Dactyloa frenata (Cope 1899)

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 : 27-29

Dactyloa frenata (Cope 1899)

Figures 2 ; 12; 17D–F; 18E–G.

Anolis frenatus: Cope (1899) ; Dunn (1937); Breder (1946); Peters and Donoso-Barros (1970); Myers (1971); Scott et al. (1976); Myers (1977); Savage and Talbot (1978); Rand and Myers (1990); Arosemena et al. (1992); Auth (1994); Martínez and Rodríguez (1994); Martínez et al. (1995); Ibáñez et al. (1995, 1996, 1997); Young et al. (1999); Ibáñez et al. (2001); Poe (2004); Nicholson et al. (2005); Hamad (2009); Fläschendräger and Wijffels (2009); Poe et al. (2009); Ugueto et al. (2009); Carrizo (2010); Jaramillo et al. (2010); Lotzkat et al. (2010b); Stadler (2010); Castañeda & de Queiroz (2011).

Anolis purpurescens: Taylor (1956) and references therein.

Holotype. Lost; type locality in Colombia, but exact location uncertain according to Savage and Talbot (1978).

Diagnosis. A large species (maximum SVL 143 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. ginaelisae , D. ibanezi , D. insignis , D. kunayalae , and D. microtus ). Dactyloa frenata can readily be distinguished from these six species by the presence of a pronounced light interorbital bar with dark anterior and posterior borders, and by its unique color pattern on dorsum and flanks consisting of dark blotches or ocelli arranged to form oblique bands ( Fig. 12 ). Moreover, D. frenata has 9 or more SPL (vs. usually 8 or fewer) to the level below center of eye. In addition, D. frenata differs from D.ginaelisae , D. insignis , D. kunayalae , and D. microtus in having long legs (tip of fourth toe of adpressed hind limb reaching to eye or beyond in D. frenata ), and from D. ginaelisae , D. ibanezi , D. insignis , D. kunayalae , and D. microtus in having the suboculars separated from the SPL by at least one scale row (vs. suboculars and SPL in contact).

Description. Total length to 430 mm; SVL to 143 mm in males, to 118 mm in females; tail long, about two times SVL, compressed, without dorsal crest; legs long, tip of fourth toe of adpressed hind limb reaching to eye or beyond; internasals, canthals, and loreals keeled; scales of frontal and prefrontal area mostly smooth, some weakly keeled; IP distinct, surrounded by smaller, mostly smooth, scales; parietal eye distinct; scales of SS not distinctly enlarged, keeled; scales of supraorbital disk only slightly enlarged, keeled; an elongate, keeled anterior superciliary scale, at least half as long as horizontal eye diameter, often followed by one or two similarly keeled, but much shorter scales; anterior sublabials not distinctly enlarged, keeled; scales of temporal arch slightly larger than those above and below; ear opening large, higher than SPL and INL together, at least as high as eye, much larger than IP; nuchal and dorsal crests pronounced in males, weakly developed in female; 2–4 rows of keeled middorsal scales slightly enlarged; other dorsal scales as well as lateral scales granular, smooth to slightly keeled; ventrals slightly larger than largest dorsals, smooth to very slightly keeled; scales on anterodorsal surface of thigh mostly unicarinate, some bi- or tricarinate; scales on dorsal surface of forearm unicarinate, becoming multicarinate towards wrist; fourth toe with well-developed dilated pad, about three times width of distal phalanx; male dewlap very large, extending posteriorly to almost one-half of the distance between axilla and groin in large specimens; female dewlap moderate, extending posteriorly to slightly beyond axilla; dewlap in both sexes with welldemarcated gorgetal-sternal scale rows and scattered scales in their interspaces.

The completely everted hemipenis of SMF 89467 ( Figs. 17 D–F) 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; a small asulcate ridge present; lobes finely calyculate, truncus with transverse folds.

Coloration in life. Dorsal and lateral surfaces green, usually involving brownish, yellowish, or bluish tonalities; series of darker green to dark brown blotches or ocelli form oblique transverse bands on dorsum, flanks, limbs, and anterior portion of tail; posterior portion of tail with dark crossbands; alternating yellow and dark stripes radiate more or less distinctly from eye, the upper ones meeting on dorsal surface of head to form a pronounced light-colored interorbital bar usually with dark anterior and posterior borders; ventral surfaces largely unicolor cream or gray, becoming lighter towards head; iris brown; male dewlap white, female dewlap brown ( Figs. 2 ; 12). We did not observe metachrosis in Dactyloa frenata . Color photographs of D. frenata have been published by Savage (2002), Köhler (2003, 2008), Fläschendräger and Wijffels (2009), Stadler (2010), Carrizo (2010), and Uetz (2013), among others.

The coloration in life of an adult male (SMF 89467, Figs. 12 A, E) was recorded as follows (translated from Stadler 2010): Dorsal surfaces of body, limbs, and anterior portion of tail Opaline Green (162D) with numerous Parrot Green (60) ocelli, partly bordered by Straw Yellow (56) and Spectrum Yellow (55), which are arranged in a pattern of transverse bands; middle portion of tail Sepia (119), posterior portion Olive-Yellow (52); anterior dorsal surface of head with a heterogeneous pattern of Salmon Color (6), Olive-Gray (42), and Cyan (164); a Salmon Color (6) interorbital bar bordered anteriorly and posteriorly by Greenish Olive (59) stripes; posterior dorsal surface of head with a random pattern of Flesh Color (5), Paris Green (63), and Salmon Color (6) scales; lateral surfaces of head from snout to posterior border of orbit Olive Gray (42), those posterior to orbit Cyan (164); Spectrum Yellow (55) and Grayish Olive (43) stripes extending from eyes radially in all directions; ventral surfaces largely dirty white, grading into Spectrum Yellow (53) laterally; ventral surfaces of hind limbs and anterior portion of tail Straw Yellow (53), those of middle and posterior portions of tail Sepia (119) and Olive-Yellow (52), respectively; dewlap Cream Color (54) with Sulphur Yellow (157) scales on the upper and Glaucous (80) scales in the lower half; margin dirty white anteriorly and Cream Color (54) posteriorly; iris Antique Brown (37).

Coloration in preservative. After 21–45 months of preservation in 70% ethanol, the greenish tonalities have turned to brown, gray, or bluish gray (the latter especially on head and flanks), venters are cream-colored ( Figs. 18 E–G).

Geographic distribution. Dactyloa frenata is distributed from the Caribbean versant of Costa Rica to Colombia, at low and premontane elevations of 2–900 m asl. In Costa Rica and extreme western Panama, it is restricted to the Caribbean versant of the Talamancan highlands. From the Fortuna depression eastwards, D. frenata is also found in the Pacific drainage. In Panama, the species has been recorded from the provinces of Bocas del Toro, Chiriquí, Coclé, Colón, Darién, Panamá, and Veraguas, and from the Comarcas Kuna Yala and Ngöbe-Buglé ( Fig. 20 ).

Natural history notes. All specimens were encountered at night while they were sleeping on branches or lianas 1.5–4 m above ground. Around our collection sites of Dactyloa frenata , this species shares its habitat with D. ibanezi , D. insignis , and D. kunayalae .

Our automized temperature recordings at collection localities of Dactyloa frenata (700–900 m asl) range between 16.1–26.2°C. According to our combined dataset of 75 georeferenced occurrences, the species inhabits LMF, LWF, PMMF, and PMWF, with temperatures between 15.2–32.4°C, mean annual temperatures of 21.2–26.8°C and a total annual precipitation of 1735–4417 mm.

Conservation. Jaramillo et al. (2010) calculated an EVS of 8 for Dactyloa frenata , and assigned the species to the IUCN category LC. We calculated the species’ EVS as 1 (range) + 3 (persecution) + 4 (ecological distribution) = 8. Its extent of occurrence of more than 49000 km 2 does not qualify D. frenata for any of the “Threatened” IUCN categories. Considering the continuing deforestation we observed in the region, we place the species in the category “Near Threatened” (NT).

Remarks. The adult male in our sample raised the upper vertical limit for this species to 900 m asl (Stadler 2010). As confirmed by Ugueto et al. (2009), Dactyloa frenata does definitely not occur in Venezuela.

Figures

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 12. Specimens of Dactyloa frenata from western Panama: (A) adult male SMF 89467 from Cerro Mariposa, photo by Nadim Hamad and Leonhard Stadler; (B) juvenile male SMF 91459 and (C) young male SMF 91460 from Willie Mazú; (D) female SMF 89897 from Cerro Negro; (E – G) dewlaps of (E) adult male SMF 89467, photo by Nadim Hamad and Leonhard Stadler, (F) young male SMF 91460, and (G) female SMF 89897; (H) lateral, (I) dorsal, and (J) ventral views of head of SMF 91460; enlarged postcloacal scales of (K) SMF 91460 and (L) SMF 91459; ventral views of (M) left foot and (N) left hand of SMF 91460.

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 20. Distribution of Dactyloa casildae and D. frenata 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 (*). Records of D. frenata from Colombia, such as the type locality, are not shown.