one head length usually> 27 in males; legs shorter, ratio shank length/HL usually <0.93............................ 21

21a Hind legs slightly longer, ratio shank length / SVL usually 0.23-0.24, mean 0.23, in males, 0.20-0.24, mean 0.22, in females; hemipenis with rudimentary lobes, sulcus spermaticus opening into a single apical field void of ornamentation; subdigital lamellae on Phalanges II–IV of Toe IV 19–22, mean 20.4.................................... Anolis sacamecatensis

21b Hind legs slightly longer, ratio shank length / SVL usually 0.22-0.26, mean 0.24, in both sexes; hemipenis with well-developed lobes, sulcus spermaticus opening into two apical fields void of ornamentation, one on each lobe; subdigital lamellae on Phalanges II–IV of Toe IV 19–26, mean 21.8.................................................... Anolis quercorum

Figures

FIGURE 158. Dewlaps of Anolis unilobatus in life. (a) SMF 96471; (b) IBH 26570; (c) IBH 26609; (d) SMF 96278. (a–b) are males, (c–d) are females.

FIGURE 159. Anolis unilobatus in life. (a) IBH 26992; (b) SMF 96471; (c) IBH 26570; (d) IBH 26609. (a–c) are males, (d) is a female.

FIGURE 160. Anolis unilobatus: head scalation in (a,c,e) SMF 84066; (b,d,f) SMF 96278. Scale bars equal 1.0 mm.

FIGURE 161. Anolis unilobatus: nasal region in (a) SMF 96470; (b) SMF 84066; superciliary region in (c) SMF 96470; (d) SMF 84066; chin region in (e) SMF 96471; (f) SMF 96470; cloacal region in (g) SMF 96471; (h) SMF 96277. Scale bars equal 1.0 mm.

FIGURE 162. Anolis unilobatus: dorsal scales in (a) SMF 96471; (b) SMF 96470; flank scales in (c) SMF 96471; (d) SMF 96470; ventral scales in (e) SMF 96278; (f) SMF 84066. Scale bars equal 1.0 mm.

FIGURE 163. Hemipenis of Anolis unilobatus (SMF 84457): (left) sulcate view; (right) asulcate view. Scale bar equals 1.0 mm.

FIGURE 164. Map indicating collecting localities of Anolis unilobatus in the Mexican state of Oaxaca. Each symbol can represent one or more adjacent localities.