Dactyloa ginaelisae, 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 : 14-20

Dactyloa ginaelisae sp. nov.

Figures 2 ; 8; 9; 17G–I; 18R–S.

Anolis microtus: Dunn (1937: in part.); Slevin (1942); Taylor (1956: in part.); Peters and Donoso-Barros (1970: in part.); Savage and Talbot (1978: in part.); Arosemena et al. (1992: in part.); Auth (1994: in part); Young et al. (1999); Ibáñez et al. (2001); Köhler et al. (2008); Fläschendräger and Wijffels (2009: in part.); Hamad (2009: in part.); Lotzkat et al. (2010a); Jaramillo et al. (2010); Stadler (2010); Castañeda & de Queiroz (2011: in part.).

Dactyloa microtus: Savage and Guyer (1989: in part.); Savage (2002: in part.); Köhler (2003, 2008: in part.); Uetz (2013: in part.).

Holotype. SMF 91504 ( Figs. 2 ; 8; 9S–T), adult male, from the banks of Quebrada Juglí ( Fig. 19 F) on the southeastern slope of Cerro Saguí (also known as Cerro Ratón; locality 11 in Fig. 1 ) at Finca Alto Cedro, about 2 km north-northeast of the village Ratón, 8.5576°N, 81.8262°W, 1710 m asl, Corregimiento de Piedra Roja, Distrito de Kankintú, Comarca Ngöbe-Buglé, Panama; collected by Andreas Hertz and Sebastian Lotzkat on 0 8 July 2010; original field number SL 660.

Paratypes. Collected by Sebastian Lotzkat and Andreas Hertz, if not indicated otherwise. All from the Comarca Ngöbe-Buglé, Panama. Southeastern slope of Cerro Saguí ( Fig. 1 : loc. 11): MHCH 2240, juvenile female, same collecting data as holotype; MHCH 2239 and SMF 91503, females, near type locality, 8.5561°N, 81.8252°W, 1700 m asl, 0 7 July 2010; SMF 91502, juvenile female, about 850 m NNE of type locality, 8.5636°N, 81.8217°W, 1960 m asl, 0 8 July 2010. Western slope of Cerro Santiago, La Nevera ( Fig. 1 : loc. 14): SMF 89496 and 89497, juvenile female and adult female, 8.4997°N, 81.7724°W, 1700 m asl, 11 May and 17 August 2008; SMF 85069, juvenile female, 8.5000°N, 81.7722°W, 1600 m asl, collected by Abel Batista, Gunther Köhler, Marcos Ponce, and Javier Sunyer on 22 January 2006; SMF 89498, male, 8.5011°N, 81.7694°W, 1580 m asl, 14 August 2008; MHCH 2235, juvenile female, 8.5018°N, 81.7689°W, 1560 m asl, 10 August 2008; MHCH 2238, female, 8.4989°N, 81.7682°W, 1620 m asl, 11 November 2009; MHCH 2234, female, 8.5032°N, 81.7675°W, 1530 m asl, 10 August 2008; SMF 90133, female, 8.4954°N, 81.7673°W, 1810 m asl, 11 November 2009. Eastern slope of Cerro Santiago, Quebrada Ardilla ( Fig. 1 : loc. 15): MHCH 1338, female, 8.4974°N, 81.7228°W, 1600 m asl, 24 March 2009.

Referred specimens. Apart from the material collected by ourselves and listed in the Appendix, all specimens reported as Dactyloa microtus that have been collected in Panama to our knowledge are referrable to D. ginaelisae in view of their collection localities in the vicinities of Boquete, where we also collected three specimens of this species: ANSP 22418 and 22419 from above Boquete (Dunn 1937; Savage & Talbot 1978); CAS 79598 from the north slope of Volcán Barú (Slevin 1942); MVUP 966 from Cerro Horqueta and MVUP 969 from Bajo Mono near Boquete (MVUP catalogue); CHP 1038 from Cerro Horqueta (CHP catalogue).

Diagnosis. A large species (maximum SVL 112 mm) of the genus Dactyloa (sensu Nicholson et al. 2012) that is most similar in external morphology to the other members of this genus found in western Panama ( D. casildae , D. frenata , D. ibanezi , D. insignis , D. kunayalae , and D. microtus ). These species share a moderate to large adult size (SVL> 70 mm), a large dewlap and enlarged postcloacal scales in males, and smooth or faintly keeled ventrals. Dactyloa ginaelisae can readily be distinguished from these six species by its color pattern described below and shown in Figs. 2 , 8 , 9 , and 18R–S. It further differs from all mentioned species except D. microtus by its low numbers of horizontal loreal rows (4 or fewer in D. ginaelisae vs. 5 or more) and total loreal scales (25 or fewer in D. ginaelisae vs. 39 or more), and by its low number of scales around midbody (100 or usually much fewer in D. ginaelisae vs. 110 or more). Moreover, D. ginaelisae differs from D. casildae , D. frenata , and D. ibanezi in having short legs (tip of fourth toe of adpressed hind limb reaching to a point between tympanum and eye, very rarely to posterior border of eye, in D. ginaelisae vs. beyond eye; shank length/SVL = 0.22 or less in D. ginaelisae vs. 0.25 or more). Among the short-legged species of Dactyloa in western Panama, D. ginaelisae further differs from D. insignis in having fewer subdigital lamellae under the fourth toe (50 or fewer in D. ginaelisae vs. 52 or more) as well as under the fourth finger (36 or fewer in D. ginaelisae vs. 40), and from D. kunayalae in having more subdigital lamellae under the fourth toe (41 or more in D. ginaelisae vs. 35 or fewer) as well as under the fourth finger (29 or more in D. ginaelisae vs. 25 or fewer). Dactyloa ginaelisae is very similar to D. microtus , from which it differs in having longer legs (tip of fourth toe of adpressed hind limb reaching to a point between tympanum and eye in D. ginaelisae vs. to a point between shoulder and tympanum in D. microtus ; shank length/ SVL = 0.19 or more in D. ginaelisae vs. 0.183 or less) and by its conspicuous and clear-cut coloration pattern between eye and shoulder (a prominent light stripe extending from supralabials posteriorly above or a across the ear before bending down towards shoulder, delineating a dark preaxillary blotch above and posteriorly, and paralleled above by a dark postorbital stripe with darker borders that extends at least to a level above the preaxillary blotch in D. ginaelisae vs. light postsupralabial and dark postorbital stripe oriented more ventrally and losing their conspicuousness around ear).

Description of the holotype. Adult male as indicated by everted hemipenes ( Fig. 8 G), a pair of enlarged postcloacal scales ( Fig. 8 E), and presence of large dewlap ( Fig. 8 D); snout-vent length 107 mm; tail complete; tail length 244 mm, tail length/SVL ratio 2.28; tail compressed in cross section, tail height 7.0 mm, tail width 4.3 mm; axilla to groin distance 45.0 mm; head length 29.1 mm, HL/SVL ratio 0.27; snout length 14.2 mm; head width 16.7 mm; longest toe of adpressed hind limb reaching to a point between tympanum and eye; shank length 22.3 mm, shank length/SVL ratio 0.21, shank length/HL ratio 0.77; longest finger of extended forelimb reaching well beyond tip of snout; longest finger of adpressed forelimb reaching to anterior insertion of hind limbs; prefrontal ridges distinct, parietal ridges conspicuously protruding; scales on snout mostly rugose to wrinkled; 5 postrostrals; 6 scales between nasals; scales in distinct prefrontal depression wrinkled; supraorbital semicircles differentiated, composed of very rugose and partly wrinkled scales, separated by a minimum of 2 scales; supraorbital disc composed of 15 enlarged rugose scales; one slightly elongated, keeled anterior superciliary, followed posteriorly by a much smaller, keeled, elongate scale; about 2 rows of small keeled scales extending between enlarged supraorbitals and superciliary; interparietal plate not distinct, no parietal eye visible; canthal ridge distinct, composed of 5 large (posterior) and 1 small (anterior) canthal scales; 7 scales present between second canthals; 8 scales present between posterior canthals; 19/20 (right side/left side) wrinkled loreal scales in a maximum of 3 horizontal rows; subocular scales flat, wrinkled, subocular row well-defined; 7 supralabials to level below center of eye; ear opening 1.1 x 2.2 mm (length x height); mental distinctly wider than long, almost completely divided medially, bordered posteriorly by 6 postmentals; 7 infralabials to level below center of eye; sublabials enlarged, about as high as INL anterior to level of orbit, first two sublabials posterior to mental greatly enlarged, higher than INL; keeled granular to elongate scales present on chin and throat; dewlap large, extending well onto body, anterior insertion at a level halfway between orbit and tip of snout, posterior insertion at a level between one-fourth and one-third the distance between axilla and groin, with about 5 gorgetal-sternal rows 2–4 scales wide, becoming more diffuse posteriorly; low nuchal and dorsal crests present, dorsum of body with elevated, wrinkled scales, 2 middorsal rows of prominently protruding, keeled, but not otherwise enlarged scales, largest dorsal scales about 0.8 x 0.8 mm (length x width); about 42 medial dorsal scales in one HL; about 72 medial dorsal scales between axilla and groin; lateral scales raised, rugose to wrinkled, average size 0.7 mm in diameter, with minute granules occupying varying portions of the interspaces between them; ventrals at midbody smooth, subimbricate, about 0.5 x 0.5 mm (length x width); about 57 ventral scales in one HL; about 88 ventral scales between axilla and groin; about 94 scales around midbody; caudal scales strongly keeled, without whorls of enlarged scales, subcaudal scales with a single prominent keel; a pair of greatly enlarged postcloacal scales, larger one about 1.8 x 3.1 mm (length x width); tube-like axillary pocket not developed; scales on anterodorsal surface of thigh and on dorsal surface of forearm multicarinate, imbricate; digital pads dilated, dilated pad about 3 times width of non-dilated scales under distal phalanx; distal phalanx narrower than and raised from dilated pad; 33/35 lamellae under phalanges ii to iv of 4th toe; 12/13 scales under distal phalanx of 4th toe; 24 lamellae under phalanges ii to iv of 4th finger; 12/11 scales under distal phalanx of 4th finger.

The completely everted hemipenis is a medium-sized, bilobate organ; sulcus spermaticus bordered by welldeveloped sulcal lips, opening at base of apex into two broad concave areas, one on each lobe; large asulcate processus and ridge present; a knob-like processus present on each lateral side of truncus below base of apex; lobes finely calyculate, truncus with transverse folds.

Coloration in life. The coloration in life, including two stages of metachrosis, is shown in Figs. 8 A–E. The holotype represents the most contrastingly colored morph known for the species, with the dark crossbands on the dorsal and lateral surfaces of body, limbs, and anterior portion of tail well-delineated against the very light ground color by darker bands. Equally well-demarcated are the dark postorbital stripe extending posteriorly to above shoulder, and the large, elongate preaxillary blotch. The dirty to bright white supralabial stripe extends below the dark postorbital one, over and across the ear, to above the posterior portion of the preaxillary blotch, where it curves down towards the shoulder. Otherwise, no detailed notes of the holotype were taken. Coloration after approximately two years of preservation in 70% ethanol ( Figs. 8 F–M) is similar to that in life, apart from that all reddish, greenish, and bluish tonalities have faded.

Variation. The paratypes and referred specimens agree well with the holotype in terms of general morphometrics and pholidosis (see Tables 2 and 3), but are very variable in coloration ( Fig. 9 ). The following account of variation among our sample of Dactyloa ginaelisae is congruent with the standardized description sections in the subsequent species accounts for the other six species of Dactyloa found in western Panama.

Total length to 362 mm; SVL to 112 mm in males, to 108 mm in females; tail long, about 1.7–2.4 times SVL, compressed, with a low dorsal crest on the anterior portion; legs short, tip of fourth toe of adpressed hind limb reaching to a point between anterior border of tympanum and posterior border of eye; dorsal and lateral head scales generally large; internasals, canthals, and loreals rugose to wrinkled; scales of frontal and prefrontal area mostly rugose to wrinkled; IP indistinct in most specimens, if discernable, then usually surrounded by scales of both smaller and equal size; scales of parietal area generally rugose to wrinkled; parietal eye indistinct in most specimens; scales of SS distinctly enlarged, rugose; scales of supraorbital disk conspicuously enlarged, rugose; one or sometimes two usually only slightly elongate, keeled anterior superciliary scale(s), one-fourth to half as long as horizontal eye diameter, usually followed by several similarly keeled, but much shorter scales; one, two, or more anterior sublabials greatly enlarged, higher than INL; scales of temporal arch usually larger than those above and below; ear opening small, by far not as high as eye, less high than SPL and INL together, usually about as high as SPL; low nuchal and dorsal crests present; two rows of middorsal scales strongly keeled and projecting upwards, but not laterally enlarged; other dorsal scales as well as lateral scales elevated and rugose to wrinkled in adults, smooth in very small juveniles, with very small granules more or less densely filling the interspaces between them; ventrals not or only slightly larger than largest dorsals, smooth; scales on anterodorsal surface of thigh multicarinate; scales on dorsal surface of forearm multicarinate; 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, with well-demarcated gorgetal-sternal scale rows of densely arranged scales and widely spaced scales in the broad interspaces between the rows; female dewlap moderate, extending posteriorly to slightly beyond axilla, with more diffuse gorgetal-sternal rows owing to the less widely spaced scales in their interspaces.

The completely everted hemipenis of SMF 89498 ( Figs. 17 G–I) is a medium-sized, 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; a knob-like processus present on each lateral side of truncus just below base of apex; lobes finely calyculate, truncus with transverse folds.

Coloration in life. Dactyloa ginaelisae exhibits a very variable coloration. Among the typical pattern elements are dark crossbands around the tail and, mostly with light centers, on dorsal surfaces of limbs and digits; a light stripe extending posteriorly from below the eye over the ear opening before bending down towards shoulder, paralleled above by a dark postorbital stripe with darker borders; a dark preaxillary blotch between tympanum and shoulder. Otherwise, dorsal and lateral surfaces with spots, blotches, reticula, or solid bars or chevrons that are lighter and/or darker than ground color; ventral surfaces usually comparably unicolor white or yellow; ground and marking colors of dorsal and lateral surfaces ranging from bright white over different shades of yellow, green, blue, and brown to black; iris gray to blue; male dewlap light salmon color; female dewlap pinkish to salmon color, in some individuals with dark blotches ( Figs. 2 ; 8; 9). Apart from its highly variable pattern, this species is capable of overwhelming metachrosis (compare Figs. 9 B and C, D and E) as already noticed by Dunn (1937). The green phase, usually shown while the animal is sleeping, can culminate in restricting all colors to white, green, and blue, lightening up to invisibility some or all of the otherwise contrasting markings. Similarly, the dark or brown phase, often assumed when the animal is handled, can cause the whole animal to appear solid dark brown at its climax. In between, most coloration pattern elements of a given individual can apparently assume almost every color from the palette of this species. The only individual spotted at daytime was in the brown phase and showed no green at all. Color photographs of D. ginaelisae have been published by Köhler (2008), Köhler et al. (2008); Fläschendräger and Wijffels (2009), Hamad (2009), and Uetz (2013).

The coloration in life of an adult female (SMF 89501, Figs. 9 B–C, M–P) was recorded in the brown phase as follows: Dorsal and lateral ground color of body and limbs Straw Yellow (56) suffused with Tawny Olive (223D); dorsal and lateral surfaces of body and limbs with Jet Black (89) irregular, sometimes broken, lines forming a reticulum suggesting transverse bands that enclose Robin Rufous (340) blotches, are suffused with Emerald Green (163) spots and disintegrate into mottles lateroventrally; dorsal surface of head Light Russet Vinaceous (221D) with a reticulum of Sepia (219) bordered by shadings of Brick Red (132A) and Leaf Green (146); ventral surface of head Cream Color (54) with Tawny (38) shadings and Maroon (31) mottling posteriorly; a Buff (124) supralabial stripe originating anterior to eye with shadings of Paris Green (63) (also present on borders of infralabials) and grading to dirty white posterior to eye continues bordered by Sepia (119) above ear to level above shoulder; a Brick Red (132A) postorbital stripe bordered by Sepia (119) almost reaching to above shoulder; an elongate blotch of the same color behind ear opening; tail Drab-Gray (119D) with Walnut Brown (221B) transverse bands and various Vandyke Brown (221) scales; iris Sky blue (66); dewlap Pink (7) with dirty white scales and longitudinal series of Olive-Brown (28) flecks that fade away posteriorly. The coloration in life of another female (MHCH 2234, Figs. 9 G, Q) was recorded as follows: Dorsal ground color Lime Green (159), grading into Chartreuse (158) laterally; dorsal surfaces of body and limbs with partly broken Burnt Sienna (132) transverse bands, Spectrum Yellow (55) transverse stripes and Burnt Sienna (132), Sepia (119) as well as Shamrock Green (162B) mottling; dorsal surface of head Tawny Olive (223D) with a Sepia (219) V-shaped marking and some scale margins shaded in the same color; a Burnt Sienna (132) postocular stripe and blotch anterior to shoulder bordered by Sepia (119); a Spectrum Yellow (55) supralabial stripe extending above ear to above shoulder; supralabial Region Spectrum Yellow (55) with a suggestion of Yellow-Green (58), grading into Opaline Green (162D) towards ventral surface of head; ventral surfaces of body, limbs and base of tail Straw Yellow (56) with sparse Cinnamon (123A) mottling; tail grading into Olive-Yellow (52) with broad Sepia (219) transverse bands grading into Army Brown (219B) ventrally; iris Sky Blue (66); dewlap Pink (7) with a suggestion of Salmon Color (6), with Opaline Green (162D) scales aggregating to form longitudinal rows, and a suggestion of Yellowish Olive-Green (50) mottling.

Coloration in preservative. After 22–76 months of preservation in 70% ethanol, colors are largely reduced to white, cream, different shades of brown, and black. Some individuals have retained bluish-gray tonalities in certain places, often between eye and shoulder ( Figs. 8 F–M; 18R–S).

Geographic distribution. As far as we know, Dactyloa ginaelisae is endemic to the Cordillera Central of western Panama, where it occurs at premontane and lower montane elevations on both Caribbean and Pacific versants. The species has been recorded from Volcán Barú in Chiriquí province along 95 airline km to the eastern slopes of Cerro Santiago in the Comarca Ngöbe-Buglé, at 1370–2130 m asl ( Fig. 22 ).

Natural history notes. The holotype was encountered at 00:19 sleeping 6 m above ground on a branch in gallery forest. Most other specimens were encountered at night while they were sleeping on leaves, branches, or lianas 0.5–6 m above ground. An individual spotted fleeing on the ground at night had probably been driven from its sleeping site in the vegetation by our disturbance. One adult male was spotted around noon as it moved, trying to secure a flying moth, about 4 m above ground.

Our automatized temperature recordings at collection localities of Dactyloa ginaelisae (1430–1830 m asl) range between 12.6–22.2°C. According to our combined dataset of 36 georeferenced occurrences, the species inhabits PMWF and LMWF, with temperatures between 8.6–24.7°C, mean annual temperatures of 14.8–18.7°C and a total annual precipitation of 2208–2524 mm.

Dactyloa ginaelisae is a typical anole species of premontane and lower montane forests, particularly of cloud forests (sensu Myers 1969). Especially in the so-called elfin forest on ridgetops and summits, with its low canopy and exuberant epiphytic vegetation ( Fig. 19 G), D. ginaelisae can locally be found in high numbers at certain times. At RFLF and on the western slopes of Cerro Santiago ( Fig. 1 : localities 10 and 14), we found this species to occur syntopically with D. casildae , and in one case even to share a sleeping branch with that species.

Conservation. Jaramillo et al. (2010) calculated an EVS of 11 for Dactyloa microtus , which at the time comprised the populations described herein as D. ginaelisae , and assigned that species to the IUCN category LC. We calculated the new species’ EVS as 5 (range) + 3 (persecution) + 4 (ecological distribution) = 12. Its extent of occurrence of just 530 km 2 and the continuing deforestation we observed in the region qualify D. ginaelisae for the IUCN category “Endangered” (EN).

Etymology. Sebastian Lotzkat dedicates this exceptionally beautiful new species to his even more enchanting fiancée Gina Elisa Moog, who has made more than a third of his life worthwhile by now, in deepest gratitude for that wonderful time and pleasant anticipation of a mutual future.

Remarks. All previous records of Dactyloa microtus from Panama are from the area around Boquete and thus must have been based on individuals of D. ginaelisae , considering the distribution of the two species as documented herein. All previous authors have reported the absence of a distinct IP and parietal eye as a diagnostic character for Dactyloa microtus , and most individuals of D. ginaelisae share this characteristic. However, exceptions exist: three of our 26 specimens have a well-discernable IP with parietal eye (also see remarks concerning this matter for D. insignis and D. microtus ). Apart from the diagnostic differences used here to distinguish D. ginaelisae from D. microtus , we observed other differences among our material. Without having examined more specimens of the latter species, preferably from both Costa Rica and Panama, we refrain from including these discrepancies in the formal diagnosis, but provide a brief summary below.

Regarding general morphology, individuals of Dactyloa microtus seem to be able to raise a much higher nuchal crest than representatives of D. ginaelisae . To a lesser extent, this also seems to apply to the dorsal crest, at least along its anterior portion.

Regarding scalation, three additional differences deserve mention: First, most individuals of Dactyloa ginaelisae have at least two greatly enlarged (at least as high as the adjacent INL) anterior sublabials following the mental plate on each side of the head ( Figs. 8 I, J; 9J–N, P). The three individuals of D. microtus in our collection have only the first sublabial greatly enlarged, which is followed posteriorly by a much narrower, rather elongate sublabial. This second sublabial (and usually the similarly shaped third one following it) is bordered medially by an enlarged scale which may approximate the size of the greatly enlarged first sublabial ( Figs. 16 E–H, J), but is not, or only very narrowly, in contact with the INL. Second, the dorsal and lateral scales of adult and subadult D. ginaelisae are prominently protruding from the skin and very rugose to wrinkled or clearly keeled ( Fig. 8 K), but rather flat, smooth, and not protruding in the holotype of D. microtus ( Fig. 7 G). However, our subadult males of D. microtus have the dorsal and lateral body scales less flat than the holotype, as it seems to be the case with the individuals photographed in Costa Rica. Third, the median subcaudal scales of D. ginaelisae bear one very prominent keel ( Fig. 8 L), whereas those of the holotype ( Fig. 7 H) and our subadult males of D. microtus are bi-, tri-, or multicarinate.

Regarding coloration, we observed two additional discrepancies. First, individuals of Dactyloa ginaelisae typically have an iris color ranging from gray over bluish-gray to bright blue in life ( Figs. 8 ; 9), whereas the representatives of D. microtus of which we have color photos in life ( Figs. 7 ; 16) have the iris dominated by brownish or purplish tonalities. Secondly, the dewlap is salmon to pinkish already in the young males of D. ginaelisae ( Fig. 9 K), whereas the dewlap is yellowish in the two young males of D. microtus ( Figs. 16 E–F).

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 7. Specimens of Dactyloa microtus from Costa Rica: (A – H) holotype USNM 31282, adult female, SVL = 106 mm: entire specimen photographed by James Pointdexter in (A) lateral and (B) ventral view; (C) dorsal, (D) lateral, and (E) ventral view of head; (F) ventral view of cloacal region showing lack of enlarged postcloacal scales; (G) detail of dorsal scales at midbody, arrows indicate middorsal rows; (H) detail of subcaudal scales, each bearing more than one prominent keel; (I, J) individual photographed by Mason Ryan at Parque Nacional Tapantí, Cartago province; (K, L) individual photographed by Daniel Cascante at Cerro Dantas, Heredia province. Scale bars = 1 mm, all arrows point posteriorly.

FIGURE 8. Holotype of Dactyloa ginaelisae sp. nov. (SMF 91504, adult male, SVL = 107 mm), (A – E) in life: (A) lateral view at time of capture; (B) portrait the next day; (C) dorsal view of entire specimen; (D) dewlap (width of forceps = 3 mm); (E) ventral view of cloacal region showing enlarged postcloacal scales; (F – M) specimen after approximately two years in preservative: (F) dorsal and (G) ventral view of entire specimen; (H) dorsal, (I) lateral, and (J) ventral view of head; (K) detail of dorsal scales at midbody, arrows indicate middorsal rows; (L) detail of subcaudal scales with one prominent keel; (M) detail of multicarinate scales on upper surface of right forearm. All arrows point posteriorly, scale bars = 1 mm.

FIGURE 9. Specimens of Dactyloa ginaelisae from Chiriquí and the Comarca Ngöbe-Buglé: (A) adult male MHCH 2237 from RFLF; (B) female SMF 89501 from RFLF in sleeping position and (C) the next day; (D) female SMF 91503 from Cerro Saguí in sleeping position and (E) the next day; (F) female SMF 89497 and (G) female MHCH 2234 from La Nevera; (H) uncollected female juvenile and (I) uncollected female from La Nevera; (J – M) dewlaps of (J) adult male SMF 89738 from RFLF, (K) male juvenile SMF 89737 from PNVB, (L) female SMF 89499 from RFLF, and (M) female SMF 89501; (N) lateral, (O) dorsal, and (P) ventral views of head of SMF 89501; (Q) ventral view of MHCH 2234; (R) enlarged postcloacal scales of SMF 89738; ventral views of (S) left foot and (T) left hand of male holotype SMF 91504 from Cerro Saguí.

FIGURE 16. Specimens of Dactyloa microtus from Río Changena, Bocas del Toro: young male SMF 91499 in (A) sleeping position and (B) the next day; (C) and (D) young male SMF 91500; (E – G) dewlaps of (E) young male SMF 91499, (F) young male SMF 91500, and (G) female hatchling SMF 91501; (H) lateral, (I) dorsal, and (J) ventral views of head of SMF 91500; enlarged postcloacal scales of (K) SMF 91500 and (L) SMF 91499; ventral views of (M) right foot and (N) right hand of SMF 91500.

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 19. Habitats of Dactyloa in western Panama. From west to east (collected species in parentheses): (A) Lower Montane Wet Forest (LMWF) at Río Changena, 1640 m asl (D. microtus); (B) LMWF at Sendero La Cascada, 1830 m asl (D. ginaelisae); (C) Premontane Wet Forest (PMWF) at headwaters of Río Chiriquí Malí, BPPS, 1050 m asl (westernmost and highest locality for D. kunayalae); (D) PMWF at Willie Mazú, 640 m asl (D. frenata, D. ibanezi, and D. insignis); E) PMWF at Cerro Pata de Macho, RFLF, ca. 1500 m asl (D. casildae and D. ginaelisae); (F) looking north from pasture at ca. 1800 m asl onto primary and secondary LMWF along valley of Quebrada Juglí, ca. 1710 m asl (type locality of D. ginaelisae); (G) LMWF at Cerro Santiago near La Nevera, 1810 m asl (D. ginaelisae); (H) looking from PMWF at Cerro Negro, 700 m asl (D. frenata), towards PMWF on Cerro Mariposa (D. frenata, D. ibanezi, D. insignis, and D. kunayalae).

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 (*).