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Panama is a hotspot of amphibian and reptile diversity and many areas of the country still remain underexplored. Here we present results of our field work carried out through several years in western Panama, further increasing the numbers of amphibian and reptile species known from the country. We provide the first country records for Smilisca manisorum and Lepidophyma reticulatum. Additionally, we extend the known distributional ranges of Pristimantis taeniatus, Pleurodema brachyops, Leptodactylus fuscus, Bachia blairi, Basiliscus plumifrons, Anolis auratus, A. capito, A. cryptolimifrons, A. humilis, A. kemptoni, A. pseudopachypus, Geophis godmani, Mastigodryas pleei, and Bothriechis supraciliaris.

DIASPORUS SAPO SP. NOV. FIGS 13, 14G, H ELEUTHERODACTYLUS SP. – MYERS 1969: FIG. 19C. Holotype SMF 97329 (original field number AB 429), an adult female (Fig. 13) collected by Abel Batista & Gustavo Dojirama at the top of Cerro Sapo, PND, Distrito de Garachine, Darien, Panama, on 4 December 2011, at 20:00 h (7.97618 N, 78.36263 W; 1169 m a.s.l.). Paratypes MHCH 2853 – 58, SMF 97328, SMF 97330 – 32; same collecting data as for holotype. Diagnosis Diasporus sapo sp. nov. is characterized by the following combination of characters (see Tables 4 – 6): (1) dorsal skin texture slightly tuberculate, venter smooth; (2) tympanum indistinguishable, annulus tympanicus and tympanic membrane absent; (3) snout rounded in dorsal view and in profile; (4) conical supraocular tubercle and cranial crests absent; (5) dentigerous processes of vomers with between seven and 11 teeth each, straight in outline, from the centre of the orbit to the centre of the roof of mouth; (6) vocal sac and vocal slits not differentiated, only a slightly differentiated fold beside the tongue, no nuptial pads; (7) finger II longer than finger I, ungual flap expanded, spadate, more evident on fingers II – IV; (8) no fringes or webbing on fingers; (9) palmar tubercle ovoid or rounded, flattened and almost the same size as thenar tubercle; thenar tubercle elongate; subarticular tubercles rounded and globular; two or three supernumerary tubercles; (10) heel smooth; (11) no fringes or webbing on toes, ungual flap on toes expanded, spadate, more evident on toes IV and V; (12) plantar tubercle indistinguishable, subarticular tubercles rounded and globular (one on toes I and II, two on toes III and V, and three on toe IV); foot without supernumerary tubercles; inner metatarsal tubercle elongated, outer metatarsal tubercles rounded and globular, smaller than inner; tarsal ridge absent; (13) dorsal ground color in life reddish and patternless, venter translucent, vocal sac not visible (Fig. 14H); (14) SVL 22.6 ƚ 2.86 (18.8—29.1, N = 11), males 22.6 ƚ 2.59 (19.9—29.1, N = 9), females 22.6 ƚ 5.3 (18.8—26.3, N = 2); (15) advertisement call unknown. Description of the holotype An adult female (SVL 28.91), with slender body; dorsal skin texture slightly tuberculate, venter skin smooth, discoidal fold not evident; protuberant eyes 1.78 times longer than snout; tympanum small, ratio TD/EL 23%; tympanum indistinguishable, annulus tympanicus and tympanic membrane absent, positioned 2.6 mm behind the orbit; head as wide as long (HL/ HW 0.95), greatest head width between angles of jaw 35% of SVL; snout rounded from above and in profile; nares situated near tip of snout and slightly dorsolaterally directed, visible in frontal view, also visible dorsally but not ventrally; canthus rostralis rounded; loreal region concave; dentigerous processes of vomers with ten (right) and eight (left) teeth each side, straight in outline, from the centre of the orbit to the centre of the roof of mouth, and separated by a space of half of its total length; vocal slits absent; tongue long (26% of SVL) and broadening to the end, first third attached to floor of mouth; hands moderate in size, 22% of SVL; relative lengths of adpressed fingers I Coloration of holotype in life Coloration recorded as follows (Fig. 13): iris medium neutral gray (298) with reticulations sepia (286), iris periphery jet black (300), eye periphery sky blue (192); dorsal ground color uniform Pratt’s ruby (68), becoming darker to the front as dark carmine (61); venter and limbs chrome orange (74), throat pale buff (1). Coloration in preservative Dorsal ground color cinnamon – drab (50), becoming darker to warm sepia (40) to the tip of snout; limbs and venter cream color (12), throat buff (5), hand and foot drab (19). Measurements of holotype (mm) SVL 28.91; HL 9.63; HW 10.19; IOD 2.93; EL 4.29; TD 0.98; FL 11.58; TL 12.33; HAL 6.37; 3FW 0.52; 3FD 1.8; 3TW 0.61; 3TD 1.12; 4TW 0.57; 4TD 1.20; BW 9.99 (for variation in the species, see Tables 4 – 6). Natural history This species is known only from the top of Cerro Sapo, which is covered by elfin forest. The vegetation predominantly consists of small trees (roughly 10 m in height) fully covered with moss and bromeliads. Diasporus sapo sp. nov. was most often found at 1 – 2 m above ground during the night; individuals were seen walking over tree branches and tree bark. Etymology The species name is derived from the name of Cerro Sapo, where the species was found.

Batista, Abel, Kohler, Gunther, Mebert, Konrad, Hertz, Andreas, Vesely, Milan (2016): An integrative approach to reveal speciation and species richness in the genus Diasporus (Amphibia: Anura: Eleutherodactylidae) in eastern Panama. Zoological Journal of the Linnean Society 178 (2): 267-311, DOI: 10.1111/zoj.12411, URL: http://dx.doi.org/10.1111/zoj.12411

Pseudogonatodes lunulatus (Roux, 1927) fue descrito de un único ejemplar recolectado en la población El Mene, municipio Acosta, al noreste del estado Falcón, Venezuela. Aunque la especie ha sido señalada para otras localidades de Venezuela y Colombia, un estudio extenso acerca de su taxonomía y distribución geográfica en el país no había sido efectuado. En este sentido, se pudo examinar y comparar una muestra mayor de 20 especímenes depositados en varios museos nacionales y extranjeros. Adicionalmente, se extiende sustancialmente la distribución conocida de la especie al norte del río Orinoco. Pseudogonatodes lunulatus (Roux, 1927) was described from a single specimen collected in the village El Mene, Acosta municipality, northeastern Falcón state, Venezuela. Although the species has been reported for other Venezuelan and Colombian localities, an extensive study on taxonomy and geographic distribution in the country had not been performed. In this sense, we examined and compared a larger sample of 20 specimens deposited in several national and foreign museums. Additionally, we substantially expand the known distribution of the species to the north of the Orinoco River.

Lepidoblepharis victormartinezi sp. nov. Figs. 3 ���6, 12 Lepidoblepharis sp.: Mart��nez & Rodriguez (1994: possibly); Mart��nez et al. (1995: possibly). Lepidoblepharis xanthostigma: all in part. (referring to certain populations in Col��n and Veraguas): Auth (1994); Young et al. (1999); Ib����ez et al. (2001); K��hler (2008); Carrizo (2010: referring to SMF 89963); Jaramillo et al. (2010); Lotzkat et al. (2010: referring to SMF 89963). Holotype. Adult female SMF 50951, original field number AB 1241 (Fig. 12) collected in leaf-litter 500 m east of the village Chicheme (8.865278 ��N, 80.643829 ��W, 100 m elev.), San Jos�� del General, Donoso district, Col��n province, Panama, collected on 25 January 2013 at 11: 40 hrs by Abel Batista, Lester V��squez, and Leysi D��az. Paratypes. Four adult males, all from Panama: SMF 89963 from Cerro Negro, Veraguas, on 28 July 2008; SMF 50950 (collected on 20 July 2011), 50952, and MHCH 2954 from Petaquilla, Cocl�� del Norte, Donoso, Col��n; see Appendix I for locality details. Diagnosis. Lepidoblepharis victormartinezi (our sp. nov. 3) is characterized by the following combination of characters: (1) dorsal scales small, granular, and juxtaposed, ventral scales large, cycloid, flat, and imbricate; (2) scales on head small and granular; (3) 3���5 (4.4 �� 0.89) postrostrals; (4) two paramedian clefts demarcate the slightly convex median portion of the posterior mental border, rendering the latter vaguely M-shaped in its totality; (5) 5���7 (6.0 �� 0.71) postmentals; (6) lamellae under fourth toe 6���8 (6.6 �� 0.89), lamellae under fourth finger 4���5 (4.8 �� 0.45), the subdigital lamellae under each digit showing a peculiar morphology that is unique within the genus, with 1���3 proximal one(s) longitudinally greatly enlarged, each about 3���4 times longer than any of the remaining lamellae; (7) median subcaudals only slightly larger than the neighboring scales, about as long as wide, with rounded posterior margins, arranged in a regular tail sequence of 1 ' 1 '' (Fig. 3 D); (8) ventral escutcheon consisting of 61���68 (63.3 �� 3.30) scales, 6���8 (6.8 �� 0.96) scales long and 12���15 (13.8 �� 1.26) wide; (9) lack of a discernible subfemoral escutcheon; (10) ventral scales at midbody 15���19 (17.4 �� 1.52); (11) dorsal scales at midbody 53���67 (59.8 �� 5.36); (12) bilobate hemipenis, with a third lobule rising from the pedicel; (13) SVL 25���27 (25.6 �� 0.89) mm. Comparison with other species of the genus. Lepidoblepharis victormartinezi can be differentiated from many species in the genus by its small size and its low number of lamellae under the fourth toe and finger (Figs. 3 D; 4). Moreover, the conspicuous morphology of its subdigital lamellae is unique within the genus and immediately distinguishes it from any described congener. In the following, we provide further comparisons to all other species within the genus, with the characteristics for L. victormartinezi in parentheses. Lepidoblepharis victormartinezi can be distinguished from the Panamanian species L. xanthostigma, L. sanctaemartae, L. rufigularis, and L. emberawoundule by a genetic p-distance of 10���21 % in 16 S mtDNA between individuals. Lepidoblepharis xanthostigma has greatly enlarged median subcaudal scales (slightly enlarged) and 12���16 lamellae under its fourth toe (6���8). Lepidoblepharis sanctaemartae has large, flat, imbricate dorsal body scales (small, granular, juxtaposed). Lepidoblepharis emberawoundule has 5���8 lamellae under its fourth finger (4���5), a ventral escutcheon consisting of 38���61 scales (57���68), and a well-discernible subfemoral escutcheon consisting of 4���5 scales in a single row under each thigh (no discernible subfemoral escutcheon). Lepidoblepharis rufigularis has 13 lamellae under its fourth toe (6���8) and 11 under its fourth finger (4���5). To date, seven species of the genus Lepidoblepharis have been reported to possess ten or fewer lamellae under the fourth toe, i.e., to be short-toed. Two of these, L. miyatai and L. sanctaemartae, possess large, flat, and imbricate dorsal scales (dorsal scales small, granular, and juxtaposed). Lepidoblepharis buchwaldi, L. microlepis, L. montecanoensis, L. peraccae, and L. williamsi can confidently be ruled out as conspecifics since the inspection of the illustrations and photographs available for the holotypes showed no subdigital lamellae to be as conspicuously enlarged as in L. victormartinezi. Moreover, in the holotype of L. peraccae the plantar and palmar scales have the posterior border ovoid and imbricated (plantar and palmar scales small rounded and juxtaposed), and in the holotype of L. microlepis all of the slightly enlarged median subcaudal scales are bordered laterodistally by one scale, forming a regular tail sequence of 1 ' 1 ' (the slightly more enlarged median subcaudals bordered laterodistally by two scales, the slightly smaller ones by one, forming a regular tail sequence of 1 ' 1 ''). The remaining species of the genus, i.e., L colombianus, L conolepis, L. duolepis, L. festae, L. grandis, L. heyerorum, L. hogmoedi, L. intermedius, and L. ruthveni are longtoed with eleven or more lamellae under the fourth toe (6���8 in L. victormartinezi). Description of holotype. Variation among the entire type series is given in parentheses (see Table 1 for details). Adult female as indicated by absence of hemipenes and escutcheon scales; SVL 25 mm (25���27 mm), TL 27.2 mm (27���31), HL 5.4 mm (4.9���6.1 mm), HW 3.7 mm (3.7���4.2 mm), forelimbs 5.7 mm (6.8 ��� 5.7), hind limbs 8.3 mm (9.8 ��� 7.4), shank 3.5 mm (3.5���4.3); rostral large, clearly visible from above, with a shallow, horseshoeshaped posterior depression and a long median cleft; postrostrals four (3���5) including supranasals, median ones smaller than supranasals, and indenting the rostral, at least one median postrostral slightly larger than posteriorly adjacent scales on snout; postnasals two (1���2), both about same size as posteriorly adjacent loreal scales; scales on snout small and smooth, 17 (16���22) scales across snout between anterior sutures of second SPLs; loreal scales subimbricate, elevated towards posterior and dorsal directions, 6 (5���9) loreal scales on a longitudinal line between postnasals and orbit; scales on top of head small, granular, and juxtaposed, generally pointing upward, about half as large as those on the central snout; superciliary flap with two enlarged scales on anterior border, of which the first is slightly longer than the second; followed by 2���5 small and globular scales; supralabials three (3���4), posteriormost one below center of eye; ear-opening small, oval, in oblique orientation; mental large, posterior margin vaguely M-shaped and slightly convex in the middle, with two short paramedian clefts; postmentals five (5���7), flat and slightly larger than the posteriorly adjacent chin scales, the two median ones slightly larger than the others (1���2 slightly larger than the others); scales on chin small, rounded, and juxtaposed; on posterior chin region granular to flat-granular, approximately vertical in position or slightly inclined, pointing posteriorly, becoming smaller posteriorly; scales near posterior infralabials flat, subimbricate, and larger than in median area of chin; infralabials four, first largest and almost reaching anterior level of orbit, fourth below center of eye; throat with a short transition between the anterior region with small granular scales and the posterior region with larger, granular, and posteriorly pointing scales; dorsal scales on neck and body small, granular, and juxtaposed, at midbody and posterior portion of trunk pointed-granular or, in frontal view, triangular, mostly directed posteriorly, some flatgranular on flanks and in lumbar region; 38 (36���42) middorsal scales in one HL, 78 (78���96) between levels of axilla and groin; ventrals flat, smooth, imbricate, with an ovoid posterior margin, increasing moderately in size from gular region to belly, posterior ventral scales longer than wide; 13 (10���15) midventral scales in one HL, 27 (24���27) between levels of axilla and groin, 33 (29���33) to border of cloaca; transition between ventrals and scales on flanks abrupt; scales around midbody 77 (71���84), of which 19 are ventrals (15���19); scales on precloacal plate similar to ventrals, except for those on border of cloaca, which are smaller; tail dorsally and laterally with flattened, smooth, and imbricate scales, less elongated than ventrals (with a transitional zone at base of tail); underside of tail with a median row of slightly enlarged scales, mostly with a repeated series of one median scale bordered laterodistally by one scale, followed by a slightly larger median scale in contact laterodistally with two scales, constituting a regular tail sequence of 1 ' 1 ''; dorsal scales on forelimbs granular to flat-granular; scales on hind limbs flat, smooth, and imbricate on anteroventral femoral and shank surfaces, granular elsewhere; fingers, from longest to shortest, IV-III-II-V-I; toes IV-III-II-V-I, fourth and third toes about the same length; lamellae under fourth finger five (4���5), with 2 (2���3) proximal ones greatly enlarged, each about 3 (3���4) times as long as the remaining distal ones; lamellae under fourth toe six (6���8), with 2 (2���3) proximal ones greatly enlarged; claws enclosed by an ungual sheath composed of six scales, as typical for the genus. Hemipenis morphology. The everted hemipenis of SMF 50950 (Fig. 6 E���F) is a small, bilobate organ, divided for around one third of its length, with a naked base; sulcus spermaticus bordered by well-developed, smooth sulcal lips; lips opening into two broad concave areas, one on each lobe; ornamentation of papillate calyces present on each lobe; asulcate area of the truncus covered by papillate calyces and some barely visible spinulate calyces; a third lobule like rising from the pedicel, not connected to the sulcus spermaticus, and covered by spinulate calyces. Coloration in preservative (alcohol 70 %; variation among the paratypes in parentheses). Dorsal ground color Hair Brown (277); occipital marks Grayish Horn Color (268) (Drab-Gray (256) in males), posteriorly bordered with Sepia (279); dorsum of head Army Brown (46); infra- and supralabials with alternating Sepia (279) and white transverse bars; chin and throat with Sepia (286) reticulations on a dirty white background; venter strongly suffused with Vandyke Brown (281) (escutcheon of males unpigmented in the center, with Olive Brown (278) borders). Coloration in life. Dorsal ground color Mahogany Red (34), with small scattered Lavender Blue (195) and Sepia (286) dots; neck region Glaucous (272); a Sepia (286) line from the tip of the snout to the anterior border of the eye; two postorbital Sepia (286) lines, one directed to the occipital region and the other towards the ear; infra- and supralabials with alternating Sepia (279) and dirty white transverse bars; two occipital M-shaped Smoke Gray (267) marks (Chamois (84) in males), posteriorly bordered with Sepia (279); chin and throat with Sepia (286) reticulations on a dirty white background; ventral ground color Lavender Blue (195), strongly suffused with Vandyke Brown (281); a pale dorsolateral line from above the hind limbs to mid tail, a Sepia (286) line from above groin to mid tail; toes and fingers Vandyke Brown (282). Distribution and habitat. Lepidoblepharis victormartinezi is an endemic species of the Isthmian-Atlantic moist forests in west-central Panama (Fund 2011), known from around 100 m elev. in the province of Col��n and 700 m elev. in Veraguas province. Most probably, L. victormartinezi lives in the leaf-litter and feeds on small invertebrates like other Lepidoblepharis (Vitt et al. 2005). Most specimens have been found on top of small hills, giving the impression that this species prefers drier environments on the hills rather than the more wet flat areas around the same locality. However, the specimen SMF 89963 was found in a wet flat area. Etymology. The specific epithet victormartinezi is a patronym for Victor Mart��nez Cort��s, who has pioneered Panamanian herpetology among native researchers, and was the first Panamanian herpetologist ever to publish his results in scientific journals. Since the 1980 s, he has conducted herpetological inventory work at biogeographically significant localities throughout western Panama. The now unfortunately lost (V. Mart��nez, personal communication) specimens of " Lepidoblepharis sp." mentioned in his species lists of the region around Santa F�� de Veraguas (Mart��nez et al. 1995, Mart��nez & Rodriguez 1994), which includes Cerro Negro as the provenance of one of our paratypes, might have been the first specimens of this new species that were ever collected. We dedicate this species to our friend and colleague Victor Mart��nez in due recognition of his passionate dedication to, and great achievements for, Panamanian herpetology., Published as part of Batista, Abel, Ponce, Marcos, Vesely, Milan, Mebert, Konrad, Hertz, Andreas, K��hler, Gunther, Carrizo, Arcadio & Lotzkat, Sebastian, 2015, Revision of the genus Lepidoblepharis (Reptilia: Squamata: Sphaerodactylidae) in Central America, with the description of three new species, pp. 187-221 in Zootaxa 3994 (2) on pages 201-204, DOI: 10.11646/zootaxa.3994.2.2, http://zenodo.org/record/243281, {"references":["Martinez, V. & Rodriguez, A. (1994 \" 1992 \") Del primer inventario en \" Cerro Tute \". Amphibia: Caudata y Anura. Reptilia: Squamata. Sauria y Serpentes. Scientia (Panama), 7, 29 - 53.","Martinez, V., Pimentel, N. & Hurdaneta, A. (1995 \" 1994 \") Diversidad herpetofaunistica en los cerros \" Narices \" y \" La Anselma \": Provincia de Veraguas. Distrito de Santa Fe. Scientia (Panama), 9, 59 - 79.","Auth, D. L. (1994) Checklist and bibliography of the amphibians and reptiles of Panama. Smithsonian Herpetological Information Service, 98, 1 - 59. http: // dx. doi. org / 10.5479 / si. 23317515.98.1","Young, B. E., Sedaghatkish, G., Roca, E. & Fuenmayor, Q. D. (1999) El estatus de la conservacion de la herpetofauna de Panama. Resumen del primer taller internacional sobre la herpetofauna de Panama. The Nature Conservancy y Asociacion Nacional para la Conservacion de la Naturaleza (ANCON), Arlington, Virginia, 40 pp.","Kohler, G. (2008) Reptiles of Central America. Second edition. Herpeton, Offenbach, 400 pp.","Jaramillo A., C. A., Wilson, L. D., Ibanez D., R. & Jaramillo, F. E. (2010) The herpetofauna of Panama: distribution and conservation status. In: Wilson, L. D., Townsend, J. H., Johnson, J. D. & Murphy, J. B. (Eds.) Conservation of Mesoamerican amphibians and reptiles. Eagle Mountain Press, Eagle Mountain, Utah, pp. 604 - 671.","Lotzkat, S., Hertz, A., Stadler, L., Hamad, N., Carrizo Diaz, A. R. & Kohler, G. (2010) Noteworthy distribution records records of reptiles from Western Panama. Herpetological Review, 41, 520 - 523.","Fund, W. (2011) Isthmian-Atlantic moist forests. Available from http: // www. eoearth. org / view / article / 153927 (accessed 24 October 2013)","Vitt, L. J., Sartorius, S. S., Avila-Pires, T. C., Zani, P. A. & Esposito M. C. (2005) Small in a big world: Ecology of leaf-litter geckos in new world tropical forests. Herpetological Monographs, 19, 137 - 152. http: // dx. doi. org / 10.1655 / 0733 - 1347 (2005) 019 [0137: SIABWE] 2.0. CO; 2"]}

Lepidoblepharis emberawoundule sp. nov. Figs. 3 ���7, 10. Lepidoblepharis xanthostigma: all in part. (referring to certain populations in eastern Panama): Auth (1994); Young et al. (1999); Ib����ez et al. (2001); K��hler (2001: Fig. 172; 2008: Fig. 136); Jaramillo et al. (2010). Holotype. Adult male SMF 50968, original field number AB 963 (Fig. 10), collected from leaf-litter at La Cascada trail, Burbayar private reserve (9.31837 ��N, 79.00266 ��W, 360 m elev.), Cart��, Nargan��, Comarca Guna Yala, Panama, on 26 November 2013 at 23: 40 hrs, by Abel Batista and Konrad Mebert. Paratypes. Three adult males, two adult females, and one juvenile, all from Panama. Three males (SMF 81950 ��� 52) from Nusagand�� field station and two females (SMF 81953 ��� 54) from the nearby Sendero Nusagand��, Comarca Guna Yala, collected 14���17 April 2000; one juvenile (MHCH 2952) from R��o Terable, El Llano, Chepo, Panama, collected on 27 November 2012; see Appendix I for locality details. Referred specimens. MHCH 2951, 2957; SMF 50969 ��� 70; FMNH 170029, 170042��� 45; see Appendix I for locality details. Diagnosis. Lepidoblepharis emberawoundule (our sp. nov. 1) is characterized by the following combination of characters: (1) dorsal body scales small, granular, and juxtaposed, ventral scales large, cycloid, flat, and imbricate; (2) scales on head small and granular; (3) 3���4 (3.1 �� 0.3) postrostral scales; (4) a vaguely M-shaped posterior mental border with two paramedian clefts; (5) 3���7 (5.1 �� 1.04) postmentals, larger than the posteriorly adjacent scales on chin; (6) lamellae under fourth toe 6���9 (7.9 �� 0.85), lamellae under fourth finger 5���8 (6.7 �� 1.01); (7) median subcaudals conspicuously wider than long, but their width less than twice the width of the laterally adjacent scales or their own length, with straight or rounded posterior margins, arranged in a regular tail sequence of 1 ' 1 ''; (8) ventral escutcheon consisting of 38���61 (48.4 �� 8.85) scales, 6���7 (6.8 �� 0.45) scales long and 10���13 (11.6 �� 1.14) wide; (9) subfemoral escutcheon consisting of 4���5 (4.4 �� 0.52) well-discernible scales per thigh arranged in a single row (Fig. 5 A); (10) 16���20 (18.2 �� 1.17) longitudinal rows of ventral scales at midbody; (11) 52���74 (61.8 �� 7.61) longitudinal rows of dorsal scales at midbody; (12) bilobate hemipenis, with a third lobule rising from the pedicel (Fig. 6 A���B); (13) SVL 21���30 (25 �� 2.07) mm. Comparison with other species of the genus. Lepidoblepharis emberawoundule can be differentiated from many species in the genus by its small size and its low number of lamellae under the fourth toe and finger (Figs. 3��� 4). In the following, we provide comparisons to all other species within the genus, with the characteristics for L. emberawoundule in parentheses. Lepidoblepharis emberawoundule can be distinguished from the Panamanian species L. xanthostigma, L. sanctaemartae, Lepidoblepharis sp. nov. 2 (described below), and Lepidoblepharis sp. nov. 3 (described below) by uncorrected genetic p-distance (10���26 % in 16 S mtDNA between individuals). Lepidoblepharis xanthostigma has greatly enlarged median subcaudal scales (slightly enlarged), and 12���16 lamellae under its fourth toe (6���9). Lepidoblepharis sanctaemartae has large, flat, imbricate dorsal body scales (small granular scales). Lepidoblepharis sp. nov. 2 (described below) has 13 lamellae under its fourth toe (6���9) and 11 under its fourth finger (5���8). Lepidoblepharis sp. nov. 3 (described below) has 4���5 lamellae under its fourth finger (5���8) and a unique lamellar configuration with 1���3 proximal lamellae per digit greatly enlarged, i.e., about 3���4 times longer than any of the remaining lamellae, the ventral escutcheon consisting of 61���68 scales (38���61), and no discernible subfemoral escutcheon (4���5 discernible subfemoral escutcheon scales per thigh). To date, seven species of the genus Lepidoblepharis have been reported to possess ten or fewer lamellae under the fourth toe, i.e., to be short-toed. Two of these, L. miyatai Lamar 1985 and L. sanctaemartae, possess large, flat, and imbricate dorsal scales (small, granular, and juxtaposed dorsals). Three others, Lepidoblepharis buchwaldi Werner 1910, L. montecanoensis Markezich & Taphorn 1994, and L. williamsi Ayala & Serna 1986, can be readily distinguished from L. emberawoundule because the inspection of the illustrations and photographs available for the respective holotypes showed clear differences between the species: In L. buchwaldi, the enlarged subcaudals are much wider than long and at least twice as wide as the laterally adjacent subcaudals (less than twice as wide as they are long or as the neighboring subcaudals are wide), the dorsal tail scales are small, i.e., less than twice the size of the dorsal body scales (twice or more the size of the dorsal body scales), and the posterior border of the mental has a single median cleft (two paramedian clefts). Lepidoblepharis montecanoensis is a very small species with a SVL of 18���21 mm (21���30), and lacks defined occipital marks in males (two well defined occipital marks in males) as well as distinctly enlarged median subcaudals (median subcaudals distinctly enlarged). Lepidoblepharis williamsi also lacks enlarged median subcaudal scales (median subcaudals distinctly enlarged), and has only 25���40 ventral escutcheon scales (38���61). The holotype of L. peraccae Boulenger 1908 has eight lamellae under the fourth finger (5���8) and ten under the fourth toe (6���9), its plantar and palmar scales have ovoid and strongly imbricate posterior borders (those scales small, rounded, and juxtaposed; Fig. 7). The holotype of L. microlepis (Noble 1923) is very similar to L. emberawoundule, but differs in the scalation of the chin region and the ventral tail surface (Fig. 7). The posterior margin of its mental is V-shaped and lacks conspicuous clefts (posterior margin M-shaped, i.e., slightly convex in the middle, with two conspicuous paramedian clefts), there are six postmentals, with one medial postmental greatly enlarged and two neighboring scales slightly enlarged (3���7 postmentals, median scales slightly larger than the others), and the posteriorly adjacent chin scales are small and conical (small and flat, some slightly pointed, and juxtaposed, Figs. 3, 7). Most decisively, each of the slightly enlarged subcaudal scales of the holotype of L. microlepis is bordered laterodistally by only one scale, leading to a regular tail sequence of 1 ' 1 ' (the larger of the enlarged subcaudals bordered laterodistally by two scales, the smaller ones by one, forming a regular tail sequence of 1 ' 1 ''; see Fig. 7 E���F). The remaining species of the genus, i.e., L colombianus Mechler 1968, L conolepis Avila-Pires 2001, L. duolepis Ayala & Castro 1983, L. festae Peracca 1897, L. grandis Miyata 1985, L. heyerorum Vanzolini 1978, L. hoogmoedi Avila-Pires 1995, L. intermedius Boulenger 1914, and L. ruthveni Parker 1926 are long-toed with eleven or more lamellae under the fourth toe (6���9 in L. emberawoundule). Description of the holotype. Variation among the entire type series is given in parentheses for selected characters (see Table 1 for details and variation among all examined specimens). Adult male as judged by everted hemipenes; SVL 26 mm (21���27 mm), TL 27 mm (measured while the now broken tail was still intact) (18���27 mm), HL 5.6 mm (5.0��� 5.6 mm), HW 3.8 mm (3.2���4.4 mm), forelimbs 5.2 mm, hind limbs 8.6 mm, shank 3.7 mm; rostral large, clearly visible from above, with a shallow, horseshoe-shaped posterior depression and a long median cleft; postrostrals including supranasals three (3���4), the median ones smaller than supranasals, and indenting the rostral, the median postrostral about the same size as the posterior scales on snout; postnasals two (1���2), both about the same size as posteriorly adjacent loreal scales; scales on snout small and smooth, 20 (18���21) scales across snout between anterior sutures of second SPLs; loreal scales juxtaposed, elevated, and rounded, 7 (6���7) on a longitudinal line between postnasals and orbit; scales on top of head small, granular, juxtaposed, generally pointing upward, about as half as large as those on the middle area of snout; superciliary flap with two enlarged scales on anterior border, of which the first is slightly longer than the second, followed by four small scales; supralabials three (3���4), posteriormost one below center of eye; ear-opening small, oval, in oblique orientation; mental large, posterior margin slightly convex in the middle, with two small clefts bordering this convexity, resulting M-shaped; postmentals 7 (4���7), median scales slightly larger than the others, postmentals larger than the posteriorly adjacent chin scales (Figs. 3, 7); scales on chin small and juxtaposed, most of them flat but some slightly pointed, on posterior region granular, approximately vertical in position, with a slight reduction in size towards posterior portion of chin; scales near posterior infralabials flat, subimbricate, and larger than scales in median area of chin; infralabials four, first largest, fourth below center of eye; throat with small granular scales, the posterior region with larger, granular, and pointed scales directed upward (some directed posteriorly); dorsal scales on neck and body small, granular, and juxtaposed; dorsals around midbody and on posterior portion of trunk pointed, granular, or, in frontal view, triangular, mostly directed posteriorly, some scales on flanks and in lumbar region slightly flattened; 40 (36���50) middorsal scales in one HL, 89 between levels of axilla and groin; ventrals flat, smooth, imbricate, with an ovoid posterior margin, increasing moderately in size from gular region to belly, posterior ventral scales longer than wide; 12 (11���15) midventral scales in one HL, 27 between levels of axilla and groin, 32 to border of cloaca; ventral escutcheon patch with 53 (38���46) scales, some of which have slightly pointed posterior margins, 7 (6���7) scales long and 13 (10���13) wide, escutcheon long/wide ratio 54 % (50���70 %); subfemoral escutcheon scales five (4���5) per thigh, arranged in a single row; transition between ventrals and scales on flanks abrupt; scales around midbody 92 (69���92), of which 18 are ventrals (16���19); scales on precloacal plate similar to ventrals, except for those on border of cloaca, which are smaller; tail dorsally and laterally with flattened, smooth, and imbricate scales, less elongated than ventrals (with a transitional zone at base of tail); underside of tail with a median row of moderately enlarged scales, mostly with a repeated series of one median scale bordered laterodistally by one scale, followed by a slightly larger median scale that is in contact laterodistally with two scales (Figs. 3, 7), constituting a regular tail sequence of 1 ' 1 ''; dorsal scales on forelimbs granular; scales on hind limbs flat, smooth, imbricate on anteroventral thigh and shank surfaces, granular elsewhere; fingers, from longest to shortest, IV-III-II-V-I; toes IV- III-II-V-I, fourth and third toes about the same length; lamellae under fourth finger six (5���8), under fourth toe eight (8���9), proximal lamellae slightly larger than distal ones; claws enclosed by an ungual sheath composed of six scales, as typical for the genus. Hemipenis morphology. The everted hemipenis of SMF 50968 (Fig. 6 A���B) is a small, bilobate organ, divided for around one third of its length, with a naked base; sulcus spermaticus bordered by well-developed, smooth sulcal lips; ornamentation of papillate calyces present on each lobe, asulcate area of the truncus covered by small spines; a third lobule-like rising from the pedicel, not connected to the sulcus spermaticus, and covered with papillate calyces. Coloration in preservative (alcohol 70 %; variation among the paratypes in parentheses). Dorsal ground color Hair Brown (277); occipital marks Beige (254) (Gray Horn Color (268) in females), posterior margin of orbit bordered with Sepia (279); dorsum of head with small Beige (254) blotches; infra- and supralabials with alternating Sepia (279) and white bars; chin and throat with Sepia (286) marks on a dirty white background; venter slightly pigmented with Vandyke Brown (281); escutcheon scales unpigmented in the center, with Olive Brown (278) borders. Coloration in life (Fig. 10; variation among the paratypes in parentheses). Dorsal ground color Glaucous (272), with small scattered Lavender Blue (195) and Sepia (286) dots; neck region Olive Brown (278); an indistinct Sepia (286) line from tip of snout to anterior border of eye; two diffuse postorbital Sepia (286) lines, one directed towards the occipital region and the other towards the ear; infra- and supralabials with alternating Sepia (279) and white bars; a vaguely M-shaped dirty white (Smoke Gray (267) in females) occipital mark bordered with Burnt Sienna (38); top of head suffused with Fawn Color (258) and Lavender Blue (195); chin and throat with Sepia (286) reticulations on a Chamois (84) background; venter Lavender Blue (195) suffused with Cinnamon-Rufous (31); an indistinct dorsolateral pale line from behind the ear to mid tail; tail Cinnamon-Rufous (31); a Sepia (286) line from above groin to mid tail. Distribution and habitat. Lepidoblepharis emberawoundule is currently known from a few sites in eastern Panamanian montane forests and Choc��-Dari��n moist forests (Fund 2011), from 227 to 773 m elevation in Dari��n and Panam�� provinces as well as in the Comarcas Ember�� and Guna Yala. Most probably, L. emberawoundule lives in the leaf-litter and feeds on small invertebrates like other Lepidoblepharis (Vitt et al. 2005). Etymology. The name emberawoundule is a compound word in honor to ���the forest guardians���, the three indigenous peoples inhabiting eastern Panama; embera: Ember�� Indians from the foothills of Jingurud��, Bagre, Sapo, Dari��n, and Pirre mountain ranges; woun: Wounaan Indians, mainly from the Tuira basin and Maj�� mountain range; dule: meaning people in the language of the Guna Indians from the Caribbean and Pacific versants of the San Blas and Dari��n mountain ranges., Published as part of Batista, Abel, Ponce, Marcos, Vesely, Milan, Mebert, Konrad, Hertz, Andreas, K��hler, Gunther, Carrizo, Arcadio & Lotzkat, Sebastian, 2015, Revision of the genus Lepidoblepharis (Reptilia: Squamata: Sphaerodactylidae) in Central America, with the description of three new species, pp. 187-221 in Zootaxa 3994 (2) on pages 194-196, DOI: 10.11646/zootaxa.3994.2.2, http://zenodo.org/record/243281, {"references":["Auth, D. L. (1994) Checklist and bibliography of the amphibians and reptiles of Panama. Smithsonian Herpetological Information Service, 98, 1 - 59. http: // dx. doi. org / 10.5479 / si. 23317515.98.1","Young, B. E., Sedaghatkish, G., Roca, E. & Fuenmayor, Q. D. (1999) El estatus de la conservacion de la herpetofauna de Panama. Resumen del primer taller internacional sobre la herpetofauna de Panama. The Nature Conservancy y Asociacion Nacional para la Conservacion de la Naturaleza (ANCON), Arlington, Virginia, 40 pp.","Kohler, G. (2001) Anfibios y reptiles de Nicaragua. Herpeton Verlag, Offenbach, 208 pp.","Jaramillo A., C. A., Wilson, L. D., Ibanez D., R. & Jaramillo, F. E. (2010) The herpetofauna of Panama: distribution and conservation status. In: Wilson, L. D., Townsend, J. H., Johnson, J. D. & Murphy, J. B. (Eds.) Conservation of Mesoamerican amphibians and reptiles. Eagle Mountain Press, Eagle Mountain, Utah, pp. 604 - 671.","Lamar, W. W. (1985) A new Lepidoblepharis (Sauria: Gekkonidae) from the north coast of Colombia. Herpetologica, 41, 128 - 132.","Werner, F. (1910) Uber neue oder seltene Reptilien des Naturhistorischen Museums in Hamburg. ii. Eidechsen. Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 27, 1 - 46.","Markezich, A. L. & Taphorn, D. C. (1994) A new Lepidoblepharis (Squamata: Gekkonidae) from the Paraguana Peninsula, Venezuela, with comments on its conservation status. Herpetologica, 50, 7 - 14.","Ayala, S. C. & Serna, M. A. (1986) Una nueva especie de Lepidoblepharis (Sauria, Gekkonidae) de la Cordillera Central de Colombia. Caldasia, 15, 649 - 654.","Boulenger, G. A. (1908) Descriptions of new South-American reptiles. The Annals and Magazine of Natural History, 8, 111 - 115. http: // dx. doi. org / 10.1080 / 00222930808692365","Noble, G. K. (1923) A new gekkonid lizard and a new brachycephalid frog from Colombia. American Museum Novitates, 88, 1 - 3.","Mechler, B. (1968) Les Geckonides de la Colombie. Revue Suisse de Zoologie, 75, 305 - 371.","Avila-Pires, T. C. S. (2001) A new species of Lepidoblepharis (Reptilia: Squamata: Gekkonidae) from Ecuador, with a redescription of Lepidoblepharis grandis Miyata, 1985. Occasional Papers of the Sam Noble Oklahoma Museum of Natural History, 11, 1 - 11.","Ayala, S. C. & Castro, F. (1983) Dos nuevos gecos (Sauria: Gekkonidae, Sphaerodactylinae) para Colombia: Lepidoblepharis xanthostigma (Noble) y descripcion de una nueva especie. Caldasia, 13, 743 - 753.","Peracca, M. G. (1897) Viaggio del Dr. Enrico Festa nell'Ecuador e regioni vicine. IV. Rettili. Bollettino dei Musei di Zoologia e di Anatomia Comparata della R. Universita di Torino, 12, 1 - 20.","Miyata, K. (1985) A new Lepidoblepharis from the Pacific slope of the Ecuadorian Andes (Sauria: Gekkonidae). Herpetologica, 41, 121 - 126.","Vanzolini, P. E. (1978) Lepidoblepharis in Amazonia (Sauria, Gekkonidae). Papeis Avulsos de Zoologia (Sao Paulo), 31, 203 - 211.","Avila-Pires, T. C. S. (1995) Lizards of Brazilian Amazonia. (Reptilia: Squamata). Zoologische Verhandelingen Leiden, 299, 1 - 706.","Boulenger, G. A. (1914) On a second collection of batrachians and reptiles made by Dr. H. G. F. Spurrell, F. Z. S. in the Choco, Colombia. Proceedings of the Zoological Society London, 1914, 813 - 817.","Parker, H. W. (1926) The neotropical lizards of the genera Lepidoblepharis, Pseudogonatodes, Lanthrogecko, and Sphaerodactylus, with the description of a new genus. The Annals and Magazine of Natural History, 17, 291 - 301. http: // dx. doi. org / 10.1080 / 00222932608633413","Fund, W. (2011) Isthmian-Atlantic moist forests. Available from http: // www. eoearth. org / view / article / 153927 (accessed 24 October 2013)","Vitt, L. J., Sartorius, S. S., Avila-Pires, T. C., Zani, P. A. & Esposito M. C. (2005) Small in a big world: Ecology of leaf-litter geckos in new world tropical forests. Herpetological Monographs, 19, 137 - 152. http: // dx. doi. org / 10.1655 / 0733 - 1347 (2005) 019 [0137: SIABWE] 2.0. CO; 2"]}

Lepidoblepharis rufigularis sp. nov. Figs. 3 –5, 11. Lepidoblepharis xanthostigma: all in part. (referring to populations in extreme southeastern Panama): Auth (1994); Young et al. (1999); Ibáñez et al. (2001); Köhler (2008); Jaramillo et al. (2010). Holotype. Adult male SMF 50659, original field number AB 527 (Figs. 3 –5, 11), collected on a hill 1 km north of Río Púcuro (8.057501 °N, 77.370217 °W, 1043 m elev.), Pinogana, Darién, Panama, on 0 8 July 2012 at 22: 40 hrs by Abel Batista. Diagnosis. Lepidoblepharis rufigularis (our sp. nov. 2) is characterized by the following combination of characters: (1) dorsal scales small, granular, and juxtaposed, ventral scales large, cycloid, flat, and imbricate; (2) scales on head small and granular; (3) four postrostral scales; (4) two short, barely discernible paramedian clefts in the more or less U-shaped posterior mental border; (5) five postmentals, the two median ones larger than the posteriorly adjacent chin scales; (6) 13 lamellae under fourth toe, 11 lamellae under fourth finger; (7) median subcaudals conspicuously wider than long, almost twice as wide as the laterally adjacent scales, with straight posterior margins arranged in a regular tail sequence of 1 ' 1 ''; (8) ventral escutcheon consisting of 62 scales, almost twice as wide (13 scales) as long (7 scales); (9) subfemoral escutcheon consisting of 3–4 scales per thigh; (10) 17 longitudinal rows of ventral scales at midbody; (11) bilobate hemipenis; (12) SVL 25 mm. Comparison with other species of the genus. Lepidoblepharis rufigularis can be differentiated from all species in the genus by its small size, number of lamellae under the fourth toe and finger, the reddish throat in males (Fig. 11), and the configuration of the ventral escutcheon. In the following, we present comparisons to all other species within the genus, with the characteristics for L. rufigularis in parentheses. Lepidoblepharis rufigularis can be distinguished from the Panamanian species L. xanthostigma, L. sanctaemartae, L. emberawoundule, and Lepidoblepharis sp. nov. 3 (described below) by a genetic p-distance of 14–23 % between individuals in 16 S mtDNA. Lepidoblepharis xanthostigma is the most similar species, but has a different chin and throat coloration with dark reticulations on a pale background (orange background), and greatly enlarged median subcaudal scales which are more than two times as wide as the laterally adjacent subcaudal scales (enlarged but less than two times the width of laterally adjacent subcaudals, Fig. 3), usually 21 or more, very rarely 18, scales across snout (19), usually 18 or more, rarely 16 or 17, ventral scales a midbody (17), and an escutcheon long/wide ratio of 67–120 % (54 %). Lepidoblepharis emberawoundule, L. sanctaemartae, and Lepidoblepharis sp. nov. 3 (described below) have fewer than 10 lamellae under the fourth toe (13) and under the fourth finger (11). Additionally, L. sanctaemartae has large, flat, imbricate dorsal body scales (small, granular, and juxtaposed). To date, seven species of the genus Lepidoblepharis have been reported to possess ten or fewer lamellae under the fourth toe, i.e., to be short-toed (L. miyatai, L. sanctaemartae, L. buchwaldi, L. montecanoensis, L. williamsi, L. peraccae, and L. microlepis), and are therefore readily differentiable from the long-toed L. rufigularis (13 lamellae under the fourth toe). Of the remaining members of the genus, L. colombianus, L. conolepis, L. duolepis, L. festae, L. grandis, L. heyerorum, L. hoogmoedi, L. intermedius, and L. ruthveni are relatively to very large lizards for this genus with adult SVLs between 33 and 56 mm (25 mm). Additionally, L. conolepis and L. grandis have 14–20 lamellae under the fourth toe (13). The dorsal ground color in males of L. heyerorum is black with yellow dorsal markings (no yellow dorsal markings). The two long-toed specimens with granular dorsals reported as L. xanthostigma from Colombia by Ayala & Castro (1983) are similar to L. rufigularis in the number of ventral scales, but they have 22–25 scales across snout (19), an escutcheon with only 25 scales (62), the gular region with blotches (gular region with longitudinal bars), and an occipital pale W-shaped mark (no occipital mark at all). Description of the holotype. Adult male as judged by everted hemipenes; SVL 25 mm, TL 33.0 mm (measured while the now broken tail was still intact), HL 5.6 mm, HW 4.1 mm, forelimbs 5.0 mm, hind limbs 9.3, shank 3.6 mm; rostral large, clearly visible from above, with a shallow, horseshoe-shaped posterior depression and a long median cleft; postrostrals four, including supranasals, one median postrostral slightly larger than posteriorly adjacent scales on snout and indenting the rostral; postnasals two, both about the same size as posteriorly adjacent loreal scales; scales on snout rounded and pointed backward; loreal scales subimbricate, elevated toward posterior and dorsal directions, six loreal scales on a longitudinal line between postnasals and orbit; 19 scales across snout between anterior sutures of second SPLs; scales on top of head small, granular, juxtaposed, generally pointing upward, about half the size of those on the middle area of snout; superciliary flap with two enlarged scales on anterior border, of which the first is slightly longer than the second, followed by 2–5 small and globular scales; supralabials three, posteriormost one below center of eye; ear-opening small, oval, in oblique orientation; mental large, posterior margin somewhat U-shaped, with two barely discernible, very short paramedian clefts; five enlarged postmentals, slightly larger than the posteriorly adjacent chin scales (Fig. 3 B); scales on chin small, flat, rounded, and juxtaposed, on posterior region flat to flat-granular, slightly inclined backward; scales near posterior infralabials flat, subimbricate, and larger than scales in median area of chin; infralabials four, first largest and almost reaching anterior level of orbit, fourth below center of eye; throat with small granular scales, pointed and directed upward; dorsal scales on neck and body small, granular, and juxtaposed, at midbody and on posterior trunk pointed, granular, or, in frontal view, triangular, mostly directed posteriorly, some scales on flanks and in lumbar region pointed; dorsal skin on body too damaged to conduct longitudinal counts of middorsal scales; ventral scales flat, smooth, imbricate, with an ovoid posterior margin, increasing moderately in size from gular region to belly, posterior ventral scales longer than wide; 13 midventral scales in one HL, 23 between levels of axilla and groin, 28 to border of cloaca; ventral escutcheon patch with 62 scales, 7 long and 13 wide, escutcheon long/wide ratio 54 %; subfemoral escutcheon with 3 scales in a single row under each thigh, and a barely discernible fourth scale in a second row under the right thigh; transition between ventrals and lateral scales abrupt; 17 longitudinal rows of ventrals at midbody; scales on precloacal plate similar to ventrals, except for those on border of cloaca, which are smaller; tail dorsally and laterally with flattened, smooth, and imbricate scales that are less elongate than ventrals (with a transitional zone at base of tail); underside of tail with a median row of moderately enlarged scales (usually no more than two times wider than the laterally adjacent subcaudal scales; Fig. 3 B), mostly with a repeated series of one median scale bordered laterodistally by one scale, followed by a slightly larger median scale in contact laterodistally with two scales, constituting a regular tail sequence of 1 ' 1 ''; dorsal scales on forelimbs granular to conical; scales on hind limbs flat, smooth, and imbricate on anteroventral femoral and shank surfaces, granular elsewhere; fingers, from longest to shortest, IV-III-II-V-I; toes IV-III-II-V-I, fourth and third toes about the same length; 11 lamellae under fourth finger, 13 under fourth toe; claws enclosed by an ungual sheath composed of six scales, as typical for the genus. Hemipenis morphology. The partially everted hemipenis of SMF 50659 (Fig. 5 B) is a small organ; sulcus spermaticus bordered by well-developed and smooth sulcal lips; asulcate area of the truncus covered by papillate calyces; proximal portion of apex covered by small spinulate calyces. Due to its incomplete eversion, it is not possible to determinate whether the hemipenis is as bilobate as those of other species of the genus described and/or pictured herein (Fig. 6). Nevertheless, it is apparent that the hemipenis of Lepidoblepharis rufigularis does not bear a conspicuous basal third lobule, in contrast to the other two species described herein. Coloration in preservative (alcohol 70 %). Dorsal ground color Hair Brown (277); posterior margin of orbit with a pale Beige (254) line; dorsum of head suffused with Beige (254); infra- and supralabials with alternating Sepia (279) and white bars; chin and throat with Sepia (286) reticulations on a dirty white background; venter pigmented with Vandyke Brown (281); escutcheon scales unpigmented in the center, with Olive Brown (278) borders. Coloration in life (Fig. 11). Dorsal ground color Grayish Horn Color (268), with patches of Lavender Blue (195) and Vandyke Brown (281) scales, same pattern on neck and head; a straight Light Sky Blue (191) postorbital line, running up to level of ear; infra- and supralabials with alternating Sepia (279) and Burnt Orange (10) bars; chin and throat with Sepia (279) reticulations on a Burnt Orange (10) background; ventral parts suffused with Dusky Brown (285) and Light Sky Blue (191); escutcheon scales Smoke Gray (267), with Dusky Brown (285) scale tips; tail with a dorsolateral Salmon Color (59) line; toes and fingers suffused with Brick Red (36). Distribution and habitat. Lepidoblepharis rufigularis is known only from the type locality in southeastern Darién province. Given the close proximity of the type locality to Colombia, it likely occurs there as well. The habitat at the type locality of L. rufigularis is part of the eastern Panamanian montane forests (Fund 2011), in the Darién mountain range, around 1000 m elev. Most probably, L. rufigularis lives in the leaf-litter and feeds on small invertebrates like other Lepidoblepharis spp. (Vitt et al. 2005). The specimen probably resided between the leaf-litter on a large fallen log, and was uncovered and collected after one of the local supporters in the group stepped over the log. Etymology. The name rufigularis is a compound word that comes from the Latin rufus (red) and gula (throat) referring to the bright orange throat color in this species in life.

Batista, Abel, Ponce, Marcos, Vesely, Milan, Mebert, Konrad, Hertz, Andreas, Köhler, Gunther, Carrizo, Arcadio, Lotzkat, Sebastian (2015): Revision of the genus Lepidoblepharis (Reptilia: Squamata: Sphaerodactylidae) in Central America, with the description of three new species. Zootaxa 3994 (2): 187-221, DOI: http://dx.doi.org/10.11646/zootaxa.3994.2.2

Amphibien bevölkern die Erde seit über 300 Millionen Jahren und sind heute, mit über 7000 bekannten Arten und vielen weiteren die noch zu entdecken sind, eine der artenreichsten Wirbeltierklassen der Welt. Wie verschiedene Studien herausgefunden haben ist etwa ein Drittel der bekannten Amphibienarten unmittelbar vom Aussterben bedroht, was sie zur meist bedrohten Wirbeltierklasse macht. Panama hat im Verhältnis zu seiner relativ kleinen Landesfläche eine der artenreichsten Amphibienfaunen weltweit. Dennoch sind die Rolle im Ökosystem und die Lebensraumansprüche vieler Arten bislang nur unzureichend bekannt, was Gegenstand laufender Forschung ist. Moderne taxonomische Forschung in anderen tropischen Regionen der Welt zeigte außerdem, dass Amphibien weitaus artenreicher sind als vormals angenommen, was mutmaßlich auch auf Panama zutrifft. Gleichzeitig ist das Sammeln von Daten und die Beschreibung neuer Arten ein Wettlauf gegen die Zeit, da in den letzten 30 Jahren die Amphibienvielfalt der Welt im Allgemeinen und Panamas im Besonderen in beispielloser Weise abgenommen hat. Die Gründe dafür sind vielfältig und hauptsächlich durch Zerstörung, Veränderung und Zerstückelung natürlicher Amphibienlebensräume verschuldet, aber auch die tödliche Amphibienkrankheit Chytridiomykose, die durch den Pilz Batrachochytrium dendrobatidis (Bd) verursacht wird, hat ihren Anteil daran. In Panama und Costa Rica hat sich diese Emerging Infectious Disease (EID) (engl. für Neue Infektionskrankheit) wellenartig von Westen nach Osten ausgebreitet und selbst in geschützten Biotopen zu Massenausterbeereignissen und Biodiversitätsverlust unter den Amphibien geführt. Von der Krankheit waren in erster Linie an Bäche gebundene Hochlandarten betroffen. Die letzte großangelegte Bewertung des Erhaltungszustands der Amphibien Panamas, die von der Weltnaturschutzorganisation IUCN 2004 durchgeführt wurde ergab, dass etwa 30% der bekannten Arten vom Aussterben bedroht sind. Außerdem mangelte es bei 17% der damals bekannten Arten an genügend Daten für eine Bewertung. In Anbetracht von Panamas ohnehin schon überwältigenden Amphibienvielfalt und der Vielfalt an Lebensräumen und Orte an denen zuvor nicht nach Amphibien gesucht worden war, startete ich diese Arbeit mit der Überzeugung, dass die Inventarisierung der Amphibienarten noch lange nicht abgeschlossen ist. Außerdem war zu Beginn unklar, ob es überlebende Amphibienarten in Gebieten gäbe in denen Chytridiomykose aufgetreten war. Der Totalzusammenbruch ganzer Amphibiengemeinschaften im Hochland Westpanamas führte zu einer örtlichen Verschiebung der Amphibienforschung hin zu Tieflandgebieten in Zentral- und Ostpanama. Dabei wurde sich verstärkt auf das Eintreffen des Pathogens, mit anschließender Dokumentation des Ausbruchs der Epizootie und dem folgenden Rückgang der Populationen konzentriert. Darum war die Situation der Amphibiengemeinschaften in Gebieten, in denen es bereits zu Populationsrückgängen gekommen war, größtenteils unbekannt. Folglich waren die Hauptziele meiner Arbeit, erstens die taxonomische Inventarisierung der Amphibienarten Panamas voran zu treiben und zweitens die Situation der Amphibiengemeinschaft in Gebieten zu bewerten, in denen Rückgänge der Populationen aufgrund von Bd beobachtet worden waren. Um diese Aufgaben anzugehen führte ich Feldforschung im Westen Panamas durch und legte dabei ein besonderes Augenmerk auf Höhenstufen von 1000 m NN bis auf Panamas höchsten Gipfel, den 3475 m hohen Volcán Barú. Zusätzlich besammelte ich verschiedene Orte im Tiefland zwischen 0 und 1000 m NN um Vergleichsproben zu erhalten. Im Zeitraum zwischen 2008 und 2013 unternahm ich fünf Sammelreisen nach Panama, die zusammengerechnet etwa 13 Monate im Feld bedeuteten. Dabei habe ich neun Regionen im Westen Panamas besucht und gemeinsam mit Studenten 767 Belegexemplare gesammelt, von denen 531 unter meiner Feldnummer gesammelt wurden. Begleitende Daten zu diesen Belegexemplaren beinhalten 68 Rufaufnahmen männlicher Anuren, 102 standardisierte Beschreibungen der Lebendfärbung und 259 Gewebeproben von denen bislang 185 16S mtDNA Fragmente sequenziert wurden. Dieser Datensatz ist der bislang größte für panamaische Amphibien und das erste großangelegte DNA Barcoding der Amphibien Westpanamas. Nachdem ich die ersten Barcodinguntersuchungen durchgeführt und die Ergebnisse mit morphologischen und bioakustischen Daten verglichen hatte, war die Anzahl der näher zu bearbeitenden taxonomischen Probleme sehr viel höher als ich zunächst annahm. Um zu endgültigen Ergebnissen zu kommen mussten die meisten genetischen Linien sehr viel genauer taxonomisch bearbeitet werden. In Anbetracht des hohen finanziellen und zeitlichen Aufwands musste also eine Auswahl getroffen werden mit welchen Arten eine komplette taxonomische Bearbeitung zu machen sei. Ich entschied mich dafür jeweils eine Gattung aus den drei Panamaischen Amphibienordnungen genauer zu analysieren. Die Auswahl dieser Gattungen wurde maßgeblich davon abhängig gemacht, ob genügend Material zur Verfügung steht und ob die Bearbeitung der jeweiligen Gattung wissenschaftliche Neuerungen versprach. Aus der Ordnung Anura wählte ich die Gattung Diasporus. Diese kleinen Frösche sind in vielen Biotopen allgegenwärtig und daher relativ leicht zu finden. Außerdem sind sie nur unzureichend taxonomische bearbeitet worden. Die vorliegende Arbeit ist daher die erste die sich auf die Gattung Diasporus bezieht und sowohl eine molekulare Phylogenie, als auch eine vergleichende Analyse der Anzeigerufe verschiedener westpanamaischer Populationen beinhaltet. Insgesamt sammelte ich 67 Diasporus Belegexemplare in ganz Westpanama und verglich diese mit 49 weiteren mittelamerikanischen Belegexemplaren aus Sammlungen, einschließlich der namenstragenden Typen von D. diasporus und D. hylaeformis. Weitere Vergleichsdaten entnahm ich der Literatur. Die DNA Barcoding Analyse eines Fragments des 16S rRNA Gens beinhaltete 43 eigene Sequenzen, die mit 15 relevanten Sequenzen von GenBank verglichen wurden. Außerdem verglich ich die Anzeigerufe von 26 männlichen Individuen untereinander und mit Angaben aus der Literatur. Die DNA Barcoding Analyse ließ mehrere unbenannte genetische Linien erkennen, aber legte auch einige Individuen zusammen, die sich ansonsten morphologisch und bioakustisch gut unterscheiden ließen. Im Großen und Ganzen ergab die morphologische Untersuchung des Materials keine eindeutigen Merkmale um genetische Linien zu unterscheiden. Allerdings ist es möglich Arten zu unterscheiden, indem man eine Kombination verschiedener Merkmale heranzieht. Welche das sind muss aber von Art zu Art neu entschieden werden. Meine Umfassende Sammlung von Rufaufnahmen machte es möglich zum ersten Mal den Einfluss verschiedener Parameter auf den Ruf von D. hylaeformis zu testen. Die Analyse ergab, dass es eine signifikante Abhängigkeit zwischen der Dominanzfrequenz und der Körpergröße des rufenden Frosches gibt, dabei ist die Frequenz umso höher, je kleiner der Frosch ist. Ein ähnliches Verhältnis konnte zwischen Rufrate und Temperatur beobachtet werden, wobei die Rufrate mit abnehmender Temperatur ebenfalls abnimmt. Ich vermute, dass dieses allgemeingültige Muster, welches bereits für andere Anurengattungen beschrieben wurde, auch auf andere Diasporus Arten zutrifft. Unter Berücksichtigung dieser innerartlichen Variation betrachte ich die vergleichende Rufanalyse als die beste Methode um Arten auseinanderzuhalten. Das gilt insbesondere für syntopische Arten. Die Einbindung aller drei Beweislinien, also der Morphologie, des DNA Barcodings und der bioakustischen Analyse, führte zur Entdeckung von vier unbeschriebenen Arten von denen zwei (D. citrinobapheus und D. igneus) bereits formal von Kollegen und mir beschrieben wurden. In der Ordnung Caudata führte ich eine integrative taxonomische Analyse der westpanamaischen Vertreter der Gattung Bolitoglossa, der größeren der beiden panamaischen Salamandergattungen, durch. Bolitoglossa ist eine sehr artenreiche Gattung mit einem Artbildungszentrum in den Höhenlagen von Costa Rica und Westpanama. Ich sammelte 53 Bolitoglossa Belegexemplare und verglich diese mit zwölf Belegen aus Sammlungen, einschließlich des Holotypus und eines Paratypen von B. gomezi. Der Datensatz wurde dann mit Daten aus der Literatur vervollständigt. Unter den gesammelten Belegen befanden sich zwei Arten, die als gefährdet eingestuft werden und die jeweils seit mehreren Jahrzehnten weder gesammelt noch gesichtet worden waren. Bolitoglossa magnifica wurde seit 34 Jahren nicht gesehenen und B. anthracina seit 22 Jahren. Außerdem fand ich Salamander an mehreren zuvor nicht besammelten Orten. Mein 16S mtDNA Barcoding, das aus 21 eigenen Sequenzen kombiniert mit 47 GenBank Sequenzen besteht, stellt das bislang umfangreichste Taxon-Sampling für panamaische Bolitoglossa dar. Auch wenn diese molekulare Phylogenie nur auf einem Marker basiert, so konnte ich doch weitgehende Übereinstimmungen mit vorangegangen Studien feststellen und die Knoten des Baumes sind statistisch gut unterstützt. Ich erhielt in meinen Bäumen alle zuvor definierten Untergattungen und Artengruppen. Auf Grundlage der molekularen Phylogenie ordne ich B. anthracina, welcher hier erstmals molekular untersucht wurde, der B. subpalmata Artengruppe zu. Wegen seiner Erscheinung als großer, dunkel gefärbter Salamander wurde dieser vormals ohne weitere Untersuchungen in der B. schizodactyla Artengruppe geführt, in der sich die großen, schwarzen Salamander aus dem B. nigescens Artenkomplex befinden. Außerdem fand ich mehrere weit abweichende genetische Linien zwischen geografisch getrennten Populationen von B. minutula. Trotzdem konnte ich bislang keine zusätzlichen morphologischen Merkmale finden, die es erlauben würden diese Linien auseinanderzuhalten. Außerdem konnten meine Kollegen und ich eine neue genetische Linie innerhalb der B. robinsoni Artengruppe als neue Art mit Namen B. jugivagans beschreiben. Im Gegensatz dazu fand ich nur geringfügige genetische Unterschiede zwischen Belegen von B. sombra und B. nigrescens. Ich kombinierte die morphometrischen Messungen und die Anzahl der Zähne, die ich an selbst gesammelten, neuen Belegen von nahe der Typuslokalität von B. sombra genommen hatte, mit Angaben aus der Literatur zu beiden Arten. Dabei verschwammen die Unterscheidungsmerkmale die zur Artabgrenzung verwendet worden waren. Im Besonderen zeigte die Einbeziehung wesentlich größerer Individuen von B. sombra in die Analyse, die zum Zeitpunkt der Artbeschreibung nicht zur Verfügung standen, dass die Unterschiede in der Anzahl der Zähne mit dem Alter und der Größe des Individuums zusammenhängen. So haben größere Individuen mehr Zähne im Oberkiefer und Gaumendach. Auf Grund dieser Beweise sehe ich B. sombra als Juniorsynonym von B. nigrescens an. Ich habe weiterhin die panamaische Verbreitung der relativ häufigen Salamanderarten des Tieflands B. colonnea und B. lignicolor überarbeitet. Dabei untersuchte ich die molekulare und morphologische Variation beider Arten in Panama und schloss die Verbreitungslücken in den nur bruchstückhaft bekannten Verbreitungsgebieten. Während nur wenig Variation innerhalb der Art B. lignicolor zu beobachten war, fand ich weit auseinanderlaufende genetische Linien zwischen geographisch getrennten Populationen von B. colonnea, was weitere Untersuchungen nach sich ziehen wird. Blindwühlen (Ordnung Gymnophiona) gehören zu den am wenigsten untersuchten terrestrischen Wirbeltieren. Nachdem ich ein erstes Individuum aus der vornehmlich südamerikanischen Gattung Oscaecilia (Familie Caeciliidae) erhalten hatte, begann ich mich intensiver mit der Taxonomie der Caeciliidae in Mittelamerika zu beschäftigen. Die Belegexemplare aus Westpanama waren nicht gleich einer beschriebenen Art zuzuordnen, sondern zeigten sowohl Merkmale von O. elongata als auch O. osae. Während O. osae nur anhand des Holotypus bekannt war, ist das Typusmaterial von O. elongata im Zweiten Weltkrieg vernichtet worden. Nach der Originalbeschreibung ist O. elongata von anderen Arten der Gattung durch das einzigartige Fehlen von subdermalen Schuppen im hinteren Teil des Körpers zu unterscheiden. Dieses besondere Merkmal kann in einem der Art zugeordneten Beleg aus dem Osten Panamas nicht überprüft werden, da besagtem Exemplar die hinteren zwei Drittel des Körpers fehlen. Aus diesem Grund fertigte ich hochauflösende Mikro CT Bilder mittels Synchrotron Strahlung (HRμCT) von verschiedenen Belegen an, um Schädelmerkmale vergleichen zu können. Zudem machte ich gewöhnliche Röntgenbilder um Wirbel zu zählen und subdermale Schuppen sichtbar zu machen. Dabei ergab sich, dass das bruchstückhafte Exemplar aus Ostpanama keine Gemeinsamkeiten mit O. osae oder den Oscaecilia aus dem Westen Panamas aufweist und am ehesten der relativ gut bekannten Art O. ochrocephala zuzuordnen ist. Im Gegensatz dazu haben die Belegexemplare aus dem Westen Panamas eine Reihe von Gemeinsamkeiten mit O. osae aber zeigen auch einige Unterschiede auf. Ein genetisches Barcoding ergab zwar, dass beide Arten nahe Verwandte sind, jedoch konnte die genetische Distanz nicht zufriedenstellend erhoben werden, da die 16S Sequenzen von O. osae, die aus Blutproben lebender Individuen gewonnen wurden, von schlechter Qualität waren. Darum vergleiche ich zwar die westpanamaischen Oscaecilia mit O. osae in dieser Arbeit, kann aber derzeit keine taxonomischen Schlüsse ziehen bis weiteres Material zur Verfügung steht. Als weiteres Ergebnis, designiere ich den Namen O. elongata als nomen dubium, da das Typusmaterial verloren ist, die Typuslokalität nur mit „Panama“ angegeben ist und die Erstbeschreibung keine eindeutige Zuordnung zulässt. Vorherige molekulare Untersuchungen hatten bislang nur O. ochrocephala zur Verfügung, daher konnte die Monophylie der Gattung nie getestet werden. Bislang ist die Gattung Oscaecilia weitgehend an einem einzigen Schädelmerkmal, nämlich dass sich die Augen unter dem Schädelknochen befinden, festgemacht. In dieser Arbeit habe ich die 16S mtDNA Sequenzen von O. osae aus Costa Rica und zwei Sequenzen von O. sp. aus Westpanama mit zwei O. ochrocephala Sequenzen und zehn Sequenzen von vier Arten aus der Schwestergattung Caecilia kombiniert. Die daraus resultierende Phylogenie weist zwei gut unterstützte monophyletische Gruppen auf. Eine Gruppe beinhaltet neben den Oscaecilia Arten auch eine Caecilia aus Panama und eine aus dem Westen Ecuadors. Die zweite Gruppe besteht aus zwei Caecilia Arten aus dem Amazonas Becken. Ich vermute daher, dass die Trennung beider Gruppen auf die Hebung der Anden zurückgeht die zur heutigen cis- und trans-andinen Verbreitung beider Gruppen geführt hat. Um die Monophylie vorerst wieder herzustellen schlage ich vor Oscaecilia in die Synonymie von Caecilia zu überführen bis weitere Taxa zur Verfügung stehen. Um den Erhaltungszustand der Amphibien Arten im Hochland Westpanamas zu bewerten, habe ich zunächst eine Liste der Arten erstellt, die ich potentiell hätte finden können. Anhand der IUCN Kategorien bestimmte ich den Anteil gefährdeter Arten, wie viele ich davon gefunden habe und welchen Familien und Artengruppen die gefundenen Arten zuzuordnen sind. Überraschenderweise, sind die wiedergefundenen Arten nicht gleichmäßig über die vier Amphibienfamilien mit den meisten gefährdeten Arten (Bufonidae, Craugastoridae, Hylidae, und Plethodontidae) verteilt. Während ich zehn der elf gefährdeten Laubfroscharten und sechs der neun Lungenlosen Salamander finden konnte, fand ich nur eine von vier gefährdeten Krötenarten und keinen der neun gefährdeten Craugastoriden. Ich glaube, dass die versteckt lebenden Salamander, bei denen auch keine Chytrid bezogenen Bestandsrückgänge beobachtet wurden, in den letzten Jahrzehnten einfach übersehen wurden. Im Gegensatz dazu vermute ich, dass die Laubfrösche, bei denen mit Chytridiomykose verbundene Bestandsrückgänge hinreichend dokumentiert wurden, einen evolutionären Weg gefunden haben mit dem Pathogen zu leben. Das steht anscheinend im Gegensatz zu Bufoniden und Craugastoriden, bei denen ich keine Hinweise auf eine Bestandserholung finden konnte. Die einzigen momentan bekannten Populationen von Arten aus diesen Familien überleben anscheinend nur in klimatischen Nischen, in denen sie vor Bd geschützt sind. Meine Daten bestätigen daher die derzeitige Sichtweise, dass die Wahrscheinlichkeit für Populationszusammenbrüche in einer Amphibiengemeinschaft, die zuvor keinen Kontakt mit dem Erreger hatte, für eine Art umso höher ist je höher deren Verbreitungsgebiet liegt und umso stärker die Art ans Wasser gebunden ist. Also spielt während einer Epizootie die taxonomische Zugehörigkeit eher eine untergeordnete Rolle. Hingegen haben anscheinend nur bestimmte Amphibienfamilien (z.B. Hyliden und Centroleniden) die Fähigkeit Abwehrmechanismen zu entwickeln, um während der enzootischen Phase mit dem Erreger zu koexistieren. Das ist eine sehr neue Sichtweise auf die schlimmste Amphibienkrankheit der Welt, die es erlaubt in neue Richtungen zu forschen, um die Wirt-Erreger-Beziehungen besser zu verstehen. Desweiteren untersuchte ich in welchen Gebieten im Hochland Westpanamas der Anteil an überlebenden, gefährdeten Arten besonders hoch ist. Wie zu erwarten war fanden sich die meisten gefährdeten Arten innerhalb von Schutzgebieten. Eine Ausnahme bildet die ungeschützte Cerro Colorado Region in dem Autonomiegebiet der Ngöbe-Buglé, wo ich eine Vielzahl gefährdeter und unbeschriebener Arten nachweisen konnte. Trotzdem ist hier ein Tagebau geplant, dem große Teile der Wälder zum Opfer fallen würden. Das zeigt wieder einmal, dass der Mensch die größte Gefahr für die Amphibien Panamas und der Welt darstellt. Amphibians have existed on the planet for over 300 million years and are today one of the most diverse vertebrate classes in the world with over 7000 known species and still many more to be discovered. However, several studies assume that approximately one third of the world´s known living amphibians are directly threatened with extinction, making it the most endangered vertebrate class. In relation to the relatively small land mass that is occupied by the state of Panama, it supports one of the most diverse amphibian faunas. However, in many cases the ecological role of single species in a wider context and their habitat preferences are still poorly understood and subject to ongoing research. Modern taxonomic approaches in other tropical regions have shown that former assumptions of amphibian diversity were distinct underestimations of the actual species diversity; a situation that is probably also true for Panama. Concurrently, the collection of amphibian diversity data and the description of new species is a race against time. The amphibian fauna of the world and that of Panama in particular, has suffered from an unprecedented loss of diversity over the last 30 years. The reasons are manifold and include destruction, alteration, and fragmentation of their natural habitats as the main causes, but also the deadly amphibian disease chytridiomycosis caused by the fungal pathogen Batrachochytrium dendrobatidis (Bd). In Panama and Costa Rica, this Emerging Infectious Disease (EID) spread in a wave-like manner from west to east causing mass die-offs and reduced amphibian diversity even in well-preserved habitats. The disease has primarily affected stream-associated highland species. The last large-scale evaluation of the conservation status of Panama´s amphibians through the IUCN Red List of Threatened Species in 2004 concluded that approximately 30% of the known species are acutely threatened with extinction. Furthermore, around 17% of the amphibian species that have been known back then lacked adequate data to be assessed. In view of Panama´s already overwhelming amphibian diversity, as well as the variety of habitats and the large number of sites that have not been examined with regard to amphibians before, I started this study with the conviction that the inventory of Panama´s amphibian diversity is far from being completed. Furthermore, when I started this study, it was uncertain if there would be any surviving amphibian species in areas where chytridiomycosis had emerged. The loss of whole amphibian communities in upland western Panama following Bd arrival led to a shift of amphibian research to lowland sites in central and eastern Panama aiming primarily on pathogen arrival and the documentation of epizootic outbreak and subsequent population decline. The situation of amphibian communities in areas post-decline was therefore largely unknown. Accordingly, the main goals of my study were to add to the taxonomic inventory of amphibians in Panama and to assess the situation of amphibian populations in habitats where chytrid-driven declines have been observed. To address these tasks I conducted fieldwork in western Panama with a focus on mountainous elevations between 1000 and 3475 m asl. Additionally, I visited different lowland sites between sea level and 1000 m asl to collect comparative material. In the period between 2008 and 2013, I conducted five collection trips to Panama that add up to a total of approximately 13 months in the field. I have sampled nine regions in western Panama and collected 767 specimens together with student collaborators, 531 of which were collected under my personal field number. Additional data obtained from those specimens include 68 male anuran call recordings, 102 standardized color descriptions of specimens in life, and 259 tissue samples that to date yielded 185 16S mtDNA sequences. This comprises the most comprehensive data set for amphibians of Panama and the first large-scale DNA barcoding approach for western Panama to date. After a preliminary DNA barcoding and subsequent comparative examination of morphological und bioacoustic data of all specimens collected, the number of taxonomic problems that needed to be addressed was higher than I previously anticipated. For most genetic lineages deeper taxonomic analyses were required to reach conclusive results. A selection had to be made with which lineages to proceed in the analyses, in view of the substantial financial and time expenditure that would be needed for a complete taxonomic revision. Therefore, I chose to run deeper analyses on one genus from each of the three amphibian orders in Panama. The genera selection depended largely on the availability of sufficient material and the scientific relevance of the respective genus. I selected the genus Diasporus from the order Anura. These small frogs are omnipresent in many habitats and thus relatively easy to find. In addition, the genus is underrepresented in taxonomic studies. This is the first taxonomic study on the genus Diasporus to include a molecular phylogeny and the first comparison of advertisement calls between several populations from western Panama. In total, I collected 67 Diasporus specimens throughout western Panama and compared them morphologically with 49 additional specimens from Central America in collections, including the primary types of D. diasporus and D. hylaeformis. Additional comparative data were taken from literature. The DNA barcoding analysis of a fragment of the 16S rRNA gene included 43 own sequences that were complemented with 15 relevant GenBank sequences. In addition, I compared the advertisement calls of 26 male individuals among each other and with call descriptions from the literature. The DNA barcoding approach revealed several unnamed genetic lineages, but in some cases also resulted in the lumping of morphologically and bioacoustically distinct specimens. Generally, the morphological examination of the collected material revealed almost no specific characters that could be used to distinguish between genetic lineages. However, it was possible to identify species using a combination of several morphological characteristics. Which ones are relevant in the individual case depends on the respective species. My extensive collection of call recordings made it possible to test for the first time the intraspecific call variation of D. hylaeformis in dependency of various parameters. This analysis showed that the dominant frequency depends significantly on the body size of the calling male; the smaller the calling male, the higher the frequency of the call. A similar relationship was observed between the call rate and temperature: the lower the temperature during calling, the lower the call rate. I suppose that these general patterns, which have already been observed in other anuran genera, are also true in other Diasporus species that could not be tested in this study. Taking into account the intraspecific variation of Diasporus advertisement calls, I consider comparative call analyses to be the best way to distinguish between species. This is especially true in syntopic species. Integration of the three lines of evidence (i.e., morphology, DNA barcoding, and bioacoustics) led to the identification of four new species, two of which (i.e., D. citrinobapheus and D. igneus) colleagues and I have already formally described. I conducted an integrative taxonomic analysis of the western Panamanian representatives of the genus Bolitoglossa from the order Caudata, the larger of the two Panamanian salamander genera. Bolitoglossa is very species-rich with a centre of diversification in the high mountains of Costa Rica and western Panama. I collected 53 Bolitoglossa specimens and compared them to twelve specimens in collection, including the holotype and one paratype of B. gomezi. The dataset was complemented with information from the literature. Among the sampled specimens were two species considered to be endangered that have not been collected or observed for several decades; B. magnifica has not been seen for 34 years and B. anthracina has not been seen for 22 years. Further, I collected salamanders at several new locations. To date, my 16S mtDNA barcoding analysis represents the densest taxon sampling for Panamanian Bolitoglossa composed of 21 own sequences that were combined in the final alignment with 47 GenBank sequences. Even though the molecular phylogeny is based only on a single marker, the received trees largely coincide with previous studies and the nodes received high statistical support. In these trees, I retrieve all previously defined subgenera and species groups. On the basis of this molecular phylogeny, I placed B. anthracina, here sequenced for the first time, in the B. subpalmata species group. Due to the fact that B. anthracina is a large and dark colored species it had previously been placed by implication in the B. schizodactyla species group along with other large black salamanders of the B. nigrescens species complex. Moreover, I found deep divergent genetic lineages among geographically separated populations of B. minutula. However, until now there were no additional morphological characteristics detectable to distinguish between these lineages. Additionally, my colleagues and I described a new deep divergent lineage in the B. robinsoni species group as B. jugivagans, a species new to science. In contrast, I found only minor genetic differences between specimens of B. sombra and B. nigrescens. After combining morphometric data and tooth counts from literature of both species with additional data from specimens of B. sombra that I collected near the type locality, the distinguishing features blurred. In particular, including much larger specimens of B. sombra, not yet known at the time of its description, showed that the tooth count difference is dependent on the size and age of the specimen examined. Larger specimens have more maxillary and vomerine teeth. Based on this evidence I regard B. sombra as a junior synonym of B. nigrescens. Further, I revised the Panamanian distribution of the two relatively common lowland salamanders, B. colonnea and B. lignicolor. Besides filling the gaps in the fragmentary known distributions of these species, I assessed the molecular and morphological variation of both species among populations in Panama. While there was little variation in B. lignicolor, I found divergent genetic lineages among geographically distinct populations of B. colonnea that require further taxonomic examination. Caecilians (order Gymnophiona) are among the least investigated terrestrial vertebrates. After I received a first specimen of the predominantly South American genus Oscaecilia (family Caeciliidae) in western Panama, I started to work more extensively on the taxonomy of Caeciliidae in Central America. The specimens from western Panama were not readily assignable to a single described species, but shared characters with O. elongata and O. osae. While O. osae was only known from the holotype, the type material of O. elongata was destroyed during World War II. On the basis of the original description, the unique feature in O. elongata within Oscaecilia is the absence of subdermal scales in the posterior part of the body. In a referred specimen of O. elongata mentioned in the original description from eastern Panama, this characteristic cannot be examined as it consists of head and neck only. Therefore, I used non-destructive high-resolution, synchrotron-based X-ray micro CT imaging (HRμCT) to examine cranial characters in the specimens in question and took normal radiographs to count vertebrae and to make subdermal scales visible. I found that the fragmented specimen from eastern Panama likely belongs to the well-sampled species O. ochrocephala and has not much in common with O. osae or the specimens from western Panama. Contrarily, O. osae and the specimens from western Panama share many morphological characters, but also show some differences. Genetic barcoding revealed that both species are close relatives, but the genetic distance could not be finally resolved, because 16S sequences obtained from blood samples of living O. osae were of poor quality. Thus, I compare the Oscaecilia from western Panama to O. osae in this study, but postpone a taxonomic decision until further material becomes available. Further, I designate O. elongata a nomen dubium, because the type material is lost, the type locality is not defined in more detail than “Panama”, and the original description does not allow for a definite assignment. Since previous molecular studies only considered O. ochrocephala, the monophyly of Oscaecilia was never tested before. So far, the genus Oscaecilia is based largely on a single cranial character, the eyes covered with bone. Here, I combined two 16S mtDNA sequences of O. osae from Costa Rica and two sequences from O. sp. from western Panama with two sequences of O. ochrocephala and ten sequences of four species of the genus Caecilia, the sister genus of Oscaecilia. The resulted phylogeny contains two well-supported clades, one clade containing two species of Caecilia, one from Panama and one from western Ecuador and all species of Oscaecilia tested. The other clade consists of two species of Caecilia from the Amazon basin. I therefore assume that the split in both clades is due to the rise of the Andes, what led to today’s cis-trans-Andean distribution of the two clades. For now, to restore monophyly, I suggest to place Oscaecilia within the synonymy of Caecilia until more taxa have been tested. When assessing the conservation status of the amphibian species in mountainous western Panama, I first compiled a list of known species that I potentially could have found during my fieldwork. Using the IUCN categories, I analyzed how many of the endangered species I actually found and how these are distributed over families and species groups. Surprisingly, my rediscoveries of lost species were not equally distributed among the four families that comprise most endangered amphibian species (i.e., Bufonidae, Craugastoridae, Hylidae, and Plethodontidae). While I discovered ten of eleven endangered hylids and six of nine endangered plethodontids, I found only one of four endangered bufonids and none of the nine endangered craugastorids. I assume that the secretive living plethodontids, for which no Bd related declines have been documented, were just overlooked in the past decades. In contrast, I propose that hylids, in which Bd related population decline is well documented, developed distinct evolutionary solutions permitting coexistence with the pathogen. The situation is obviously different in bufonids and craugastorids, where I found no signs of population recoveries at present. So far, the only surviving populations of species from these families exist in climatic or physiographic niches that have probably shielded them from Bd. My data confirm the current view that the risk for naïve amphibian populations to decline during Bd epizootics is predicted by ecological traits (e.g., aquatic index, vertical distribution) and not dependent on taxonomic affiliation. However, I propose that only certain amphibian families (e.g., hylids and centrolenids) have the ability to acquire immunity solutions to coexist with the pathogen during enzootic stages. This is a very new perspective on the worst infectious disease in amphibians worldwide, allowing for new research approaches to understand the host-pathogen dynamics. Moreover, I examined where the share of surviving endangered amphibian species is particularly high in mountainous western Panama. As was to be expected, most of the endangered species are found within the boundaries of protected areas. One exception is the unprotected Cerro Colorado region in the Comarca Ngöbe-Buglé that provides habitat for a wide variety of endangered and undiscovered amphibian species. Nonetheless, planned open pit mining would destroy the forests in a large part of the area. This demonstrates once again that human activities are the biggest threat to amphibians in Panama and elsewhere.

Ecnomiohyla veraguensis sp. nov. Ecnomiohyla rabborum – Köhler 2011: p. 224 Fig. 537; p. 226 Fig. 541 b. Holotype. SMF 89877 (original field number AH 210) an adult male (Figs. 7–8) collected near Cerro Negro (8.5533 °N, - 81.09261 °W, 540 m a.s.l.), Santa Fé National Park, Veraguas, Panama, on 31 March 2009 at 12:00 hrs, collected by Smelin Abrego, Arcadio Carrizo, Andreas Hertz, and Sebastian Lotzkat. Diagnosis. A medium-sized species of Ecnomiohyla. The single known specimen is an adult male, 57.8 mm in SVL) differing from other known species of the genus by following combination of characters: 1) finger webbing extensive, web touching the finger disk on at least one side on Fingers II–IV; 2) toes extensively webbed; web reaching the toe disk at least on one side on four toes; 3) skin on dorsum finely tuberculate with scattered minute keratin tipped tubercles posteriorly; 4) cranial and dorsal osteoderms present; 5) skin on upper surface of head not co-ossified with underlying cranial elements; 6) humerus without enlarged crista lateralis; 7) prepollex distinct, recurved, with distinct bony prepollical projection, spadelike and directed laterally; 8) 6–8 widely spaced, keratinized black spines present bordering the outer side of the thumb; 9) a distinct scalloped fringe without pointed tubercles on its ventral surface, arising at the heel and continuing on the outer side of Toe V and reaching almost to the disk of Toe V (Fig. 8); 10) dorsal coloration in life smoke gray, with upper surface of forearms bearing a suggestion of lime green (Fig. 7). Comparison with other species of Ecnomiohyla. Ecnomiohyla veraguensis can be distinguished from other species of Ecnomiohyla by the following characters (with contrasting features for E. veraguensis in parentheses; see Table 3 for more details): E. echinata, E. minera, E. rabborum, E. salvaje, E. thysanota, and E. valancifer can be distinguished from the new species by the lack of cranial and dorsal osteoderms (both present); E. rabborum and E. minera are further distinct in having a humeral projection in males (no humeral projection); E. rabborum has substantial finger webbing, web reaching base of disk on one finger (extensive webbing, web touching the finger disk on at least one side on Fingers II–IV); E. fimbrimembra (Fig. 7 E-F), E. miliaria and E. phantasmagoria lack scalloped fleshy fringes on heels and have pointed heel tubercles instead (scalloped fleshy fringes present, no heel tubercles); E. fimbrimembra and E. salvaje have the skin on the head co-ossified with the cranium, (skin not coossified with cranium); males of E. miliaria and E. phantasmagoria have a sharp prepollical spine protruding from the prepollex (prepollex recurved, no protruding spine); E. bailarina has a strongly tuberculate dorsum (finely tuberculate) and two clusters of numerous, small nuptial spines at the distal end of the prepollex and the base of the pollex (only 6–8 larger, widely spaced nuptial spines along the outer side of the pollex; Fig. 10); E. tuberculosa lacks an enlarged prepollical bony projection or keratinized black spines on the prepollex in adult males (enlarged prepollical bony projection and 6–8 widely spaced, keratinized black spines presents); E. miotympanum lacks of scalloped dermal fringes on the outer margin of the forearm and foot, large digital disks, and enlarged prepollices (present in E. veraguensis); in terms of general appearance, E. sukia is most similar to the new species, but differs by a genetic distance in the 16 S gene of 7 % and the lack of nuptial spines in adult males (6–8 widely spaced nuptial spines on the outer side of the pollex; see Fig. 9 (B, D) and 10); further, E. sukia lacks keratin tipped tubercles on the dorsum (presence of keratin tipped tubercles on the dorsum). Description of the holotype. An adult male, as determinated by the presence of nuptial spines, and vocal slits. Measurements of the holotype are indicated in Table 4. Head rounded in dorsal view, wider than long (HL/HW= 85.6 %); snout truncate in dorsal and lateral views; nostrils directed laterally; top of head flat; canthus rostralis concave; loreal region concave; a well-developed supratympanic fold running from above the upper margin of tympanum, slightly curved around its upper posterior edge; tympanum prominent, smooth, same color as dorsum, and 60.0% of ED; separated from eye by 3.04 mm; upper surface of body finely tuberculated, scattered tubercles present on dorsal surfaces of the limbs; arms robust, hypertrophied; a fleshy scalloped fringe extends from the elbow along the ventrolateral margin of forearm and continues along the outer edge of Finger IV to base of disk; scallops of fringe largest on forearm, weak scallops along finger; hands moderate in length (HAL /SVL= 36.0%); finger lengths II 1 3 / 4 – 2 II 3 / 4 – 1 1 / 4 III 1 1 / 4 – 3 / 4 IV; legs relatively long and slender (FL/ SVL= 48.4 %), heels of adpressed limbs overlapping about 1 / 3 length of tibia, thigh 26.70 mm long; distinct fleshy, scalloped fringe begins on heel and extends along ventrolateral margin of tarsus and outer margin of Toe V to base of disk; scallops sinuously serrated, widest on tarsus, smaller along toe; tarsal fold slightly evident; outer metatarsal tubercle barely distinct, inner metatarsal tubercle large (1.12 times 3 TD), ovoid, slightly elevated, and spadelike distally; toe lengths IVI 3 / 4 – 1 1 / 2 II 3 / 4 – 1 1 / 4 III 3 / 4 – 3 / 4 IV 1 / 4 – 3 / 4 V; gular area and venter granulate, fine granulation on undersides of arms and proximal thighs, smooth skin on anterior surfaces of thighs and ventral parts of legs; cloacal opening directed posteriorly at mid-level of thighs, a distinct granular dermal fold under the vent. Tongue slightly cordiform, broader at the base; vomerine ridges large and transverse, well separated medially, placed between the posterior margins of choanae; vomerine teeth 10–14; paired vocal slits extending posteriorly from posterior lateral base of tongue toward angle of jaws. Coloration of holotype in life (Fig. 7). Coloration in life was recorded at daytime: Dorsal ground color Smoke Gray (44); snout, canthus rostralis, and supraorbital regions Brownish Olive (29) suffused with Olive Green (Auxiliary 47); upper surfaces of forearms with a suggestion of Lime Green (59); dorsal surfaces of finger webbing like dorsal coloration on body, but toe webbing Vandyke Brown (121); ventral surfaces of chin and body Cream Color (54), spotted with Raw Sienna (136); ventral surfaces of hindlimbs True Cinnamon (139); ventral coloration of toe and finger webbings Vandyke Brown (121). Coloration in preservative (Fig. 8). Dorsal surfaces Grayish Horn Color (268); snout, canthus rostralis, and supraorbital regions Medium Plumbeus (294); darker bars on upper surfaces of limbs Medium Plumbeus (294); cloacal region Pale Buff (1), suffused with Medium Plumbeus (294); groin and posterior surfaces of thighs mottled with Maroon (39) on a Pale Buff (1) ground; ventral regions Pale Buff (1); chin suffused with Maroon (39); toe and finger webbing Burnt Umber (48). Distribution and natural history. Ecnomiohyla veraguensis is known only from the type locality in the Isthmian-Pacific moist forests (Fund & Hogan 2012). The holotype was found at noon on a sunny day at the end of the dry season. Relative air humidity at the moment of encounter was 68 % at a temperature of 21.8 °C and it was slightly windy. The frog was sitting in a water conserving posture (Fig. 7 D) on a fern leaf approximately 0.5 m above the ground, next to a water tube that is used by local people to obtain drinking water. Other amphibian species that were observed at Cerro Negro on this expedition conducted between March 31 and April 0 3 2009 include Atelopus varius (Lichtenstein & Martens, 1856), Bolitoglossa colonnea (Dunn, 1924), Craugastor gollmeri (Peters, 1863), C. megacephalus (Cope, 1875 “ 1876 “), Diasporus citrinobapheus Hertz, Hauenschild, Lotzkat & Köhler, 2012, Lithobates warszewitschii (Schmidt, 1857), Pristimantis caryophyllaceus (Barbour, 1928), P. cerasinus (Cope, 1875 “ 1876 “), P. cruentus, P. museosus (Ibáñez, Jaramillo & Arosemena, 1994), P. pardalis (Barbour, 1928), Rhaebo haematiticus Cope, 1862, and Sachatamia albomaculata (Taylor, 1949). Since E. veraguensis is only known from a single specimen from a single locality, the distribution is unknown. It is expected to occur along mid-elevations of the Serranía de Tabasará. Etymology. The species name is derived from the province name Veraguas where the holotype was found, with the Latin suffix - ensis donating a place or locality. The species name has been chosen to accentuate the particular role the province of Veraguas plays in terms of amphibian conservation. It is the only Panamanian province with Atlantic and Pacific coasts, thus encompassing a great variety of habitats for many amphibian species. Conservation status. As other Ecnomiohyla species, E. veraguensis could be considered as a rare species, due to the habitats it uses, this fact makes it difficult to assess its populations. Like E. bailarina, the data deficient (DD) criterion, according the IUCN (IUCN 2013), seems appropriate for E. veraguensis too, until data on its population trend become available.

Ecnomiohyla bailarina sp. nov. Holotype. SMF 97398 (original field number AB 297; Fig. 3–4), an adult male from the north slope of the Jingurudó mountain range (Fig. 5), about 14.6 km S from Pavarandó village (7.70903 °N, - 78.04882 °W, 750 m a.s.l.), Sambú, Comarca Emberá-Wounaan N° 2, Darién, Panama, collected by Abel Batista and Milan Vesely on 25 September 2011 at 21: 27 hrs. Diagnosis. A medium-sized Ecnomiohyla (single known specimen is an adult male 68.1 mm in SVL; Figs. 3–4), differing from other known species in the genus by the following combination of characters: 1) finger webbing extensive, web reaching the finger disk on at least one side on two fingers (Fig. 4); 2) toes extensively webbed as well, web reaching the toe disk at least on one side of four toes (Fig. 4); 3) skin on dorsum strongly tuberculate; 4) cranial and dorsal osteoderms present; 5) skin on upper surface of head not co-ossified with underlying cranial elements; 6) humerus without enlarged crista lateralis; 7) prepollex distinct, obtuse, with bony prepollical projection rounded distally, bluntly pointed at side adjacent to thumb; 8) two clusters of nuptial spines at the distal end of prepollical tubercle and at the end of the first phalanx of the thumb; 9) a distinct scalloped fringe with pointed tubercles on a ventral surface of heel flaps, continuing almost to the disc of the 5 th toe; 10) dorsal coloration in life green with scattered brownish or black flecks. Comparison with other species of Ecnomiohyla. Ecnomiohyla bailarina can be distinguished from other species of Ecnomiohyla by the following characters (with contrasting features for E. bailarina in parentheses, see Table 3 for more details): Ecnomiohyla minera, E. thysanota (see Fig. 6) and E. rabborum are easily distinguished from the new species by having smooth heels without a scalloped fringe (triangular serrate fringe with pointed tubercles on a ventral surface of heel flaps); E. rabborum and E. minera are further distinct in having a humeral projection in males (no humeral projection); E. rabborum has a substantial webbing on one finger only, reaching base of disk on one finger (webbing extensive reaching base of disk on two fingers); E. echinata, E. fimbrimembra, E. minera, E. salvaje and E. valancifer lack of cranial or dorsal osteoderms (well developed cranial and dorsal osteoderms); the type locality of the only known specimen of E. thysanota, a female collected at Cerro Malí, Darién (Duellman 1966), is only 100 km northeast of the type locality of E. bailarina, but E. thysanota lacks cranial and dorsal osteoderms (well developed cranial and dorsal osteoderms, see Figs. 3−4, and 6), skin on dorsum is granular (strongly tuberculate), coloration in life is reported to be uniformly green (green with scattered brown or blackish flecks); in addition, these potentially sympatric species would probably differ also in size, as the E. thysanota specimen is a female that is much larger (95 mm vs 68.1 mm SVL, see Table 3) than our male E. bailarina; males in Ecnomiohyla spp. tend to be bigger or at least the same size as females (Table 3, Savage & Kubicki 2010), and hence, an adult male E. thysanota is presumed to be considerably larger than the adult male holotype of E. bailarina; E. fimbrimembra (see Fig. 9), E. miliaria, and E. phantasmagoria also lack a fringe on heels (present), but have pointed heel tubercles; in addition, males of E. miliaria and E. phantasmagoria have a sharp prepollical spine directed laterally (prepollical spine vestigial, bluntly pointed and directed to the thumb); E. fimbrimembra and E. salvaje have the skin on the head co-ossified with the cranium, (Fig. 9 E–H) (skin not coossified with cranium); males of E. miliaria, E, phantasmagoria, E. sukia, E. tuberculosa and E. valancifer have no nuptial black spines on prepollex (numerous small black keratinized spines present on prepollex); E. miotympanum lacks of scalloped dermal fringes on the outer margin of the forearm and foot, large digital disks, and enlarged prepollices (present in E. bailarina); E. tuberculosa does not have a prepollical projection in adult males (prepollical projection present); in E. sukia, the prepollical spine has a similar size and direction, but is rather spade-like, not forming a sharp spine as in E. bailarina; E. veraguensis (sp. nov., see below) can be distinguished from E. bailarina by having only a few large, widely spaced nuptial black keratinized spines, dorsolaterally on the base of the pollex and none on the prepollex in adult males (thickly clustered smaller spines on prepollex and pollex; Fig. 10); further, it has a finely tuberculated dorsum (strongly tuberculated dorsum), and keratinized tubercles on the ventral side of the scalloped fringe on the heels are absent (present in E. bailarina). Description of the holotype. An adult male, as indicated by the presence of keratinized nuptial spines. Measurements of the holotype are shown in Table 4. Head rounded in dorsal view, slightly wider than long (HL/ HW = 91.3 %); snout truncate in dorsal and lateral views; nearly terminal nostrils directed laterally; top of head flat; canthus rostralis concave; loreal region concave; skin on dorsal surface of head and body tuberculate, tubercles formed by osteoderms; tubercles on upper lip, loreal and supraorbital area tipped with tiny blunt keratinous spines; lower eyelid with transparent upper part; a well-developed supratympanic fold running from midpoint of posterior margin of eye above the upper margin of tympanum, slightly curved around its upper posterior edge, tympanum prominent, opaque, smooth, 51.5 % of ED, separated from eye by 3.20 mm; upper surfaces of body and limbs tuberculate, intermixed with scattered larger tubercles, cluster of tubercles above the insertion of arms; a triangular serrate-like fringe extends from the elbow along the ventrolateral margin of the forearm and continues along the outer edge of Finger IV to the base of the disk; serrate fringe largest on forearm, less evident serration along fingers; hands moderate in length (HAL /SVL = 31.1 %); Finger lengths II 1 3 / 4 – 2 II 3 / 4 – 1 1 / 2 III 1 1 / 2 – 1 1 / 4 IV; legs relatively long and slender (TL/SVL = 52.6 %), heels of adpressed limbs overlapping about 1 / 4 length of tibia, thigh 30.00 mm long; distinct fleshy, triangular serrate like fringe begins on heel by a striking flap and extends along ventrolateral margin of tarsus and outer margin of Toe V to base of disk; scallops deeply incised and pointed, largest on tarsus, smaller along toe; small tubercles with keratinized tips present on dorsal and ventral surface of fringe on heel; tarsal fold and outer metatarsal tubercle absent, inner metatarsal tubercle moderately large (same size as 3 TD), ovoid, flat, and spadelike distally; toe lengths II 3 / 4 – 1 1 / 4 II 3 / 4 – 1 1 / 4 III 3 / 4 – 1 IV 1 1 / 4 –¾ V; gular area and venter strongly granulate, fine granulation on undersides of arms and proximal thighs, smooth skin on anterior surfaces of thighs and ventral parts of legs; cloacal opening directed posteriorly at mid-level of thighs, two distinct granular dermal folds under the vent; tongue slightly cordiform; vomerine ridges transverse, narrowly separated medially, placed between the posterior margins of the moderately large ovoid choanae; vomerine teeth 12–13; vocal slits not present. Coloration of holotype in life (Fig. 3). Dorsal ground colour Light Grass Green (color 109 of Köhler 2012) with irregular Vandyke Brown (281) flecks scattered all over the head and body giving the animal a “moss cryptic” appearance; Raw Umber (22) bands present on dorsal surfaces of arms and legs, edges of scalloped fringes on arms and fleshy flaps on heels Cream Color (12); toe webbing Tawny Olive (17); tops of some dorsal granules and tubercles Orange-Rufous (56). After metachrosis (day and night coloration), ground coloration faded to Pale Emerald Green (141), brown areas to Dark Salmon color (59), pattern did not change; throat, chest, venter and ventral surfaces of arms and legs Cream Color (12) grading into Salmon (83) ventrolaterally and Orange Yellow (8) on anterior surface of thigh; a few small dark blotches on the edge of lower lip; iris Light Yellow Ocher (13), finely reticulated with Dark Brownish Olive (127); tympanum Pale Mauve (204) with scattered irregular Vinaceous Pink (245) blotches. Coloration in preservative (Fig. 4). Dorsal surfaces Glaucous (272) with Sepia (279) mottling on upper surfaces of hind limbs; tympanum Pratt’s Payne’s Gray (293) with scattered irregular Maroon (39) blotches, cloacal region Pratt’s Payne’s Gray (293) dorsally and Cream (12) ventrally; posterior surfaces of thighs Light Yellow Ocher (13); ventral surfaces of body and limbs Cream (12); toe webbing Amber (51). Distribution and natural history. Ecnomiohyla bailarina is known only from the type locality, in the eastern Panamanian montane forest (Fund & Hogan 2012; Fig. 5 A-D). The potential area of distribution of E. bailarina comprises the vicinities of Jingurudó and Sapo mountain ranges, between 400 to 1400 m a.s.l. (Fig. 1). Although the type locality is in a primary forest, there are some open areas with successional secondary forest. The area is on a ridge, so the trees could be affected by strong winds. In the surroundings we saw four fallen large trees probably overthrown by the wind that left clearings in the otherwise pristine forest. The largest trees in this area reached more than 20 m in height having branches in the canopy covered by bromeliads and other epiphytes (e. g., orchids and Lorantaceae), Tree trunks were almost bare or with just a little epiphytic growth. In the understory, palms and vines were predominant. The holotype was found on a ridge in a water conserving posture (see Fig 1 B in Pough et al. 1983) on the bark of a small tree (Fig. 5 A), approximately 1.5 m above the ground. The day before the night of the capture was dry except for a drizzle that had fallen in the afternoon between 14:00– 15:00 hrs. During the encounter, a slight breeze was blowing. Other amphibian species observed in the area that day were: Colostethus aff. pratti (Boulenger, 1899), Craugastor opimus (Savage and Myers, 2002), Pristimantis cruentus (Peters, 1873), P. taeniatus (Boulenger, 1912), Rhinella alata (Thominot, 1884), and Sachatamia ilex (Savage, 1967). Etymology. The name bailarina is a noun in apposition in reference to the hill where the specimen was found. The indigenous people of the Embera call it “Cerro Bailarín”, in addition, the English translation of “ bailarina ” is ballerina, so the name also refers to the resemblance of the fringes on arms and feet of the frog to the tutu skirt that a ballerina wears. Conservation status. The secretive habits of Ecnomiohyla bailarina make the assessment of the population size difficult, as in other Ecnomiohyla species. Considering that the status of the E. bailarina population is unknown, the data deficient (DD) criterion, according the IUCN (IUCN 2013), seems appropriate for this species, until data on its population trend become available. Moreover, due to fact that E. bailarina and E. thysanota occur in a region affected by social problems and political conflicts along the border between Panama and Colombia, it is unlikely that there will be sufficient opportunity to visit the region to assess population sizes in the near future.

Forest canopy-dwelling frogs are usually among the rarest anuran species observed in the neotropical forest, mainly because they fall outside of the scope of the standard search methods used by herpetologists. During field explorations undertaken in western and eastern Panama in recent years, we discovered two species belonging to the genus Ecnomiohyla, which showed significant differences in genetic distances (16S mtDNA gene) and morphological characteristics different from any known Ecnomiohyla species. The first specimen originates from the Serranía de Jingurudó, Darién province, southeastern Panamá, and is described herein as E. bailarina sp. nov., and the second specimen was found at Santa Fe National Park, Veraguas province, central-western Panama, and is described as E. veraguensis sp. nov. We provide a detailed description of both new species, including comparisons of morphological and molecular characters of almost all members of the genus in lower Central America, as well as an identification key for the entire genus. Forest canopy-dwelling frogs are usually among the rarest anuran species observed in the neotropical forest, mainly because they fall outside of the scope of the standard search methods used by herpetologists. During field explorations undertaken in western and eastern Panama in recent years, we discovered two species belonging to the genus Ecnomiohyla, which showed significant differences in genetic distances (16S mtDNA gene) and morphological characteristics different from any known Ecnomiohyla species. The first specimen originates from the Serranía de Jingurudó, Darién province, southeastern Panamá, and is described herein as E. bailarina sp. nov., and the second specimen was found at Santa Fe National Park, Veraguas province, central-western Panama, and is described as E. veraguensis sp. nov. We provide a detailed description of both new species, including comparisons of morphological and molecular characters of almost all members of the genus in lower Central America, as well as an identification key for the entire genus.

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

Bolitoglossa jugivagans sp. nov. Continental Divide Salamander Figures 4–7 Holotype. SMF 94467 (original field number AH 314), an adult female from the headwaters of Río Chiriquí Malí, approximately 5 km N of the La Fortuna dam reservoir (8.7890 °N, 82.2154 °W, 1060 m), Bosque Protector Palo Seco, Comarca Ngöbe-Buglé (formerly province of Bocas del Toro), Panama (Fig. 1); collected by Andreas Hertz and Sebastian Lotzkat on 28 October 2009. Diagnosis. We assign the new species to Bolitoglossa because it has fewer than 14 costal grooves and lacks a sublingual fold (Savage 2002), and to the subgenus Eladinea based on mtDNA sequence data. It differs from all described species by the following combination of characters: Bolitoglossa jugivagans is a small species with short legs and a long tail. The hands and feet are moderately webbed, with the two terminal phalanges of the longest digits free of webbing. The prehensile tail is considerably longer than the body. Both maxillary and vomerine tooth counts are comparatively high in relation to SVL. Bolitoglossa jugivagans is well differentiated from all described species of the subgenus Eladinea by 16 S mtDNA distances (Fig. 2; Table 1). It is further distinguished morphologically by the following characters (condition for B. jugivagans in parentheses): It differs from B. biseriata Tanner, B. bramei Wake, Savage, and Hanken, B. colonnea (Dunn), B. lignicolor, B. minutula Wake, Brame, and Duellman, B. schizodactyla Wake and Brame, and B. epimela Wake and Brame by its less extensively webbed hands and feet. All these species have fully webbed hands and feet, or at least less than two phalanges free of webbing (slightly more than two phalanges free of webbing). Bolitoglossa jugivagans is easily distinguishable by coloration (described below in coloration section) from the large, almost entirely black species B. anthracina Brame, Savage, Wake, and Hanken, B. compacta Wake, Brame, and Duellman, B. copia Wake, Hanken, and Ibañez, B. magnifica Hanken, Wake, and Savage, B. nigrescens (Taylor), B. robusta (Cope), and B. sombra Hanken, Wake, and Savage. It can be differentiated from other small salamanders with high tooth counts from the Talamancan Mountains as follows: Bolitoglossa pygmaea Bolaños and Wake is almost unpigmented and has a short tail that never exceeds SVL (B. jugivagans has a normal reddishbrown pigmentation and the tail is much longer than SVL). Bolitoglossa gracilis Bolaños and Robinson is bright yellowish to golden tan with dark streaks and spots (reddish-brown with light streaks), has longer legs with a limb interval of 3 to 3.5 (4) and fewer vomerine teeth VT: 10–22 (27; see also Fig. 3). Bolitoglossa pesrubra Taylor, with which the new species shares high vomerine and maxillary tooth counts, has longer legs with a limb interval of 0 to 3 folds (4) and a shorter tail TL/SVL≈ 1 (TL/SVL= 1.24). Bolitoglossa subpalmata (Boulenger) has fewer maxillary and vomerine teeth at the same size (Fig. 3) and a limb interval of 1 to 3 (4). Bolitoglossa gomezi Wake, Savage, and Hanken has a shorter tail TL/SVL: 1.15–1.19 in females (1.24), limb interval of 1.5–3 (4), and fewer maxillary and vomerine teeth at a comparable SVL (Fig. 3). Bolitoglossa kamuk Boza-Oviedo, Rovito, Chaves, García-Rodríguez, Artavia, Bolaños, and Wake has a dark ground coloration with golden flecking on the posterior part of the body and the tail (reddish-brown with light streaks), as well as fewer maxillary and vomerine teeth MT: 34 (51), VT: 13–16 (27), and a shorter tail TL/SVL≈ 1 (TL/SVL= 1.24). Bolitoglossa splendida Boza-Oviedo, Rovito, Chaves, García-Rodríguez, Artavia, Bolaños, and Wake is peculiarly colored with a bright-red dorsal band and bright-yellow spots laterally and ventrolaterally on jet black ground coloration (reddish-brown with light streaks), has only 18 vomerine teeth at a SVL of 47.8 mm (27 at SVL= 31.2 mm), has more webbing on feet that extends to between first and second phalangeal articulations of longest digits (two terminal phalanges free on the longest digits), and a broader head HW/SVL= 0.17 (HW/SVL= 0.14). Based on molecular data, Bolitoglossa jugivagans is most closely related to B. robinsoni and B. aureogularis. Both differ from B. jugivagans by the following morphological characters (condition for B. jugivagans in parentheses): Bolitoglossa robinsoni is a large and stout species with a broad head HW/SVL= 0.16–0.17 (0.14) and a tail that is shorter than the standard length TL/SVL≤ 1 (TL/SVL= 1.24). The second species B. aureogularis is superficially very similar to B. jugivagans, primarily in dorsal coloration. However, B. aureogularis has fewer maxillary and vomerine teeth at a comparable size (Fig. 3) and much shorter legs with a limb interval of 6.5 in females and 5 in males (4 in the female holotype of B. jugivagans). Bolitoglossa aureogularis differs most markedly from B. jugivagans in ventral coloration, having a bright yellow gular region and a prominent midventral dark stripe on dirty white ground (gular region mustard yellow; ventral coloration of body grey to brownish speckled with blue, no conspicuous dark midvental stripe). Description of the holotype. This is a small and slender Bolitoglossa. The holotype is an adult female of 31.2 mm SVL; tail long and prehensile, TL/SVL is 1.24; head short (HL about 21 % of SVL) and narrow (HW slightly less than 14 % of SVL); hands and feet narrow (HAW = 7 %, HFW = 9 % of SVL), and limbs short (HLL about 20 % of SVL, limb interval 4 costal folds between adpressed limbs); webbing moderate with the two terminal phalanges free on the longest digits (Fig. 4); order of increasing length of digits I-IV-II-III in hands, and I-V-II-IV- III in feet; subterminal pads present on longest digits; small eyes that do only slightly protrude beyond lateral margins of head in ventral view; labial protuberances weakly developed; teeth moderate in size and numerous (PMT 4, MT 51, VT 27), maxillary teeth extending to about level of center of eye; prevomer clearly projecting from level of palate, bearing vomerine teeth in long, slightly arched, irregular series extending laterally well beyond outer margin of choanae. Measurements (in mm), tooth counts and limb interval of the holotype. SVL 31.2, TL 38.8, HW 4.3, SG 6.5, HD 2.5, EW 0.7, EL 1.5, ES 1.8, ED 1.3, IC 2.2, IO 1.6, SF 8.4, IN 1.0 SP 0.6, SW 3.6, SAV 28.7, AX 17.8, LI 4, FLL 4.8, HLL 6.2, HAW 2.2, FW 2.8, T 3 1.3, T 5 1.0, Numbers of teeth: PMT 4, MT 23 / 28, VT 12 / 15. Coloration of the holotype in life (Fig. 5). Dorsal ground color Chestnut (32), interspersed with Fuscous (21) and Salmon Color (106) broken longitudinal lines. A Salmon Color (106) dorsolateral line running from eye to groin separates the middorsal coloration from the lateral and ventral ground coloration; ventral ground coloration Sulfur Yellow (157) flecked with gray at night and Fuscous (21) at daytime. Ventral surfaces speckled with Sky Blue (66). Gular region, hands, and feet Sulfur Yellow (157) suffused with Fuscous (21) on limbs. Coloration of the holotype in alcohol (Fig. 6). After three years in ethanol (60 %), its coloration has not changed much. Yellowish colors have faded to cream-colored shades as well as the reddish tones in the brown dorsal coloration. Osteology (Fig. 7). The radiographs of the holotype show the typical osteological features of the genus Bolitoglossa: Head well ossified; vertebral column consists of one atlas, 14 trunk vertebrae, one sacral and two caudosacral vertebrae, and 35 caudal vertebrae; digits well developed on all limbs; slightly more than two distal phalanges of the longest fingers and toes free of webbing, phalangeal formulae of the hand 1–2 – 3 – 2, and the foot 1–2 – 3 – 3 – 2. Terminal phalanges of longer fingers and toes slightly expanded distally (Fig. 4). Habitat and natural history notes. Bolitoglossa jugivagans is only known from the type locality, the upper section of the valley of a feeder stream of Río Chiriquí Malí, on the Caribbean slope of the Cordillera Central, not far from the continental divide in the Fortuna depression of the Cordillera Central of western Panama (Fig. 1). The site is easily reachable on a paved road that serves as a maintenance access way to the Trans-Panama Oil Pipeline. The riverbank next to the street is an old clearing, overgrown with grass, herbaceous plants, and bushes, while the opposite bank is predominantly covered by broadleaf, evergreen mature forest. Mean annual precipitation is approximately 3000 mm and mean annual temperature approximately 20.6 ° C, making the site assignable to the Premontane Wet Forest life zone according to the Holdridge (1967) classification. The holotype was encountered on October 28, 2009 at 23:00 h active on the large leaf of an Araceae plant growing out of the long grass on the cleared side of the river. That day, it had been raining in the afternoon and evening; our pluviometer registered 44.5 mm of precipitation between 16:00 and 24:00 h. The automatically recorded temperature was 19.6 ° C between 18:00 and 23:00 h, and then gradually decreased to a minimum of 17.6 ° C at 06:00 h. Relative humidity was constantly high at close to 100 %. Bolitoglossa jugivagans was the only salamander encountered at this site. Other amphibian species collected in the vicinity include Craugastor fitzingeri (Schmidt), Diasporus sp., Duellmanohyla uranochroa (Cope), Espadarana prosoblepon (Boettger), Incilius coniferus (Cope), Pristimantis cruentus (Peters), P. pardalis (Barbour), and Smilisca phaeota (Cope). Additionally this site is the type locality of the recently described snake species Sibon noalamina Lotzkat, Hertz, and Köhler. Lotzkat et al. (2012) give additional information on the reptile fauna of the surroundings. Etymology. The species name is a compound adjective derived from the locative of the Latin noun jugum meaning “mountain ridge” and the present participle of the Latin verb vagare meaning “to wander”. The species epithet signifies “wandering on the mountain ridge” referring to the type locality next to the continental divide of the Cordillera Central.

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 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

Hertz, Andreas, Lotzkat, Sebastian, Köhler, Gunther (2013): A new species of Bolitoglossa (Caudata, Plethodontidae) from the continental divide of western Panama. Zootaxa 3636 (3): 463-475, DOI: http://dx.doi.org/10.11646/zootaxa.3636.3.5

Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix, Köhler, Gunther (2013): Distribution and variation of the giant alpha anoles (Squamata: Dactyloidae) of the genus Dactyloa in the highlands of western Panama, with the description of a new species formerly referred to as D. microtus. Zootaxa 3626 (1): 1-54, DOI: http://dx.doi.org/10.11646/zootaxa.3626.1.1

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

Diasporus igneus sp. nov. Holotype. MVUP 2301 (original field number MHCH 1327), an adult male (Fig. 3���5) collected on the southeastern slope of Cerro Santiago, Serran��a de Tabasar�� about ca. 4.6 Km from Llano Tugr�� (Buabit��), capital of the Comarca Ng��be Bugl��, Distrito de M��na, Corregimiento de Pe��a Blanca, Panama, on March 25, 2009 at 19: 35 (8.5079 ��N, 81.7168 ��W, 1710 m a.s.l.). Paratypes. Three adult males from La Nevera, southwestern slope of Cerro Santiago, Serran��a de Tabasar��, Comarca Ng��be-Bugl��, Distrito de Nole Duima, Corregimiento de Jadeberi, Panama. MHCH 1388 and MHCH 2072 collected by A. Batista April 11, 2009, at 20: 42 (8.4996 ��N, 81.7716 ��W, 1700 m a.s.l.) and April 12, 2009, at 02:05 (8.4998 ��N, 81.7719 ��W, 1700 m a.s.l.), respectively, and SMF 89821 collected by A. Hertz November 11, 2009 (8.4955 ��N, 81.7672 ��W, 1820 m a.s.l.). Diagnosis. Diasporus igneus is characterized by the following combination of characters (see Table 1): (1) dorsal skin texture smooth with rounded scattered tubercles, edge of tibia and forearm with a series of low conical tubercles; (2) tympanic membrane not differentiated, tympanic annulus partially visible, 33.7 �� 4.6 % (27.8���38.5) of eye length; (3) snout rounded in dorsal view and in profile; (4) an enlarged and conical supraocular tubercle, cranial crests absent; (5) dentigerous processes of vomers oblique, behind the choanae and posteriorly separated about half of their total length from each other; (6) vocal sac well developed, vocal slits present, no nuptial pads; (7) Finger II longer than Finger I, ungual flap mostly expanded, palmate to truncate, more evident on fingers II���IV, the III finger disk is 2.0 �� 0.2 (1.8���2.2) times wider than distal end of adjacent phalanx (Fig. 5); (8) no fringes or webbing on fingers; (9) a row of rounded ulnar tubercles, most evident in life (Fig. 3 and 5), palmar tubercle rounded, flattened and slightly larger than thenar tubercle; thenar tubercle low and elongate; subarticular tubercles rounded and globular; no supernumerary tubercles, palmar and plantar accessory tubercles small and rounded; (10) heel with two or three small and rounded tubercles; a row of rounded outer tarsal tubercles; (11) no fringes or webbing on toes, ungual flap mostly expanded, more evident on toes II���V, IV disc of toe is 1.8���2.3 (2.0 �� 0.2) times wider than distal end of adjacent phalanx; (12) plantar tubercle rounded, one to three non-protuberant subarticular tubercles present (one on toes I and II, two on toes III and V, three on toe IV); inner metatarsal tubercle elongated; outer metatarsal tubercles pointed and smaller than inner; tarsal ridge absent; (13) dorsal ground color in life brown with yellow to orange reticulations, groin and axilla red, vocal sac yellow; (14) SVL 26.1 �� 0.5 mm (25.5���26.6); (15) advertisement call composed of a single, amplitude-modulated short note (50.0���100.0 ms) with harmonic structure, and with most energy emitted with the first harmonic (2.2���2.5 kHz). Similar species. Diasporus igneus can be distinguished from other species of the group as follows (see Table 1 for more details): from D. quidditus (Lynch 2001), D. tinker (Lynch 2001) and D. vocator (Taylor 1955) by its larger size and palmate to truncate ungual flap instead of lanceolate to papillate ungual flap. It differs from D. diastema (Cope 1875), D. tigrillo (Savage 1997), D. hylaeformis (Cope 1875), and D. ventrimaculatus (Chavez et al. 2009) by its larger body size, by having one to several enlarged supraocular tubercles, by having the outer edge of the tibia and forearm covered with a series of tubercles, and by having the groin and axilla colored in red. It is distinguished from D. anthrax (Lynch 2001), D. citrinobapheus (Hertz et al. 2012) and D. gularis (Boulenger 1898) by its larger size, dorsal skin with scattered tubercles, and its color pattern. Description of the holotype. An adult male, with slender body; smooth dorsal skin with low widely spaced pustules, venter coarsely areolate, discoidal fold not evident; one enlarged supraocular tubercle, edge of tibia and forearm with a series of tubercles; eye slightly longer than snout; ratio SL/ED 77 %; tympanum of moderate size, ratio TD/ED 38 %; tympanum indistinguishable, annulus tympanicus partially visible through skin anteriorly and ventrally, positioned closely behind orbit and lower jaw; head slightly wider than long, greatest head width between angles of jaw 38 % of SVL; snout rounded from above and in profile; nares situated near tip of snout and slightly dorsolaterally directed, barely visible in front view, also visible dorsally but not ventrally; canthus rostralis rounded; loreal region feebly concave; dentigerous processes of vomer 1.5 mm of length, and clearly visible, posterior to choana in an oblique outline, each with seven teeth; vocal slits present; tongue length 5.4 mm and 0.4 of width, first half attached to floor of mouth; hands moderate in size, 29 % of SVL; relative lengths of adpressed fingers IColoration of holotype in life. Holotype (MVUP 2301) (Fig. 3) recorded as follows: iris Yellow Ocher (123) with a longitudinal Army Brown (219 B) bar in the middle; dorsal ground color Verona Brown (223 B), suffused with Clay Color (123 D) and Burnt Orange (116); interorbital region with a Burnt Orange (116) line; groin and axilla immaculate Scarlet (14); posterior thigh surface Scarlet (14) suffused with Raw Umber (23), and with Buff (24) bars; venter immaculate Olive Yellow (52) suffused with Scarlet (14); vocal sac immaculate Orange Yellow (18). Coloration in preservative (including paratypes, Fig. 4 ���5 ). Dorsal ground color Buff (124), with reticulations Dark Brownish Olive (129), groin Warm Buff (118), vocal sac Trogon Yellow (153), ventral surface Chamois (123 D) pointed with Sepia (119). Measurements of holotype (mm). SVL 25.8; HL 8.2; HW 9.7; IOD 3.2; ED 3.3; TD 1.3; FL 11.6; TL 11.3; HAN 7.5; 3 FW 0.7; 3 FD 1.3; 3 TW 0.6; 3 TD 1.3; 4 TW 0.8; 4 TD 1.4; BW 13.8 (see table 2). Vocalization. The vocalizations produced by the holotype (environmental temperature = 16.9 ��C; humidity 78 %; 19: 30) and two other males (MHCH 1388, environmental temperature = 16.1 ��C; humidity 86 %; 20: 30 and MHCH 2072, environmental temperature = 15.9 ��C; humidity 93 %; 02: 10) were analyzed. The calls consist of single, short, monophasic notes that are reminiscent of a ���whistle��� (Fig. 6). Note duration is 0.07 �� 0.01 s (0.05���0.10) with an interval between calls of 13.5 �� 3.5 s (9.0��� 17.5) and a call rate of approximately four calls per minute. The peak frequency band ranges from 2.0 to 2.7 kHz. The fundamental frequency is also the dominant frequency, at 2.4 �� 0.08 kHz (2.2���2.5), followed by five main harmonic components at 4.6 �� 0.2 kHz (4.5 ���5.0), 6.8 �� 0.4 kHz (6.2���7.5), 9.3 �� 0.4 kHz (8.7���10.1), 11.8 �� 0.5 kHz (10.1���12.1), 13.6 �� 0.6 kHz (12.1���14.6)(Fig. 6), with most energy emitted in the first harmonic, followed by the second. Natural history. This species is found in the lower - montane rain forest (1699���1815 m a.s.l.) in the western portion of the Serran��a de Tabasar��. The vegetation predominantly consists of big trees (roughly 30 m in height) covered with moss, bromeliads (Werauhia spp. and Guzmania spp.), giant ferns (Cyathea spp.) and orchids. The holotype was found calling inside a hole filled with dry leaves in a branch about two meters above the ground. The specimen MHCH 1388 was found calling from inside a bromeliad around five and a half meters above ground. The specimen MHCH 2072 was found calling from a moss-covered bark on a branch approximately five meters above the ground. The specimen SMF 89821 was not calling and sat exposed on a leaf about 0.7 m above the ground. Although the holotype and SMF 89821 were found at low heights, most calling individuals were sitting more than four meters above the ground. Calling individuals are well covered and hard to locate (after searching for approximately three hours at La Nevera, A. Batista could not detect a single individual, although he heard several of them calling). One paratype (MHCH 1388) was stimulated to regurgitate the contents of its stomach (technique provided by A. Am��zquita), that contained a woodlouse (Isopoda: Oniscidea) of 2.9 mm length and 8.4 mm width. Etymology. The species epithet igneus is the Latin word for fire, and is here used as a noun in apposition. It refers to the reticulated dorsum in contrast to the red coloration in the axilla and groin that is evocative of fire. Remarks. All in all, the calls of most species of Diasporus appear to be very similar to one another to the human ear, and thus it is difficult to distinguish among them. Subjective description of the call types of the species D. anthrax, D. gularis, D. quidditus, D. tigrillo, and D. tinker have been mentioned (Chavez et al. 2009; Ib����ez et al. 1999; Lynch 2001; Savage 2002), but there are no data available for the acoustic properties of these species. Diasporus vocator from the western Pacific side of Panama (R��o Chiriqu�� Viejo, 200 m a.s.l) calls at a highest frequency between 4.6 to 4.9 kHz (A. Batista, unpublished data), very different from that of D. igneus, which ranges from 2.0 to 2.7 kHz. The call of D. diastema from central Panama ranges between 2.8 to 4.0 kHz (Fouquette 1960, Wilczynski & Brenowitz 1988), and appears to be most similar to that of D. hylaeformis, which also calls at highest frequency between 2.4 to 3.1 kHz (Chavez et al. 2009), rather than to that of D. igneus. The call of D. igneus is also different to that of D. citrinobapheus which call in bouts (8 to 22 calls per bout) at 2.9 kHz (Hertz et al. 2012). The call most similar to that of D. igneus is that of D. ventrimaculatus, which range between 2.5 to 2.6 kHz, but they differ from one another in call rate, 11 calls/min in D. ventrimaculatus (Chavez et al. 2009) and 4 calls/min in D. igneus. Moreover, most species of the genus Diasporus diastema call at low elevations, between the ground and the understory (Ib����ez et al. 1999, Lynch 2001, Savage 2002), but D. igneus call at higher elevations, from the understory to the mid-canopy. Diasporus igneus has thus far only been encountered on the western and eastern slopes of Cerro Santiago. We suspect it to be also present on the southern and northern slopes. The estimated distribution area, the slopes of Cerro Santiago above 1500 m a.s.l., comprises an area of about 40 km 2. The whole area previously has been called Cerro Colorado by Myers and Duellman (1982). This poorly investigated area comprises, besides Cerro Santiago, another prominent peak to the west, the Cerro Sagui, and the surrounding parts of the continental divide between them. The discovery of D. igneus increases the number of endemic vertebrate species in this region of the Serran��a de Tabasar�� to ten (Myers & Duellman 1982; K��hler et al. 2007; Mendelson III et al. 2008; Birdlife international 2011, Hertz et al. 2012). The potential high levels of endemism in this unprotected part of the central-Panamanian mountain range is a reason important enough to create the first protected area for the mountains of the Comarca Ng��be-Bugl��, especially when considering that between 1992 and 2000 the Comarca Ng��be-Bugl�� lost about 21.8 % of its forest, which amounts to the highest deforestation rate in Panama (ANAM 2009)., Published as part of Batista, Abel, Ponce, Marcos & Hertz, Andreas, 2012, A new species of rainfrog of the genus Diasporus (Anura: Eleutherodactylidae) from Serran��a de Tabasar��, Panama, pp. 51-60 in Zootaxa 3410 on pages 54-58, DOI: 10.5281/zenodo.215009, {"references":["Lynch, J. D. (2001) Three new rainfrogs of the Eleutherodactylus diastema group from Colombia and Panama. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales, 25, 287 - 297.","Taylor, E. H. (1955) Addition to the known herpetofauna of Costa Rica. The University of Kansas Science Bulletin, 37, 499 - 757.","Cope, E. D. (1875) On the Batrachia and Reptilia of Costa Rica. Journal of the academy of Natural sciences Philadelphia series, 2, 93 - 157.","Savage, J. M. (1997) A new species of rainfrog of the Eleutherodactylus diastema group from the Alta Talamanca region of Costa Rica. Amphibia-Reptilia, 18, 241 - 247.","Hertz, A., Hauenschild, F., Lotzkat, S. & Kohler, G. (2012) A new golden frog species of the genus Diasporus (Amphibia, Eleutherodactylidae) from the Cordillera Central, western Panama. ZooKeys, 196, 23 - 46.","Boulenger, G. A. (1898) An account of the reptiles and batrachians collected by Mr. F. H. Rosenberg in western Ecuador. Proceedings of the Zoological Society of London, 1898, 107 - 126.","Ibanez, R., Rand, A. S. & Jaramillo, C. L. (1999) Los anfibios del Monumento Natural Barro Colorado, Parque Nacional Soberania y areas adyacentes / The amphibians of Barro Colorado Nature Monument, Soberania National Park and adjacent areas. Editorial Mizrachi & Pujol, S. A., Panama, 187 pp.","Savage, J. M. (2002) The Amphibians and Reptiles of Costa Rica: A Herpetofauna between two Continents, between two Seas. University of Chicago Press, Chicago, 934 pp.","Fouquette, M. J., Jr. (1960) Call structure in frogs of the family Leptodactylidae. The Texas Journal of Science, 12, 201 - 215.","Wilczynski, W. & Brenowitz, E. A. (1988) Acoustic cues mediate inter-male spacing in a neotropical frog. Animal Behavior, 36, 1054 - 1063.","Myers, C. W. & Duellman, W. E. (1982) A new species of Hyla from Cerro Colorado, and other tree frog records and geographic notes from western Panama. American Museum Novitates, 2752, 1 - 32.","Kohler, G., Ponce, M., Sunyer, J. & Batista, A. (2007) Four new species of anoles (genus Norops) from the Serrania de Tabasara, west-central Panama (Squamata: Polychrotidae). Herpetologica, 63, 375 - 391.","Mendelson III, J. R., Savage, J. M., Griffith, E., Ross, H., Kubicki, B. & Gagliardo, R. (2008) A spectacular new gliding species of Ecnomiohyla (Anura: Hylidae) from Central Panama. Journal of Herpetology, 42, 750 - 759."]}

Key to the Lower Central American species of Sibon 1 Seventeen rows of dorsal scales at midbody and 15 rows one head length anterior to cloaca; dorsal scale rows changing orientation from the sides of body towards middorsum................................. Sibon perissostichon (Fig. 5 E) - Thirteen or 15 rows of dorsal scales at midbody, usually without reduction on posterior portion of body; dorsal scale rows oriented longitudinally................................................................................. 2 2 Thirteen dorsal rows, both at midbody and one head length anterior to cloaca (very rarely 15 rows at midbody, but then 13 both one head length posterior to head and anterior to cloaca); coral snake-like pattern of alternating light and dark rings................................................................................................ Sibon anthracops - Fifteen dorsal rows, both at midbody and one head length anterior to cloaca....................................... 3 3 Five supralabials; only one supralabial posterior to orbit, this ultimate one with a higher anterior and a lower posterior portion (Fig. 2 A); coral snake-like pattern of alternating light and dark rings (Fig. 1)......................... Sibon noalamina - Six or (in most cases) more supralabials, two supralabials posterior to orbit, with the penultimate one greatly enlarged; coloration variable, but no conspicuous coral snake-like pattern of alternating light and dark rings (for example Figs. 5 A���D) 4 4 Enlarged penultimate supralabial usually in contact with eye; dorsal ground color greenish, with an ocellated or blotched rather than a banded or striped pattern; these dark dorsal markings reach at most to outer edges of ventrals, although dark shading or mottling may be present on venter........................................................................ 5 - Enlarged penultimate supralabial usually not in contact with eye; dorsal color pattern banded or striped rather than ocellate or blotched; these dark dorsal markings reach well onto venter or even across venter.................................. 6 5 181���201 ventrals; 112���121 subcaudals; body very slender; head blunt and very distinct from neck; eyes conspicuously large and protuberant.............................................................................. Sibon argus - 147���173 ventrals; 80���106 subcaudals; body rather stout; snout acuminate, head gradually narrowing towards neck; eyes not conspicuously large, nonprotuberant................................................. Sibon longifrenis (Fig. 5 C) 6 First infralabials in contact with each other behind the mental.............................. Sibon nebulatus (Fig. 5 D) - First infralabials separated from each other by one or two postmentals behind the mental............................. 7 7 Bands dark brown ventrally; laterally, their centers are of a paler brown, which is paler than their borders or the interspaces between the bands; usually a single postmental................................................. Sibon dimidiatus - Bands dark brown ventrally and laterally, their centers not paler brown than their borders laterally, and darker than interspaces; one or two postmentals present........................................................................... 8 8 Dorsal surface of head with green or yellow as well as red and black markings; ground color of body green; 162���171 ventrals; 77���119 subcaudals........................................................................... Sibon lamari - Dorsal surface of head with pale and dark brown markings; ground color of body usually olive to pale brown; 161���193 ventrals; 103���135 subcaudals..................................................... Sibon annulatus (Figs. 5 A���B), Published as part of Lotzkat, Sebastian, Hertz, Andreas & K��hler, Gunther, 2012, A new species of Sibon (Squamata: Colubroidea: Dipsadidae) from the Cordillera Central of western Panama, with comments on other species of the genus in the area, pp. 26-40 in Zootaxa 3485 on page 37, DOI: 10.11646/zootaxa.3485.1.2, http://zenodo.org/record/208860

Sibon noalamina sp. nov. Figs. 1���3 Dipsas articulata: K��hler (2008: p. 219: Fig. 592); Stadler (2010); Carrizo (2010: in part, referring to Stadler 2010) Holotype. SMF 91539 (original field number SL 775; Figs. 1 A���C, 2���3), adult male, from headwaters of R��o Chiriqu�� Mal��, approximately 6.4 km NW Fortuna dam (8.7891 ��N, 82.2155 ��W, 1050 m), Bosque Protector Palo Seco, Comarca Ng��be-Bugl�� (formerly province of Bocas del Toro), Panama; collected by Andreas Hertz and Sebastian Lotzkat on 10 August 2010. Paratypes. SMF 90180 (original field number SL 494; Fig 1 D), juvenile male, same locality as holotype; collected by Andreas Hertz and Sebastian Lotzkat on 29 October 2009; SMF 89550 (original field number LSt 015; Fig 1 E), juvenile, from Cerro Mariposa near Alto de Piedra, approx. 3.5 km W of Santa F��, 8.5001 ��N, 81.1170 ��W, 1260 m, province of Veraguas, Panama; collected by Sebastian Lotzkat and Andreas Hertz on 28 May 2008. Diagnosis. Sibon noalamina differs from all described species of Sibon, and from all other Central American snail-eaters, in its slight keeling on the third to fifth dorsal row at midbody in adults, and, most obviously, in having only five supralabials, with the fifth and ultimate one being the only supralabial posterior to the orbit (versus two supralabials posterior to orbit in all other species of Sibon) and exhibiting a peculiar shape: Its anterior portion is almost twice as high as the remaining supralabials, resembling the enlarged penultimate supralabial of other Sibon. Then it decreases in height towards the posterior portion that is about as high as the third supralabial, resembling the ultimate, usually moderately-sized, supralabial of other Sibon. Additionally, its contrasting color pattern of complete dark rings on light background distinguishes S. noalamina from all Lower Central American Sibon except S. anthracops (Cope), from which it differs by having 15 dorsal rows throughout the body (vs. 13 in S. anthracops). At first sight, especially the more contrastingly colored juveniles of the new species might be confused with the coral snake-mimics Dipsas articulata, D. bicolor (G��nther), D. temporalis (Werner), or D. viguieri (Bocourt). From these, S. noalamina is distinguished by the presence of a mental groove (lacking in the genus Dipsas), lower ventral counts (164���177 vs. 196���217 in D. articulata, 186���199 in D. bicolor, 170���208 in D. temporalis, and 190���203 in D. viguieri), the unique supralabial condition (5 supralabials with only the ultimate, peculiarly shaped one posterior to orbit versus 9���10 supralabials in D. articulata, 10���11 in D. bicolor, 7���8 in D. temporalis, and 9���10 in D. viguieri, all shaped similarly, with three or more, rarely two, posterior to orbit), and fewer infralabials (6���7 infralabials versus 11���12 in D. articulata, 10���11 in D. bicolor, 8���13 in D. temporalis, and 9���11 in D. viguieri). Furthermore, S. noalamina differs from D. temporalis in retaining the contrast between light and dark rings throughout body and tail (Fig. 1), whereas in D. temporalis the light portions grade into medium to dark brown posteriorly (Fig. 5 H). Description of the holotype. Adult male (Figs. 1 A���C, 2���3), as indicated by everted hemipenes; SVL 385 mm, TL 161 mm (but tail incomplete); TOL 546 mm; HL 12.4 mm; HW 8.2 mm; SL 3.5 mm. Head distinct from neck; snout short, blunt in dorsal and lateral outline; rostral wider (2.1 mm) than high (1.3 mm), not extending posteriorly between internasals, its length visible from above one fifth length of median internasal suture (1.0 mm); internasals shorter (1.1 mm) than wide (1.7 mm), their length between one half and two-thirds length of prefrontal suture (1.9 mm); prefrontals shorter (2.5 mm) than wide (2.8 mm), their median suture (2.1 mm) about three-fifths length of frontal; prefrontals bordering eye above loreal; frontal longer (3.6 mm) than wide (3.0 mm), widest anteriorly, length almost two-thirds that of parietal, in contact with prefrontals, supraoculars, and parietals; supraoculars longer (3.4 mm) than wide (1.7 mm), nearly as long as frontal; parietals longer (5.8 mm) than wide (3.8 mm), widest anteriorly, their median suture (4.1 mm) slightly longer than frontal; parietals in contact with frontal, supraoculars, upper postoculars, anterior and posterior temporals, and nine nuchal scales; nasal divided, in contact with first two supralabials, loreal, prefrontal, internasal, and rostral; loreal single, longer (1.9 mm) than high (1.0 mm), its length more than one-half length of snout, and two-thirds length of eye (2.8 mm); preoculars absent; postoculars 2; temporals 1 + 2 / 2 + 2, all above fifth supralabial; supralabials five, with third and fourth bordering eye (enlarged anterior portion of ultimate supralabial separated from eye by lower postocular); fifth and ultimate supralabial conspicuously long (3.9 mm), longer than combined lengths of third and fourth supralabials, with a higher (1.7 mm) anterior and a lower (1.2 mm) posterior portion; pupil vertically elliptical; eye length one and a half times length of loreal; infralabials 6, first pair in contact behind mental; no postmental; infralabials 1���4 in contact with anterior chin shields; anterior chin shields paired, longer (2.8 mm) than wide (1.3 mm), longer than posterior chin shields; posterior chin shields longer (1.9 mm) than wide (1.4 mm), paired, in broad contact medially; mental groove present; ventrals 170; cloacal scute single; subcaudals 80 (but tail incomplete), divided; ventrals plus subcaudals 250 (but tail incomplete); dorsal scales mostly smooth, but slightly keeled on dorsal rows 3���5 around midbody, striate, in 15 ��� 15 ��� 15 rows, without apical pits or supracloacal ridges; vertebral row slightly enlarged. The fully everted hemipenis (Fig. 3) is a stout unilobed capitate organ; the capitulum is covered by calyces bearing spinules; no naked pocket; the sulcus spermaticus is bordered by well-developed sulcal lips, bifurcates at the base of apex with the branches continuing onto the apex; truncus with two series of large recurved spines. Coloration in life. Dorsal, lateral, and ventral surfaces of body and tail Pale Horn Color (92); body with 14, tail with eight broad Sepia (219) rings, body rings extending over 6���12 middorsal scales, narrowing towards and on venter; interspaces between these rings extending over 3���6 middorsal scales, speckled with Sepia (219) except for narrow portions immediately adjacent to the dark rings; tip of (incomplete) tail Sepia (119); dorsal and lateral surfaces of head Pale Horn Color (92), grading into Flesh Ocher (132 D) anteriorly, heavily mottled with Sepia (219); parietals, frontal, and central portions of prefrontals almost completely Sepia (219), as well as the middorsal scales of the first two transverse rows of dorsals posterior to parietals, connecting Sepia (219) coloration to first body ring; ventral surface of head Pale Horn Color (92) mottled with Sepia (219), especially on infralabials; iris Sepia (119). Color after ten months in preservative (70 % ethanol) is similar to that in life, apart from that all yellow and orange shades have faded. Variation. The juvenile paratypes (SMF 89550, Fig. 1 E; and SMF 90180, Fig. 1 D) agree well with the holotype in terms of general morphology and pholidosis (even in their tails being incomplete), differing mostly in their lack of any keels on dorsal rows 3���5 and in their somewhat more contrasting coloration. Scale counts that differ from that of the holotype are as follows: ventrals 164 (SMF 89550), 177 (SMF 90180); subcaudals 93, 96 (both tails are incomplete); ventrals plus subcaudals 257, 273 (but both tails are incomplete); temporals 1 + 2 / 1 + 2 in both, anterior temporals of both sides entering orbit between upper and lower postoculars (SMF 89550); parietals bordered posteriorly by 8 nuchal scales in both; infralabials 6 /7, 1��� 5 in contact with first chin shield on left side (SMF 89550). Body measurements of the juvenile specimens are SVL 178 mm, TL 68 mm (SMF 89550), SVL 161 mm, TL 64 mm (SMF 90180). As illustrated in Fig. 1, the coloration in life of the juveniles presents a stronger contrast than that of the adult holotype. Moreover, the paratypes show variation in the relative width of dark and light rings. The latter appear most narrow in the juvenile from Veraguas (SMF 89550), which was recorded as follows (modified from Stadler 2010): Body and tail with alternating, complete dark and light rings; broader rings Jet Black (89), light rings Chamois (123 D), about half as wide as, and becoming lighter toward, the dark rings; dorsal and lateral ground color of head Jet Black (89); crescent-shaped lateral markings on posterior portion of head and small blotches on snout Chamois (123 D); ventral surfaces of head and neck Chamois (123 D) mottled with Jet Black (89); iris black. The color after 37 months in preservative (70 % ethanol) is similar to that in life, apart from that all yellow and orange shades have faded. Natural history notes. All three specimens were encountered at night on vegetation. The juvenile SMF 89550 (Fig. 1 E) from Cerro Mariposa, Veraguas, was moving through epiphytic liverworts growing on a small tree-trunk about 0.5 m above ground at 18: 40 hrs, after a rainy afternoon. The habitat may be characterized as ridgetop cloud forest with abundant epiphytic vegetation. Annual precipitation is approx. 2500 mm, mean annual temperature approx. 20.2 ��C, indicating the Premontane Wet Forest life zone according to the Holdridge (1967) classification. Other snail-eaters found at lower elevations on the slopes of Cerro Mariposa are Dipsas temporalis, Sibon annulatus, S. longifrenis, and S. nebulatus. A complete list of the herpetofauna documented around Cerro Mariposa was given by Stadler (2010), who also provided extensive data on climate and vegetation. Additionally, the herpetofauna of the general Santa F�� region was treated by Mart��nez and Rodriguez (1994 ��� 1992 ���), Mart��nez et al. (1995 ��� 1994 ���), and Carrizo (2010), and updated by Lotzkat et al. (2010). Figure 1 D shows the juvenile SMF 90180 as it was encountered at 19: 40 hrs and 19.1 ��C air temperature at the type locality: It was lying in loose coils upon a large leaf about 0.5 m above ground, in herbaceous secondary vegetation below the floodline of, and less than 10 m away from, a small stream. The holotype was found close to this stream while moving about in secondary forest, 2 m above ground at 23:00 hrs. The type locality is the valley of the mentioned stream, with rather undisturbed forest on the slope ascending northwards, minor clearings overgrown with grass and solitary trees directly by the riverside, and a more secondary forest on the slope ascending southward. Reptiles collected in this valley include Anolis capito Peters, A. humilis Peters, A. limifrons Cope, A. lionotus Cope, Lepidoblepharis xanthostigma (Noble), Imantodes cenchoa (Linnaeus), Sibon annulatus, and Bothrops asper (Garman). Annual total precipitation at the type locality is approx. 3000 mm, mean annual temperature approx. 20.6 ��C, indicating the Premontane Wet Forest life zone according to the Holdridge (1967) classification. The datalogger recordings yielded a temperature range of 17.6���22.6 ��C, with a mean of 19.4 ��C and standard deviation �� 1.3 ��C. Geographic distribution. Sibon noalamina is hitherto only known from the highlands of western Panama (Fig. 4). The two known localities lie some 125 airline km apart, roughly at either end of the easternmost uninterrupted portion of the Cordillera Central, known as Serran��a de Tabasar��, which is approximately delimited to the west by the Fortuna depression and has its eastern limit in the region around Santa F��. This indicates, as a minimum, a continuous distribution of the new taxon along the Caribbean versant of the Serran��a de Tabasar��, at premontane elevations between 1050 and 1260 m, most of it located in the Comarca Ng��be-Bugl��. Etymology. The specific epithet is a contraction of the exclamation ���no a la mina!���, Spanish for ���no to the mine���, in the sense of ���no mining���. This affirmation was and is used by members of the indigenous Ng��be communities living in the Serran��a de Tabasar�� in the course of their protests against mining interests aiming to exploit their territory, especially around Cerro Colorado. The specific name is given in recognition and support of the Ng��be���s struggle to protect their territory and environment, which is home to the new species described herein and many others, from profit-driven destructive interventions., Published as part of Lotzkat, Sebastian, Hertz, Andreas & K��hler, Gunther, 2012, A new species of Sibon (Squamata: Colubroidea: Dipsadidae) from the Cordillera Central of western Panama, with comments on other species of the genus in the area, pp. 26-40 in Zootaxa 3485 on pages 27-32, DOI: 10.11646/zootaxa.3485.1.2, http://zenodo.org/record/208860, {"references":["Kohler, G. (2008) Reptiles of Central America. Herpeton, Offenbach [Germany], 400 pp.","Stadler, L. (2010) Diversitat, Taxonomie und Zoogeographie der Herpetofauna der Umgebung von Alto de Piedra (Veraguas, Panama). Diploma thesis. Justus-Liebig-Universitat, Giessen, 290 pp. Available from http: // www. senckenberg. de / files / content / forschung / abteilung / terrzool / herpetologie / herpetofauna _ alto _ de _ piedra _ veraguas _ panama _ stadler _ 2010. pdf (accessed 26 June 2011).","Carrizo, A. (2010) Riqueza y abundancia de la herpetofauna de la cuenca alta del Rio Santa Maria, Santa Fe, Veraguas. Master thesis. Universidad Autonoma de Chiriqui, David, 123 pp. Available from http: // www. senckenberg. de / files / content / forschung / abteilung / terrzool / herpetologie / herpetofauna _ cuenca _ alta _ rio _ santa _ maria _ veraguas _ panama _ carrizo _ 2010. pdf (accessed 26 June 2011).","Holdridge, L. R. (1967) Life zone ecology. Tropical Science Center, San Jose, Costa Rica, 206 pp.","Martinez V. C. & Rodriguez, A. (1994 \" 1992 \") Del primer inventario en \" Cerro Tute \". Amphibia: Caudata y Anura. Reptilia: Squamata. Sauria y Serpentes. Scientia (Panama), 7, 29 - 53.","Martinez, V. C., Pimentel, N. & Hurdaneta, A. (1995 \" 1994 \") Diversidad herpetofaunistica en los cerros \" Narices \" y \" La Anselma \": Provincia de Veraguas. Distrito de Santa Fe. Scientia (Panama), 9, 59 - 79.","Lotzkat, S., Hertz, A., Stadler, L., Hamad, N., Carrizo, A. & Kohler, G. (2010) Noteworthy distribution records of reptiles from Western Panama. Herpetological Review, 41, 520 - 523"]}

We report on new localities for Anolis gruuo Köhler, Ponce, Sunyer and Batista, 2007 along the Serranía de Tabasará in the Comarca Ngöbe-Buglé and Veraguas province of western Panama. These records extend the known geographic distribution of this lizard about 80 km eastward, and the known vertical distribution approximately 40 m lower and 630 m higher. We provide photos of specimens from different localities and comment on their morphology. Only the easternmost populations of this Panamanian endemic live inside a protected area.

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."]}

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