Anolis, Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix & Köhler, Gunther, 2013

Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix & Köhler, Gunther, 2013, Distribution and variation of the giant alpha anoles (Squamata: Dactyloidae) of the genus Dactyloa in the highlands of western Panama, with the description of a new species formerly referred to as D. microtus, Zootaxa 3626 (1), pp. 1-54 : 36-38

Dactyloa kunayalae (Hulebak, Poe, Ibáñez and Williams 2007)

Figures 2 ; 15; 17M– O; 18N–Q.

Anolis kunayalae: Hulebak et al. (2007) ; Poe et al. (2009); Jaramillo et al. (2010); Castañeda & de Queiroz (2011). Anolis new species 1: Nicholson et al. (2005).

Holotype. MSB 72605, male, from along the trails of Parque National General de División Omar Torrijos Herrera, 5 km North of El Cope, Coclé Province, Panama (8°40.315'N, 80°35.518'W) ( Fig. 1 : loc. 31; Fig. 22 ).

Diagnosis. A large species (maximum SVL 109 mm) of the genus Dactyloa (sensu Nicholson et al. 2012) that is most similar in external morphology to the other members of this clade found in western Panama ( D. casildae , D. frenata , D. ginaelisae , D. ibanezi , D. insignis , and D. microtus ). Dactyloa kunayalae can readily be distinguished from these six species by its coloration (described below, shown in Figs. 2 and 15 ), and by the morphology of its toes: In contrast to the other species, D. kunayalae has narrow lamellar pads that are barely dilated under all toes and fingers, a distal phalanx (including claw) of the fourth toe that is longer than phalanges ii and iii combined, and the lowest number of subdigital lamellae under the fourth toe (35 or fewer in D. kunayalae vs. 38 or more). Moreover, it has fewer subdigital lamellae under the fourth finger than the other species except D. ibanezi (25 or fewer in D. kunayalae vs. 29 or more). In addition, D. kunayalae differs from D. casildae , D. frenata , and D. ibanezi in having short legs (tip of fourth toe of adpressed hind limb reaching to tympanum in D. kunayalae ).

Description. Total length to 354 mm (in our material); SVL to 109 mm in males, to 81 mm in females; tail long, about 2–2.5 times SVL, compressed, without dorsal crest; legs short, tip of fourth toe of adpressed hind limb reaching to tympanum; internasals, canthals, and loreals keeled; scales of frontal and prefrontal area mostly rugose to keeled; IP distinct, surrounded by smaller, rugose scales; parietal eye distinct; scales of SS slightly but distinctly enlarged, rugose to keeled; scales of supraorbital disk less conspicuously enlarged, rugose or usually keeled; an elongate, keeled anterior superciliary scale (or two or three accordingly shorter ones, then together) at least half as long as horizontal eye diameter, usually followed by 1–3 similarly keeled, but much shorter scales; anterior sublabials slightly enlarged, but less high than INL; scales of temporal arch much larger than those above and below; ear opening moderate to large, higher than SPL and INL together, slightly lower to higher than eye, much larger than IP; nuchal and dorsal crests present; females without, males with two rows of keeled middorsal scales slightly enlarged; other dorsal scales as well as lateral scales granular, conical to keeled; ventrals larger than largest dorsals, smooth to slightly keeled; scales on anterodorsal surface of thigh unicarinate; scales on dorsal surface of forearm unicarinate, becoming multicarinate towards wrist; all fingers and toes with very weakly developed dilated pad, not even twice the width of distal phalanx; distal phalanx including claw of fourth toe longer than phalanges ii and iii combined; male dewlap large, extending posteriorly to one-third of the distance between axilla and groin; female dewlap moderate to large, extending posteriorly to up to one-fourth of the distance between axilla and groin in large specimens; interspaces between well-demarcated gorgetal-sternal scale rows almost free of scales in both sexes.

The completely everted hemipenis of SMF 91484 ( Figs. 17 M– O) is a small, slightly bilobate organ; sulcus spermaticus bordered by well-developed sulcal lips, opening at base of apex into two broad concave areas, one on each lobe; no asulcate processus or ridge present; lobes finely calyculate, truncus with transverse folds.

Coloration in life. Dactyloa kunayalae presents a pronounced sexual dichromatism and apparently also ontogenetic changes in coloration. Dorsal and lateral surfaces in males largely green, turquoise, and blue, with darker green or blue as well as brown and yellow blotches that tend to form transverse and reticulate bands and lines ( Figs. 15 A, F); females chiefly green with dark green to brown blotches usually suggesting transverse bands, and interspersed light markings ( Figs. 15 B–D, G–L); juvenile females with conspicuous dark vertical lip bars that extend onto ventral surfaces of head; larger females and the adult male with rather immaculate, yellowish green lips; similarly, lateral pattern extending further onto venter in juveniles, whereas adults have a largely unicolor venter. Both sexes with crossbanded limbs, a diffuse postorbital blotch or stripe, more or less pronounced dark crossbands throughout the tail, and a reddish brown iris; male dewlap white with an orange-yellow margin; female dewlap white with a narrower margin of lighter yellow ( Fig. 15 ). In the course of metachrosis, the ground color may assume a reddish brown similar to iris color, approximate the tonality of the dark markings, or the dark markings lighten up to become diffuse or barely visible. Color photographs of D. kunayalae have been published by Hulebak et al. (2007) and Köhler (2008).

The coloration in life of the adult male (SMF 91484, Figs. 15 A, F, M) was recorded as follows: Dorsal ground color Bunting Green (150) with a suggestion of Lime Green (159), suffused with Cyan (164) on the flanks and bearing a few scattered Natal Brown (219A) areas; body with mottling partly fusing to form a reticulum of Cream Color (54), grading into Raw Sienna (136) anteriorly and on the head; dorsal surfaces of body and legs mottled with Sepia (119); tail Light Drab (119C), grading into Drab-Gray (119D) ventrally, with Sepia (119) crossbands; lips and ventral surface of head Chartreuse (158); ventral surfaces of limbs, body and base of tail Sulphur Yellow (57); iris Mahogany Red (132B); dewlap Sulphur Yellow (57) on basal portions, grading into dirty white towards the center, and a broad Spectrum Orange (17) with a suggestion of Orange Yellow (18) margin; dewlap scales dirty white and Chartreuse (158). The coloration in life of a young female (SMF 91485, Figs. 2 ; 15B, H–L, N, O) was recorded as follows: Dorsal and lateral ground color of body, limbs, base of tail, and head Lime Green (159), grading into Pistachio (161) laterally, with Sepia (119) mottling except on snout that partly fuses to form broken crossbands on body, limbs, and base of tail; head, body, and limbs with scattered, diffuse blotches of Mahogany Red (132B); tail Ground Cinnamon (239) with Warm Sepia (221A) crossbands that grade into Sepia (119) dorsally, their Sepia (119) portions ventrally bordered by dirty white mottling; ventral surfaces of body, head, and limbs Tawny Olive (223D), mottled with Warm Sepia (221A) and dirty white, giving the impression of transverse stripes especially laterally and under head; iris Mahogany Red (132B); dewlap Yellowish Olive-Green (50) with a Buff Yellow (53) margin and broad, partly broken gorgetal-sternal rows of small Buff-Yellow (53) scales.

Coloration in preservative. After 22–74 months of preservation in 70% ethanol all yellow, green, and blue colors have given way to brown, cream, and white. Only a very faint suggestion of yellow is retained on the margin of the male dewlap after 23 months ( Figs. 18 N–Q).

Geographic distribution. Dactyloa kunayalae is distributed from eastern Panama to western Panama, mostly along the Caribbean versant. This Panamanian endemic has been recorded from the Comarcas Kuna Yala and Ngöbe-Buglé as well as from the provinces of Chiriquí, Coclé, Colón, Veraguas, and probably also Darién (see remarks). It is a denizen of low and premontane elevations at 320–1050 m asl ( Fig. 22 ).

Natural history notes. Our specimens were encountered at night while they were sleeping on branches and lianas 2–8 m above ground. We found this species to occur in syntopy with Dactyloa frenata , D. ibanezi , and D. insignis at two sites ( Fig. 1 : locs. 9 and 16), and with D. casildae at one locality ( Fig. 1 : loc. 13).

Our automatized temperature recordings at localities where we encountered Dactyloa kunayalae (480–1050 m asl) range between 17.6–27.6°C. According to our combined dataset of 26 georeferenced occurrences, the species inhabits LMF, PMMF, and PMWF, with temperatures between 14.7–30.1°C, mean annual temperatures of 20.6–25.4°C and a total annual precipitation of 1735–3452 mm.

Conservation. Jaramillo et al. (2010) calculated an EVS of 12 for Dactyloa kunayalae , and assigned the species to the IUCN category LC. We calculated the species’ EVS as 5 (range) + 3 (persecution) + 4 (ecological distribution) = 12. Its extent of occurrence of 19441 km 2 and the continuing deforestation we observed in the region qualify D. kunayalae for the IUCN category “Vulnerable” (VU).

Remarks. Our findings of Dactyloa kunayalae raise the altitudinal maximum by 250 m, originate from up to 180 km west of the documented range, and constitute the first records for Chiriquí, Veraguas, and the Comarca Ngöbe-Buglé (Hulebak et al. 2007). Moreover, GK had the privilege to examine the specimen FMNH 170034 (collected in August 1967 by Michael Duever at "border of Darien, Summit site, 8°55'N 77°51'W, 320m ") and identified it as an adult male D. kunayalae , probably the first specimen of this species ever collected. This specimen further extends the documented range some 135 km east-southeast from the nearest record in Nusagandí (Hulebak et al. 2007), may be regarded as the first record for Darién province, and sets the new lower vertical limit for the species.

Hulebak et al. (2007) described Dactyloa kunayalae to be long-legged, with the fourth toe reaching “to eye when hindleg pressed against body.” This trait was implied as diagnostic in the key of Köhler (2008). Strictly adhering to that key characteristic, all our specimens would have been identified as D. insignis or D. microtus , since their hind legs are short with the fourth toe merely reaching to tympanum. Although we do not know if there is really so much geographical variation in relative hind limb length in D. kunayalae , we consider it rather unlikely and prefer to attribute this discrepancy to different ways of extending or adpressing hind limbs, or bending the specimens’ bodies while doing so.

Figures

FIGURE 1. Collection localities of Dactyloa specimens in western Panama (main map) and Lower Central America (inset). Black circles represent examined specimens and reliable photo records listed in the Appendix, gray circles stand for other localities that are mentioned in the text. One symbol may summarize several localities close to each other. See text for abbreviations and details. (1) Río Changena; (2) Bajo Mono, Sendero La Cascada; (3) Alto Chiquero; (4) Sendero El Pianista, Casa de Calixto; (5) Sendero Pianista, 1500 m; (6) Cerro Guayabo; (7) headwaters of Río Chiriquí Malí; (8) Quebrada Arena; (9) Willie Mazú; (10) Cerro Pata de Macho; (11) Cerro Saguí; (12) Río Flor; (13) Río Hacha; (14) W slope Cerro Santiago, La Nevera; (15) E slope Cerro Santiago, Quebrada Ardilla; (16) Cerro Mariposa; (17) Cerro Negro; (18) Río Chilagres; (19) Distrito de Donoso: between Botija, Brazo, Petaquilla, and Rio del Medio; (20) Camp Summit; (21) La Palma; (22) Cerro Dantas; (23) Parque Nacional Tapantí; (24) Sendero Quetzales on N slope Volcán Barú, 7000 ft = 2130 m; (25) Cerro Horqueta; (26) Boquete; (27) BPPS, transect of Hofer and Bersier (2001); (28) Smithsonian station at RFLF; (29) Quebrada Frank; (30) road to Almirante, 400 m; (31) PNGDOTH north of El Copé; (32) Valle de Antón; (33) Altos de Campana; (34) Isla Barro Colorado; (35) Nusagandí.

FIGURE 2. (A) Maximum likelihood consensus tree of 19 Panamanian anoles. Black bars demarcate smallest well-supported clusters. (B) Sampled individuals in life, with locality numbers referring to Fig. 1. (C) Taxonomic identity derived from morphology on the basis of available literature.

FIGURE 15. Specimens of Dactyloa kunayalae from western Panama: (A) adult male SMF 91484 from BPPS; (B) young female SMF 91485 from Río Hacha; (C) female SMF 96009 from Cerro Mariposa; (D) SMF 96009 in sleeping position; (E) uncollected, probably male specimen from Río Chilagres; (F – H) dewlaps of (F) male SMF 91484, (G) female SMF 96009, and (H) young female SMF 91485; (I) lateral, (J) dorsal, and (K) ventral views of head of SMF 91485; (L) ventral view of SMF 91485; (M) enlarged postcloacal scales of SMF 91484; ventral views of (N) left foot and (O) left hand of SMF 91485.

FIGURE 17. Hemipenes of Dactyloa from western Panama: (A – C) Dactyloa casildae, SMF 91454: (A) sulcate, (B) asulcate, and (C) lateral view; (D – F) Dactyloa frenata, SMF 89467: (D) sulcate, (E) asulcate, and (F) lateral view; (G – I) Dactyloa ginaelisae, SMF 89498: (G) sulcate, (H) asulcate, and (I) lateral view; (J – L) Dactyloa ibanezi, MHCH 2184: (J) sulcate, (K) asulcate, and (L) lateral view; (M – O) Dactyloa kunayalae, SMF 91484: (M) sulcate, (N) asulcate, and (O) lateral view; all scale bars equal 1 mm.

FIGURE 18. Preserved specimens of Dactyloa from western Panama: (A – D) Dactyloa casildae, (A) and (B) young male MHCH 2121, SVL = 69 mm; (C) and (D) female SMF 89453, SVL = 81 mm; (E – G) D. frenata, (E) male SMF 89467, SVL = 132 mm, (F) and (G) young male SMF 91460, SVL = 72 mm; (H – J) D. ibanezi, (H) male paratype SMF 89459, SVL = 78 mm, (I) and (J) female SMF 91475, SVL = 78 mm; (K – M) D. insignis, (K) adult female SMF 89482, SVL = 124 mm, (L) and (M) juvenile female SMF 91477, SVL = 64 mm; (N – Q) D. kunayalae, (N) and (O) male SMF 91484, SVL = 103 mm; (P) and (Q) young female SMF 91485, SVL = 64 mm; (R) and (S) D. ginaelisae, female SMF 91503, SVL = 100 mm; (T) and (U) D. microtus, young male SMF 91499, SVL = 65 mm.

FIGURE 22. Distribution of Dactyloa ginaelisae, D. kunayalae, and D. microtus in western Panama (main map) and Lower Central America (inset). Localities of specimens with 16 S barcodes included in the molecular analyses are marked with an asterisk (*).