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Dactyloa ginaelisae sp. nov. Figures 2; 8; 9; 17 G–I; 18 R–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. (2010 a); 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; 9 S–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 18 R–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 4 th toe; 12 / 13 scales under distal phalanx of 4 th toe; 24 lamellae under phalanges ii to iv of 4 th finger; 12 / 11 scales under distal phalanx of 4 th 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 (223 D); 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 (221 D) with a reticulum of Sepia (219) bordered by shadings of Brick Red (132 A) 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 (132 A) postorbital stripe bordered by Sepia (119) almost reaching to above shoulder; an elongate blotch of the same color behind ear opening; tail Drab-Gray (119 D) with Walnut Brown (221 B) 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 (162 B) mottling; dorsal surface of head Tawny Olive (223 D) 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 (162 D) towards ventral surface of head; ventral surfaces of body, limbs and base of tail Straw Yellow (56) with sparse Cinnamon (123 A) mottling; tail grading into Olive-Yellow (52) with broad Sepia (219) transverse bands grading into Army Brown (219 B) ventrally; iris Sky Blue (66); dewlap Pink (7) with a suggestion of Salmon Color (6), with Opaline Green (162 D) 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; 18 R–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

Dactyloa insignis (Cope 1871) Figures 2; 14; 18 K���M. Anolis microtus: Peters and Donoso-Barros (1970: in part.: referring to the holotype of Diaphoranolis brooksi). Anolis insignis: Cope (1871, 1876); Boulenger (1885); Dunn (1937); Taylor (1956); Peters and Donoso-Barros (1970); Savage (1974); Myers (1977); Savage and Talbot (1978); Arosemena et al. (1992); Auth (1994); Mart��nez and Rodr��guez (1994); Young et al. (1999); Ib����ez et al. (2001); Poe (2004); Hamad (2009); Fl��schendr��ger and Wijffels (2009); Poe et al. (2009); Carrizo (2010); Jaramillo et al. (2010); Lotzkat et al. (2010 b); Stadler (2010); Casta��eda & de Queiroz (2011). Diaphoranolis brooksi: Barbour (1923). Holotype. Lost according to Savage and Talbot (1978); from Costa Rica: Provincia de San Jos��: near Ciudad San Jos��; probably from near La Palma (Savage 1974; Fig. 1: loc. 21; Fig. 21). Diagnosis. A large species (maximum SVL 160 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. kunayalae, and D. microtus). Dactyloa insignis can readily be distinguished from these six species by its unique coloration of fine reticulate lines forming transverse crossbands on dorsum and flanks, and a preaxillary dark blotch (Fig. 14). Moreover, D. insignis has all dorsal and lateral body scales smooth (vs. conical, keeled, rugose or wrinkled in the other species), and the highest number of scales around midbody (190 or more vs. 178 or fewer). In addition, D. insignis differs from D. casildae, D. frenata, and D. ibanezi in having short legs (tip of fourth toe of adpressed hind limb reaching at most to a point between shoulder and tympanum in D. insignis), and from D. ginaelisae, D. ibanezi, D. kunayalae, and D. microtus in having more subdigital lamellae under the fourth toe (52 or more in D. insignis vs. 50 or fewer) as well as under the fourth finger (40 in D. insignis vs. 37 or fewer). Description. Total length to 464 mm; SVL to 160 mm in males, to 135 mm in females; tail long, about two times SVL, compressed, without dorsal crest; legs short, tip of fourth toe of adpressed hind limb reaching at most to a point between shoulder and tympanum; internasals, canthals, and loreals keeled; scales of frontal and prefrontal area mostly flat, smooth; IP usually distinct, surrounded by not much smaller scales (indistinct from surrounding scales in our juvenile); parietal eye usually distinct (barely visible in our juvenile); scales of SS barely enlarged, smooth; scales of supraorbital disk not conspicuously enlarged, smooth; two or three only slightly elongate, keeled anterior superciliary scales, none reaching a length of one-fourth of horizontal eye diameter; anterior sublabials distinctly enlarged, about as high as INL, keeled; scales of temporal arch conspicuously 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; 0���2 rows of smooth middorsal scales slightly enlarged; other dorsal scales as well as lateral scales flat, smooth; ventrals much larger than largest dorsals, smooth; scales on anterodorsal surface of thigh mostly smooth, only on anterior edge unicarinate with a few bi- or tricarinate; scales on dorsal surface of forearm smooth, becoming unicarinate towards wrist and anterior edge; fourth toe with well-developed dilated pad, about three times width of distal phalanx; female dewlap moderate, extending posteriorly to slightly beyond axilla, with broad, diffuse gorgetal-sternal rows leaving very narrow interspaces with less densely packed scales. The hemipenis of Dactyloa insignis remains undescribed. Coloration in life. Dorsal and lateral surfaces white to brown; dorsum, flanks, limbs, and tail with fine reticulate lines conglomerating to form light-centered crossbands that become solid only on posterior portion of tail; lips with dark vertical bars; a pronounced, light-centered blotch between tympanum and axilla, and a dark postorbital stripe extending to above tympanum; ventral surfaces largely white, except for dark crossbands forming complete rings under tail and dark flank markings and lip bars extending onto ventral surfaces; iris brown; female dewlap brown, with dark and light blotches and lines (Figs. 2; 14). Dactyloa insignis exhibits considerable metachrosis with ground color ranging from white to dark brown, and the dark markings being green, brown, or almost black. Most notably, in certain situations numerous beige to orange blotches appear on body and head, and even the color of the iris can range from almost white to dark reddish brown (compare Figures 14 B, C, and F). Color photographs of D. insignis have been published by Savage (2002), K��hler (2003, 2008), Fl��schendr��ger and Wijffels (2009), Stadler (2010), and Uetz (2013). The coloration in life of the adult female (SMF 89482, Figs. 2; 14 B���D, F, G) was recorded as follows: Ground color Sulphur Yellow (57) with Sepia (219) broken crossbands on dorsum which contain Pale Pinkish Buff (121 D) blotches; vertical series of Salmon Color (106) blotches between crossbands; several Spectrum Orange (17) blotches on neck; a Raw Umber (23) postorbital stripe; a Burnt Umber (22) blotch with Raw Umber (23) center anterior to shoulder; tail with Sepia (119) crossbands, anterior ones suffused with Natal Brown (219 A); ventral surfaces of body and tail dirty white; Dorsal surfaces of limbs with Dark Brownish Olive (129) and Lime Green (59) transverse lines as well as transverse series of Pale Pinkish Buff (121 D) flecks; iris Cinnamon-Rufous (40); dewlap Sayal Brown (223 C) with Indigo (73) horizontal lines and horizontal series of dirty white flecks. The coloration in life of the juvenile female (SMF 91477, Figs. 2; 14 A, E, H���L) was recorded as follows: Dorsal ground color Opaline Green (162 D), grading into dirty white with a suggestion of Pale Horn Color (92) ventrolaterally; head and body with a reticulum of fine Sepia (119) lines that are suffused with, and partly turn into, Emerald Green (163), reach onto venter and accumulate along middorsum to give the impression of four blotches; limbs, toes, and fingers with pairs of transverse Sepia (119) lines; tail with six Sepia (119) transverse bands, the anterior three composed of dense reticula of lines similar to the middorsal blotches; head, body, and limbs with irregularly distributed Orange-Rufous (132 C) and Cream Color (54) blotches; ventral ground color dirty white with a suggestion of Pale Horn Color (92); iris Pale Pinkish Buff (121 D); tongue Olive-Gray (42); dewlap dirty white with a suggestion of Pale Horn Color (92) at base, grading into Ground Cinnamon (239) mottled with Buff (24) towards apex, with Sepia (119) longitudinal lines, longitudinal series of dirty white spots, and a Pale Horn Color (92) margin. Coloration in preservative. After 23���48 months of preservation in 70 % ethanol, the ground color has lightened to dirty white, and all markings have assumed grayish or brownish tonalities. Only slight suggestions of blue where green was present in life in our juvenile are still discernable after 23 months (Figs. 18 K���M). Geographic distribution. Dactyloa insignis is distributed throughout Costa Rica and Panama at low and premontane elevations of 0���1600 m asl. In Panama, the species has been recorded from the provinces of Bocas del Toro, Chiriqu��, Cocl��, Col��n, Dari��n, Panam��, and Veraguas, and from the Comarca Ng��be-Bugl�� (Fig. 21). According to Young et al. (1999), it is also present in the Comarca Kuna Yala. Natural history notes. Both specimens were encountered at night while they were sleeping on branches 5���6 m above ground. Around our collection sites, Dactyloa insignis co-occurs with D. frenata, D. ibanezi, and D. kunayalae. Our automatized temperature recordings around our collection sites of Dactyloa insignis (640 and 880 m asl) range between 18.2���26.2 ��C. According to our combined dataset of 36 georeferenced occurrences, the species inhabits LMF, LWF, PMMF, and PMWF, with temperatures between 12.4���33.7 ��C, mean annual temperatures of 18.0��� 26.4 ��C and a total annual precipitation of 1696���4373 mm. Conservation. Jaramillo et al. (2010) calculated an EVS of 11 for Dactyloa insignis, and assigned the species to the IUCN category LC. We calculated the species��� EVS as 4 (range) + 3 (persecution) + 4 (ecological distribution) = 11. Its extent of occurrence of more than 63000 km 2 does not qualify D. insignis 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. Our record from Willie Maz�� (Fig. 1: loc. 9) constitutes the first record for the Comarca Ng��be- Bugl��. In the maps of K��hler (2003, 2008), the distribution area of Dactyloa insignis extends into Colombia. Although it is very likely that the species ranges at least into the Colombian Dari��n, no specimens from this country are known to date. The occurrence of D. insignis in the Comarca Kuna Yala as stated by Young et al. (1999) seems plausible considering that the specimen FMNH 170087 was collected at Paradise Camp, which lies less than 4 km west of Camp Summit (Fig. 1: loc. 20) that is situated on the border between Dari��n province and the Comarca Kuna Yala. Unfortunately, our sample is composed of just two females. Savage (2002) pictured male individuals from Monteverde in Costa Rica (plates 256 and 257), and described the male dewlap as dark red (key on p. 451) or as ���primarily orange-red, with or without several horizontal green or white bars and dark spotting��� (p. 455) with a white to greenish free margin. Both Savage (2002) and K��hler (2008) stated that Dactyloa insignis has 51 or more lamellae under the fourth toe, and used this as a key characteristic to distinguish this species from D. microtus, which has at most 49 according to these authors. Nevertheless, Savage and Talbot (1978) counted just 50 lamellae on the fourth toe of MCZ 16297, a value that is also reached by one of our specimens of D. ginaelisae (which formerly was included in D. microtus). All previous authors have reported the presence of a distinct IP and parietal eye as a diagnostic character to distinguish D. insignis from D. microtus. However, the IP is indistinct and the parietal eye barely visible in one of our two specimens (Fig. 14 H), suggesting that strict adherence to this key characteristic might not always be recommendable (also see remarks concerning this matter for D. ginaelisae and D. microtus). Savage and Talbot (1978) pointed out differences in anterior head scale surface and coloration between individuals from Costa Rica and Pacific west-central Panama on the one hand and examples from northwestern, central, and eastern Panama on the other hand, concluding that additional material was required to finally evaluate whether or not two species are involved. Our specimens have smooth anterior dorsal head scales and, just as the individuals photographed by Michael Castillo in Donoso district in Col��n province, agree in exhibiting dark lip bars, a dark postorbital marking (in form of a short stripe rather than a spot or blotch), a dark preaxillary blotch, and in having their transverse bands formed by reticula. Thus, our specimens would fall somewhere between Savage and Talbot���s (1978) specimens from Costa Rica and their specimen from Valle de Ant��n (Cocl�� province). Since we have no male from our study area available, we cannot contribute further to the issue of geographical variation in male dewlap coloration., Published as part of Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix & K��hler, 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, pp. 1-54 in Zootaxa 3626 (1) on pages 32-34, DOI: 10.11646/zootaxa.3626.1.1, http://zenodo.org/record/283424

Dactyloa ginaelisae sp. nov. Figures 2; 8; 9; 17 G���I; 18 R���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); Ib����ez et al. (2001); K��hler et al. (2008); Fl��schendr��ger and Wijffels (2009: in part.); Hamad (2009: in part.); Lotzkat et al. (2010 a); Jaramillo et al. (2010); Stadler (2010); Casta��eda & de Queiroz (2011: in part.). Dactyloa microtus: Savage and Guyer (1989: in part.); Savage (2002: in part.); K��hler (2003, 2008: in part.); Uetz (2013: in part.). Holotype. SMF 91504 (Figs. 2; 8; 9 S���T), adult male, from the banks of Quebrada Jugl�� (Fig. 19 F) on the southeastern slope of Cerro Sagu�� (also known as Cerro Rat��n; locality 11 in Fig. 1) at Finca Alto Cedro, about 2 km north-northeast of the village Rat��n, 8.5576 ��N, 81.8262 ��W, 1710 m asl, Corregimiento de Piedra Roja, Distrito de Kankint��, Comarca Ng��be-Bugl��, 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 Ng��be-Bugl��, Panama. Southeastern slope of Cerro Sagu�� (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 K��hler, 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 Volc��n Bar�� (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 18 R���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 4 th toe; 12 / 13 scales under distal phalanx of 4 th toe; 24 lamellae under phalanges ii to iv of 4 th finger; 12 / 11 scales under distal phalanx of 4 th 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 K��hler (2008), K��hler et al. (2008); Fl��schendr��ger 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 (223 D); 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 (221 D) with a reticulum of Sepia (219) bordered by shadings of Brick Red (132 A) 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 (132 A) postorbital stripe bordered by Sepia (119) almost reaching to above shoulder; an elongate blotch of the same color behind ear opening; tail Drab-Gray (119 D) with Walnut Brown (221 B) 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 (162 B) mottling; dorsal surface of head Tawny Olive (223 D) 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 (162 D) towards ventral surface of head; ventral surfaces of body, limbs and base of tail Straw Yellow (56) with sparse Cinnamon (123 A) mottling; tail grading into Olive-Yellow (52) with broad Sepia (219) transverse bands grading into Army Brown (219 B) ventrally; iris Sky Blue (66); dewlap Pink (7) with a suggestion of Salmon Color (6), with Opaline Green (162 D) 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; 18 R���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, Published as part of Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix & K��hler, 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, pp. 1-54 in Zootaxa 3626 (1) on pages 14-20, DOI: 10.11646/zootaxa.3626.1.1, http://zenodo.org/record/283424

Dactyloa casildae (Arosemena, Ib����ez and de Sousa 1992) Figures 2; 10���11; 17 A���C; 18 A���D. Anolis casildae: Arosemena et al. (1992); Auth (1994); Young et al. (1999); Hofer and Bersier (2001); Ib����ez et al. (2001); Nicholson et al. (2001, 2005); Hamad (2009); Fl��schendr��ger and Wijffels (2009); Carrizo (2010); Chun (2010); Jaramillo et al. (2010); Stadler (2010); Casta��eda & de Queiroz (2011). Holotype. MVUP 755, male, from Quebrada Frank, 1100 m, 8 �� 44 ���N, 82 �� 13 ���W, Reserva Forestal de Fortuna, Chiriqu�� province, Panama (Fig. 1: loc. 29; Fig. 20). Diagnosis. A large species (maximum SVL 114 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. frenata, D. ginaelisae, D. ibanezi, D. insignis, D. kunayalae, and D. microtus). Dactyloa casildae can readily be distinguished from these six species by its coloration (described below, shown in Figs. 2, 10, and 11), and from all except D. ibanezi by its very long tail (more than 2.5 times SVL). Moreover, D. casildae 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, usually to nostril or beyond, in D. casildae vs. at most to posterior border of eye), and from D. ginaelisae, D. insignis and D. microtus in having more horizontal loreal rows (6 or more in D. casildae vs. 5 or fewer). Additionally, D. casildae has more subdigital lamellae under the fourth toe (42 or more in D. casildae vs. 41 or fewer) as well as under the fourth finger (29 or more in D. casildae vs. 28 or fewer) than D. ibanezi and D. kunayalae. Description. Total length to 446 mm; SVL to 114 mm in males, to 99 mm in females; tail very long, about 2.5���3 times SVL, compressed (only very slightly in many individuals), without dorsal crest; legs long, tip of fourth toe of adpressed hind limb reaching at least to eye, in most individuals well beyond eye, and in some even beyond snout; internasals, canthals, and loreals keeled; scales of frontal and prefrontal area mostly keeled, some rugose or almost smooth (especially in juveniles); IP distinct, surrounded by smaller, keeled scales; parietal eye distinct; scales of SS enlarged, keeled; scales of supraorbital disk distinctly enlarged, keeled; a very elongate (in several specimens two, rarely three, accordingly shorter scales), keeled anterior superciliary scale, more than half as long (in some specimens almost as long) as horizontal eye diameter, usually followed by up to three similarly keeled but much shorter scales; anterior sublabials slightly enlarged, but never as high as INL, keeled; temporal arch weakly defined, with enlarged scales usually only in its anterior portion; ear opening large, higher than SPL and INL together, at least as high as eye, much larger than IP; nuchal and dorsal crests present in males; 2 rows of keeled middorsal scales slightly enlarged only in few specimens; other dorsal scales as well as lateral scales granular, conical to keeled; ventrals larger than largest dorsals, smooth; 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 halfway 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 gorgetalsternal rows owing to the less widely spaced scales in their interspaces. The completely everted hemipenis of SMF 91454 (Figs. 17 A���C) is a small, bilobate organ; sulcus spermaticus bordered by well-developed sulcal lips, opening at base of apex into two broad concave areas, one on each lobe; an asulcate ridge present; a knob-like processus present on asulcate side of truncus; lobes finely calyculate, truncus with transverse folds. Coloration in life. Dorsal and lateral surfaces green to bluish green, either almost unicolor, or with brown, blue, and yellow spots or blotches, or additionally with oblique transverse bands that may appear solid or composed of blotches; some females of the unicolor and spotted variants with a dark-bordered light middorsal stripe; blue and yellow mottling usually most contrasting around chin, gular area, throat, and shoulders; light and dark lip bars often present; usually a postorbital stripe with dark borders; postorbital stripes of both sides often vaguely connected by a diffuse nuchal band; a less prominent preorbital stripe extending to snout in many specimens; tail with dark mottling suggesting diffuse crossbands; ventral surfaces unicolor yellow or yellowish green, in some individuals with dark mottling; iris brown; male dewlap cream-colored with gorgetal-sternal rows of yellow scales and interspersed blue or green scales, scales in interspaces between gorgetal-sternal rows widely spaced; female dewlap with contrasting yellow and green or blue mottling suggesting longitudinal stripes or a reticulate pattern, with only weakly demarcated gorgetal-sternal rows (Figs. 2; 10���11). Apart from the extremely variable color pattern, Dactyloa casildae is capable of considerable metachrosis (compare Figs. 10 A and I, or 10 E and F). While the description above refers to the colorful or green phase usually shown while the animal is sleeping, the dark or brown phase, typically assumed when the animal is handled, can cause all colors to turn dark brown, making most or all elements of a possible spotted or banded pattern disappear. Specimens spotted at daytime showed either of the color phases. Color photographs of D. casildae have been published by K��hler (2003, 2008), Hamad (2009), Fl��schendr��ger and Wijffels (2009), and Chun (2010). The coloration in life of a subadult male (MHCH 2121, Figs. 9 B, J) was recorded as follows: Dorsal and lateral ground color Apple Green (61); body with a series of broad transverse Greenish Olive (49) bands, interrupted by a Buff-Yellow (52) reticulate mottling covering dorsal and lateral surfaces of body, and mottled with Dark Grayish Brown (20); dorsal surface of head densely mottled with Buff (124) and Sepia (219); lateral and ventral surfaces of head Robin���s Egg Blue (93) mottled with Spectrum Yellow (55); a Sulphur Yellow (57) postorbital stripe bordered by Sepia (219) extending from eye to above ear opening; chin region with a Sulfur Yellow (157) reticulate mottling bordered by Sepia (119); dorsal surfaces of limbs and base of tail with Greenish Olive (49) mottling forming transverse bands; posterior portion of tail Citrine (51) with mostly ill-defined Sepia (219) transverse bands; ventral surface of body Cream Color (54) mottled with Citrine (51) and a suggestion of Lime Green (59); ventral surfaces of limbs and base of tail Citrine (51) mottled with Cream Color (54) and Lime Green (59); iris Clay Color (26); dewlap with a continuum of the chin coloration in anterior portion, grading into Pale Horn Color (92) longitudinal lines of scales suffused with Leaf Green (146); dewlap skin between these lines Lime Green (59) with solitary Leaf Green (146) scales; dewlap margin Cream Color (54). The coloration in life of an adult male (SMF 89455) was recorded as follows (translated from Hamad 2009): Dorsal surfaces with alternating crossbands from anterior head to tip of tail; transverse bands Yellowish Olive- Green (50), alternating with Olive-Yellow (52) bands with Straw Yellow (56) mottling, which grade into Turquoise Blue (65) and Straw Yellow (56) laterally; dorsal surface of head Spectrum Yellow (55), Greenish Olive (49), and Paris Green (63); lateral surfaces of head Spectrum Yellow (55), Turquoise Blue (65), and Sulphur Yellow (57), extending over ventral surfaces of head and body posteriorly to midventer; then grading into Beige (219 D) that continues to ventral surface of base of tail; tail thereafter with Sepia (219) and Raw Umber (23) crossbands; dewlap Cream (84) with longitudinal Orange-Yellow (18) stripes and Cerulean Blue (67) scales. Coloration in preservative. After 21���48 months of preservation in 70 % ethanol, all green and blue elements have turned to brown or, especially around head, to bluish gray; yellow throat markings are dirty white to yellowish brown (Figs. 18 A���D). After 10 years (SMF 85370), also the bluish shades have vanished almost completely. Geographic distribution. As currently understood, Dactyloa casildae is an endemic of the Panamanian Cordillera Central in Chiriqu�� province and the Comarca Ng��be-Bugl��. The species ranges from the eastern extreme of the Serran��a de Talamanca at Cerro Guayabo (RFLF, Chiriqu��) and nearby BPPS (Comarca Ng��be- Bugl��) over 60 airline km east to Cerro Santiago. It occurs on both Caribbean and Pacific slopes, at premontane and lower montane elevations of 990���1720 m asl (Fig. 20). Natural history notes. All collected specimens were encountered at night while they were sleeping on branches, lianas, or leaves 0.3���4 m above ground. Hamad (2009) spotted individuals of this species sleeping as high as 8 m above the R��o Hornito at RFLF. The only two individuals seen at daytime were perched head down on treetrunks 1.5 and 5 m above ground, respectively. During some surveys, Dactyloa casildae appeared very abundant at certain localities both at RFLF and Cerro Santiago in the Comarca Ng��be-Bugl��, with many sleeping individuals encountered very close to each other in some nights. In these surroundings, we also found this species to occur syntopically with D. ginaelisae, and in one case even to share a sleeping branch with that species. At the lowest collection site at R��o Hacha, we found D. casildae to occur in syntopy with D. kunayalae. Our automatized temperature recordings at collection localities of Dactyloa casildae (990���1560 m asl) range between 12.6���23.1 ��C. According to our combined dataset of 41 georeferenced occurrences, the species inhabits PMWF and LMWF, with temperatures between 11.8���27.4 ��C, mean annual temperatures of 17.8���21.2 ��C and a total annual precipitation of 2385���3376 mm. Conservation. Jaramillo et al. (2010) calculated an EVS of 13 for Dactyloa casildae, and assigned the species to the IUCN category NT. We calculated the species��� EVS as 5 (range) + 3 (persecution) + 4 (ecological distribution) = 12. Its extent of occurrence of just 426 km 2 and the continuing deforestation we observed in the region qualify D. casilde for the IUCN category ���Endangered��� (EN). Remarks. Our records expand the known range about 56 km southeastwards and 320 m uphill from previously reported localities (Nicholson et al. 2001; Chun 2010). Hofer and Bersier (2001) and consequently Nicholson et al. (2001) stated the westernmost specimens from BPPS to come from Bocas del Toro province. However, the region in question (Fig. 1: loc. 27) is presently situated within the Comarca Ng��be-Bugl��. As far as we know, Dactyloa casildae has not been reported from Bocas del Toro yet. Based on a sample of eight specimens, Nicholson et al. (2001) restricted the color morphs without transverse bands to females. Among our sample, there are also male specimens which exhibit a non-banded pattern (Figs. 10 G; 11 E), just like the specimen LACM FS 1084 pictured by Chun (2010: Fig. 4). Likewise, the middorsal stripe (Fig. 10 C) is not present in all specimens, not even in all females, that lack transverse bands (Fig. 10 D). One of our adult male specimens (SMF 89673, Figs. 10 A, I, Q) has no enlarged postcloacal scales. The same is true for the young male MHCH 2121 (Figs. 10 B, J; 18 A, B), that was erroneously pictured as a female by K��hler (2008: p. 99: Fig. 197) and could only be confidently sexed by verifying the presence of testes through an incision in the flanks. Considering the lack of postcloacal scales in some males, we assume that those allegedly female specimens having a chiefly white dewlap similar to the male dewlap (Nicholson et al. 2001) are actually males lacking enlarged postcloacal scales, whereas the female dewlap in reality always is contrastingly mottled., Published as part of Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix & K��hler, 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, pp. 1-54 in Zootaxa 3626 (1) on pages 23-27, DOI: 10.11646/zootaxa.3626.1.1, http://zenodo.org/record/283424

Dactyloa kunayalae (Hulebak, Poe, Ib����ez and Williams 2007) Figures 2; 15; 17 M��� O; 18 N���Q. Anolis kunayalae: Hulebak et al. (2007); Poe et al. (2009); Jaramillo et al. (2010); Casta��eda & de Queiroz (2011). Anolis new species 1: Nicholson et al. (2005). Holotype. MSB 72605, male, from along the trails of Parque National General de Divisi��n Omar Torrijos Herrera, 5 km North of El Cope, Cocl�� 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 K��hler (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 (219 A) 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 (119 C), grading into Drab-Gray (119 D) 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 (132 B); 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; 15 B, 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 (132 B); tail Ground Cinnamon (239) with Warm Sepia (221 A) 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 (223 D), mottled with Warm Sepia (221 A) and dirty white, giving the impression of transverse stripes especially laterally and under head; iris Mahogany Red (132 B); 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 Ng��be-Bugl�� as well as from the provinces of Chiriqu��, Cocl��, Col��n, Veraguas, and probably also Dari��n (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 Chiriqu��, Veraguas, and the Comarca Ng��be-Bugl�� (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 Nusagand�� (Hulebak et al. 2007), may be regarded as the first record for Dari��n 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 K��hler (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., Published as part of Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix & K��hler, 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, pp. 1-54 in Zootaxa 3626 (1) on pages 36-38, DOI: 10.11646/zootaxa.3626.1.1, http://zenodo.org/record/283424

Dactyloa frenata (Cope 1899) Figures 2; 12; 17 D���F; 18 E���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); Mart��nez and Rodr��guez (1994); Mart��nez et al. (1995); Ib����ez et al. (1995, 1996, 1997); Young et al. (1999); Ib����ez et al. (2001); Poe (2004); Nicholson et al. (2005); Hamad (2009); Fl��schendr��ger and Wijffels (2009); Poe et al. (2009); Ugueto et al. (2009); Carrizo (2010); Jaramillo et al. (2010); Lotzkat et al. (2010 b); Stadler (2010); Casta��eda & 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), K��hler (2003, 2008), Fl��schendr��ger 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 (162 D) 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, Chiriqu��, Cocl��, Col��n, Dari��n, Panam��, and Veraguas, and from the Comarcas Kuna Yala and Ng��be-Bugl�� (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., Published as part of Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix & K��hler, 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, pp. 1-54 in Zootaxa 3626 (1) on pages 27-29, DOI: 10.11646/zootaxa.3626.1.1, http://zenodo.org/record/283424

Dactyloa microtus (Cope 1871) Figures 2; 7; 16; 18 T���U. Anolis microtus: Cope (1871, 1876); Boulenger (1885); Dunn (1937: in part.); Taylor (1956: in part.); Peters and Donoso- Barros (1970: in part.); Savage (1974); Savage and Talbot (1978: in part.); Arosemena et al. (1992: in part.); Auth (1994: in part.); Poe (2004); Nicholson et al. (2005); Fl��schendr��ger and Wijffels (2009: in part.); Hamad (2009: in part.); Casta��eda & de Queiroz (2011: in part.); Lotzkat et al. (2011). Holotype. USNM 31282, female, from Costa Rica: Provincia de San Jos��: near San Jos��; probably from near La Palma (Savage 1974; Fig. 1: loc. 21; Fig. 22). Diagnosis. A large species (maximum SVL 111 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. ginaelisae, D. ibanezi, D. insignis, and D. kunayalae). Dactyloa microtus can readily be distinguished from these six species by its color pattern described below and shown in Figs. 2, 7, 16, and 18 T���U. It further differs from all mentioned species except D. ginaelisae by its low numbers of horizontal loreal rows (4 or fewer in D. microtus vs. 5 or more) and total loreal scales (26 or fewer in D. microtus vs. 39 or more), and by its low number of scales around midbody (106 or fewer in D. microtus vs. 110 or more). Moreover, D. microtus differs from D. casildae, D. frenata, and D. ibanezi in having short legs (tip of fourth toe of adpressed hind limb reaching at most to tympanum in D. microtus vs. beyond eye; shank length/SVL = 0.183 or less in D. microtus vs. 0.25 or more). Among the short-legged species of Dactyloa in western Panama, D. microtus further differs from D. insignis in having fewer subdigital lamellae under the fourth toe (47 or fewer in D. microtus vs. 52 or more) as well as under the fourth finger (37 or fewer in D. microtus vs. 40), and from D. kunayalae in having more subdigital lamellae under the fourth toe (44 or more in D. microtus vs. 35 or fewer) as well as under the fourth finger (32 or more in D. ginaelisae vs. 25 or fewer). Dactyloa microtus is very similar to D. ginaelisae, from which it differs in having shorter legs (tip of fourth toe of adpressed hind limb reaching to a point between shoulder and tympanum in D. microtus vs. to a point between tympanum and eye in D. ginaelisae; shank length/SVL = 0.183 or less in D. microtus vs. 0.19 or more) and by its less conspicuous and clear-cut coloration pattern between eye and shoulder (light postsupralabial and dark postorbital stripe oriented rather ventrally and losing their conspicuousness around ear in D. microtus vs. a prominent light stripe extending from supralabials posteriorly above or 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). For more characters that might help to distinguish these two very similar species, see remarks section in the species account of D. ginaelisae. Description. Total length to 331 mm; SVL to 111 mm in males, to 104 mm in females; tail long, about 1.4���1.7 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 shoulder and tympanum; 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 either distinct or indistinct, if discernable, then usually surrounded by scales of both smaller and equal size; parietal area generally with flat, rugose scales; parietal eye either distinct or indistinct; scales of SS distinctly enlarged, rugose; scales of supraorbital disk conspicuously enlarged, rugose; usually only one slightly elongate, keeled anterior superciliary scale, one-fourth to half as long as horizontal eye diameter, usually followed by several similarly keeled, but much shorter scales; one or two 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; prominent nuchal and lower dorsal crest present; no rows of middorsal scales slightly enlarged; other dorsal scales as well as lateral scales smooth to slightly rugose or wrinkled in two young males, smooth in a very small juvenile, with very small granules scattered in 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; dewlap of young males large, extending posteriorly to about one-fourth of the distance between axilla and groin, with gorgetal-sternal scale rows of densely arranged scales and widely spaced scales in the broad interspaces between the rows; dewlap of a small juvenile female moderate, extending posteriorly to slightly beyond axilla. The hemipenis of Dactyloa microtus remains undescribed. Coloration in life. Dactyloa microtus exhibits a rather variable coloration. Among the more reliable features usually present are dark crossbands around the tail and, mostly with light centers, on dorsal surfaces of limbs and digits; a light postsupralabial and a dark postorbital stripe extending to near the ear opening; a dark preaxillary blotch. Otherwise, dorsal and lateral surfaces with spots, blotches, and reticula that are are lighter and/or darker than ground color and partly arranged in transverse bars; dorsolateral pattern elements may reach well onto the otherwise comparably unicolor white or yellow ventral surfaces; ground and marking colors of dorsal and lateral surfaces ranging from white over different shades of yellow, green, blue, and brown to black; iris purplish blue to brown, or gray with suffusions of these colors; dewlap of young males yellowish; dewlap of a hatchling female pinkish to salmon color (Figs. 2; 7; 16; see remarks section for adult dewlap coloration reported from Costa Rica). This species is capable of considerable metachrosis (compare Figs. 16 A and B, C and D, F and H). The green phase, usually shown while the animal is sleeping, can culminate in restricting most colors to white, green, and blue, blurring some or all of the otherwise contrasting markings. Similarly, the dark or brown phase, often assumed when the animal is handled, can cause all pattern elements to assume different shades of brown. In between, most pattern elements of a given individual can apparently assume most colors from the palette of this species. A female hatchling was mostly gray when found at daytime. Color photographs of D. microtus have been published by Savage (2002). No standardized description of the coloration in life was recorded. Coloration in preservative. After approximately two years of preservation in 70 % ethanol, colors are largely reduced to white, cream, different shades of brown, and black (Figs. 18 T���U). Geographic distribution. Dactyloa microtus is endemic to the Lower Central American, or Talamancan, highlands of Costa Rica and extreme western Panama. It occurs on both Caribbean and Pacific versants, at premontane and lower montane elevations of 1100 to probably above 2000 m asl (see remarks). In Panama, D. microtus has only been recorded from R��o Changena, Bocas del Toro province, at 1640 m asl (Fig. 22). Natural history notes. Two young males were encountered at night while they were sleeping on a leaf and a branch, 1.5 and 3 m above ground, respectively. A hatchling female virtually dropped into camp on 18 July 2010 while we detached tent strings from a tree. Our automatized temperature recordings at R��o Changena (1640 m asl) range between 13.7���19.2 ��C. According to our combined dataset of 12 georeferenced occurrences (after the exclusion of two localities below 1000 m asl, see remarks), the species inhabits PMMF, PMWF, LMWF, and LMRF, with temperatures between 7.2���26.4 ��C, mean annual temperatures of 12.2���20.4 ��C and a total annual precipitation of 1660���4556 mm. Conservation. Jaramillo et al. (2010) calculated an EVS of 11 for Dactyloa microtus, which at the time also comprised the populations described herein as D. ginaelisae, and assigned that species to the IUCN category LC. We calculated the EVS for D. microtus as 4 (range) + 3 (persecution) + 5 (ecological distribution) = 12. Its extent of occurrence of just 3346 km 2 and the continuing deforestation we observed in the region qualify the species for the IUCN category ���Endangered��� (EN). Remarks. All previous records of Dactyloa microtus from Panama must have been based on individuals of D. ginaelisae, considering the distribution of the two species as documented herein. Thus, our record from R��o Changena is not only the first record of this species from Bocas del Toro province, but also the first and only true record from Panama. Savage (2002) and consequently K��hler (2008) gave the species��� lower distributional limit as 425 m asl. Re-examination of the underlying specimens showed them to be misidentified D. insignis, raising the lower elevational limit to about 1100 m asl in Costa Rica and thus for the species as a whole (J. M. Savage, pers. comm.). The high elevation of Cerro Dantas, provenance of the individual photographed by Daniel Cascante (Figs. 7 K���L), suggests that the species occurs well above the upper elevational limit of 1500 m asl given by Savage (2002) and K��hler (2008). All previous authors have reported the absence of a distinct IP and parietal eye as a diagnostic character for Dactyloa microtus. However, two of our three specimens do have a well-discernable IP with parietal eye (Fig. 16 I; also see remarks concerning this matter for D. ginaelisae and D. insignis). Savage (2002) described the dewlap of adult males as ���pink with white scales��� or ���pale salmon with several vertical rows of white scales,��� and that of females as ���pinkish.��� Inasmuch as they are visible, the dewlaps of the adult Costa Rican specimens (Figs. 7 J and L) comply with these descriptions. We strongly oppose against the common name ���Tiny anole��� assigned to this species by Frank and Ramus (1995). Since Dactyloa microtus is one of the largest mainland anoles, this common name apparently is derived from a mistranslation of the specific epithet microtus, a compound noun in apposition translating to ���small ear.��� If one really needs to use common instead of scientific names for whatever reason, then the correct translation ���Small-ear anole��� would be much more appropriate., Published as part of Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix & K��hler, 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, pp. 1-54 in Zootaxa 3626 (1) on pages 39-41, DOI: 10.11646/zootaxa.3626.1.1, http://zenodo.org/record/283424

Dactyloa ibanezi (Poe, Latella, Ryan and Schaad 2009) Figures 2; 13; 17 J���L; 18 H���J. Anolis chocorum: Talbot (1974), Myers (1971: in part.); Auth (1994: in part.); Mart��nez and Rodr��guez (1994); Mart��nez et al. (1995); Young et al. (1999: in part.); Ib����ez et al. (2001: in part.); Fl��schendr��ger 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); Casta��eda and de Queiroz (2011). Dactyloa chocorum: Savage and Guyer (1989: in part.); Savage (2002: in part.); K��hler (2003, 2008: in part.). Holotype. MSB 72574, male, from the trails of Parque Nacional General de Divisi��n Omar Torrijos Herrera, 5 km North of El Cop��, Cocl�� 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 K��hler (2008), Poe et al. (2009), Fl��schendr��ger 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 (123 B) 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 (123 A); 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; 13 E, 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 (Lim��n 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, Cocl��, Col��n, Veraguas, and the Comarca Ng��be Bugl�� (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 Col��n province and the Comarca Ng��be-Bugl��. 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. K��hler (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 Ng��be-Bugl��, and is ubicated more to the north than the corresponding points mapped by K��hler (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 F��, Veraguas., Published as part of Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix & K��hler, 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, pp. 1-54 in Zootaxa 3626 (1) on pages 31-32, DOI: 10.11646/zootaxa.3626.1.1, http://zenodo.org/record/283424

Anolis benedikti sp. nov. Figures 1 F���H, 5���7 Anolis pachypus: Poe and Iba��ez 2007 (in part); Norops pachypus: Savage 2002 (in part); K��hler 2008 (in part) Holotype. SMF 90149 (Figs. 1 H, 5 A and F, 6), adult male, from the north slope of Cerro Pando, leaving the cattle trail to the right after following it to about 1000 m airline north of the large border monument (Fig. 8 D), 8.9333 ��N, 82.7131 ��W, 2310 m asl, Parque Internacional La Amistad (PILA), Bocas del Toro Province, Panama, close to the border with Costa Rica; collected by Andreas Hertz and Sebastian Lotzkat on 19 November 2009; original field number SL 529. Paratypes. Collected by Sebastian Lotzkat and Andreas Hertz, if not indicated otherwise. All from the PILA in Panama. Bocas del Toro: north slope Cerro Pando: SMF 89744, male, near type locality, 8.9314 ��N, 82.7137 ��W, 2390 m asl, 19 April 2009; SMF 89745 ��� 6, male and female, same locality as holotype, 2330 and 2310 m asl, 20 April 2009; SMF 90148, juvenile male, near type locality, 8.9354 ��N, 82.7128 ��W, 2280 m asl, 19 November 2009; SMF 91508, male, SL 677 ��� 8, subadult male and female, along tributary to R��o Changena, 8.9474 ��N, 82.7098 ��N, 1980���2000 m asl, 13 July 2010; SMF 91507, female, along tributary to R��o Changena, 8.9524 ��N, 82.7093 ��N, 1960 m asl, 14 July 2010; SMF 91505 ��� 6, SL 686, 695, females, R��o Changena, 8.9785 ��N, 82.6901 ��N, 1640���1660 m asl, 14���17 July 2010; SMF 91509, juvenile, between R��o Clarito and R��o Changena, 8.9887 ��N, 82.6749 ��N, 1820 m asl, 17 July 2010; Chiriqu��: south slope Cerro Pando, Jurutungo: SMF 85276 ��� 7, males, 8.9083 ��N, 82.7168 ��W, 2010��� 2100 m asl, collected by Gunther K��hler on 11 January 2006; SMF 85272 ��� 3, subadult males, 8.9109 ��N, 82.7144 ��W, 2060 m asl, collected by Gunther K��hler on 10 January 2006; SL 202, female, 8.9119 ��N, 82.7095 ��W, 2250 m asl, 6 July 2008; SMF 89505, female, 8.9111 ��N, 82.7159 ��W, 2020 m asl, 8 July 2008; SMF 89506, male, 8.9121 ��N, 82.7096 ��W, 2200 m asl, 8 July 2008; SMF 89507, male, 8.9114 ��N, 82.7129 ��W, 2070 m asl, 15 July 2008. Referred specimens. See Appendix. Diagnosis. A medium-sized species (maximum SVL 48.6 mm in males, 48.1 mm in females) of the genus Anolis sensu Poe (2004), that is most similar in external morphology to A. magnaphallus, A. pachypus, A. pseudopachypus, and A. tropidolepis. These four species and A. benedikti share narrow toe pads, long legs (tip of fourth toe of adpressed hind limb reaches to a point anterior to eye), a dark interorbital bar, and usually at least the indication of a lyriform marking on the occipital region, keeled dorsal scales on head and body, weakly keeled ventral scales at midbody, and the lack of enlarged postcloacal scales in males. Anolis benedikti can be distinguished readily from all four aforementioned species by the coloration of the male dewlap which is red with a yellow anterior portion comprising just the anterior margin or, at most, the anterior third of the dewlap (versus solid purplish red in A. magnaphallus and A. tropidolepis, red with a central yellow blotch in A. pachypus, and solid yellow, sometimes grading into orange-yellow on the posterior portion in A. pseudopachypus). In addition, A. benedikti differs from A. magnaphallus, A. pachypus, and A. tropidolepis in having more scale rows between the supraorbital semicircles (usually five or more in A. benedikti, versus four or fewer in A. magnaphallus, A. pachypus, and usually four or fewer, rarely five, in A. tropidolepis), and more scales between supraorbital semicircles and interparietal plate (usually five or more, rarely four, in A. benedikti, versus four or fewer in A. magnaphallus, A. tropidolepis, and usually four or fewer, rarely five, in A. pachypus). Furthermore, A. benedikti differs from A. magnaphallus, A. pachypus, and A. tropidolepis in having a very dark (almost blackish) gray tongue (versus a very light gray tongue in A. magnaphallus, A. pachypus, and A. tropidolepis). At first sight, individuals of A. benedikti might be confused with similarly-colored A. humilis. Yet, regardless of sex or age, individuals of the latter species have at least eight longitudinal rows of greatly enlarged, visibly keeled median dorsal scales that are larger than the ventrals, whereas A. benedikti has a maximum of two slightly enlarged middorsal rows, the scales of which are smaller than the ventrals. Males of A. humilis have a red dewlap with a complete (i.e. bordering the whole dewlap) yellow margin, whereas males of A. benedikti exhibit yellow coloration only on the anterior portion of their otherwise solid red dewlap. Description of the holotype. Adult male as indicated by everted hemipenes and presence of moderate-sized dewlap (Fig. 1 H); snout-vent length 48.6 mm; tail length 81.7 mm, tail complete; tail slightly compressed in cross section, tail height 2.2 mm, tail width 1.8 mm; axilla to groin distance 18.7 mm; head length 11.9 mm, head length/ snout-vent length ratio 0.24; snout length 5.3 mm; head width 8.1 mm; longest toe of adpressed hind limb reaching to snout; shank length 12.6 mm, shank length/head length ratio 1.1; longest finger of extended forelimb reaching about 4.2 mm beyond tip of snout; longest finger of adpressed forelimb reaching to anterior insertion of hind limbs; scales on snout keeled; 5 postrostrals; 5 scales between nasals; scales in distinct prefrontal depression distinctly keeled; supraorbital semicircles differentiated, separated by a minimum of 7 scales; supraorbital disc composed of 14 slightly enlarged keeled scales; 3 elongated superciliaries; about 5 rows of small keeled scales extending between enlarged supraorbitals and superciliaries; interparietal plate slightly enlarged, 0.6 x 0.3 mm (length x width), surrounded by small scales; 10 scales present between interparietal plate and supraorbital semicircles; canthal ridge distinct, composed of 5 large (posterior) and 3 small (anterior) canthal scales; 15 scales present between second canthals; 17 scales present between posterior canthals; 93 / 95 keeled loreal scales in a maximum of 12 horizontal rows; subocular scales granular, subocular row ill-defined; 8 supralabials to level below center of eye; ear opening 0.8 x 1.5 mm (length x height); mental distinctly wider than long, almost completely divided medially, bordered posteriorly by 7 postmentals; 8 infralabials to level below center of eye; sublabials undifferentiated; slightly keeled granular scales present on chin and throat; dewlap moderate-sized, extending well onto chest, anterior insertion at level of anterior border of orbit, posterior insertion at level of axilla, with about 10 somewhat irregular, partially interrupted gorgetal-sternal rows of 20���26 scales per row; dorsum of body with keeled scales, 2 medial rows of slightly enlarged scales, largest dorsal scales subimbricate, about 0.40 x 0.35 mm (length x width); about 66 medial dorsal scales in one head length; about 112 medial dorsal scales between axilla and groin; lateral scales pointed, granular and homogeneous, average size 0.18 mm in diameter; ventrals at midbody weakly keeled, slightly mucronate, and subimbricate, about 0.30 x 0.40 mm (length x width); about 42 ventral scales in one head length; about 58 ventral scales between axilla and groin; about 162 scales around midbody; caudal scales strongly keeled, without whorls of enlarged scales; no enlarged postcloacal scales; tubelike axillary pocket not developed; scales on dorsal surface of forelimb strongly keeled, mucronate and imbricate, about 0.30 x 0.45 mm (length x width); digital pads dilated, dilated pad about 2 times width of non-dilated scales on distal phalanx; distal phalanx narrower than and raised from dilated pad; 23 / 22 lamellae under phalanges II���IV of fourth toe; 9 / 10 scales under distal phalanx of fourth toe; 15 lamellae under phalanges II���IV of fourth finger; 9 scales under distal phalanx of fourth toe. The completely everted hemipenis is a stout bilobate organ; sulcus spermaticus bordered by well-developed sulcal lips and opening at base of apex into two broad concave areas, one on each lobe; a large asulcate ridge-like processus present; lobes strongly calyculate, truncus with transverse folds. Coloration in life. The dewlap (Fig. 1 H) was recorded as follows: Flame Scarlet (15) with a suggestion of Chrome Orange (16) on posterior three fourths, Orange Yellow (18) on anterior fourth and anterior margin; dewlap scales Sepia (119) and dirty white. Otherwise, no detailed notes of the holotype were taken; see Figures 5 A and F for photographs of the holotype in life. Coloration after 18 months of preservation in 70 % ethanol is similar to that in life, apart from that all reddish shades have faded, especially on the dewlap, which now appears dirty white. Variation. The paratypes and referred specimens agree well with the holotype in terms of general morphology and pholidosis (see Table 1). Four of the 32 examined specimens (SMF 89507, 91505; SL 686, 695) have only four scales between the supraorbital semicircles. Two specimens (SMF 89507, 91505) have only four scales between the supraorbital semicircles and the interparietal plate. Among the 17 adult males included in statistical analyses, the variability in the nasal region (Fig. 7) was recorded as follows: Type A one specimen (5.9 %); Type B two specimens (11.8 %); Type C four specimens (23.5 %); Type D five specimens (29.4 %); Type E three specimens (17.6 %); Type F two specimens (11.8 %). Similar to what we observed in the other four members of the Anolis pachypus complex, the coloration of A. benedikti is rather variable among individuals, mostly concerning the extent and shape of color patterns, i.e., stripes, blotches, and bands on dorsal and lateral surfaces (compare Figs. 5 F��� O). Common patterns include a broad middorsal band which is often interspersed with blotches or chevrons, flanked by lateral markings, or disintegrates into diamond-shaped markings posteriorly (Figs. 5 F���J); a narrow light middorsal stripe, often bordered by narrow dark stripes (Figs. K���L); and series of triangular markings pointing laterally from middorsum (with or without narrow medial light stripe) which are either offset to form a zig-zag pattern or arranged symmetrically to form a series of diamonds (Figs. 5 M��� O). Most specimens exhibit contrasting markings on dorsal and lateral surfaces of the head, with postocular stripes and an interorbital bar being the most common. Furthermore, these anoles are capable of metachrosis and, therefore, the coloration of a given individual is subject to physiological change according to time or situation. Most individuals encountered at night exhibited light tonalities, but could turn very dark during the day or when handled (Figs. 5 K���L). Individuals spotted during the day all appeared dark. In the course of metachrosis, the conspicuousness of individual color patterns may vary considerably (as in the postocular stripes between the two females in Figs. 5 C���D). The coloration of the male paratype SMF 89506 (Figs. 1 G, 5 B and G) was recorded as follows: Dorsal and lateral ground color Antique Brown (37) suffused with Light Drab (119 C) and Tawny (38), laterally grading into Olive-Gray (42) mottled with Burnt Umber (22); a Mars Brown (223 A) postorbital stripe mottled with Sepia (219) continues to form a dorsolateral band, which disintegrates into a series of blotches around midbody; a few Sepia (119) middorsal blotches present; dorsal and lateral surfaces of head Light Drab (119 C); occipital region Mars Brown (223 A) mottled with Sepia (219); a Mars Brown (223 A) interorbital bar and a preorbital stripe bordered by Sepia (219); a dirty white with a suggestion of Yellow-Green (58) lateral postorbital stripe extending to shoulder; dorsal and lateral surfaces of limbs and tail Antique Brown (37), with irregular Sepia (219), Tawny (38), and Olive- Gray (42) transverse stripes; ventral surface of body and head Pearl Gray (81) suffused with Cinnamon-Rufous (40); ventral surface of tail with a Cinnamon-Rufous (40) midventral stripe; iris Brick Red (132 A); tongue Plumbeous (78) with a suggestion of Indigo (73); gular region Flame Scarlet (15); dewlap Flame Scarlet (15) with a small Pearl Gray (81) region at posterior insertion and an Orange Yellow (18) anterior margin; dewlap scales dirty white bordered by Sepia (119) or Sepia (219). continued next page The dewlap coloration of the juvenile male SMF 90148 from the vicinity of the type locality was recorded as follows: Flame Scarlet (15), with a suggestion of Chrome Orange (16), grading into Orange Yellow (18) on anterior margin; dewlap scales Sepia (119). As illustrated in Figs. 1 F���H, the extent of the yellow coloration on the anterior portion of the male dewlap of A. benedikti varies from covering just the basal part of the anterior margin (Fig. 1 F) to covering the anterior third of the dewlap (Fig. 1 H). The female dewlap (Figs. 5 C���E) is much smaller than in males, and usually of a whitish color (Fig. 5 C). Nevertheless, some female individuals bear reddish coloration around their gular area. These females tend to exhibit a reddish dewlap (Figs. 5 D���E), sometimes even resembling that of males, like the female paratype SMF 89746 (Fig. 5 E), that was recorded as follows: Flame Scarlet (15) posteriorly, grading into Orange Yellow (18) on anterior half and into dirty white with a suggestion of Chamois (123 D) on posterior base; dewlap scales mostly Sepia (119), some dirty white. Natural history notes. By far, most specimens were encountered at night sleeping on vegetation, i.e., on the upper surface of leaves or on twigs, usually 0.5���2 m above ground. During daytime, specimens were spotted when fleeing on the ground, mostly seeking shelter between tree buttresses or bushes. The vegetation at the type locality (Figs. 8 E���F) is a lush ridgetop cloud forest rich in palms and with abundant growth of epiphytes that cover virtually every surface. Descending either slope of Cerro Pando, the trees become higher and the canopy cover denser. A detailed description of the forest environments around Cerro Pando was given by Myers (1969). Presently, especially below 2000 m elevation on either slope, major clearings for cattle farming disrupt the forest (Figs. 8 A and G). In these pastures, Anolis benedikti can be found associated with bushes, single trees, or remnants of cut trees. Annual total precipitation at the type locality is approximately 2600 mm and mean annual temperature approximately 13.6 ��C, indicating the presence of the Lower Montane Wet Forest life zone according to the Holdridge (1967) classification. Our datalogger recordings (N = 205) at the type locality yielded a temperature range of 11.1���16.7 ��C, with a mean of 12.7 ��C and standard deviation �� 1.1 ��C. Without exception, our collecting localities of A. benedikti lie above 1600 m asl on the Caribbean slope and above 1900 m asl on the Pacific slope and receive more than 2000 mm of annual precipitation, i.e., are assignable to the Lower Montane Wet Forest life zone. In the immediate surroundings of the type locality, A. benedikti was the only reptile species we could find. Reptile species collected on the Caribbean versant along the trail descending the north slope of Cerro Pando (Figs. 8 G���H) between 1640 and 2000 m asl were Anolis capito (Peters), A. humilis (Peters), A. kemptoni Dunn, A. microtus (Cope), Ptychoglossus plicatus (Taylor), and Sibon annulatus (G��nther). Reptile species collected on the Pacific versant between 2000 and 2400 m asl, in the high valley known as Jurutungo (Figs 8 A���B) and along the trail ascending the south slope of Cerro Pando from Jurutungo, were A. kemptoni, A. pachypus, Sceloporus malachiticus Cope, Geophis godmani Boulenger, Rhadinaea calligaster (Cope), and Bothriechis nigroviridis Peters. Geographic distribution. As presently understood, Anolis benedikti inhabits montane elevations along the Talamancan highlands of eastern Costa Rica and extreme western Panama (Fig. 9). All individuals we examined originated from between 82.728 ��W and 82.425 ��W. The species seems to dwell predominantly on the Caribbean versant of the Serran��a de Talamanca, where we found it from the continental divide at up to 2390 m asl (Cerro Pando). Descending the Caribbean slope, we encountered it down to 1640 m asl at the R��o Changena, 1740 m asl along the Culebra trail, and 1650 m asl along the Pianista trail. However, in at least three regions, this species is also found on the Pacific versant, where it occurs next to its close relatives: At Las Tablas, in the Costa Rican Province of Puntarenas next to the Panamanian border, SMF 92134 was found at 1960 m asl, less than a kilometer from the nearest collected A. pachypus at 1880 m asl. Just a few kilometers east, in Jurutungo (the headwaters of R��o Candela in Chiriqu�� Province, Panama; Fig. 8 A), we found A. benedikti as low as 2020 m asl at certain places, less than 50 m away from the nearest collection site of A. pachypus. Further east, on the slopes of Volc��n Bar�� north of Boquete, about one kilometer west of the ranger station of Alto Chiquero close to the upper R��o Caldera, we collected four males of A. benedikti at 1920 m asl, as well as a single female farther south. These two sites are close to different collection sites (at both lower and higher elevations) as well as to the type locality of A. magnaphallus. Etymology. The specific name is a patronym for Benedikt St��kl, Germany, in recognition of the financial support of taxonomic research provided by Karsten Lutz through the BIOPAT initiative., Published as part of Lotzkat, Sebastian, Bienentreu, Joe-Felix, Hertz, Andreas & K��hler, Gunther, 2011, A new species of Anolis (Squamata: Iguania: Dactyloidae) formerly referred to as A. pachypus from the Cordillera de Talamanca of western Panama and adjacent Costa Rica, pp. 1-21 in Zootaxa 3125 on pages 8-18, DOI: 10.5281/zenodo.200703, {"references":["Savage, J. M. (2002) The amphibians and reptiles of Costa Rica. A herpetofauna between two continents, between two seas. University of Chicago Press, Chicago, xx + 934 pp.","Poe, S. (2004) Phylogeny of anoles. Herpetological Monographs, 18, 37 - 89.","Myers, C. W. (1969) The ecological geography of cloud forest in Panama. American Museum Novitates, 2396, 1 - 52.","Holdridge, L. R. (1967) Life zone ecology. Tropical Science Center, San Jose, Costa Rica, 206 pp."]}

Anolis benedikti sp. nov. Figures 1 F–H, 5–7 Anolis pachypus: Poe and Ibañez 2007 (in part); Norops pachypus: Savage 2002 (in part); Köhler 2008 (in part) Holotype. SMF 90149 (Figs. 1 H, 5 A and F, 6), adult male, from the north slope of Cerro Pando, leaving the cattle trail to the right after following it to about 1000 m airline north of the large border monument (Fig. 8 D), 8.9333 °N, 82.7131 °W, 2310 m asl, Parque Internacional La Amistad (PILA), Bocas del Toro Province, Panama, close to the border with Costa Rica; collected by Andreas Hertz and Sebastian Lotzkat on 19 November 2009; original field number SL 529. Paratypes. Collected by Sebastian Lotzkat and Andreas Hertz, if not indicated otherwise. All from the PILA in Panama. Bocas del Toro: north slope Cerro Pando: SMF 89744, male, near type locality, 8.9314 °N, 82.7137 °W, 2390 m asl, 19 April 2009; SMF 89745 – 6, male and female, same locality as holotype, 2330 and 2310 m asl, 20 April 2009; SMF 90148, juvenile male, near type locality, 8.9354 °N, 82.7128 °W, 2280 m asl, 19 November 2009; SMF 91508, male, SL 677 – 8, subadult male and female, along tributary to Río Changena, 8.9474 °N, 82.7098 °N, 1980–2000 m asl, 13 July 2010; SMF 91507, female, along tributary to Río Changena, 8.9524 °N, 82.7093 °N, 1960 m asl, 14 July 2010; SMF 91505 – 6, SL 686, 695, females, Río Changena, 8.9785 °N, 82.6901 °N, 1640–1660 m asl, 14–17 July 2010; SMF 91509, juvenile, between Río Clarito and Río Changena, 8.9887 °N, 82.6749 °N, 1820 m asl, 17 July 2010; Chiriquí: south slope Cerro Pando, Jurutungo: SMF 85276 – 7, males, 8.9083 °N, 82.7168 °W, 2010– 2100 m asl, collected by Gunther Köhler on 11 January 2006; SMF 85272 – 3, subadult males, 8.9109 °N, 82.7144 °W, 2060 m asl, collected by Gunther Köhler on 10 January 2006; SL 202, female, 8.9119 °N, 82.7095 °W, 2250 m asl, 6 July 2008; SMF 89505, female, 8.9111 °N, 82.7159 °W, 2020 m asl, 8 July 2008; SMF 89506, male, 8.9121 °N, 82.7096 °W, 2200 m asl, 8 July 2008; SMF 89507, male, 8.9114 °N, 82.7129 °W, 2070 m asl, 15 July 2008. Referred specimens. See Appendix. Diagnosis. A medium-sized species (maximum SVL 48.6 mm in males, 48.1 mm in females) of the genus Anolis sensu Poe (2004), that is most similar in external morphology to A. magnaphallus, A. pachypus, A. pseudopachypus, and A. tropidolepis. These four species and A. benedikti share narrow toe pads, long legs (tip of fourth toe of adpressed hind limb reaches to a point anterior to eye), a dark interorbital bar, and usually at least the indication of a lyriform marking on the occipital region, keeled dorsal scales on head and body, weakly keeled ventral scales at midbody, and the lack of enlarged postcloacal scales in males. Anolis benedikti can be distinguished readily from all four aforementioned species by the coloration of the male dewlap which is red with a yellow anterior portion comprising just the anterior margin or, at most, the anterior third of the dewlap (versus solid purplish red in A. magnaphallus and A. tropidolepis, red with a central yellow blotch in A. pachypus, and solid yellow, sometimes grading into orange-yellow on the posterior portion in A. pseudopachypus). In addition, A. benedikti differs from A. magnaphallus, A. pachypus, and A. tropidolepis in having more scale rows between the supraorbital semicircles (usually five or more in A. benedikti, versus four or fewer in A. magnaphallus, A. pachypus, and usually four or fewer, rarely five, in A. tropidolepis), and more scales between supraorbital semicircles and interparietal plate (usually five or more, rarely four, in A. benedikti, versus four or fewer in A. magnaphallus, A. tropidolepis, and usually four or fewer, rarely five, in A. pachypus). Furthermore, A. benedikti differs from A. magnaphallus, A. pachypus, and A. tropidolepis in having a very dark (almost blackish) gray tongue (versus a very light gray tongue in A. magnaphallus, A. pachypus, and A. tropidolepis). At first sight, individuals of A. benedikti might be confused with similarly-colored A. humilis. Yet, regardless of sex or age, individuals of the latter species have at least eight longitudinal rows of greatly enlarged, visibly keeled median dorsal scales that are larger than the ventrals, whereas A. benedikti has a maximum of two slightly enlarged middorsal rows, the scales of which are smaller than the ventrals. Males of A. humilis have a red dewlap with a complete (i.e. bordering the whole dewlap) yellow margin, whereas males of A. benedikti exhibit yellow coloration only on the anterior portion of their otherwise solid red dewlap. Description of the holotype. Adult male as indicated by everted hemipenes and presence of moderate-sized dewlap (Fig. 1 H); snout-vent length 48.6 mm; tail length 81.7 mm, tail complete; tail slightly compressed in cross section, tail height 2.2 mm, tail width 1.8 mm; axilla to groin distance 18.7 mm; head length 11.9 mm, head length/ snout-vent length ratio 0.24; snout length 5.3 mm; head width 8.1 mm; longest toe of adpressed hind limb reaching to snout; shank length 12.6 mm, shank length/head length ratio 1.1; longest finger of extended forelimb reaching about 4.2 mm beyond tip of snout; longest finger of adpressed forelimb reaching to anterior insertion of hind limbs; scales on snout keeled; 5 postrostrals; 5 scales between nasals; scales in distinct prefrontal depression distinctly keeled; supraorbital semicircles differentiated, separated by a minimum of 7 scales; supraorbital disc composed of 14 slightly enlarged keeled scales; 3 elongated superciliaries; about 5 rows of small keeled scales extending between enlarged supraorbitals and superciliaries; interparietal plate slightly enlarged, 0.6 x 0.3 mm (length x width), surrounded by small scales; 10 scales present between interparietal plate and supraorbital semicircles; canthal ridge distinct, composed of 5 large (posterior) and 3 small (anterior) canthal scales; 15 scales present between second canthals; 17 scales present between posterior canthals; 93 / 95 keeled loreal scales in a maximum of 12 horizontal rows; subocular scales granular, subocular row ill-defined; 8 supralabials to level below center of eye; ear opening 0.8 x 1.5 mm (length x height); mental distinctly wider than long, almost completely divided medially, bordered posteriorly by 7 postmentals; 8 infralabials to level below center of eye; sublabials undifferentiated; slightly keeled granular scales present on chin and throat; dewlap moderate-sized, extending well onto chest, anterior insertion at level of anterior border of orbit, posterior insertion at level of axilla, with about 10 somewhat irregular, partially interrupted gorgetal-sternal rows of 20–26 scales per row; dorsum of body with keeled scales, 2 medial rows of slightly enlarged scales, largest dorsal scales subimbricate, about 0.40 x 0.35 mm (length x width); about 66 medial dorsal scales in one head length; about 112 medial dorsal scales between axilla and groin; lateral scales pointed, granular and homogeneous, average size 0.18 mm in diameter; ventrals at midbody weakly keeled, slightly mucronate, and subimbricate, about 0.30 x 0.40 mm (length x width); about 42 ventral scales in one head length; about 58 ventral scales between axilla and groin; about 162 scales around midbody; caudal scales strongly keeled, without whorls of enlarged scales; no enlarged postcloacal scales; tubelike axillary pocket not developed; scales on dorsal surface of forelimb strongly keeled, mucronate and imbricate, about 0.30 x 0.45 mm (length x width); digital pads dilated, dilated pad about 2 times width of non-dilated scales on distal phalanx; distal phalanx narrower than and raised from dilated pad; 23 / 22 lamellae under phalanges II–IV of fourth toe; 9 / 10 scales under distal phalanx of fourth toe; 15 lamellae under phalanges II–IV of fourth finger; 9 scales under distal phalanx of fourth toe. The completely everted hemipenis is a stout bilobate organ; sulcus spermaticus bordered by well-developed sulcal lips and opening at base of apex into two broad concave areas, one on each lobe; a large asulcate ridge-like processus present; lobes strongly calyculate, truncus with transverse folds. Coloration in life. The dewlap (Fig. 1 H) was recorded as follows: Flame Scarlet (15) with a suggestion of Chrome Orange (16) on posterior three fourths, Orange Yellow (18) on anterior fourth and anterior margin; dewlap scales Sepia (119) and dirty white. Otherwise, no detailed notes of the holotype were taken; see Figures 5 A and F for photographs of the holotype in life. Coloration after 18 months of preservation in 70 % ethanol is similar to that in life, apart from that all reddish shades have faded, especially on the dewlap, which now appears dirty white. Variation. The paratypes and referred specimens agree well with the holotype in terms of general morphology and pholidosis (see Table 1). Four of the 32 examined specimens (SMF 89507, 91505; SL 686, 695) have only four scales between the supraorbital semicircles. Two specimens (SMF 89507, 91505) have only four scales between the supraorbital semicircles and the interparietal plate. Among the 17 adult males included in statistical analyses, the variability in the nasal region (Fig. 7) was recorded as follows: Type A one specimen (5.9 %); Type B two specimens (11.8 %); Type C four specimens (23.5 %); Type D five specimens (29.4 %); Type E three specimens (17.6 %); Type F two specimens (11.8 %). Similar to what we observed in the other four members of the Anolis pachypus complex, the coloration of A. benedikti is rather variable among individuals, mostly concerning the extent and shape of color patterns, i.e., stripes, blotches, and bands on dorsal and lateral surfaces (compare Figs. 5 F– O). Common patterns include a broad middorsal band which is often interspersed with blotches or chevrons, flanked by lateral markings, or disintegrates into diamond-shaped markings posteriorly (Figs. 5 F–J); a narrow light middorsal stripe, often bordered by narrow dark stripes (Figs. K–L); and series of triangular markings pointing laterally from middorsum (with or without narrow medial light stripe) which are either offset to form a zig-zag pattern or arranged symmetrically to form a series of diamonds (Figs. 5 M– O). Most specimens exhibit contrasting markings on dorsal and lateral surfaces of the head, with postocular stripes and an interorbital bar being the most common. Furthermore, these anoles are capable of metachrosis and, therefore, the coloration of a given individual is subject to physiological change according to time or situation. Most individuals encountered at night exhibited light tonalities, but could turn very dark during the day or when handled (Figs. 5 K–L). Individuals spotted during the day all appeared dark. In the course of metachrosis, the conspicuousness of individual color patterns may vary considerably (as in the postocular stripes between the two females in Figs. 5 C–D). The coloration of the male paratype SMF 89506 (Figs. 1 G, 5 B and G) was recorded as follows: Dorsal and lateral ground color Antique Brown (37) suffused with Light Drab (119 C) and Tawny (38), laterally grading into Olive-Gray (42) mottled with Burnt Umber (22); a Mars Brown (223 A) postorbital stripe mottled with Sepia (219) continues to form a dorsolateral band, which disintegrates into a series of blotches around midbody; a few Sepia (119) middorsal blotches present; dorsal and lateral surfaces of head Light Drab (119 C); occipital region Mars Brown (223 A) mottled with Sepia (219); a Mars Brown (223 A) interorbital bar and a preorbital stripe bordered by Sepia (219); a dirty white with a suggestion of Yellow-Green (58) lateral postorbital stripe extending to shoulder; dorsal and lateral surfaces of limbs and tail Antique Brown (37), with irregular Sepia (219), Tawny (38), and Olive- Gray (42) transverse stripes; ventral surface of body and head Pearl Gray (81) suffused with Cinnamon-Rufous (40); ventral surface of tail with a Cinnamon-Rufous (40) midventral stripe; iris Brick Red (132 A); tongue Plumbeous (78) with a suggestion of Indigo (73); gular region Flame Scarlet (15); dewlap Flame Scarlet (15) with a small Pearl Gray (81) region at posterior insertion and an Orange Yellow (18) anterior margin; dewlap scales dirty white bordered by Sepia (119) or Sepia (219). continued next page The dewlap coloration of the juvenile male SMF 90148 from the vicinity of the type locality was recorded as follows: Flame Scarlet (15), with a suggestion of Chrome Orange (16), grading into Orange Yellow (18) on anterior margin; dewlap scales Sepia (119). As illustrated in Figs. 1 F–H, the extent of the yellow coloration on the anterior portion of the male dewlap of A. benedikti varies from covering just the basal part of the anterior margin (Fig. 1 F) to covering the anterior third of the dewlap (Fig. 1 H). The female dewlap (Figs. 5 C–E) is much smaller than in males, and usually of a whitish color (Fig. 5 C). Nevertheless, some female individuals bear reddish coloration around their gular area. These females tend to exhibit a reddish dewlap (Figs. 5 D–E), sometimes even resembling that of males, like the female paratype SMF 89746 (Fig. 5 E), that was recorded as follows: Flame Scarlet (15) posteriorly, grading into Orange Yellow (18) on anterior half and into dirty white with a suggestion of Chamois (123 D) on posterior base; dewlap scales mostly Sepia (119), some dirty white. Natural history notes. By far, most specimens were encountered at night sleeping on vegetation, i.e., on the upper surface of leaves or on twigs, usually 0.5–2 m above ground. During daytime, specimens were spotted when fleeing on the ground, mostly seeking shelter between tree buttresses or bushes. The vegetation at the type locality (Figs. 8 E–F) is a lush ridgetop cloud forest rich in palms and with abundant growth of epiphytes that cover virtually every surface. Descending either slope of Cerro Pando, the trees become higher and the canopy cover denser. A detailed description of the forest environments around Cerro Pando was given by Myers (1969). Presently, especially below 2000 m elevation on either slope, major clearings for cattle farming disrupt the forest (Figs. 8 A and G). In these pastures, Anolis benedikti can be found associated with bushes, single trees, or remnants of cut trees. Annual total precipitation at the type locality is approximately 2600 mm and mean annual temperature approximately 13.6 °C, indicating the presence of the Lower Montane Wet Forest life zone according to the Holdridge (1967) classification. Our datalogger recordings (N = 205) at the type locality yielded a temperature range of 11.1–16.7 °C, with a mean of 12.7 °C and standard deviation ± 1.1 °C. Without exception, our collecting localities of A. benedikti lie above 1600 m asl on the Caribbean slope and above 1900 m asl on the Pacific slope and receive more than 2000 mm of annual precipitation, i.e., are assignable to the Lower Montane Wet Forest life zone. In the immediate surroundings of the type locality, A. benedikti was the only reptile species we could find. Reptile species collected on the Caribbean versant along the trail descending the north slope of Cerro Pando (Figs. 8 G–H) between 1640 and 2000 m asl were Anolis capito (Peters), A. humilis (Peters), A. kemptoni Dunn, A. microtus (Cope), Ptychoglossus plicatus (Taylor), and Sibon annulatus (Günther). Reptile species collected on the Pacific versant between 2000 and 2400 m asl, in the high valley known as Jurutungo (Figs 8 A–B) and along the trail ascending the south slope of Cerro Pando from Jurutungo, were A. kemptoni, A. pachypus, Sceloporus malachiticus Cope, Geophis godmani Boulenger, Rhadinaea calligaster (Cope), and Bothriechis nigroviridis Peters. Geographic distribution. As presently understood, Anolis benedikti inhabits montane elevations along the Talamancan highlands of eastern Costa Rica and extreme western Panama (Fig. 9). All individuals we examined originated from between 82.728 °W and 82.425 °W. The species seems to dwell predominantly on the Caribbean versant of the Serranía de Talamanca, where we found it from the continental divide at up to 2390 m asl (Cerro Pando). Descending the Caribbean slope, we encountered it down to 1640 m asl at the Río Changena, 1740 m asl along the Culebra trail, and 1650 m asl along the Pianista trail. However, in at least three regions, this species is also found on the Pacific versant, where it occurs next to its close relatives: At Las Tablas, in the Costa Rican Province of Puntarenas next to the Panamanian border, SMF 92134 was found at 1960 m asl, less than a kilometer from the nearest collected A. pachypus at 1880 m asl. Just a few kilometers east, in Jurutungo (the headwaters of Río Candela in Chiriquí Province, Panama; Fig. 8 A), we found A. benedikti as low as 2020 m asl at certain places, less than 50 m away from the nearest collection site of A. pachypus. Further east, on the slopes of Volcán Barú north of Boquete, about one kilometer west of the ranger station of Alto Chiquero close to the upper Río Caldera, we collected four males of A. benedikti at 1920 m asl, as well as a single female farther south. These two sites are close to different collection sites (at both lower and higher elevations) as well as to the type locality of A. magnaphallus. Etymology. The specific name is a patronym for Benedikt Stökl, Germany, in recognition of the financial support of taxonomic research provided by Karsten Lutz through the BIOPAT initiative.

Lotzkat, Sebastian, Bienentreu, Joe-Felix, Hertz, Andreas, Köhler, Gunther (2011): A new species of Anolis (Squamata: Iguania: Dactyloidae) formerly referred to as A. pachypus from the Cordillera de Talamanca of western Panama and adjacent Costa Rica. Zootaxa 3125: 1-21, DOI: 10.5281/zenodo.200703