Dactyloa ibanezi (Poe, Latella, Ryan and Schaad 2009)

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 : 31-32

Dactyloa ibanezi (Poe, Latella, Ryan and Schaad 2009)

Figures 2 ; 13; 17J–L; 18H–J.

Anolis chocorum: Talbot (1974) , Myers (1971: in part.); Auth (1994: in part.); Martínez and Rodríguez (1994); Martínez et al. (1995); Young et al. (1999: in part.); Ibáñez et al. (2001: in part.); Fläschendräger and Wijffels (2009: in part.); Hamad (2009); Stadler (2010: in part.); Carrizo (2010: in part.).

Anolis ibanezi: Poe et al. (2009) ; Jaramillo et al. (2010); Castañeda and de Queiroz (2011).

Dactyloa chocorum: Savage and Guyer (1989: in part.); Savage (2002: in part.); Köhler (2003, 2008: in part.).

Holotype. MSB 72574, male, from the trails of Parque Nacional General de División Omar Torrijos Herrera, 5 km North of El Copé, Coclé province, Panama ( Fig. 1 : loc. 31; Fig. 21 ).

Diagnosis. A large species (maximum SVL 81 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. insignis , D. kunayalae , and D. microtus ). Dactyloa ibanezi can immediately be distinguished from these six species by its unique coloration consisting of immaculate green dorsal and lateral surfaces with several parallel, narrow, diagonal dark stripes on the flanks and bright yellow surrounding the eye ( Fig. 13 ). In addition, D. ibanezi 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 D. ibanezi ), and from D. casildae , D. frenata , and D. insignis by having fewer subdigital lamellae under the fourth toe (41 or fewer in D. ibanezi vs. 42 or more) as well as under the fourth finger (28 or fewer in D. ibanezi vs. 29 or more).

Description. Total length to 278 mm (in our material); SVL to 81 mm in males, to 78 mm in females; tail very long, about 2.5 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 rugose or keeled, almost flat in some specimens; IP distinct, surrounded by smaller, rugose scales; parietal eye distinct; scales of SS conspicuously enlarged, rugose; scales of supraorbital disk less conspicuously enlarged, rugose to slightly keeled; an elongate, keeled anterior superciliary scale, less than half as long as horizontal eye diameter, usually followed by a single similarly keeled, but much shorter scale; anterior sublabials slightly enlarged, but not as high as INL, keeled; scales of temporal arch much larger than those above and below; ear opening moderate, about as high as SPL and INL together, not as high as eye, about the size of IP; low nuchal and dorsal crests present; 0–2 rows of keeled middorsal scales slightly enlarged; other dorsal scales as well as lateral scales granular, keeled; ventrals larger than largest dorsals, very slightly keeled; scales on anterodorsal surface of thigh unicarinate; 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 large, extending posteriorly to between one-fourth and one-third of the distance between axilla and groin in large specimens; female dewlap moderate, extending posteriorly to slightly beyond axilla; interspaces between the conspicuously demarcated gorgetal-sternal scale rows almost free of scales in both sexes.

The completely everted hemipenis of MHCH 2184 ( Figs. 17 J–L) is a medium-sized, 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; large asulcate processus and ridge present; lobes finely calyculate, truncus with transverse folds.

Coloration in life. Dorsal and lateral surfaces homogeneously green, grading to yellowish green laterally; three to five parallel, narrow, dark, usually slightly curved lines descending diagonally on flanks; eye surrounded by bright yellow; ventral surfaces white, light gray, or light greenish yellow; iris brown; male dewlap orange with an uninterrupted white margin in young males, in older males only anterior margin white; female dewlap violet ( Fig. 13 ). Our specimens showed little metachrosis, their green color only occasionally changing to greenish brown (compare SMF 89459 in Figs. 2 and 13 E), but never throughout the whole animal. However, the intensity and width of the lateral stripes can alter considerably (compare Figures 13 A and B). Color photographs of Dactyloa ibanezi have been published by Köhler (2008), Poe et al. (2009), Fläschendräger and Wijffels (2009), and Stadler (2010).

The coloration in life of an adult female (SMF 91475, Figs. 13 A, B, G–I) was recorded as follows: Dorsal ground color of body, base of tail, head, and limbs Parrot Green (260) with a suggestion of Lime Green (159), grading into Lime Green (159) with a suggestion of Chartreuse (158) laterally; flanks with a series of Hooker’s Green (162) diagonal stripes; scarce Clay Color (123B) and Hooker’s Green (162) mottling scattered on dorsal and lateral surfaces; tail posterior to base Sepia (219); a Sulfur Yellow (157) ring around eye; ventral surface of head Lime Green (159), those of body and base of tail Pratt’s Payne’s Gray (88) mottled with dirty white; iris Cinnamon (123A); dewlap True Violet (172) with a dirty white margin and gorgetal-sternal stripes where scales of the same color are concentrated. The coloration in life of a male paratype (SMF 89459; Figs. 2 ; 13E, K, M, N) was recorded as follows (translated from Stadler 2010): Dorsal ground color of body, head, limbs, and tail Lime Green (159) with a suggestion of Yellow-Green (58); flanks with four diagonal, parallel dark green stripes; ventral ground color of body, head, limbs, and tail Chartreuse (158); a dirty white longitudinal stripe running from tip of snout to throat; ventral surface of thighs speckled with dirty white; eyes encircled by a Sulfur Yellow (157) line; dewlap Spectrum Orange (17) with Chartreuse (158) scales.

Coloration in preservative. After 22 months of preservation in 70% ethanol, all green has turned to blue; male dewlaps are cream white with blue scales, female dewlap dark lilac ( Figs. 18 H–J). In the male paratype SMF 89459, after four years in preservative, the blue has largely turned to lilac ( Fig. 18 H).

Geographic distribution. Dactyloa ibanezi is known from the Caribbean versant of eastern Costa Rica (Limón province near border with Panama) to the type locality at PNGDOTH in western Panama, at low and premontane elevations of 400–1070 m asl. It has been found in the Panamanian provinces of Bocas del Toro, Coclé, Colón, Veraguas, and the Comarca Ngöbe Buglé ( Fig. 21 ).

Natural history notes. All specimens were encountered at night while they were sleeping on leaves 1–2.5 m above ground. The female SMF 91475 and, one night later, the male SMF 91476 were found at virtually the same place, within less than 5 m distance between their respective sleeping sites. Around our collection sites of this species, we also found Dactyloa frenata , D. insignis , and D. kunayalae .

Our automatized temperature recordings at localities where we encountered Dactyloa ibanezi (480–900 m asl) range between 17.7–27.6°C. According to our combined dataset of 18 georeferenced occurrences, the species inhabits LMF, LWF, PMMF, and PMWF, with temperatures between 15.6–31.8°C, mean annual temperatures of 21.6–26.2°C and a total annual precipitation of 2494–3581 mm.

Conservation. Jaramillo et al. (2010) calculated an EVS of 8 for Dactyloa chocorum , which at the time comprised the populations described by Poe et al. (2009) as D. ibanezi , and assigned the species to the IUCN category LC. For D. ibanezi , we calculated the EVS as 4 (range) + 3 (persecution) + 4 (ecological distribution) = 11. Its extent of occurrence of 7336 km 2 and the continuing deforestation we observed in the region qualify the species for the IUCN category “Vulnerable” (VU).

Remarks. The female dewlap was described and pictured by Poe et al. (2009: Fig. 3 B) as “mostly dark orange.” Although the female pictured by these authors appears short-snouted and thus rather young, it remains unclear whether the difference in dewlap coloration between their and our female (with a violet dewlap, Fig. 13 G) is due to individual variation or ontogenetic change. Also, neither of the two possibilities can at this time be ruled out to explain the difference in dewlap coloration among our adult and young males (compare Figs. 13 E and F).

The new records provided herein document for the first time the occurrence of Dactyloa ibanezi in Colón province and the Comarca Ngöbe-Buglé. According to the catalogue of CHP, the specimen CHP 3529 from “Bocas del Toro” was collected at an elevation of 1067 m, which is the highest elevation reported so far for Dactyloa ibanezi . Köhler (2008, p. 99: Fig. 196) pictured an individual from “Reserva Forestal Fortuna, Panama, 400 m asl.” Since the photo was taken on the Caribbean versant (B. Akeret, pers. comm.), the correct locality is BPPS in the Comarca Ngöbe-Buglé, and is ubicated more to the north than the corresponding points mapped by Köhler (2008) and Poe et al. (2009). On their distribution map, the latter authors plotted a point in the Pacific drainage in southwestern Veraguas province, well south of the Cordillera Central. However, no corresponding specimen is mentioned, and we are unaware of any specimen or observation underlying this distributional record. To our knowledge, the species has not yet been reported from the Pacific versant west of the region around Santa Fé, Veraguas.

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 3 Results of molecular analyses. (A) Consensus tree from Bayesian analysis. Scale bar refers to number of substitutions per site. (B) Parsimony network with a 95 % parsimony probability resulting in a connection limit of 10 steps; each node represents a unique haplotype separated from the next by one substitution step. Small circles represent unsampled haplotypes, rectangles the probably ancestral haplotypes. Locality numbers refer to Fig. 1. (C) Consensus tree from Maximum Parsimony analysis. Scale bar refers to number of changes over whole sequence.

FIGURE 13. Specimens of Dactyloa ibanezi from western Panama: (A) female SMF 91475 from Willie Mazú in sleeping position and (B) the next day; (C) male MHCH 2184 from Cerro Mariposa; (D) juvenile from Sendero Pianista, photo by Johannes J. Köhler; (E – G) dewlaps of (E) male paratype SMF 89459 from Cerro Mariposa, (F) male SMF 91476 from Willie Mazú, and (G) female SMF 91475; (H) lateral, (I) dorsal, and (J) ventral views of head of SMF 91475; enlarged postcloacal scales of (K) SMF 89459 and (L) MHCH 2184; ventral views of (M) left foot and (N) right hand of SMF 89459.

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 21. Distribution of Dactyloa ibanezi and D. insignis 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 (*).