Search

Myanophis thanlyinensis gen. nov. , sp. nov. Gunther K��hler, Khin Pa Pa Khaing, Ni Lar Than ZOOBANK urn:lsid:zoobank.org: act: 68106D4B-425B-4094-A6D3-81C4C42FEFB7 Figs. 5���6; Table 2 Holotype. SMF 100707, an adult male from wetlands near campus of East Yangon University (16.74255, 96.28307; 6 m a.s.l.), Thanlyin, Yangon, Myanmar, collected on 5 of November 2019 by Ni Lar Than and Gunther K��hler. Original field tag GK-7730. Paratypes. EYU PaPa-0023 and SMF 100709 (original field tag PaPa-0019), two adult females from wetlands near campus of East Yangon University (16.740297, 96.285184; 6 m a.s.l.), Thanlyin, Yangon, Myanmar, collected on 14 of September 2018 by a local fishermen and received by Khin Pa Pa Khaing. SMF 100708 (original field tag GK-7731), an adult male from wetlands near campus of East Yangon University (16.74275, 96.31250; 6 m a.s.l.), Thanlyin, Yangon, Myanmar, collected on 15 of October 2019 by a local fishermen and received by Ni Lar Than. Diagnosis. A species of the genus Myanophis that differs from all other homalopsid species by the following combination of characters: (1) dorsal scales smooth, row formula 21���21���19 or 21���21���17; (2) tail short, ratio TaL/SVL 0.185 ���0.204 in males, 0.160 ���0.167 in females; (3) nasal scales separated; (4) 125���126 ventral scales in males, 120���122 in females; (5) 38���39 subcaudal scales in males, 32���34 in females; and (6) hemipenis bilobed. Phylogenetically, (based on analyses of 16S and MT-CYB sequences), Myanophis thanlyinensis gen. nov. , sp. nov. differs from its closest relatives, Myrrophis and Gyiophis, by having a bilobed hemipenis (vs. unilobed). Further, Myanophis thanlyinensis gen. nov. , sp. nov. differs from the species of Myrrophis by having 125���126 ventral scales in males, 120���122 in females (vs. 137���162, 137���164, respectively), and 38���39 subcaudal scales in males, 32���34 in females (vs. 39���55, 37���52, respectively). Myanophis thanlyinensis gen. nov. , sp. nov. differs further from the species of Gyiophis by lacking dark blotches along flank (vs. having blotches), and by having 21 dorsal scales rows at midbody (vs. 25 scales). The following other homalopsid species share with M. thanlyinensis gen. nov., sp. nov. the combination of having smooth dorsal scales in 21 rows at midbody, reducing to 17 or 19 rows anterior to vent and fewer than 130 ventral scales: Enhydris chanardi, E. innominata, E. jagorii, and E. longicauda. However, all species have more than 50 subcaudal scales in males and more than 40 in females (vs. 38���39, 32���34, respectively, in M. thanlyinensis gen. nov., sp. nov.), and a TaL/SVL of> 0.22 in males, and> 0.17 in females (vs. 0.185���0.204, 0.160���0.167, respectively).Also, Enhydris jagorii and E. longicauda have 23���27 dorsal scales rows on neck (vs. 21 in Myanophis Thanlyinensis gen. nov. , sp. nov.). Description of the holotype (Figs. 5 & 6). Adult male, indicated by fully developed everted hemipenes; head depressed, distinct from neck; snout short and rounded, rostrum tapers downwards; rostral scale pentagonal, nearly as broad as tall, visible from above; eye moderate in size, pupils round; nasals divided with nasal groove contacting first supralabial; internasal squarish, not in contact with loreal; loreals 1/1, quadrangular, contacting first three supralabials; preocular 1/1; supraocular 1/1; postoculars 2/2; prefrontals 2, in broad contact with each other, frontal, internasal, loreals, preoculars, supraoculars, and nasals; frontal roughly pentagonal, 1.4 times longer than supraocular; parietals elongate; temporal scale formula 1 + 2 + 3; supralabials 8/8, 4th supralabial entering orbit; infralabials 10/10, first five in contact with anterior chin shield; anterior pair of chin shields larger than second pair, the former in contact with each other, the latter separated medially from each other by two smaller scales; 21 ASR, 21 MSR, 19 PSR; dorsal scales smooth without scale pits; 126 ventrals; cloacal plate divided; subcaudal scales 39. Body short, somewhat stocky; tail short (TaL/SVL 0.2038); SVL 260 mm; TaL 53 mm. Coloration in life (Fig. 5; colors in parentheses according to K��hler 2012): Dorsal ground color Olive Brown (color 278), scale rows 4 and 5 slightly darker (Sepia 286) and scale rows 1 to 3 Pale Cinnamon (55) with Hair Brown (277) blotch on anterior portion of each scale; ventral scales Pale Cinnamon (55) with Hair Brown (277) suffusion of anterior portion of each scale; iris Olive Brown (278). Coloration after about three months in preservation in 70% ethanol agrees well with the description of coloration in life. Variation. Paratypes agree well with the holotype in general appearance, morphometrics and scalation (Table 2). Strong sexual dimorphism is evident in the number of ventral and subcaudal scales that show non-overlapping values between males and females. Etymology. The species epithet refers to the city of Thanlyin, a major port city of Myanmar, located across the Bago River southeast of the city of Yangon, where the type series of M. thanlyinensis gen. nov., sp. nov. was collected. Natural history notes. The type locality is open grassland (Fig. 7), mostly flooded during the rainy season with only a few scattered wooden houses, adjacent to the campus of East Yangon University (EYU), Thanlyin, Myanmar. The snakes were caught at night with the help of local fishermen in shallow water (M. thanlyinensis gen. nov., sp. nov. we collected the homalopsid species Enhydris enhydris and Gyiophis vorisi (Fig. 8) as well the semiaquatic colubrids Amphiesma stolatum, Xenochrophis bellulus, and X. piscator. Of these syntopic species, E. enhydris was the by far most abundant snake at this site with>20 individuals encountered per night-search. Geographic Distribution and Conservation. As currently known, Myanophis thanlyinensis gen. nov. , sp. nov. is only known from the vicinity of its type locality. Given the little we know about this species, we classify M. thanlyinensis gen. nov., sp. nov. as Data Deficient based on the IUCN Red List Categories and Criteria (IUCN 2012)., Published as part of K��hler, Gunther, Khaing, Khin Pa Pa, Than, Ni Lar, Baranski, Damian, Schell, Tilman, Greve, Carola, Janke, Axel & Pauls, Steffen U., 2021, A new genus and species of mud snake from Myanmar (Reptilia, Squamata Homalopsidae), pp. 301-325 in Zootaxa 4915 (3) on pages 308-311, DOI: 10.11646/zootaxa.4915.3.1, http://zenodo.org/record/4456421, {"references":["Kohler, G. (2012) Color Catalogue for Field Biologists. Herpeton, Offenbach, 49 pp.","IUCN (2012) IUCN Red List Categories and Criteria, Version 3.1. 2 nd Edition. Available from: https: // portals. ucn. org / library / node / 10315 (accessed 17 November 2018)"]}

Key to species of mud snakes (family Homalopsidae) known to occur in Myanmar 1 Dorsal scales at midbody in more than 30 rows.............................................................. 2 ��� Dorsal scales at midbody in fewer than 30 rows............................................................. 3 2a Nasals separated from each other............................................................ Bitia hydroides 2b Nasals in contact with each other...................................................... Homalopsis semizonata 3a Nasals separated from each other........................................................................ 4 ��� Nasals in contact with each other........................................................................ 6 4a Dorsal scales at midbody in 23���27 rows.................................................... Fordonia leucobalia ��� Dorsal scales at midbody in 17 or 19 rows.................................................................. 5 5a Dorsal scales at midbody in 17 rows...................................................... Gerarda prevostiana ��� Dorsal scales at midbody in 19 rows........................................................ Cantoria violacea 6a Dorsal scales keeled.................................................................... Cerberus rynchops ��� Dorsal scales smooth.................................................................................. 7 7a Dorsal scales at midbody in 19���21 rows................................................................... 8 ��� Dorsal scales at midbody in 25���29 rows.................................................................. 10 8a Dorsal scales at midbody in 19 rows..................................................... Hypsiscopus plumbea ��� Dorsal scales at midbody in 21 rows...................................................................... 9 9a Subcaudal scales>50; long-tailed, ratio tail length / SVL> 0.21 in males,> 0.18 in females............ Enhydris enhydris ��� Subcaudal scales Myanophis thanlyinensis gen. nov. , sp. nov. 10a Dorsal scales at midbody in 29 rows......................................................... Ferania sieboldii ��� Dorsal scales at midbody in 25 rows..................................................................... 11 11a Dorsal scales on neck in 25 rows; scales of first three dorsal scale rows oval; a central spot on each ventral scale; no stripe on first dorsal scale row.................................................................... Gyiophis maculosa ��� Dorsal scales on neck in more than 25 rows; scales of first three dorsal scale rows squarish; no central spot on each ventral scale; stripe on first dorsal scale row present.................................................................... 12 12a Dorsal scales at posterior body in 20 rows; 129 ventrals in single known female; 30/29 subcaudals in single known female; 4 rows of spots on dorsum; rostral scale narrow............................................ Gyiophis salweenensis ��� Dorsal scales at posterior body in 21���23 rows; 142���148 ventrals in females; 41���58 subcaudals in females; 3 rows of spots on dorsum; rostral scale broad................................................................. Gyiophis vorisi, Published as part of K��hler, Gunther, Khaing, Khin Pa Pa, Than, Ni Lar, Baranski, Damian, Schell, Tilman, Greve, Carola, Janke, Axel & Pauls, Steffen U., 2021, A new genus and species of mud snake from Myanmar (Reptilia, Squamata Homalopsidae), pp. 301-325 in Zootaxa 4915 (3) on pages 311-312, DOI: 10.11646/zootaxa.4915.3.1, http://zenodo.org/record/4456421

Myanophis gen. nov. Gunther K��hler, Khin Pa Pa Khaing, Ni Lar Than ZOOBANK urn:lsid:zoobank.org: act: 286D70A4-EF16-41D3-97CD-01BA4830878F Type species. Myanophis thanlyinensis sp. nov., by present designation. Content. Monotypic having only Myanophis thanlyinensis, gen. nov, sp. nov. Distribution. Currently only known from the vicinity of East Yangon University (EYU), Thanlyin, Myanmar. Diagnosis and definition. A medium-sized Indochinese homalopsid snake with the following combination of characters: (1) dorsal scales smooth, row formula 21���21���19 or 21���21���17; (2) tail short, ratio tail length/SVL 0.185 ��� 0.204 in males, 0.160 ���0.167 in females; (3) nasal scales separated; (4) 125���126 ventral scales in males, 120���122 in females; (5) 38���39 subcaudal scales in males, 32���34 in females; (6) hemipenis bilobed. Etymology. The generic name is derived from Myanmar���s abbreviated name ���Myan��� and the Greek ���ophis���, meaning ���snake���. The generic name is feminine. Myanophis thanlyinensis gen. nov. , sp. nov. Gunther K��hler, Khin Pa Pa Khaing, Ni Lar Than ZOOBANK urn:lsid:zoobank.org: act: 68106D4B-425B-4094-A6D3-81C4C42FEFB7 Figs. 5���6; Table 2 Holotype. SMF 100707, an adult male from wetlands near campus of East Yangon University (16.74255, 96.28307; 6 m a.s.l.), Thanlyin, Yangon, Myanmar, collected on 5 of November 2019 by Ni Lar Than and Gunther K��hler. Original field tag GK-7730. Paratypes. EYU PaPa-0023 and SMF 100709 (original field tag PaPa-0019), two adult females from wetlands near campus of East Yangon University (16.740297, 96.285184; 6 m a.s.l.), Thanlyin, Yangon, Myanmar, collected on 14 of September 2018 by a local fishermen and received by Khin Pa Pa Khaing. SMF 100708 (original field tag GK-7731), an adult male from wetlands near campus of East Yangon University (16.74275, 96.31250; 6 m a.s.l.), Thanlyin, Yangon, Myanmar, collected on 15 of October 2019 by a local fishermen and received by Ni Lar Than. Diagnosis. A species of the genus Myanophis that differs from all other homalopsid species by the following combination of characters: (1) dorsal scales smooth, row formula 21���21���19 or 21���21���17; (2) tail short, ratio TaL/SVL 0.185 ���0.204 in males, 0.160 ���0.167 in females; (3) nasal scales separated; (4) 125���126 ventral scales in males, 120���122 in females; (5) 38���39 subcaudal scales in males, 32���34 in females; and (6) hemipenis bilobed. Phylogenetically, (based on analyses of 16S and MT-CYB sequences), Myanophis thanlyinensis gen. nov. , sp. nov. differs from its closest relatives, Myrrophis and Gyiophis, by having a bilobed hemipenis (vs. unilobed). Further, Myanophis thanlyinensis gen. nov. , sp. nov. differs from the species of Myrrophis by having 125���126 ventral scales in males, 120���122 in females (vs. 137���162, 137���164, respectively), and 38���39 subcaudal scales in males, 32���34 in females (vs. 39���55, 37���52, respectively). Myanophis thanlyinensis gen. nov. , sp. nov. differs further from the species of Gyiophis by lacking dark blotches along flank (vs. having blotches), and by having 21 dorsal scales rows at midbody (vs. 25 scales). The following other homalopsid species share with M. thanlyinensis gen. nov., sp. nov. the combination of having smooth dorsal scales in 21 rows at midbody, reducing to 17 or 19 rows anterior to vent and fewer than 130 ventral scales: Enhydris chanardi, E. innominata, E. jagorii, and E. longicauda. However, all species have more than 50 subcaudal scales in males and more than 40 in females (vs. 38���39, 32���34, respectively, in M. thanlyinensis gen. nov., sp. nov.), and a TaL/SVL of> 0.22 in males, and> 0.17 in females (vs. 0.185���0.204, 0.160���0.167, respectively).Also, Enhydris jagorii and E. longicauda have 23���27 dorsal scales rows on neck (vs. 21 in Myanophis Thanlyinensis gen. nov. , sp. nov.). Description of the holotype (Figs. 5 & 6). Adult male, indicated by fully developed everted hemipenes; head depressed, distinct from neck; snout short and rounded, rostrum tapers downwards; rostral scale pentagonal, nearly as broad as tall, visible from above; eye moderate in size, pupils round; nasals divided with nasal groove contacting first supralabial; internasal squarish, not in contact with loreal; loreals 1/1, quadrangular, contacting first three supralabials; preocular 1/1; supraocular 1/1; postoculars 2/2; prefrontals 2, in broad contact with each other, frontal, internasal, loreals, preoculars, supraoculars, and nasals; frontal roughly pentagonal, 1.4 times longer than supraocular; parietals elongate; temporal scale formula 1 + 2 + 3; supralabials 8/8, 4th supralabial entering orbit; infralabials 10/10, first five in contact with anterior chin shield; anterior pair of chin shields larger than second pair, the former in contact with each other, the latter separated medially from each other by two smaller scales; 21 ASR, 21 MSR, 19 PSR; dorsal scales smooth without scale pits; 126 ventrals; cloacal plate divided; subcaudal scales 39. Body short, somewhat stocky; tail short (TaL/SVL 0.2038); SVL 260 mm; TaL 53 mm. Coloration in life (Fig. 5; colors in parentheses according to K��hler 2012): Dorsal ground color Olive Brown (color 278), scale rows 4 and 5 slightly darker (Sepia 286) and scale rows 1 to 3 Pale Cinnamon (55) with Hair Brown (277) blotch on anterior portion of each scale; ventral scales Pale Cinnamon (55) with Hair Brown (277) suffusion of anterior portion of each scale; iris Olive Brown (278). Coloration after about three months in preservation in 70% ethanol agrees well with the description of coloration in life. Variation. Paratypes agree well with the holotype in general appearance, morphometrics and scalation (Table 2). Strong sexual dimorphism is evident in the number of ventral and subcaudal scales that show non-overlapping values between males and females. Etymology. The species epithet refers to the city of Thanlyin, a major port city of Myanmar, located across the Bago River southeast of the city of Yangon, where the type series of M. thanlyinensis gen. nov., sp. nov. was collected. Natural history notes. The type locality is open grassland (Fig. 7), mostly flooded during the rainy season with only a few scattered wooden houses, adjacent to the campus of East Yangon University (EYU), Thanlyin, Myanmar. The snakes were caught at night with the help of local fishermen in shallow water (M. thanlyinensis gen. nov., sp. nov. we collected the homalopsid species Enhydris enhydris and Gyiophis vorisi (Fig. 8) as well the semiaquatic colubrids Amphiesma stolatum, Xenochrophis bellulus, and X. piscator. Of these syntopic species, E. enhydris was the by far most abundant snake at this site with>20 individuals encountered per night-search. Geographic Distribution and Conservation. As currently known, Myanophis thanlyinensis gen. nov. , sp. nov. is only known from the vicinity of its type locality. Given the little we know about this species, we classify M. thanlyinensis gen. nov., sp. nov. as Data Deficient based on the IUCN Red List Categories and Criteria (IUCN 2012). Myanophis thanlyinensis gen. nov. , sp. nov. Gunther K��hler, Khin Pa Pa Khaing, Ni Lar Than ZOOBANK urn:lsid:zoobank.org: act: 68106D4B-425B-4094-A6D3-81C4C42FEFB7 Figs. 5���6; Table 2 Holotype. SMF 100707, an adult male from wetlands near campus of East Yangon University (16.74255, 96.28307; 6 m a.s.l.), Thanlyin, Yangon, Myanmar, collected on 5 of November 2019 by Ni Lar Than and Gunther K��hler. Original field tag GK-7730. Paratypes. EYU PaPa-0023 and SMF 100709 (original field tag PaPa-0019), two adult females from wetlands near campus of East Yangon University (16.740297, 96.285184; 6 m a.s.l.), Thanlyin, Yangon, Myanmar, collected on 14 of September 2018 by a local fishermen and received by Khin Pa Pa Khaing. SMF 100708 (original field tag GK-7731), an adult male from wetlands near campus of East Yangon University (16.74275, 96.31250; 6 m a.s.l.), Thanlyin, Yangon, Myanmar, collected on 15 of October 2019 by a local fishermen and received by Ni Lar Than. Diagnosis. A species of the genus Myanophis that differs from all other homalopsid species by the following combination of characters: (1) dorsal scales smooth, row formula 21���21���19 or 21���21���17; (2) tail short, ratio TaL/SVL 0.185 ���0.204 in males, 0.160 ���0.167 in females; (3) nasal scales separated; (4) 125���126 ventral scales in males, 120���122 in females; (5) 38���39 subcaudal scales in males, 32���34 in females; and (6) hemipenis bilobed. Phylogenetically, (based on analyses of 16S and MT-CYB sequences), Myanophis thanlyinensis gen. nov. , sp. nov. differs from its closest relatives, Myrrophis and Gyiophis, by having a bilobed hemipenis (vs. unilobed). Further, Myanophis thanlyinensis gen. nov. , sp. nov. differs from the species of Myrrophis by having 125���126 ventral scales in males, 120���122 in females (vs. 137���162, 137���164, respectively), and 38���39 subcaudal scales in males, 32���34 in females (vs. 39���55, 37���52, respectively). Myanophis thanlyinensis gen. nov. , sp. nov. differs further from the species of Gyiophis by lacking dark blotches along flank (vs. having blotches), and by having 21 dorsal scales rows at midbody (vs. 25 scales). The following other homalopsid species share with M. thanlyinensis gen. nov., sp. nov. the combination of having smooth dorsal scales in 21 rows at midbody, reducing to 17 or 19 rows anterior to vent and fewer than 130 ventral scales: Enhydris chanardi, E. innominata, E. jagorii, and E. longicauda. However, all species have more than 50 subcaudal scales in males and more than 40 in females (vs. 38���39, 32���34, respectively, in M. thanlyinensis gen. nov., sp. nov.), and a TaL/SVL of> 0.22 in males, and> 0.17 in females (vs. 0.185���0.204, 0.160���0.167, respectively).Also, Enhydris jagorii and E. longicauda have 23���27 dorsal scales rows on neck (vs. 21 in Myanophis Thanlyinensis gen. nov. , sp. nov.). Description of the holotype (Figs. 5 & 6). Adult male, indicated by fully developed everted hemipenes; head depressed, distinct from neck; snout short and rounded, rostrum tapers downwards; rostral scale pentagonal, nearly as broad as tall, visible from above; eye moderate in size, pupils round; nasals divided with nasal groove contacting first supralabial; internasal squarish, not in contact with loreal; loreals 1/1, quadrangular, contacting first three supralabials; preocular 1/1; supraocular 1/1; postoculars 2/2; prefrontals 2, in broad contact with each other, frontal, internasal, loreals, preoculars, supraoculars, and nasals; frontal roughly pentagonal, 1.4 times longer than supraocular; parietals elongate; temporal scale formula 1 + 2 + 3; supralabials 8/8, 4th supralabial entering orbit; infralabials 10/10, first five in contact with anterior chin shield; anterior pair of chin shields larger than second pair, the former in contact with each other, the latter separated medially from each other by two smaller scales; 21 ASR, 21 MSR, 19 PSR; dorsal scales smooth without scale pits; 126 ventrals; cloacal plate divided; subcaudal scales 39. Body short, somewhat stocky; tail short (TaL/SVL 0.2038); SVL 260 mm; TaL 53 mm. Coloration in life (Fig. 5; colors in parentheses according to K��hler 2012): Dorsal ground color Olive Brown (color 278), scale rows 4 and 5 slightly darker (Sepia 286) and scale rows 1 to 3 Pale Cinnamon (55) with Hair Brown (277) blotch on anterior portion of each scale; ventral scales Pale Cinnamon (55) with Hair Brown (277) suffusion of anterior portion of each scale; iris Olive Brown (278). Coloration after about three months in preservation in 70% ethanol agrees well with the description of coloration in life. Variation. Paratypes agree well with the holotype in general appearance, morphometrics and scalation (Table 2). Strong sexual dimorphism is evident in the number of ventral and subcaudal scales that show non-overlapping values between males and females. Etymology. The species epithet refers to the city of Thanlyin, a major port city of Myanmar, located across the Bago River southeast of the city of Yangon, where the type series of M. thanlyinensis gen. nov., sp. nov. was collected. Natural history notes. The type locality is open grassland (Fig. 7), mostly flooded during the rainy season with only a few scattered wooden houses, adjacent to the campus of East Yangon University (EYU), Thanlyin, Myanmar. The snakes were caught at night with the help of local fishermen in shallow water (M. thanlyinensis gen. nov., sp. nov. we collected the homalopsid species Enhydris enhydris and Gyiophis vorisi (Fig. 8) as well the semiaquatic colubrids Amphiesma stolatum, Xenochrophis bellulus, and X. piscator. Of these syntopic species, E. enhydris was the by far most abundant snake at this site with>20 individuals encountered per night-search. Geographic Distribution and Conservation. As currently known, Myanophis thanlyinensis gen. nov. , sp. nov. is only known from the vicinity of its type locality. Given the little we know about this species, we classify M. thanlyinensis gen. nov., sp. nov. as Data Deficient based on the IUCN Red List Categories and Criteria (IUCN 2012)., Published as part of K��hler, Gunther, Khaing, Khin Pa Pa, Than, Ni Lar, Baranski, Damian, Schell, Tilman, Greve, Carola, Janke, Axel & Pauls, Steffen U., 2021, A new genus and species of mud snake from Myanmar (Reptilia, Squamata Homalopsidae), pp. 301-325 in Zootaxa 4915 (3) on pages 307-311, DOI: 10.11646/zootaxa.4915.3.1, http://zenodo.org/record/4456421, {"references":["Kohler, G. (2012) Color Catalogue for Field Biologists. Herpeton, Offenbach, 49 pp.","IUCN (2012) IUCN Red List Categories and Criteria, Version 3.1. 2 nd Edition. Available from: https: // portals. ucn. org / library / node / 10315 (accessed 17 November 2018)"]}

Tropidurus catalanensis Gudynas & Skuk, 1983 Taraguira torquata: COPE 1862. Tropidurus torquatus: KOSLOWSKY 1898; BERTONI 1914, 1939; ScHOUTEN 1929; GIRAUDO & CONTRERAS 1994; DUR�� RODAS 1995; AqUINO et al. 1996; MOTTE et al. 2004; MOTTE et al. 2009; CAbRAL & WEILER 2014; CAccIALI et al. 2016. Tropidurus catalanensis GUDYNAS & SKUK, 1983. TYPE LOcALITY: 1 KM ESE OF ROUTE 30 AND ARROYO CATALAN GRANDE, ARTIGAS, URUGUAY. ��LVAREZ et al. 1995; KUNZ & BORGES-MARTINS 2013. Tropidurus torquatus torquatus: BURT & BURT 1933. Tropidurus torquatus hispidus: HELLMIcH 1960. Geographic distribution. THE DISTRIbUTION OF T. catalanensis RANGES FROM THE SOUTHERN REGION OF THE BRAZILIAN STATE OF MATO GROSSO DO SUL, THROUGH EASTERN PARAGUAY AND THE STATE OF PARAN�� (BRAZIL) TO THE PROVINcES OF CHAcO AND CORRIENTES (ARGENTINA), AND URUGUAY (FIG. 11A)., Published as part of Cacciali, Pier & K��hler, Gunther, 2018, Diversity of Tropidurus (Squamata: Tropiduridae) in Paraguay-an integrative taxonomic approach based on morphological and molecular genetic evidence, pp. 511-536 in Zootaxa 4375 (4) on page 521, DOI: 10.11646/zootaxa.4375.4.3, http://zenodo.org/record/1160960, {"references":["Gudynas, E. & Skuk, G. (1983) A new species of the iguanid lizard genus Tropidurus from temperate South America (Lacertilia: Iguanidae). Contribuciones en Biologia del Centro Educativo Don Orione, 10, 1 - 10.","Cope, E. D. (1862) Catalogue of the reptiles obtained during the exploration of the Parana, Paraguay, Vermejo and Uruguay rivers, by Capt. Thos. J. Page, U. S. N.; and of those procured by Lieut. N. Michler, U. S. Top. Eng., commander of the expedition conducting the survey of the Atrato River. Proceedings of the Academy of Natural Science of Philadelphia, 14, 346 - 359. Available from: http: // www. biodiversitylibrary. org / page / 1951818 page / 358 / mode / 1 up (Accessed 25 Jan. 2018)","Koslowsky, J. (1898) Enumeracion sistematica y distribucion geografica de los reptiles argentinos. Revista del Museo de La Plata, 8, 161 - 200.","Bertoni, A. de W. (1914) Fauna paraguaya: Catalogos sistematicos de los vertebrados del Paraguay. Descripcion Fisica y Economica del Paraguay, 59, 1 - 83.","Bertoni, A. de W. (1939) Catalogos sistematicos de los vertebrados de Paraguay. Revista de la Sociedad Cientifica del Paraguay, 4, 3 - 60.","Schouten, G. B. (1929) Notas sobre la oologia de algunos saurios del Paraguay y de los paises limitrofes. Revista Chilena de Historia Natural, 33, 518 - 521.","Giraudo, A. R. & Contreras, A. O. (1994) Lista preliminar de los reptiles registrados en el Departmento de Neembucu, Paraguay. Boletin de la Asociacion Herpetologica Argentina, 10, 1 - 4.","Dure Rodas, A. (1995) Estudio de reptiles y anfibios. Biota, 2, 20 - 24.","Cabral, H. & Weiler, A. (2014) Lista comentada de los reptiles de la Coleccion Zoologica de la Facultad de Ciencias Exactas y Naturales de Asuncion, Paraguay. Cuadernos de Herpetologia, 28, 19 - 28.","Kuntz, T. S. & Borges-Martins, M. (2013) A new microendemic species of Tropidurus (Squamata: Tropiduridae) from southern Brazil and revalidation of Tropidurus catalanensis Gudynas & Skuk, 1983. Zootaxa, 3681 (4), 413 - 439. https: // doi. org / 10.11646 / zootaxa. 3681.4.6","Burt, C. E. & Burt, M. D. (1933) A preliminary checklist of the lizards of South America. Transactions of the Academy of Science of St. Louis, 28, 1 - 104.","Hellmich, W. (1960) Die Sauria des Gran Chaco und seiner Randgebiete. Bayerische Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, 101, 1 - 131."]}

Tropidurus etheridgei Cei, 1982 Tropidurus hispidus: KOSLOWSKY 1898; BERTONI 1914, 1939; TALbOT 1978, 1979 Tropidurus etheridgei CEI, 1982. TYPE LOcALITY: MINA CLAVEROS, C��RDObA, ARGENTINA. AqUINO et al. 1996; MOTTE et al. 2009; CARVALHO 2013; CAbRAL & WEILER 2014; CAccIALI et al. 2016 Tropidurus torquatus hispidus: HELLMIcH 1960 Geographic distribution. CENTRAL AND SOUTHERN BOLIVIA, SOME AREAS IN WESTERN BRAZIL, PARAGUAYAN CHAcO (WEST OF PARAGUAY RIVER), AND A NARROW AREA IN THE DRY CHAcO OF ARGENTINA (FIG. 11B), Published as part of Cacciali, Pier & K��hler, Gunther, 2018, Diversity of Tropidurus (Squamata: Tropiduridae) in Paraguay-an integrative taxonomic approach based on morphological and molecular genetic evidence, pp. 511-536 in Zootaxa 4375 (4) on page 521, DOI: 10.11646/zootaxa.4375.4.3, http://zenodo.org/record/1160960, {"references":["Cei, J. M. (1982) A new species of Tropidurus (Sauria, Iguanidae) from the arid Chacoan and western regions of Argentina. Occasional Papers of the Museum of Natural History University of Kansas, 97, 1 - 10","Koslowsky, J. (1898) Enumeracion sistematica y distribucion geografica de los reptiles argentinos. Revista del Museo de La Plata, 8, 161 - 200.","Bertoni, A. de W. (1914) Fauna paraguaya: Catalogos sistematicos de los vertebrados del Paraguay. Descripcion Fisica y Economica del Paraguay, 59, 1 - 83.","Bertoni, A. de W. (1939) Catalogos sistematicos de los vertebrados de Paraguay. Revista de la Sociedad Cientifica del Paraguay, 4, 3 - 60.","Talbot, J. J. (1978) Ecological notes on the Paraguayan Chaco herpetofauna. Journal of Herpetology, 12, 433 - 434. https: // doi. org / 10.2307 / 1563636","Talbot, J. J. (1979) Una nueva lista sistematica de reptiles del Paraguay. Informes Cientificos del Instituto de Ciencias Basicas, 2, 76 - 94.","Carvalho, A. L. G. (2013) On the distribution and conservation of the South American lizard genus Tropidurus Wied-Neuwied, 1825 (Squamata: Tropiduridae). Zootaxa, 3640 (1), 42 - 56. https: // doi. org / 10.11646 / zootaxa. 3640.1.3","Cabral, H. & Weiler, A. (2014) Lista comentada de los reptiles de la Coleccion Zoologica de la Facultad de Ciencias Exactas y Naturales de Asuncion, Paraguay. Cuadernos de Herpetologia, 28, 19 - 28.","Hellmich, W. (1960) Die Sauria des Gran Chaco und seiner Randgebiete. Bayerische Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, 101, 1 - 131."]}

Tropidurus lagunablanca Carvalho, 2016 Tropidurus spinulosus: PERAccA 1895; ScHENKEL 1901; BERTONI 1939; HELLMIcH 1960; ELTER 1981. Tropidurus SP.: AqUINO et al. 1996 [PART.] Tropidurus guarani: FROST et al. 1998 [PART.]; CARVALHO 2013 [PART.]; CAccIALI et al. 2016 [PART.]. Tropidurus lagunablanca CARVALHO, 2016. TYPE LOcALITY: RESERVA NATURAL LAGUNA BLANcA, SANTA ROSA DEL AGUARAY, SAN PEDRO, PARAGUAY. Tropidurus tarara CARVALHO, 2016. TYPE LOcALITY: RESERVA NATURAL CERRADOS DEL TAGATIYA, CONcEPcI��N, PARAGUAY. Tropidurus teyumirim CARVALHO, 2016. TYPE LOcALITY: PARqUE NAcIONAL YbYcUI, PARAGUAR��, PARAGUAY. Tropidurus SP. 1: CAccIALI et al. 2016. Geographic distribution. BROADLY DISTRIbUTED IN MATO GROSSO AND MATO GROSSO DO SUL (BRAZIL), NARROWING THE DISTRIbUTION SOUTHWARDS, REAcHING THE YbYcU�� NATIONAL PARK (PARAGUAR��, PARAGUAY) AS THE SOUTHERNMOST REcORD FOR THE SPEcIES (FIG. 11D)., Published as part of Cacciali, Pier & K��hler, Gunther, 2018, Diversity of Tropidurus (Squamata: Tropiduridae) in Paraguay-an integrative taxonomic approach based on morphological and molecular genetic evidence, pp. 511-536 in Zootaxa 4375 (4) on page 521, DOI: 10.11646/zootaxa.4375.4.3, http://zenodo.org/record/1160960, {"references":["Peracca, M. G. (1895) Viaggio del dott. Alfredo Borelli nella Republica Argentina e nel Paraguay. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Universita di Torino, 10, 1 - 32.","Schenkel, E. (1901) Achter Nachtrag zum Katalog der herpetologischen Sammlung des Basler Museums. Verhandlungen der Naturforschenden Gesellschaft in Basel, 13, 142 - 199.","Bertoni, A. de W. (1939) Catalogos sistematicos de los vertebrados de Paraguay. Revista de la Sociedad Cientifica del Paraguay, 4, 3 - 60.","Hellmich, W. (1960) Die Sauria des Gran Chaco und seiner Randgebiete. Bayerische Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, 101, 1 - 131.","Elter, O. (1981) Cataloghi. V - La collezione erpetologica del Museo di Zoologia dell'Universita di Torino. Museo Regionale di Scienze Naturali. Torino, 116 pp.","Carvalho, A. L. G. (2013) On the distribution and conservation of the South American lizard genus Tropidurus Wied-Neuwied, 1825 (Squamata: Tropiduridae). Zootaxa, 3640 (1), 42 - 56. https: // doi. org / 10.11646 / zootaxa. 3640.1.3"]}

Leiocephalus sixtoi sp. nov. Figs. 2 a���d, 3 a,c, 4 a,c, 5���8 Holotype. SMF 99143, an adult male from Dunas de Ban��, near the village of Las Salinas, 18.21285 ��, - 70.53131 ��, 5 m asl, Provincia de Peravia, Dominican Republic, collected 22 March 2014 by Marcos Rodriguez Bobadilla. Field tag number GK- 5061. Paratypes. SMF 99144 ��� 51, same collecting data as the holotype; MNHNSD 23.2193���202, same locality as the holotype but collected 6 July 2012 by Marcos Rodr��guez, Luis M. D��az, and Nils Navarro. MNHNSD 23.202, SMF 99145, 99147, 99150 ��� 51 are adult males, the remaining specimens are adult females. Diagnosis. Leiocephalus sixtoi differs from all other congeners except L. schreibersii, L. melanochlorus, L. psammodromus, L. inaguae, and L. macropus by the presence of a lateral fold. It differs from L. melanochlorus, L. psammodromus, and L. macropus in having 3 internasals (vs. 2 in L. melanochlorus and L. macropus, and 4 in L. psammodromus, respectively). It further differs from L. melanochlorus and L. psammodromus in having 4 lorilabial scales anterior to the enlarged subocular (vs. 5���6). Leiocephalus sixtoi differs from L. inaguae in having a Ushaped bony parietal table (vs. V-shaped in L. inaguae), 3 or 4 enlarged postcloacal scales in males (vs. 2 in L. inaguae), most scales on the snout posterior to the internasal scales rugose to keeled (vs. smooth in L. inaguae), and a patternless throat in males, spots on the throat in females (vs. throat with dark streaks and bars in males and females of L. inaguae). Leiocephalus sixtoi differs from L. schreibersii in having the scales of the lateral fold only slightly smaller than adjacent scales (vs. scales of lateral fold distinctly smaller than adjacent scales; Fig. 4 a,b), having prominent caudal crest scales in adult males (vs. caudal crest scales of moderate size, even in very large males in L. schreibersii; Fig. 4 c,d), a pattern of dark gray bars on a grayish brown background in the region above the lateral body fold (vs. dense turquoise blue mottling with heavy suffusion of red pigment in L. schreibersii), a darker dorsal ground color (vs. paler in L. schreibersii), and a red iris in adult males (vs. pale grayish blue in adult male L. schreibersii). Leiocephalus sixtoi differs further from L. schreibersii in several osteological characters as follows: in L. sixtoi the nasal process of the premaxilla reaches to mid-level of the bony external nares (vs. to level of posterior margin of the bony external nares in L. schreibersii), lacking a constriction at the base of the nasal process of the premaxilla (vs. such a constriction is present in L. schreibersii), and having a reduced nasalprefrontal contact that leaves the nasal processes of the frontal bone exposed (vs. nasal and prefrontal bones in contact, thereby obscuring the nasal processes of the frontal bone in L. schreibersii). Description of the holotype (Figs. 5 and 6). Adult male, as indicated by partly everted hemipenes; SVL 87.0 mm; tail length 125.0 mm (incomplete); tail strongly compressed in cross section, tail height 12.3 mm, tail width 5.6 mm; axilla to groin distance 38.5 mm; head length 20.0 mm, head length/SVL ratio 0.23; snout length 8.2 mm; head width 14.0 mm; head height 11.1 mm; shank length 23.1 mm, shank length/head length ratio 1.16; scales on parietal and occipital regions large, rugose to wrinkled, juxtaposed; parietal eye visible; supraoculars in a single row, multicarinate, total number 6; circumorbital row complete, separating supraoculars from supraorbital semicircles; 3 overlapping superciliaries, posteriormost longest; canthals two; anteriormost canthal separated from nasal by a small scale; internasals 3, 2 in contact with rostral; postrostrals, much longer than wide; supralabials 4 to point level of mideye, supralabial series slightly sloping down after this point; 8 (right) / 6 (left) lorilabials in a single row; 4 (right) / 3 (left) lorilabials anterior to enlarged subocular; infralabials 5 to point below center of eye; 4 loreals in two rows; 1 preocular; single, large, elongate subocular scale, in contact with one supralabial; lateral temporals keeled, subimbricate; transverse count of gular scales 29 between tympanic openings; all gulars cycloid, smooth, imbricate, some slightly pointed; first pair of sublabials in contact with anteriormost infralabials; first pair of postmentals in contact medially; interparietal scale very elongate, subtriangular; parietal scales large, almost rectangular, inner pair almost equal to size of outer one; 2���3 scales between parietals and dorsal scales; dorsal scales of neck and body large, strongly keeled, imbricate, mucronate, keels mostly orientated longitudanally, with the vertebral scales forming a serrated crest that continues along body onto tail; lateral scales of neck somewhat heterogeneous in size and shape, mostly small, weakly keeled and subimbricate, granular at base lateral neck folds; lateral scales of body mostly strongly keeled, imbricate, mucronate, keels oriented obliquely upwards, becoming smaller and less keeled toward axilla and toward groin, respectively; scales of longitudinal fold mostly large, keeled, imbricate, micronate, except at lower base of fold where they are slightly smaller and less distinctly keeled; scales around midbody 68; vertebral crest scales from occiput to level of posterior margin of thigh 77; ventrals smooth, cycloid, imbricate, distinctly larger than dorsals; preauricular fringe present with 4���5 projecting scales; antegular, antehumeral, gular, longitudinal, postauricular, supra-auricular, and transverse antegular neck folds present; dorsal scales of forelimbs and hindlimbs keeled, mucronate, imbricate; scales on ventral surfaces of limbs smooth to slightly keeled, imbricate, cycloid to pointed; lamellae on Finger IV 15 / 16; lamellae on Toe IV 25 / 25; lateral caudal scales strongly keeled, mucronate, imbricate, dorsal caudal scales greatly enlarged, forming a high, serrated crest; ventral caudal scales in a double series of pointed scales, smooth on base of tail, becoming strongly keeled on middle and distal portions; posthumeral mite pocket present as 2���3 vertical folds [Type 1 of Torres- Carvajal (2007)]; postfemoral mite pocket absent; a pair of enlarged, elongate postcloacal scales present. Coloration in life was as follows (Fig. 7 a, b): Dorsal surface of head Olive Brown (278) with Pale Cinnamon (55) suffusions; lateral surfaces of head Army Brown (46) above, grading into Straw Yellow (53) in labial regions, and with Olive Brown (278) and Greenish Glaucous (271) splotches and suffusions; ventral surface of head Smoky White (261); dorsal surface of neck Drab (19) suffused with Salmon Color (58); dorsal suface of body Drab (19) with Raw Umber (22) longitudinal dorsolateral bands with Creme Color (12) speckles; lateral surfaces of body with Sepia (286) transverse bands and Light Bluish Gray (288) speckles; ventral surface of body Smoke Gray (266) with transverse bands composed of Medium Paris White (140) and Pratt���s Rufous (72) scales; dorsal surfaces of limbs Drab (19) with Olive Horn Color (16) speckles; dorsal surface of tail Drab (19) with Light Bluish Gray (288) and Creme Color (12) speckles; caudal crest scales Light Bluish Gray (288) with Glaucous (289) suffusions; ventral surface of tail Medium Fawn Color (257), heavily suffused with Carmine (64); iris Orange-Rufous (56). Variation. The paratypes agree well with the holotype in general appearance, morphometrics, and scalation. The largest male (SMF 99151) in the paratype series has a SVL of 88 mm, the largest female is 70 mm SVL. Variation in selected scalation characters: SAM in 68���79 (74.2 �� 3.4); median dorsal crest scales 72���80 (75.1 �� 2.5); number of IN is 3 in all, except SMF 99147 that has 4 IN; number of lorilabials anterior to enlarged subocular 3���4 (3.7 �� 0.4); number of SPL 4; number of IFL 5���6 (5.2 �� 0.4), number supraoculars 6���8 (7.0 �� 0.8); and subdigital lamellae on Phalanges II���IV of Toe IV 25���28 (26.3 �� 0.9). Only one specimen (SMF 99147) has a complete tail with a ratio tail length/SVL of 1.64. All specimens in the type series have a distinct series of dark grayish brown blotches on the lateral surfaces of the body (see Fig. 7). The ventral surface of head and neck is mostly immaculate in males, but with distinct dark spots in females. Coloration in life of an adult female (SMF 99144; Fig 7 d) was recorded as follows: Dorsal surface of head Grayish Horn Color (286) with a suffusion of Sepia (286), especially on supraoculars; lateral surfaces of head Drab (19) above, grading into Smoky White (261) in labial regions, and with Jet Black (300) blotches; ventral surface of head Smoky White (261); dorsal suface of Smoke Gray (267) with Sepia (286) paravertebral and lateral blotches with Pale Buff (1) speckles; ventral surface of body Smoky White (261) with transverse rows of Sepia (286) blotches; dorsal and lateral surfaces of limbs Grayish Horn Color (286) with Sepia (286) splotches and Pale Buff (1) speckles; ventral surfaces of limbs Smoky White (261); dorsal surface of tail Smoke Gray (267) anteriorly, grading into Ground Cinnamon (270) with Sepia (286) chevrons, bordered distally by Smoky White (261); ventral surface of tail Smoky White (261), heavily suffused with Pale Pinkish Buff (3); iris Warm Sepia (40). The almost completely everted hemipenis of SMF 99150 (Fig. 8) is a stout, medium-sized, slightly bilobed organ; sulcus spermaticus bordered by well-developed sulcal lips and opening into a broad concave area at base of apex; distal region of apex covered by large calyces, lower apex has 4���5 large flounces. Etymology. The name sixtoi is a patronym honoring our friend and colleague Sixto Inch��ustegui, who has contributed substantially to our knowledge of Hispaniolan amphibians and reptiles. Sixto is professor at the Universidad Aut��noma de Santo Domingo, Dominican Republic, where he teaches herpetology and history of biology. For more than 35 years, he has been involved as a major player in biological research and nature conservation on a national and international level. Geographic distribution. Leiocephalus sixtoi is presently only known from the surroundings of its type locality (Fig. 9). Natural history notes. The habitat at the type locality is sandy dunes with low Tropical Dry Forest (Fig. 10). This dune area streches about 15 km in length with a maximum width of about 3 km. The accumulation of sand for the formation of the dunes is a product of the waves from the Caribbean Sea and a more or less steady wind blowing mostly from the same direction. These dunes reach a maximum height of about 35 m. The vegetation in the vicinity of the dunes consists mostly of trees (Simarouba berteroana, Coccoloba uvifera, Ziziphus reticulata, and Acacia macracantha), cacti (Consolea moniliformis, Opuntia dillenii, Cylindropuntia caribaea, Opuntia antillana, Harrisia nashii, Lemairecereus hystrix, Pilocereus polygonus, Melocactus lemairei, and Mammillaria prolifera), shrubs, and grasses (Anomymous 2016). The type specimens of Leiocephalus sixtoi were collected during the morning hours on and under rocks in a grassy area. As the other species in this genus, this is a diurnal, terrestrial lizard that feeds mostly on a variety of arthropods. Designation of a neotype for Leiocephalus schreibersii (Gravenhorst 1838). According to Pregill (1992: 51), the type material of Leiocephalus schreibersii was unlocatable, apparently deposited in the Breslau Museum (= now Museum of Natural History at Wrocław University). Upon request by G.K., the present curator of herpetology at the Museum of Natural History at Wrocław University, Andrzej Jablonski, informed us that the type of L. schreibersii is not among the specimens in the collection under his care. Therefore, we consider it to be lost. We designate SMF 26228 (Figs. 11 and 12), an adult male from Saint Marc, Province Artibonite, Haiti, collected 19 April 1939 by Robert Mertens, as the neotype of L. schreibersii., Published as part of K��hler, Gunther, Rodr��guez Bobadilla, Marcos J. & Hedges, S. Blair, 2016, A new dune-dwelling lizard of the genus Leiocephalus (Iguania, Leiocephalidae) from the Dominican Republic, pp. 517-532 in Zootaxa 4121 (5) on pages 519-527, DOI: 10.11646/zootaxa.4121.5.2, http://zenodo.org/record/257253, {"references":["Torres-Carvajal, O. (2007) A taxonomic revision of South American Stenocercus (Squamata: Iguania) lizards. Herpetological Monographs, 21, 76 - 178. http: // dx. doi. org / 10.1655 / 06 - 001.1","Gravenhorst, J. L. C. (1838) Beitrage zur genaueren Kenntniss einiger Eidechsengattungen. Nova Acta Academiae Caesareae Leopoldino-Carolinae Germanicae Naturae Curiosorum. Verhandlungen der Kaiserlich Leopoldinisch-Carolinischen Deutschen Akademie der Naturforscher, 18, 712 - 784. [1837]","Pregill, G. K. (1992) Systematics of the West Indian lizard genus Leiocephalus (Squamata: Iguania: Tropiduridae). Miscellaneous Publication, University of Kansas, Museum of Natural History, 84, 1 - 169."]}

Pseudogekko atiorum sp. nov. (Figs. 1, 3��� 6) Holotype. PNM 9518 (CDS Field No. 954, formerly KU 302818), adult female, collected at 22:00 h on 21 February 2005, on Mt. Talinis, Barangay Bongbong, Municipality of Valencia, Negros Oriental Province, Negros Island, Philippines (N: 9.26000��, E: 123.20327 ��; WGS 84; 1100 m elev.), by CDS. Paratypes. Two adult females (CAS-SUR 19372, 21122) collected from Municipality of Cauayan, Negros Occidental Province, Negros Island, Philippines by A.C. Alcala and Q. Alcala. One adult female (CAS 138097) collected by Q. Alcala and two adult males (CAS 145793, 147491) collected by Q. Alcala and M. Pinero, respectively, from Municipality of Pamplona, Negros Oriental Province, Negros Island, Philippines. One adult female (CAS 134269) collected by A.C. Alcala and one adult male (CAS-SUR 26778) collected by D. Empeso from Municipality of Siaton, Negros Oriental Province, Negros Island, Philippines. Four adult males (CAS 128956, 128959, 128963, 128971) collected from the Municipality of Sibulan, Negros Oriental Province, Negros Island, Philippines by L.C. Alcala. Two adult females (CAS 134292, TNHC 62478) collected by A.C Alcala and RMB, respectively, and one juvenile female (KU 327770) collected by RMB from Municipality of Valencia, Barangay Bongbong, Negros Oriental Province, Negros Island, Philippines. One adult male (CAS 145710) collected from Municipality of San Juan, Siquijor Province, Negros Island, Philippines by L.C. Alcala. Other Referred Specimens. Five specimens (CAS 89812, 147523, CAS-SUR 24588���24590) collected from Municipality of Cauayan, Negros Occidental Province, Negros Island, Philippines. Three specimens (CAS 134298, 137789, CAS-SUR 26776) collected from Negros Occidental Province, Negros Island, Philippines. Twenty specimens (CAS 137653, 137672, 137790, 138096, 138134, 139258, 145791, 145792, 145804 ���6, 145875, 145876, 145976, 173760, 185991, 200539, 200540, CAS-SUR 24323, 24829) collected from Municipality of Pamplona, Negros Oriental Province, Negros Island, Philippines. One specimen (CAS-SUR 26775) collected from Municipality of Siaton, Negros Oriental Province, Negros Island, Philippines. Thirty specimens (CAS 128955, 128957, 128958, 128960���62, 128964���70, 128972���78, 131796, 132609, 142039��� 46) collected from Municipality of Sibulan, Negros Oriental Province, Negros Island, Philippines. Thirteen specimens (CAS 133055, 137791, 200538, CAS-SUR 21803, 21839, 21840, 21856, 21857, 21872, 22218 ���20, 24385) collected from Negros Oriental Province, Negros Island, Philippines. Diagnosis. Pseudogekko atiorum can be distinguished from congeners by the following combination of characters: (1) body size moderate (SVL 41.1���52.5 mm); (2) axilla���groin distance moderate (19.4���28.4 mm); (3) head length moderate (7.4���9.4 mm); (4) snout length long (4.3���5.8 mm); (5) Toe-IV scansors 14���17; (6) paravertebral scales 226���240; (7) ventral scales 119���129; (8) supralabials 15���17; (9) infralabials 12���15; (10) circumorbitals 35���38; (11) precloacal pores 13���15; (12) femoral pores absent; (13) dominant body coloration dark brown; (14) conspicuous head spotting present, cream; (15) conspicuous dorsolateral spotting present, cream; (16) conspicuous limb spotting absent; (17) tail banding absent; (18) body striping absent; (19) interorbital banding present, light brown; (20) iris ring coloration absent (Tables 1, 2; Figs. 3���6). Comparisons. Characters distinguishing Pseudogekko atiorum from all other species of Pseudogekko are summarized in Tables 1���3. Pseudogekko atiorum most closely resembles P. b rev i pe s; however, it differs from this species by having a longer snout���vent length in both males (41.1���52.5 mm vs. 39.0) and females (44.8���48.7 mm vs. 34.5���42.4), longer eye���nares distance (3.5���4.5 mm vs. 3.0��� 3.4), a tendency towards having both a longer snout length (4.3���5.8 mm vs. 3.8���4.4) and internares distance (1.5���1.9 mm vs. 1.2���1.6), and a greater number of supralabials (15���17 vs. 13 or 14), circumorbitals (35���38 vs. 33���35), paravertebrals (226���240 vs. 211���218), ventrals (119���129 vs. 96���117), and precloacal pores (13���15 vs. 12; Figs. 3���5). Pseudogekko atiorum can be distinguished from P. chavacano by having a shorter snout���vent length (41.1��� 52.5 mm vs. 54.7���55.9), longer head length (7.4���9.4 mm vs. 10.4), a tendency towards a shorter snout length (4.3��� 5.8 mm vs. 5.8 ���6.0), fewer infralabials (12���15 vs. 16 or 17), circumorbitals (35���38 vs. 46), and precloacal pores (13���15 vs. 16), greater number of paravertebrals (226���240 vs. 195���197), cream colored (vs. neon green) head and dorsolateral spotting, absence (vs. presence) of limb spotting and tail banding, and presence of interorbital banding (vs. absence); from P. compresicorpus by having a shorter snout���vent length (41.1���52.5 mm vs. 54.9���59.7), a tendency towards both a shorter axilla���groin distance (19.4���28.4 mm vs. 27.0��� 32.6) and head length (7.4���9.4 mm vs. 9.2���11.3), fewer Toe-IV scansors (14���17 vs. 18 or 19) and circumorbitals (35���38 vs. 39���45), cream colored (vs. neon green) head and dorsolateral spotting, absence (vs. presence) of limb spotting and a light blue iris ring, and presence (vs. absence) of interorbital banding; from P. di t o y by having fewer infralabials (12���15 vs. 16 or 17) and circumorbitals (35���38 vs. 40���43), greater number of paravertebrals (226���240 vs. 180���185) and ventrals (119���129 vs. 111���118), fewer precloacal pores (13���15 vs. 18), and the presence (vs. absence) of head spotting, dorsolateral spotting, and interorbital banding; from P. pungkaypinit by having a shorter snout���vent length (41.1���52.5 mm vs. 66.6���75.3), axilla���groin distance (19.4���28.4 mm vs. 37.2���41.2), total length (80.2���113.5 mm vs. 125.3���141.2), midbody width (4.8���7.7 mm vs. 7.7���9.1), head length (7.4���9.4 mm vs. 11.4���13.6), head width (5.9���8.3 mm vs. 9.3���11.2), and snout length (4.3���5.8 mm vs. 6.7���7.5), fewer Finger-III scansors (12���14 vs. 15���17), infralabials (12���15 vs. 17���19), circumorbitals (35���38 vs. 50���55), paravertebrals (226���240 vs. 265���280), and precloacal pores (13���15 vs. 17���20), a tendency towards having fewer ventrals (119���129 vs. 125���155), the presence (vs. absence) of head and dorsolateral spotting and interorbital banding, and the absence (vs. presence) of body striping; from P. smaragdinus by a tendency towards having a shorter snout���vent length (41.1���52.5 mm vs. 50.2���64.3) and axilla��� groin distance (19.4���28.4 mm vs. 26.2���35.5), fewer Finger-III scansors (12���14 vs. 15���18), paravertebrals (226��� 240 vs. 241���252), and enlarged pores (13���15 vs. 32���41), a tendency towards having fewer Toe-IV scansors (14���17 vs. 16���22) and supralabials (15���17 vs. 16���19), greater number of circumorbitals (35���38 vs. 33���35), presence of dark brown (vs. yellow to orange [undisturbed] or neon green [disturbed]) body coloration, cream colored (vs. large black and small white) head and dorsolateral spotting, presence (vs. absence) of interorbital banding, and absence (vs. presence) of limb spotting and tail banding. Description of holotype. Adult female in excellent condition; small incision in the sternal region (portion of liver removed for genetic sample). Body small, slender, SVL 44.8 mm; limbs well developed, moderately slender; tail slender; margins of limbs smooth, cutaneous flaps or dermal folds absent. Head size moderate, slightly differentiated from neck, characterized by only slightly hypertrophied temporal and adductor musculature; snout broadly rounded in dorsal view and bluntly rounded in lateral view; HW 96.9 % MBW, 83.6 % HL; HL 18.4 % SVL; SNL 70.7 % HW, 59.2 % HL; dorsal surfaces of head relatively homogeneous, with slightly to moderately pronounced concave postnasal, internasal, prefrontal, and interorbital concavities; auricular opening small, nearly round, beneath temporal swellings on either side of head; tympanum deeply sunken; orbit large; eye large, pupil vertical, margin wavy; limbs and digits relatively short; thighs moderately thicker compared to upper arms; TBL 8.0% SVL, 57.9 % FL. Rostral size moderately large, margin in wide heart shape in anterodorsal view, nearly as broad as high, sutured anterolaterally with anteriormost enlarged supralabials, projecting onto dorsal surface of head to point in line with anteriormost edge of nasal; nasal surrounded by first labial, rostral, one enlarged postnasal, and two enlarged supranasals; medial supranasals separated by 3 small median scales; enlarged supranasals moderately larger than postnasals; circumorbitals 36 in total. Total number of differentiated supralabials 15, bordered dorsally by one row of differentiated, slightly enlarged snout scales; total number of differentiated infralabials 12, bordered ventrally by 2 or 3 rows of slightly enlarged scales; chin scales enlarged; postrictal scales undifferentiated; gulars small, round, nonimbricate. Dorsal cephalic scales fairly homogeneous in size, shape, disposition, and distribution; cephalic scalation slightly convex, round to oval scales; postnasal, prefrontal, internasal, and interorbital depressions; undifferentiated posterior head scales granular, slightly convex. Axilla���groin distance 55.1 % SVL; undifferentiated dorsal body scales round to tubular, moderately to sharply convex, juxtaposed, relatively homogeneous in size, many with raised edge, irregular in placement on individual scales across body; each dorsal scale surrounded by 4���6 interstitial granules; dorsals gradually transition to imbricate ventrals along lateral body surface; paravertebrals between midpoints of limb insertions 232; ventrals between midpoints of limb insertions 128; scales on dorsal surfaces of limbs more imbricate than dorsals; scales on dorsal surfaces of hands and feet similar to dorsal limb scales, heavily imbricate; ventral body scales flat, cycloid, strongly imbricate, much larger than lateral or dorsal body scales, relatively homogeneous in size. Fourteen enlarged scales in continuous precloacal series (pore-bearing series in males), arranged in a widely obtuse, ���V���-formation; patch of slightly enlarged scales posterior to precloacal series, roughly three scale rows in width, 3���4 scale rows in length, forming an oval patch just posterior to precloacal series. Digits moderately expanded and covered on palmar and plantar surfaces by bowed, unnotched, undivided scansors; digits with minute vestiges of interdigital webbing; subdigital scansors of fingers (left/right) I (9,8), II (10,10), III (13,13), IV (14,14), V (9,9); subdigital scansors of toes (left/right) I (9,9), II (10,10), III (13,13), IV (15,15), V (12,12); mite eggs visible between distal subdigital scansors of digits on hands and feet; subdigital scansors of hands and feet bordered proximally (on palmar and plantar surfaces) by two or three slightly enlarged scales that form a near-continuous series with enlarged scansors; all digits clawed, but first (inner) claw greatly reduced; remaining terminal claw-bearing phalanges compressed, with moderately sized recurved claws. brevipes atiorum chavacano compresicorpus ditoy pungkaypinit smaragdinus sp. A (1 m, 3 f) (8 m, 7 f) (1 m, 1 f) (3 m, 5 f) (1 m, 1 f) (4 m, 2 f) (16 m, 17 f) (1 m, 1 f) individuals showing specific counts are given in parentheses. Characters summarized for adult specimens only. When coloration in life has not been documented, characters are listed as unkown. Enlarged pore series 12 (precloacal) 13���15 16 (precloacal) 10���14 (precloacal) 18 (precloacal) 17���20 (precloacal) 32���41 15 (precloacal) (precloacofemoral) (precloacal) Femoral pores ��� ��� ��� ��� ��� ��� + ��� ������continued on the next page Tail relatively long compared to body, TL 44.6 mm, 99.4 % SVL; round, not heavily depressed; TH 87.8 % TW; caudals similar in size to dorsals, subcaudals similar in size to ventrals. Coloration of holotype in preservative. Background dorsal trunk coloration light tan with intermittent small cream and dark brown speckles, continuing down the tail; dorsal region of the head with similar color pattern, except for a pronounced dark brown area above the orbits; a ridge of raised, dark brown scales present in the postnasal region; dorsal and lateral regions of the limbs with the same color pattern as the trunk; supralabials with the same pattern as the head, whereas the infralabials are a solid cream with few medium brown speckles; ventral side of the trunk has a cream background color, with the speckling pattern of the lateral side of the body wrapping around to approximately halfway to the midpoint of the venter; ventral side of the head has only sparse medium brown speckling; ventral region of the tail is solid cream with occasional light brown speckles that increase in intensity towards the tail tip; ventral regions of the limbs, hands, and feet are cream with medium to dark brown speckles scattered sparsely throughout; the space between the scansors is a medium brown. Coloration of holotype in life. Dorsal ground color of head, trunk, and tail chocolate brown; gradual transition to lighter coloration towards tip of the snout; light mottling posterior to eyes and along neck; dorsal limb surfaces colored as trunk, but with randomly distributed light blotches; tail lighter colored than trunk, but with series of longitudinally elongate, paravertebral cream blotches. Measurements and scale counts of holotype in mm. SVL 44.8; axilla���groin distance 24.7; total length 89.4; tail length 44.6; head length 8.3; head width 6.9; head height 4.7; eye diameter 3.1; snout length 4.9; eye���nares distance 4.0; internarial distance 1.6; interorbital distance 4.0; midbody width 7.1; femur length 6.2; tibia length 3.6; Toe-I length 2.1; Toe-IV length 3.9; tail width 2.5; tail height 2.2; supralabials 15; infralabials 12; circumorbitals 36; paravertebral scales 232; ventral scales 128; Finger-III scansors 13; Toe-IV scansors 15. Distribution, ecology and natural history. Pseudogekko atiorum is known only from Negros and Siquijor islands, although it is possible that it will eventually be discovered on other islands in the Western Visayan Faunal Region (i.e., Cebu, Panay; Fig. 1; Ferner et al., 2001). Pseudogekko atiorum represents the only species of Philippine false gecko distributed in the western Visayan islands, with the remaining diversity in the genus restricted to the northern, eastern, and sounthern islands of the archipelago (Siler et al., 2014 b, c). Although believed to be primarily an obligate forest species, at least one individual has been collected recently on the leaves of a banana plant at mid-elevation on Mt. Talinis in southeast Negros Island (PNM 9518). This individual was found within a small patch of disturbed forest where an agricultural plot had been established, surrounded by pristine secondary growth forest. Among all members of the genus, P. a t i or um is the best represented in museum collections, with 87 individuals housed at CAS and KU alone. Although historical collections reported this species to be quite common, recent expeditions over the last decade resulted in few observations of this Western Visayan gecko in the wild. Due to the absence of data allowing for confirmation of potential declines in population size and health resulting from widespread habitat destruction, and the lack of available information about the species ecology and intraspecific diversity, we do not find this species qualifies for Critically Endangered, Endangered, Vulnerable, or Near Threatened status under the IUCN criteria for classification (IUCN, 2013). Therefore, we recommend that the species be considered Least Concern at the current time, but strongly recommend that immediate efforts be made to study the health of wild populations of this species on Negros Island and, especially, Siquijor Island where no original forest persists (CDS, RMB, personal observations) but a small patch of protected secondary forest may protect this species. Etymology. We derive the new species name in honor of the Ati people, a Negrito ethnic group believed to be among the first to colonize the central Philippines. Similar to this new species of Pseudogekko, Inati (the native language of the Ati), is infrequently encountered and possibly threatened, and is estimated to be spoken by fewer than 2,000 people. The specific epithet is a noun of masculine gender. Suggested common name: Central Visayan False Gecko., Published as part of Davis, Drew R., Watters, Jessa L., K��hler, Gunther, Whitsett, Collin, Huron, Nicholas A., Brown, Rafe M., Diesmos, Arvin C. & Siler, Cameron D., 2015, Redescription of the rare Philippine false gecko Pseudogekko brevipes (Reptilia: Squamata: Gekkonidae) and description of a new species, pp. 357-374 in Zootaxa 4020 (2) on pages 363-371, DOI: 10.11646/zootaxa.4020.2.7, http://zenodo.org/record/242048, {"references":["Ferner, J. W., Brown, R. M., Sison, R. V. & Kennedy, R. S. (2001) The amphibians and reptiles of Panay Island, Philippines. Asiatic Herpetological Research, 9, 34 - 70.","Siler, C. D., Dececchi, T. A., Merkord, C. L., Davis, D. R., Christiani, T. J. & Brown, R. M. (2014 b) Cryptic diversity and population genetic structure in the rare, endemic, forest-obligate, slender geckos of the Philippines. Molecular Phylogenetics and Evolution, 70, 204 - 209. http: // dx. doi. org / 10.1016 / j. ympev. 2013.09.014","Siler, C. D., Welton, L. J., Davis, D. R., Watters, J. L., Davey, C. S., Diesmos, A. C., Diesmos, M. L. & Brown, R. M. (2014 c) Taxonomic revision of the Pseudogekko compresicorpus Complex (Reptilia: Squamata: Gekkonidae), with descriptions of three new species. Herpetological Monographs, 28, 110 - 139. http: // dx. doi. org / 10.1655 / herpmonographs-d- 14 - 00005"]}

Key to the species of Dactyloa occurring in Panama (modified from Lotzkat et al. 2013) 1 a. Large species SVL> 95 mm, body robust, body pattern different: with broad bars, blotches, spots, or ocelli; if homogeneously green, then without narrow dark oblique lines on flanks....................................................... 2 1 b. Relatively small species SVL Dactyloa ginaelisae 4 b. Legs very short: tip of fourth toe of hind limb (adpressed along straightened specimen) reaching to a point between shoulder and ear, shank length/SVL ratio Dactyloa microtus 5 a. Subdigital lamellar pads strongly dilated, more than three times width of distal phalanx; more than 50 lamellae under fourth toe (base of digit to claw); 190 or more scales around midbody; scales on anterodorsal surface of thigh smooth, only on anterior edge unicarinate with a few bi- or tricarinate; male dewlap orange or red, female dewlap brownish, mottled...................................................................................................... Dactyloa insignis 5 b. Subdigital lamellar pads barely dilated, less than two times width of distal phalanx; 35 or fewer lamellae under fourth toe (base of digit to claw); fewer than 150 scales around midbody; scales on anterodorsal surface of thigh unicarinate; male dewlap white with an orange-yellow margin; female dewlap white with a light yellow margin............... Dactyloa kunayalae 6 a. Series of dark blotches or ocelli form oblique transverse bands on dorsum, flanks, limbs, and anterior portion of tail; a pronounced light-colored interorbital bar with dark anterior and posterior borders; light and dark stripes radiating from eye in all directions; male dewlap cream white, female dewlap brown.................................................... 7 6 b. Coloration variable: unicolor, mottled, or with broad dark transverse bands; no pronounced interorbital bar; a dark pre- and/or postorbital stripe often present, but no light and dark stripes radiating from eye in all directions; male dewlap white with yellow and blue or green scales, female dewlap with contrasting yellow and green or blue striped or reticulate pattern............................................................................................... Dactyloa casildae 7 a. Enlarged superciliaries restricted to anterior half of eye, followed by small granular scales posteriorly, not forming a stiff ridge....................................................................................... Dactyloa frenata 7 b. All of upper margin of eye with enlarged superciliaries forming a stiff ridge......................... Dactyloa latifrons 8 a. Relatively tiny species SVL Dactyloa chloris 8 b. Somewhat larger species SVL> 60 mm, body green, with a parallel series of oblique narrow dark lines, broad bands or disperse spots on flanks; ventrals smooth; long-legged��� 4 th toe of adpressed hind limb reaches at least to anterior border of eye. 9 9 a. Flanks with a parallel series of oblique broad bands that are usually formed by ocelli; usually with 50���80 total loreal scales; species distributed in eastern Panama..................................................................... 10 9 b. Flanks with a parallel series of oblique narrow dark lines that are never formed by blotches or ocelli; usually with 38���54 total loreal scales; species distributed in central and western Panama.................................... Dactyloa ibanezi 10 a. Flanks in males with oblique rows of ocelli or blotches, in females with scattered dark green spots; male dewlap orange with white anterior margin; 12���14 (mean 12.9) scales between first canthals and 7���13 (mean 10.2) between second canthals; species distributed in the border between Panama and Colombia, in the mountain ranges of Pirre, and Jingurud��, and the southern corner of the Dari��n mountain range....................................................... Dactyloa purpurescens 10 b. Flanks in males and females with rows of ocelli in oblique broad dark green bands; male dewlap orange with continuous white margin; 12���18 (mean 14.5) scales between first canthals and 10���16 (mean 12.3) between second canthals; species distributed in eastern Panama in the mountain ranges of Maj��, Piedras-Pacora, and San Blas, and the central and northern portions of the Dari��n mountain range...................................................................... Dactyloa maia, Published as part of Batista, Abel, Vesely, Milan, Mebert, Konrad, Lotzkat, Sebastian & K��hler, Gunther, 2015, A new species of Dactyloa from eastern Panama, with comments on other Dactyloa species present in the region, pp. 57-84 in Zootaxa 4039 (1) on page 75, DOI: 10.11646/zootaxa.4039.1.2, http://zenodo.org/record/289962, {"references":["Lotzkat, S., Hertz, A., Bienentreu, J. F. & Kohler, G. (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. http: // dx. doi. org / 10.11646 / zootaxa. 3626.1.1"]}

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"]}

Dactyloa maia sp. nov. Figures 1, 4 E���F, 5 E���F, 6 D, 8, 9, 10. Holotype. SMF 97268, adult male (Figs. 4 E, 5 E, 6 D, 9), from the ridge of the Serran��a de Dari��n (Fig. 1) along the trail that connects the Comarca Wargand�� and the Comarca Guna Yala, about 10 km northeast of the village Nurra, 9.06142 �� N, 77.97961 �� W, 344 m asl., Corregimiento de Nurra, Comarca Wargand��, Panama; collected by Abel Batista and Milan Vesely on 0 3 October 2012; original field number AB 760. Paratypes. All from Panama: SMF 97269, a male, from Cerro la Javillosa Ambroya, Torti, Chepo, Panama province, collected on 28 September 2012, 19: 39 hrs, 8.92267 �� N, 78.62530 �� W, 851 m asl, collected by Abel Batista and Milan Vesely; MHCH 2782, a female, same collecting data as holotype; MHCH 2781 and MHCH 2783, females, respectively, from Cerro Pechito Parado, Bajo peque��o, Lajas blancas, C��maco, Comarca Ember��- Woun��an, on 0 7 November 2012, 8.47911 �� N, 77.52799 �� W, 718 m asl, collected by Abel Batista; SMF 97270, a female from la Cascada trail, Burbayar private reservation, Cart��, Nargan��, Comarca Guna Yala, on 26 November 2012, 9.31577 �� N, 79.00580 �� N, 322 m asl, collected by Abel Batista and Konrad Mebert. Diagnosis. A moderate-sized species (SVL 62���78 mm) of the genus Dactyloa, D. latifrons species group (sensu Nicholson et al. 2012), that is most similar in external morphology to D. pupurescens, D. limon, and D. ibanezi, and according to molecular evidence is most closely related to D. pupurescens and D. ibanezi (Fig. 7). These four species share a moderate adult size (SVL 62���88 mm); a large dewlap; a peculiar flank pattern in males, which is green with dark oblique bands, or blotches, or ocelli always arranged in oblique rows (Fig. 5); enlarged postcloacal scales in males; and smooth ventrals. Dactyloa maia can readily be distinguished from these three species by its color pattern and morphology (Figs. 3���6; Table 1���2), and from the remaining species of Dactyloa within the D. latifrons group by its moderate size (SVL Dactyloa maia can be differentiated from D. limon by its male dewlap coloration which is orange with an uninterrupted white margin (vs. yellowish near the throat and tan on distal portion, or uniformly light tan in D. limon; Fig. 4). Dactyloa maia further differs from D. purpurescens, D. limon and D. ibanezi in the color pattern of the flanks as follows: Dactyloa maia has oblique rows of turquoise ocelli or oblique dark green bands without sexual dimorphism (Fig. 8); in D. limon, males have wide dark green bands on the flanks whereas females have diffuse dark green spots distributed evenly or randomly; males of D. purpurescens exhibit oblique rows of ocelli or blotches whereas females have dark green spots arranged in oblique rows; in D. ibanezi both sexes exhibit oblique thin black lines. The hemipenis of D. maia is a small, unilobate organ (slightly bilobate in D. ibanezi; no information available for D. limon and D. purpurescens). Also, Dactyloa maia differs from D. purpurescens, D. limon, and D. ibanezi in mean values of several morphological characters as follows (mean values for D. purpurescens, D. limon, and D. ibanezi presented in that order): HL/SVL 0.27 in Dactyloa maia vs. 0.25, 0.23 and 0.26,.LST 64.7 in Dactyloa maia vs. 62.8, 43.7, and 44.5; number of scales between first and second canthals 14.5 / 12.3 (first canthals/second canthals) in Dactyloa maia vs 12.9 / 10.2, 10 / 8.6, and 11.5 / 10. Description of the holotype. Adult male as indicated by everted hemipenes, a pair of enlarged postcloacal scales, and presence of a large dewlap (Figs. 3 E, 4 E, 7 D, 9); SVL 76 mm; tail length 178 mm (tail complete), tail length/SVL ratio 2.34; tail laterally compressed in cross section, tail height 2.2 mm, tail width 2.0 mm; axilla to groin distance 35.6 mm; head length 19.2 mm, HL/SVL ratio 0.25; snout length 9.3 mm; head width 9.7 mm; longest toe of adpressed hind limb reaching posterior margin of orbit; shank length 20.7 mm, shank length/SVL ratio 0.27, shank length/HL ratio 1.08; tip of longest finger of extended forelimb reaching tip of snout; longest finger of adpressed forelimb not reaching to anterior insertion of hind limbs; prefrontal ridges distinct, parietal ridges conspicuous; scales on snout mostly keeled; 6 postrostrals; 7 scales between nasals; scales in distinct prefrontal depression smooth; supraorbital semicircles differentiated, composed of smooth scales, separated by a minimum of 2 scales; supraorbital disc composed of 7 enlarged smooth scales; two elongated, smooth anterior superciliaries, followed posteriorly by a much smaller, elongate scale; about 5 rows of small keeled scales extending between enlarged supraorbitals and superciliaries; interparietal plate distinct, parietal eye visible; canthal ridge distinct, composed of 3 large (posterior) and 7 small (anterior) canthal scales; 11 scales present between second canthals; 13 scales present between posterior canthals; 78 loreal scales arranged in 7 horizontal rows; subocular scales flat, subocular row well-defined; 9 supralabials to level below center of eye; ear opening 0.73 x 1.6 mm (length x height); mental distinctly wider than long, almost completely divided medially, bordered posteriorly by 6 postmentals; 10 infralabials to level below center of eye; third and fourth sublabials posterior to mental slightly enlarged; keeled granular to elongate scales present on chin and throat; dewlap large, extending well onto body, anterior insertion is about halfway between nose and orbit, posterior insertion at a level between one-third and one half of the distance between axilla and groin, with about 4 gorgetal-sternal rows, each 2���3 scales wide, becoming less regular posteriorly; low nuchal crest present, dorsal crest barely visible; dorsum of body with keeled scales, 1���2 middorsal rows of prominently keeled, but not otherwise enlarged scales; about 50 medial dorsal scales in one HL; about 125 medial dorsal scales between levels of axilla and groin; lateral scales small, long and keeled; ventrals at midbody smooth, subimbricate; about 66 ventral scales in one HL; about 87 ventral scales between axilla and groin; about 164 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 0.88 x 1.90 mm (length x width); tube-like axillary pocket not developed; scales on anterodorsal surface of thigh and on dorsal surface of forearm keeled; digital pads dilated, dilated pad about 3 times width of nondilated scales under distal phalanx; distal phalanx narrower than and raised from dilated pad; 31 / 31 (left/right) lamellae under phalanges ii to iv of 4 th toe; 11 / 12 scales under distal phalanx of 4 th toe; 21 / 19 lamellae under phalanges ii to iv of 4 th finger; 10 / 9 scales under distal phalanx of 4 th finger. Hemipenis description: The completely everted hemipenis (Fig. 10) of SMF 97269 is a small, unilobate organ; sulcus spermaticus bordered by well-developed sulcal lips, opening at base of apex into a small concave area; large asulcate processus and ridge present; a prominent fleshy fringe present on each lateral side of truncus; most of apex on asulcate side and distal portion finely calyculate, truncus with transverse folds.. Coloration in life of the holotype. (Fig. 4 E) Dorsal and lateral ground color of body and limbs Light Grass Green (109); lateral surface of body with three Parrot Green (121) oblique bands directed backwards as well as downwards, each with six to seven Medium Blue (168) small oval spots; tail with Dark Green (136) transverse bands; eyelids Cream Yellow (82); iris Chrome Orange (74); ventral surfaces dirty white, suffused with Pale Cyan (157); dewlap Light Pratt���s Rufous (71) with three well defined longitudinal series of Emerald Green (143) scales, free margin of dewlap dirty white, anterior insertion of dewlap suffused with Beige (254). Coloration of the holotype after approximately two years of preservation in 70 % ethanol. (Fig. 9) Dorsal and lateral ground color of body and limbs Lavender (195); lateral surfaces of body with three oblique rows of small oval Medium Water Blue (182) spots directed backwards as well as downwards; tail with Plumbeous (295) transverse bands; ventral surfaces dirty white, suffused with Medium Blue Gray (194); dewlap Pale Sulfur Yellow (92), its base suffused with Pale Neutral Gray (296). Color variation. Another male (SMF 97269, 8E���F), at the moment of encounter, was uniform green with bluish spots on flanks arranged in oblique rows; after collection it exhibited metachrosis, and the coloration recorded was as follows: the dorsal ground color of body, head and limbs Salmon Color (58), grading into Yellow Green (103) toward the flanks; tail with Russet (44) transverse bands; four well-defined Olive Green (123) oblique bands between axilla and groin, each including four to seven irregular Cyan Black (153) blotches; ventral surfaces and dewlap as in holotype. A female (MHCH 2782, Figs. 4 F, 5 F, 8 A���B) lacked the small oval spots, and only the three Parrot Green (121) transverse bands were present with three small Sulphur Yellow (80) spots between each band; dewlap Medium Chrome Orange (75) with three well defined longitudinal series of Emerald Green (143) scales, free margin of dewlap dirty white, anterior insertion of dewlap suffused with Light Neutral Gray (297). Another female (SMF 97270; Fig. 8 G) agrees in general color pattern with female MHCH 2782. A third female (MHCH 2781; Fig. 8 C���D) has the same dorsal ground color as the holotype, but with six well defined Greenish Olive (125) oblique bands in the flanks (without spots); dewlap Olive (126), with three well defined longitudinal series of Lime Green (116) scales, anterior and external border of dewlap dirty white, anterior insertion of dewlap suffused with Light Neutral Gray (297). Distribution and Natural history. As far as we know, Dactyloa maia is endemic to eastern Panama, inhabiting the foothills and ridges of the Maj��, San Blas, Dari��n (as far south and east as to R��o Tuquesa, thereafter apparently replaced by D. purpurescens), and Piedras-Pacora (two specimens photographed by Angel Sosa- Bartuano, not plotted in Fig. 1) mountain ranges, where it occurs in the eastern Panamanian montane forest (Fund 2014) and the Isthmian-Atlantic moist forests (in the Piedras-Pacora and San Blas mountain range, Hogan & Fund 2014), at 322���852 m asl. All specimens of Dactyloa maia were found during night searches sleeping on branches or leaves 2 to 3 m above the ground. Etymology. Abel Batista dedicates this beautiful new species to his recently born daughter, Maia. The name also comes from Greek mythology, where it is applied to the eldest of the Pleiades, sometimes called mountain nymphs, and are believed to live on the trees in mountains and groves as the guardians for that habitat, a role Dactyloa maia could also represent., Published as part of Batista, Abel, Vesely, Milan, Mebert, Konrad, Lotzkat, Sebastian & K��hler, Gunther, 2015, A new species of Dactyloa from eastern Panama, with comments on other Dactyloa species present in the region, pp. 57-84 in Zootaxa 4039 (1) on pages 65-70, DOI: 10.11646/zootaxa.4039.1.2, http://zenodo.org/record/289962, {"references":["Nicholson, K. E., Crother, B. I., Guyer, C. & Savage, J. M. (2012) It is time for a new classification of anoles (Squamata: Dactyloidae). Zootaxa, 3477, 1 - 108.","Hogan, C. & Fund, W. (2014) Isthmian-Atlantic moist forests. Available from: http: // www. eoearth. org / view / article / 153927 (accessed 23 September 2015)"]}

Stenocercus amydrorhytus sp. nov. Figs. 1 a, 2���4 Stenocercus cf. ornatissimus: Lehr 2002: p. 87 Holotype. MUSM 20221, an adult male from Coris, -9.83000, - 77.73383, 2785 m, Departamento Ancash, Provincia Aija, Peru, collected 11 February 1997 by Edgar Lehr. Field tag number EL 140. Paratypes. 6 (all females). All from Coris, Departamento Ancash, Provincia Aija, Peru, collected 11 February 1997 by Edgar Lehr: MUSM 20222, - 9.81183, -77.72500, 2900 m. MUSM 20224, - 9.82867, - 77.73250, 2810 m. SMF 80228, - 9.80305, - 77.73195, 2890 m. SMF 80229, - 9.81112, - 77.72750, 3015 m. MUSM 20223, SMF 80230, - 9.80305, - 77.73112, 3075 m. Diagnosis. Stenocercus amydrorhytus differs from all other congeners except S. ivitus, S. chrysopygus, S. melanopygus, S. modestus, S. orientalis, and S. stigmosus by the combination of (1) a poorly developed oblique neck fold with a shallow or no mite pocket underneath (Fig. 1); (2) a well developed antehumeral fold with a deep mite pocket underneath; (3) scales on posterior surface of thighs granular; (4) vertebral scales similar in size and shape to adjacent scale rows; (5) three caudal whorls per autotomic segment. Stenocercus amydrorhytus differs from S. ivitus in having lateral nuchals less than half the size of dorsal nuchals (vs. lateral nuchals similar in size to dorsal nuchals in S. ivitus). Stenocercus amydrorhytus differs from S. orientalis in having predominantly smooth dorsal head scales (vs. prominently keeled dorsal head scales in S. orientalis). Stenocercus amydrorhytus differs from S. modestus in having 42���47 scales around midbody (vs. fewer than 40 in S. modestus) and by having a tail length less than 70 % of total length (at least 70 % in S. modestus). Stenocercus amydrorhytus differs from S. melanopygus in having a poorly developed oblique neck fold with a shallow or no mite pocket underneath (vs. oblique neck fold absent in S. melanopygus). Stenocercus amydrorhytus differs from S. stigmosus in having a mostly yellowish venter without blotches or spots in the only known adult male (vs. males with distinct black spots on venter in S. stigmosus) and in having larger dorsal scales, 44���52, mean 47.3, vertebral scales (vs. dorsal scales smaller, 51���61, mean 56.4, vertebral scales in S. stigmosus), 42���47, mean 44.3, scales around midbody (vs. 49���57, mean 52.7, in S. stigmosus). Stenocercus amydrorhytus differs from S. chrysopygus in having larger dorsal scales, 44���52, mean 47.3, vertebral scales (vs. dorsal scales smaller, 54���86, mean 66.0, vertebral scales in S. chrysopygus), 42���47, mean 44.3, scales around midbody (vs. 48���82, mean 64.3, in S. chrysopygus), 18���21, mean 19.3, gulars (vs. 23���28, mean 22.5, in S. chrysopygus), and 23���26, mean 24.4, subdigital lamellae under fourth toe (vs. 22���30, mean 26.0, in S. chrysopygus). Definition. (1) Maximum total length in males 68 mm (n = 1); (2) maximum total length in females 60.5 mm (n = 6); (3) vertebrals 44���52; (4) paravertebrals 44���51; (5) scales around midbody 42���47; (6) supraoculars 5���6; (7) internasals 4; (8) postrostrals 4���6; (9) loreals 1���2; (10) gulars 18���21; (11) lamellae on Finger IV 17���19; (12) lamellae on Toe IV 23���26; (13) posthumeral mite pocket present as one or more vertical folds or ridges (Type 1 of Torres-Carvajal 2007); (14) postfemoral mite pocket shallow, with a posteroventrally oriented slit-like opening (Type 2 of Torres-Carvajal 2007); (15) parietal eye present; (16) occipital scales small, smooth, juxtaposed; (17) projecting angulate temporals absent; (18) enlarged supraoculars occupying most of supraocular region in one row absent; (19) scales on frontonasal region smooth, juxtaposed; (20) preauricular fringe present; (21) antegular, antehumeral, gular, longitudinal, postauricular, supra-auricular, and transverse antegular neck folds present, oblique fold ill-defined without deep mite pocket; (22) lateral nuchals smaller than dorsal nuchals; (23) posterior gulars smooth, imbricate, apical pit absent; (24) lateral body scales smaller than dorsal body scales; (25) vertebrals same size as adjacent dorsals, not forming a continuous longitudinal row; (26) dorsolateral crests absent; (27) ventrals smooth, imbricate; (28) scales on posterior surfaces of thighs granular; (29) prefemoral fold absent; (30) inguinal groove absent; (31) preanals not projected; (32) tail not compressed laterally in adult males; (33) tail length 61���67 % of total length; (34) three caudal whorls per autotomic segment; (35) caudals not spinose; (36) dark stripe that extends anterodorsally from subocular region to supraciliaries absent; (37) color pattern of gular region in adult females with dark streaks; (38) color pattern of gular region in adult males with dark streaks forming a reticulum; (39) black blotch on ventral surface of neck in adult males absent; (40) dark midventral stripe in adult males absent; (41) black patches on ventral surface of thighs in adult males absent; (42) background color of dorsum grey or brown. Description of the holotype. Adult male, as indicated by partly everted hemipenes; SVL 68.0 mm; tail length 140.0 mm (complete); tail more or less round in cross section; axilla to groin distance 30.6 mm; head length 15.6 mm, head length/SVL ratio 0.23; snout length 6.6 mm; head width 11.0 mm; head height 8.7 mm; shank length 14.9 mm, shank length/head length ratio 0.96; scales on parietal and occipital regions small, slightly wrinkled, juxtaposed; parietal eye visible; supraoculars in five rows, smooth, with the lateralmost three rows less than half the size of the medial adjacent row; distinct circumorbitals present; canthals two; anterior-most canthal separated from nasal by a small scale; internasals four; postrostrals four, much wider than long; supralabials six; infralabials seven; loreals two; lorilabials in one row; preocular divided into two scales; lateral temporals keeled, imbricate; gulars in 18 rows between tympanic openings; all gulars cycloid, smooth, imbricate, apical pit absent; second sublabial in contact with the second and third infralabials; first pair of postmentals in contact medially; mental separated from sublabials by the first pair of postmentals; dorsal and lateral scales of neck keeled, imbricate; lateral scales of body smaller than dorsals, but of similar shape, imbricate; scales around midbody 44; vertebrals not differentiated from adjacent scales, in 47 rows, not forming a continuous vertebral row; paravertebrals adjacent to vertebral row equal to vertebrals in size and shape; paravertebrals 46; ventrals smooth, imbricate, smaller than dorsals; preauricular fringe present; antegular, antehumeral, gular, longitudinal, postauricular, supra-auricular, and transverse antegular neck folds present; ventrolateral and prefemoral folds present, oblique fold ill-defined without deep mite pocket; dorsal scales of forelimbs and hind limbs keeled, imbricate; lamellae on Finger IV 19; lamellae on Toe IV 26 (right) and 25 (left); tail rounded; caudals keeled, non-mucronate, imbricate; basal subcaudals smooth, imbricate; posthumeral mite pocket present as one or more vertical folds [Type 1 of Torres-Carvajal (2007)]; postfemoral mite pocket shallow, with a posteroventrally oriented slit-like opening [Type 2 of Torres- Carvajal (2007)]. Coloration in life was recorded as follows (Fig. 2): dorsum dark brown with small greenish-yellow scales scattered along posterior half of body, anterior half with small bluish-gray scales; head dorsally blackish brown with scattered tan scales; ventral surfaces of body (except for anterior half of chest which is pale gray), hindlegs, belly, and base of tale, mustard yellow; throat dark purple brown with pale gray streaks. Coloration after 17 years of preservation in 70 % ethanol was recorded as follows: Dorsal surface of head Raw Umber (280) with Smoky White (261) scales in frontal and parietal regions and a suffusion of Clay Color (20) on snout; dorsal surface of neck Cinnamon-Drab (259) with suffusions of Pale Cyan (157) and True Cinnamon (260); lateral surface of neck Beige (254) with suffusions of Medium Neutral Gray (298); dorsal surface of body Fawn Color (258) with scattered Pale Cyan (157) scales; dorsal surfaces of limbs Sepia (279) with Smoky White (261) Pearl Gray (262) scales; dorsal surface of tail Grayish Horn Color (268) grading into Ground Cinnamon (270) towards tip and with indistinct Smoke Gray (266) bands; ventral surface of head Light Smoke Gray (263) with suffusions of Ground Cinnamon (270) and with Smoky White (261) mottling; ventral surfaces of body Smoke Gray (267) with a Grayish Horn Color (268) central area on venter; ventral surfaces of limbs Grayish Horn Color (268); ventral surface of tail Drab (19) grading into Cream White (57) towards tip. Variation. The paratypes agree well with the holotype in general appearance, morphometrics and scalation (see Table 1). Etymology. The name amydrorhytus is a compound noun in apposition derived from the Greek words amydros (���indistinct, obscure���) and rythis (���fold, wrinkle���) and refers to the poorly developed oblique neck fold characteristic for this species. Geographic distribution. Stenocercus amydrorhytus is only known from the vicinity of Coris at elevations of 2785���3075 m asl (Fig. 8). Natural history notes. The type specimens of Stenocercus amydrorhytus were collected during the morning hours on and under rocks in a grassy area. Sympatric amphibians include Rhinella limensis, and Telmatobius rimac (Lehr 2002, Lehr et al. 2002). *data taken from Cadle 1998, Published as part of K��hler, Gunther & Lehr, Edgar, 2015, Two new species of lizards of the genus Stenocercus (Iguania, Tropiduridae) from central Peru, pp. 413-427 in Zootaxa 3956 (3) on pages 414-420, DOI: 10.11646/zootaxa.3956.3.6, http://zenodo.org/record/235614, {"references":["Torres-Carvajal, O. (2007) A taxonomic revision of South American Stenocercus (Squamata: Iguania) lizards. Herpetological Monographs, 21, 76 - 178. http: // dx. doi. org / 10.1655 / 06 - 001.1","Cadle, J. E. (1998) New species of lizards, genus Stenocercus (Iguania: Tropiduridae), from western Ecuador and Peru. Bulletin of the Museum of Comparative Zoology, 155, 257 - 297."]}

Stenocercus johaberfellneri sp. nov. Figs. 1 c, 5���7 Stenocercus species: Lehr 2002: p. 87. Holotype. MUSM 20229, an adult male from Malvas, - 9.91683, - 77.65133, 3200 m, Departamento Ancash, Provincia Huarmey, Peru, collected 3 February 1997 by Edgar Lehr. Field tag number EL 99. Paratypes. 11 (5 males, 6 females). All from Malvas, Departamento Ancash, Provincia Huarmey, Peru, collected on 2���3 February 1997 by Edgar Lehr: MUSM 20225, - 9.91833, -77.65800, 3200 m. MUSM 20226, - 9.92850, - 77.64967, 3055 m. MUSM 20227, SMF 80231 ��� 33, - 9.92850, - 77.64517, 3120 m. MUSM 20228, SMF 80234, - 9.93217, -77.65500, 3130 m. MUSM 20229 ��� 30, - 9.91683, - 77.65133, 3200 m. SMF 80235, - 9.93217, - 77.65333, 3140 m. Diagnosis: Stenocercus johaberfellneri differs from all other congeners except S. latebrosus and S. ornatissimus by the combination of having (1) a well developed oblique neck fold with a deep mite pocket underneath; (2) a well developed antehumeral fold with a deep mite pocket underneath; (3) scales on posterior surface of thighs granular; (4) vertebral scales similar in size and shape to adjacent scale rows; (5) three caudal whorls per autotomic segment. Stenocercus johaberfellneri differs from S. latebrosus by having lateral nuchals less than half the size of dorsal nuchals (vs. lateral nuchals similar in size to dorsal nuchals in S. latebrosus). Stenocercus johaberfellneri differs from S. ornatissimus by having larger dorsal scales, 47���53, mean 50.0, vertebral scales (vs. dorsal scales smaller, 52���59, mean 55.4, vertebral scales in S. ornatissimus), 46���53, mean 49.2, scales around midbody (vs. 49���60, mean 52.6, in S. ornatissimus), the presence of a postfemoral pocket (Type 2 of Torres-Carvajal 2007) in adult males (vs. postfemoral pocket absent in S. ornatissimus), a throat pattern of bold black streaks or reticulations in adult males (vs. throat with small black spots or fine reticulations in S. ornatissimus), and a dusky or grayish venter without black spots or reticulations (vs. at least some black spots on pectoral region and anterior abdomen, often entire venter with small black spots in S. ornatissimus). Definition. (1) Maximum total length in males 57 mm (n = 6); (2) maximum total length in females 47.5 mm (n = 5); (3) vertebrals 47���53; (4) paravertebrals 47���52; (5) scales around midbody 46���53; (6) supraoculars 4���6; (7) internasals usually 4, rarely 2; (8) postrostrals usually 4, rarely 5; (9) loreals usually 2, rarely 1; (10) gulars 19���24; (11) lamellae on Finger IV 17���20; (12) lamellae on Toe IV 23���27; (13) posthumeral mite pocket present as one or more vertical folds or ridges (Type 1 of Torres-Carvajal 2007); (14) postfemoral mite pocket shallow, with a posteroventrally oriented slit-like opening in adult males (Type 2 of Torres-Carvajal 2007), no postfemoral mite pocket discernable in females and juveniles; (15) parietal eye present; (16) occipital scales small, smooth, juxtaposed; (17) projecting angulate temporals absent; (18) enlarged supraoculars occupying most of supraocular region in one row absent; (19) scales on frontonasal region smooth, juxtaposed; (20) preauricular fringe present; (21) antegular, oblique, antehumeral, gular, longitudinal, postauricular, supra-auricular, and transverse antegular neck folds present; (22) lateral nuchals smaller than dorsal nuchals; (23) posterior gulars smooth, imbricate, apical pit absent; (24) lateral body scales smaller than dorsal body scales; (25) vertebrals similar in size to adjacent dorsals, not forming a continuous longitudinal row; (26) dorsolateral crests absent; (27) ventrals smooth, imbricate; (28) scales on posterior surfaces of thighs granular; (29) prefemoral fold absent; (30) inguinal groove absent; (31) preanals not projected; (32) tail not compressed laterally in adult males; (33) tail length 62���68 % of total length; (34) three caudal whorls per autotomic segment; (35) caudals not spinose; (36) dark stripe that extends anterodorsally from subocular region to supraciliaries absent; (37) color pattern of gular region in adult females with dark streaks; (38) color pattern of gular region in adult males with bold black streaks or reticulations; (39) black blotch on ventral surface of neck in adult males absent; (40) dark midventral stripe in adult males absent; (41) black patches on ventral surface of thighs in adult males absent; (42) background color of dorsum grey or brown. Description of the holotype. Adult male, as indicated by partly everted hemipenes; SVL 57.0 mm; tail length 120.0 mm (complete); tail more or less round in cross section; axilla to groin distance 21.8 mm; head length 13.9 mm, head length/SVL ratio 0.24; snout length 5.7 mm; head width 10.7 mm; head height 7.8 mm; shank length 13.6 mm, shank length/head length ratio 0.98; scales on parietal and occipital regions small, slightly wrinkled, juxtaposed; parietal eye visible; supraoculars in five rows, smooth, with the lateralmost three rows less than half the size of the medial adjacent row; distinct circumorbitals present; canthals two; anterior-most canthal in contact with nasal; internasals two; postrostrals four, much wider than long; supralabials six; infralabials seven; loreals one; lorilabials in one row; preocular single; lateral temporals keeled, imbricate; gulars in 20 rows between tympanic openings; all gulars cycloid, smooth, imbricate, apical pit absent; second sublabial in contact with the second and third infralabials; first pair of postmentals in contact medially; mental separated from sublabials by the first pair of postmentals; dorsal and lateral scales of neck keeled, imbricate; lateral scales of body smaller than dorsals, but of similar shape, imbricate; scales around midbody 48; vertebrals differentiated from adjacent scales, in 48 rows, not forming a continuous vertebral row; paravertebrals adjacent to vertebral row equal to vertebrals in size and shape; paravertebrals 48; ventrals smooth, imbricate, smaller than dorsals; preauricular fringe present; antegular, oblique, antehumeral, gular, longitudinal, postauricular, supra-auricular, and transverse antegular neck folds present; ventrolateral and prefemoral folds present; dorsal scales of forelimbs and hind limbs keeled, imbricate; lamellae on Finger IV 19; lamellae on Toe IV 25 (right) and 26 (left); tail rounded in cross section; caudals keeled, non-mucronate, imbricate; basal subcaudals smooth, imbricate; posthumeral mite pocket present as one or more vertical folds (Type 1 of Torres-Carvajal 2007); postfemoral mite pocket shallow, with a posteroventrally oriented slit-like opening (Type 2 of Torres-Carvajal 2007). Coloration after 17 years of preservation in 70 % ethanol was recorded as follows: Dorsal surface of head Glaucous (289) with Vandyke Brown (282) and Dark Pearl Gray (290) mottling; dorsal surface of neck Cinnamon- Drab (50) with Sepia (279) paravertebral blotches and scattered Venetian Blue (170) scales; lateral surface of neck with a Medium Paris White (140) longitudinal stripe; dorsal surface of body Glaucous (289) with suffusions of Dark Pearl Gray (290), a few scattered Light Caribbean Blue (163) scales and indistinct Hair Brown (277) blotches; dorsal surfaces of forelimbs Dark Neutral Gray (299) with narrow Light Neutral Gray (297) bands; dorsal surfaces of hind limbs Grayish Horn Color (268) with suffusions of Pearl Gray (262); dorsal surface of tail Medium Neutral Gray (298) grading into Cinnamon (21) towards tip and with Pale Buff (1) bands; ventral surface of head Pale Greenish White (97) with contrasting Sepia (279) reticulum; ventral surfaces of body and limbs Pale Buff (1), heavily suffused with Grayish Horn Color (268) on venter; ventral surface of tail Medium Neutral Gray (298) with Smoky White (261) speckles and grading into Cinnamon (21) towards tip. Variation. The paratypes agree well with the holotype in general appearance, morphometrics and scalation (see Table 1). Coloration for one male (SMF 80235) in life was recorded as follows (Fig. 5): dorsum dark brown with blackish-brown triangles surrounded by pale gray scales, becoming irregular towards anterior half of body; mustard yellow scales scattered dorsolaterally; head dark brown dorsally, with scattered black, mustard yellow, and pale gray scales; ventral surfaces of body (except for anterior half of chest), hindlegs, and base of tale mustard yellow; anterior half of chest pale gray with dark brown flecks; throat copper with pale gray flecks and dark brown streaks which extend to upper and lower lips laterally. Etymology. The name johaberfellneri is a patronym for Johannes Haber Fellner, Germany, in recognition of his support of taxonomic studies through the BIOPAT program. Geographic distribution. Stenocercus johaberfellneri is presently only known from the vicinity of Malvas at elevations of 3055���3200 m asl (Fig. 8). Natural history notes. The type specimens of Stenocercus johaberfellneri were collected during morning hours under rocks on a grassy area. Sympatric amphibians include Telmatobius rimac, sympatric reptiles are Epictia alfredschmidti, Philodryas simonsii, and Sibynomorphus petersi (misidentified as S. vagus in Lehr 2002 and Lehr et al. 2002 as pointed out by Cadle 2007)., Published as part of K��hler, Gunther & Lehr, Edgar, 2015, Two new species of lizards of the genus Stenocercus (Iguania, Tropiduridae) from central Peru, pp. 413-427 in Zootaxa 3956 (3) on pages 420-425, DOI: 10.11646/zootaxa.3956.3.6, http://zenodo.org/record/235614, {"references":["Torres-Carvajal, O. (2007) A taxonomic revision of South American Stenocercus (Squamata: Iguania) lizards. Herpetological Monographs, 21, 76 - 178. http: // dx. doi. org / 10.1655 / 06 - 001.1","Cadle, J. E. (2007) The snake genus Sibynomorphus (Colubridae: Dipsadinae: Dipsadini) in Peru and Ecuador, with comments on the systematics of Dipsadini. Bulletin of the Museum of Comparative Zoology, 158 (5), 183 - 283. http: // dx. doi. org / 10.3099 / 0027 - 4100 (2007) 158 [183: TSGSCD] 2.0. CO; 2"]}

Comparison of Anolis boulengerianus and A. isthmicus and taxonomic conclusions A comparison of our Anolis boulengerianus and A. isthmicus samples is provided in Table 2 (also see Fig. 2). There were no obvious differences between these samples in any of the characters in the table, and we were unable to find any differences in any other characters examined. Thus, this comparison clearly demonstrates that A. boulengerianus represents the same taxonomic species that what is currently referred to as A. isthmicus. Because of the priority principle, A. isthmicus Fitch, 1978 becomes a junior synonym of A. boulengerianus Thominot, 1887. Also, A. isthmicus and Anolis subocularis Davis, 1954 were classified in the A. subocularis species group by Savage and Guyer (1989). However, because A. isthmicus is a junior synonym of A. boulengerianus and this name is older than A. subocularis, the A. subocularis group becomes the A. boulengerianus group. We provide the following synonymy and diagnosis for A. boulengerianus: ......continued on the next page Anolis boulengerianus Thominot, 1887: 182; three syntypes, MNHN 6554 and 1994.1670 ��� 71, from ���T��huant��pec (Mexique).��� Anolis isthmicus Fitch, 1978: 2; holotype, KU 176034, from ��� 19.5 km WNW Tehuantepec, Oaxaca, Mexico.��� Liner (1994), Lieb (1981, 1995), Savage and Guyer (1989). Anolis nebuloides: Guib�� (1954; in part), Brygoo (1989; in part). Anolis nebulosus: Barbour (1934; in part), Smith and Taylor (1950; in part). Diagnosis. Anolis boulengerianus is a medium-sized species (SVL of largest male examined 60.0 mm, of largest female 51.4 mm) that belongs to an unnamed group of beta anoles that occur in Mexico west of the Isthmus of Tehuantepec (except for the Atlantic versant) and differ from all of the other species of beta anoles in the lepidosis of the supraocular region; namely, in the possession of a longitudinal series of much enlarged supraocular scales, each wider than long, which combined form a broadly oval supraocular disk surrounded by relatively small scales (Fitch 1978; Figs. 1���2). The species in this unnamed group were classified in the A. gadovii, A. nebuloides, A. nebulosus, and A. boulengerianus species groups by Savage and Guyer (1989), although monophyly of these groups has not been investigated and the contents of some of them has been modified recently (Nieto-Montes de Oca et al., 2013). Anolis boulengerianus may be distinguished from all of the species in the A. gadovii group (A. dunni Smith, 1936, A. gadovii Boulenger, 1905, A. liogaster Boulenger, 1905, A. omiltemanus Davis, 1954, A. peucephilus K��hler et al., 2014, and A. taylori Smith & Spieler, 1945) in having strongly keeled midventral scales (midventral scales smooth in the other species). In addition, A. boulengerianus may be distinguished from all of the species in the above Mexican unnamed group with distinctly keeled midventral scales (i.e., those in the A. boulengerianus, A. nebuloides, and A. nebulosus groups, respectively) as follows: from the species in the A. nebuloides group (A. megapholidotus Smith, 1933 and A. nebuloides) by having dorsal scales gradually decreasing in size laterally (dorsal scales abruptly decreasing in size laterally in A. megapholidotus and A. nebuloides) and smaller, more numerous middorsal scales between the levels of the axilla and groin (52���75, x = 62.5, n = 21; versus 33���37, x = 35.0, n = 4, in the type series of A. nebuloides; and 28���32, x = 30, n = 6, in A. megapholidotus); from the species in the A. nebulosus group (A. forbesi Smith & Van Gelder, 1955, A. microlepidotus Davis, 1954, A. nebulosus, and A. quercorum Fitch, 1978) by having a larger body size (maximum SVL = 60.0 mm; versus x = 0.99, n = 20; versus 0.67���0.76, x = 0.83, n = 23, in A. forbesi [Herrera-Flores 2011]; adpressed hind limb reaching the eye or beyond in A. boulengerianus; reaching the shoulder in A. microlepidotus, the ear in A. nebulosus, and beyond the ear, but failing to reach the eye, in A. quercorum; Fitch 1978); and from the other species in the A. boulengerianus group (A. subocularis) by having more numerous middorsal and midventral scales between the levels of the axilla and groin (52���75, x = 62.5, n = 21 and 45���62, x = 50.5, n = 21 dorsals and ventrals, respectively; versus 40���55 and 30���42 dorsals and ventrals, respectively, in A. subocularis [Fitch 1978]; also 36���51, x = 41.5���44.6, n = 15���18 and 32���44, x = 37.3���38.9, n = 15���18 dorsals and ventrals, respectively, in three samples of A. subocularis [Feria-Ortiz 2001]). Geographic distribution: Slopes and foothills on Pacific versant of the Isthmus of Tehuantepec in southeastern Oaxaca, Mexico., Published as part of Oca, Adri��n Nieto-Montes De, K��hler, Gunther & Feria-Ortiz, Manuel, 2014, Anolis boulengerianus Thominot, 1887, a senior synonym of Anolis isthmicus Fitch, 1978 (Squamata: Dactyloidae), pp. 125-133 in Zootaxa 3794 (1) on pages 130-132, DOI: 10.11646/zootaxa.3794.1.5, http://zenodo.org/record/226631, {"references":["Fitch, H. S. (1978) Two new anoles (Reptilia: Iguanidae) from Oaxaca with comments on other Mexican species. Contributions in Biology and Geology of the Milwaukee Public Museum, 20, 1 - 15.","Thominot, A. (1887) Description of trois speces nouvelles d'Anolis et d'un amphisbaenien. Bulletin de la Societe Philomatique de Paris, (7), 11, 182 - 190.","Davis, W. B. (1954) Three new anoles from Mexico. Herpetologica, 10, 1 - 6.","Savage, J. M. & Guyer, C. (1989) Infrageneric classification and species composition of the anole genera, Anolis, Ctenonotus, Dactyloa, Norops and Semiurus (Sauria: Iguanidae). Amphibia-Reptilia, 10, 105 - 116. http: // dx. doi. org / 10.1163 / 156853889 x 00142","Liner, E. A. (1994) Scientific and Common Names for the Amphibians and Reptiles of Mexico in English and Spanish. SSAR Herpetological Circular, 23, 1 - 113.","Lieb, C. S. (1981) Biochemical and karyological systematics of the Mexican lizards of the Anolis gadovi and A. nebulosus species groups (Reptilia: Iguanidae). Ph. D. Dissertation, University of California, Los Angeles, USA, 308 pp.","Lieb, C. S. (1995) Preliminary key to the anole lizards of Mexico. In: Flores-Villela, O. A., Mendoza-Quijano, F. & Gonzalez- Porter, G. (Compiladores), Recopilacion de Claves para la Determinacion de Anfibios y Reptiles de Mexico. Publicaciones Especiales del Museo de Zoologia de la Facultad de Ciencias, Universidad Nacional Autonoma de Mexico, 10, pp. 146 - 157.","Guibe, J. (1954) Catalogue des types de lezards du Museum National d'Histoire Naturelle. Impremerie Colas, Bayeux, France, 119 pp.","Brygoo, E. R. (1989) Les types d´Iguanides (Reptiles, Sauriens) du Museum national d´Histoire naturelle Catalogue critique. Bulletin du Museum national d'Histoire naturelle, Series 4, 11 (Section A, no. 3), Supplement: 1 - 112.","Barbour, T. (1934) The anoles II. The mainland species from Mexico southward. Bulletin of the Museum of Comparative Zoology, 77, 119 - 155.","Smith, H. M. & Taylor, E. H. (1950) An annotated checklist and key to the reptiles of Mexico exclusive of the snakes. Bulletin of the United States Natural Museum, 199, 1 - 253.","Smith, H. M. (1936) A new Anolis from Mexico. Copeia, 1936, 9. http: // dx. doi. org / 10.2307 / 1436366","Boulenger, G. A. (1905) Descriptions of new reptiles discovered in Mexico by Dr. H. Gadow, F. R. S. Proceedings of the Zoological Society of London, 1905, 245 - 247.","Kohler, G., Gomez Trejo Perez, R., Petersen, C. B. P. & Mendez de la Cruz, F. (2014) A new species of pine anole from the Sierra Madre del Sur in Oaxaca, Mexico (Reptilia: Squamata: Dactyloidae: Anolis). Zootaxa, 3753, 453 - 468. http: // dx. doi. org / 10.11646 / zootaxa. 3753.5.4","Smith, H. M. & Spieler, R. A. (1945) A new anole from Mexico. Copeia, 1945, 165 - 168. http: // dx. doi. org / 10.2307 / 1438281","Smith, H. M. (1933) Notes on some Mexican lizards of the genus Anolis with the description of a new species, A. megapholidotus. Transactions of the Kansas Academy of Science, 36, 318 - 320. http: // dx. doi. org / 10.2307 / 3625373","Smith, H. M. & Van Gelder, R. G. (1955) New and noteworthy amphibians and reptiles from Sinaloa and Puebla, Mexico. Herpetologica, 11, 145 - 149.","Herrera-Flores, J. A. (2011) Redescripcion morfologica, posicion filogenetica y analisis de la distribucion potencial de Anolis forbesi (Squamata: Polychrotidae). M. Sc. Thesis, Instituto de Ecologia, A. C., Xalapa, Veracruz, Mexico, 62 pp.","Feria-Ortiz, M. (2001) Revision taxonomica del grupo de especies Anolis subocularis (Squamata: Polychrotidae). M. Sc. Thesis, Facultad de Ciencias, Universidad Nacional Autonoma de Mexico, Mexico, D. F., Mexico, 76 pp."]}

Atractus alytogrammus sp. nov. Figs. 1���3 Holotype. A male, SMF 88371, collected on January 9 1957 by Federico Medem in the Serrania de la Lindosa (2.46782 ��, - 72.73155 ��), south of the municipality of San Jos�� del Guaviare, department of Guaviare, Colombia. Diagnosis. A species of the genus Atractus that differs from all congeners by the following combination of characters: (1) 17 smooth dorsal scale rows; (2) preocular absent; (3) loreal present, approximately as long as high; (4) temporals 1 + 2; (5) seven supralabials; (6) seven infralabials; (7) five maxillary teeth; (8) four gular scale rows; (9) four pre-ventrals; (10) 187 ventrals in single male; (11) 29 / 31 subcaudals in single male; (12) dorsum of body with longitudinal stripes; (13) venter uniformly dark gray; (14) SVL of single male 297 mm; (15) small to moderate tail length in single male (11.8 % of SVL); (16) hemipenis morphology unknown. Comparisons. Among all currently recognized congeners, Atractus alytogrammus shares a dark venter and is morphologically similar to A. steyermarki, A. nigriventris, and A. trivittatus. It differs from A. steyermarki by having a uniform dark gray venter (vs. venter pale with a midventral stripe in A. steyermarki), 187 ventrals (vs. 160���167 in males in A. steyermarki), and 29 / 31 subcaudals (vs. 37���44 in males in A. steyermarki). Atractus alytogrammus differs from A. nigriventris and A. trivitattus by having a dorsal body pattern of longitudinal lines (vs. dorsum brown with black blotches in A. nigriventris and A. trivitattus) and 187 ventral scales in single known male (vs. 175 in single known female of A. nigriventris, 176 in single known female of A. trivitattus). Atractus alytogrammus differs further from A. trivitattus by having seven supralabials (vs. eight in A. trivitattus). Among congeners from Colombian Amazonia and surrounding lowlands, A. alytogrammus shares only with A. univittatus (Jan 1862) the combination of having 17 dorsal scale rows, seven supralabials, and a pattern with longitudinal stripes or lines on dorsum. Atractus alytogrammus differs from A. univittatus by having 187 ventrals in the single male (vs. 151���160 ventrals in males of A. univittatus), seven infralabials (vs. six infralabials in A. univittatus), and a uniform dark venter with lighter speckles on the edges (vs. a yellowish white venter in A. univittatus). Description of the holotype. Male; SVL 297 mm; moderate TL 35 mm (complete; 11.8 % SVL); TTL 332 mm; head not distinct from body; body cylindrical, diameter at midbody 5.2 mm (1.8 % of SVL); head rounded in dorsal view; HL 9.41 mm (3.2 % SVL); HW 4.9 mm (65.3 % of HL); snout rounded in dorsal view, rounded in lateral view; eye diameter 1.03 mm; pupil round; interorbital distance 3.2 mm (42.7 % HL); rostro-orbital distance 2.9 mm (38.7 % HL); naso-orbital distance 1.4 mm (18.7 % HL). 17 / 17 / 17 smooth dorsal scale rows, 16 dorsal scale rows above cloaca, 9 dorsal scale rows above second subcaudal; apical pits absent; 187 ventrals; 29 / 31 divided subcaudals; some of the subcaudals fused; 4 preventrals; cloacal scute entire; rostral well visible from above, subtriangular, wider than high (rostral height 1.3 mm, rostral width 2.24 mm); two subpentagonal internasals, height 0.7 mm, width 1.04 mm; internasal suture sinistral in respect to prefrontal suture; nostril in suture of prenasal and postnasal; 1 / 1 hexagonal postnasal, length 0.7 mm, height 0.7 mm; short subrectangular loreal, approximately as high as long (loreal length 0.7 mm; loreal height 0.6 mm; ratio length/height: 1.17); preocular lacking; two subrectangular postoculars, upper postocular larger than lower (upper postocular length 0.4 mm; upper postocular height 0.4 mm); temporal formula 1 + 2, anterior temporal larger than posterior temporals, posterior temporals equal in size (anterior temporal: height 0.9 mm, length 1.7 mm; upper posterior temporal: height 0.7 mm, length 1.0 mm); seven supralabials, first four equal in size; fifth, sixth seventh supralabials larger, sixth the largest; second and third supralabial in contact with loreal; third and fourth supralabial in contact with the eye; seven infralabials, first three infralabials in contact with the chin shields; first pair of infralabials smaller than rest; first pair of infralabials in contact, preventing contact of mental and chin shields; gular scale rows from infralabials to preventrals four; two chin shields, length 2.5 mm, width 0.7 mm; two subpentagonal prefrontals, length 1.5 mm, width 1.6 mm; pentagonal supraoculars, length 1.1 mm, width 0.6 mm; subtriangular frontal, frontal apex slightly protruding, length 2.1 mm, width 2.0 mm; parietal length 3.7 mm, width 2.0 mm. Coloration of the holotype. After 56 years preservation in 70 % ethanol: dorsal and lateral surfaces of head Hair Brown (277); most supralabials with Smoke Gray (266) speckles; surface of the chin region Fawn Color (258); ground color of the ventrals and subcaudals Dusty Brown (285) with Pale Buff (1) speckles on the edges of ventrals; dorsal ground color Sepia (279) with Verona Brown (37) mottling on the edges of the dorsal scales; a Drab (19) dashed line on the first dorsal scale row present from below level of a point four scales behind posterior tip of parietals to level of cloaca; a Drab (19) dashed line on the second dorsal scale row present from below level of a point four scales behind posterior tip of parietals to about one fourth of body where it becomes indistinct and disappears; a Fawn Color (258) longitudinal, continuous line on the fifth dorsal scale row present, from a point four scales behind posterior tip of parietals to tip of tail; a Dusty Brown (285) vertebral stripe present from posterior tip of parietals to tip of tail. Etymology. The name alytogrammus is a compound noun derived from alytos (Greek for ���unbroken���) and gramme (Greek for ���line���) referring to the continuous dorsolateral pale stripe in this species., Published as part of K��hler, Gunther & Kieckbusch, Max, 2014, Two new species of Atractus from Colombia (Reptilia, Squamata, Dipsadidae), pp. 291-300 in Zootaxa 3872 (3) on pages 292-295, DOI: 10.11646/zootaxa.3872.3.5, http://zenodo.org/record/228806, {"references":["Jan, G. (1862) Enumerazione sistematico delle specie d'ofidi del gruppo Calamaridae. Archivo per la Zoologia l´Anatomia e la Fisiologia, 2, 1 - 76."]}

Ecnomiohyla bailarina sp. nov. Holotype. SMF 97398 (original field number AB 297; Fig. 3���4), an adult male from the north slope of the Jingurud�� mountain range (Fig. 5), about 14.6 km S from Pavarand�� village (7.70903 ��N, - 78.04882 ��W, 750 m a.s.l.), Samb��, Comarca Ember��-Wounaan N�� 2, Dari��n, 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 Mal��, Dari��n (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 ����� 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 K��hler 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 Jingurud�� 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 Bailar��n���, 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., Published as part of Batista, Abel, Hertz, Andreas, Mebert, Konrad, K��hler, Gunther, Lotzkat, Sebastian, Ponce, Marcos & Vesely, Milan, 2014, Two new fringe-limbed frogs of the genus Ecnomiohyla (Anura: Hylidae) from Panama, pp. 449-474 in Zootaxa 3826 (3) on pages 458-463, DOI: 10.11646/zootaxa.3826.3.2, http://zenodo.org/record/250039, {"references":["Duellman, W. E. (1966) Taxonomic notes on some Mexican and Central American hylid frogs. University of Kansas Publications, Museum of Natural History, 17, 263 - 279.","Fund, W. & Hogan, C. (2012) Isthmian-Pacific moist forests. Available from: http: // www. eoearth. org / view / article / 153928 (accessed 25 September 2013)","Pough, H. F., Taigen, T. L., Stewart, M. M. & Brussard, P. F. (1983) Behavioral modification of evaporative water loss by a Puerto Rican frog. Ecology, 64, 244 - 252. http: // dx. doi. org / 10.2307 / 1937072","Boulenger, G. A. (1899) Descriptions of new batrachians in the collection of the British Museum (Natural History). Annals and Magazine of Natural History, Series 7, 3, 273 - 277. http: // dx. doi. org / 10.1080 / 00222939908678122","Savage, J. M. & Myers, C. W. (2002) Frogs of the Eleutherodactylus biporcatus group (Leptodactylidae) of Central America and northern South America, including rediscovered, resurrected, and new taxa. American Museum Novitates, 3357, 1 - 21. http: // dx. doi. org / 10.1206 / 0003 - 0082 (2002) 357 2.0. co; 2","Peters, W. C. H. (1873) Uber eine neue Schildrotenart, Cinosternon effeldtii und einige andere neue oder weniger bekannte Amphibien. Monatsberichte der Koniglichen Preussische Akademie des Wissenschaften zu Berlin, 1873, 603 - 618.","Boulenger, G. A. (1912) Descriptions of new batrachians from the Andes of South America, preserved in the British Museum. Annals and Magazine of Natural History, Series 8, 10, 185 - 191. http: // dx. doi. org / 10.1080 / 00222931208693215","Thominot, A. (1884) Note sur un batracien d'espece nouvelle provenant de Panama. Bulletin de la Societe Philomathique de Paris, 8, 151 - 152.","Savage, J. M. (1967) A new tree-frog (Centrolenidae) from Costa Rica. Copeia, 1967, 325 - 331. http: // dx. doi. org / 10.2307 / 1442121"]}

Ecnomiohyla veraguensis sp. nov. Ecnomiohyla rabborum ��� K��hler 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 F�� 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 & K��hler, 2012, Lithobates warszewitschii (Schmidt, 1857), Pristimantis caryophyllaceus (Barbour, 1928), P. cerasinus (Cope, 1875 ��� 1876 ���), P. cruentus, P. museosus (Ib����ez, 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 Serran��a de Tabasar��. 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., Published as part of Batista, Abel, Hertz, Andreas, Mebert, Konrad, K��hler, Gunther, Lotzkat, Sebastian, Ponce, Marcos & Vesely, Milan, 2014, Two new fringe-limbed frogs of the genus Ecnomiohyla (Anura: Hylidae) from Panama, pp. 449-474 in Zootaxa 3826 (3) on pages 463-465, DOI: 10.11646/zootaxa.3826.3.2, http://zenodo.org/record/250039, {"references":["Kohler, G. (2011) Amphibians of Central America. Herpeton Verlag Elke Kohler, Offenbach, Germany, 379 pp.","Fund, W. & Hogan, C. (2012) Isthmian-Pacific moist forests. Available from: http: // www. eoearth. org / view / article / 153928 (accessed 25 September 2013)","Lichtenstein, H. & Martens E. v. (1856) Nomenclator Reptilium et Amphibiorum Musei Zoologici Berolinensis. Namenverzeichnis der in der zoologischen Sammlung der Koniglichen Universitat zu Berlin aufgestellten Arten von Reptilien und Amphibien nach ihren Ordnungen, Familien und Gattungen. Buchdruckerei der Koniglichen akademie der wissenschaften, Berlin, 48 pp.","Dunn, E. R. (1924) New salamanders of the genus Oedipus with a synoptical key. Field Museum of Natural History Publication. Zoological Series, 12, 95 - 100.","Peters, W. C. H. (1863) Uber eine neue Schlangen Gattung, Styporhynchus, und verschiedene andere Amphibien des zoologischen Museum. Monatsberichte der Koniglichen Preussische Akademie des Wissenschaften zu Berlin, 1863, 399 - 413.","Cope, E. D. (1875 - 1876) On the Batrachia and Reptilia of Costa Rica. Journal of the Academy of Natural Sciences of Philadelphia, 8, 93 - 154.","Hertz, A., Hauenschild, F., Lotzkat, S. & Kohler, G. (2012 a) A new golden frog species of the genus Diasporus (Amphibia, Eleutherodactylidae) from the Cordillera Central, western Panama. ZooKeys, 196, 23 - 46. http: // dx. doi. org / 10.3897 / zookeys. 196.2774","Schmidt, O. (1857) Diagnosen neuer Frosche des zoologischen Cabinets zu Krakau. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Classe, 24, 10 - 15.","Barbour, T. (1928) New Central American frogs. Proceedings of the New England Zoological Club. Cambridge, Massachusetts, 10, 25 - 31.","Ibanez, R., Jaramillo, C. A. & Arosemena, F. A. (1994) A new species of Eleutherodactylus (Anura: Leptodactylidae) from Panama. Amphibia-Reptilia, 15, 337 - 341. http: // dx. doi. org / 10.1163 / 156853894 x 00371","Cope, E. D. (1862) Catalogues of the reptiles obtained during the explorations of the Parana, Paraguay, Vermejo and Uraguay Rivers, by Capt. Thos. J. Page, U. S. N.; and of those procured by Lieut. N. Michler, U. S. Top. Eng., Commander of the expedition conducting the survey of the Atrato River. Proceedings of the Academy of Natural Sciences of Philadelphia, 14, 346 - 359.","Taylor, E. H. (1949) Costa Rican frogs of the genera Centrolene and Centrolenella. University of Kansas Science Bulletin, 33, 257 - 270."]}

Dactyloa insignis (Cope 1871) Figures 2; 14; 18 K���M. Anolis microtus: Peters and Donoso-Barros (1970: in part.: referring to the holotype of Diaphoranolis brooksi). Anolis insignis: Cope (1871, 1876); Boulenger (1885); Dunn (1937); Taylor (1956); Peters and Donoso-Barros (1970); Savage (1974); Myers (1977); Savage and Talbot (1978); Arosemena et al. (1992); Auth (1994); Mart��nez and Rodr��guez (1994); Young et al. (1999); Ib����ez et al. (2001); Poe (2004); Hamad (2009); Fl��schendr��ger and Wijffels (2009); Poe et al. (2009); Carrizo (2010); Jaramillo et al. (2010); Lotzkat et al. (2010 b); Stadler (2010); Casta��eda & de Queiroz (2011). Diaphoranolis brooksi: Barbour (1923). Holotype. Lost according to Savage and Talbot (1978); from Costa Rica: Provincia de San Jos��: near Ciudad San Jos��; probably from near La Palma (Savage 1974; Fig. 1: loc. 21; Fig. 21). Diagnosis. A large species (maximum SVL 160 mm) of the genus Dactyloa (sensu Nicholson et al. 2012) that is most similar in external morphology to the other members of this clade found in western Panama (D. casildae, D. frenata, D. ginaelisae, D. ibanezi, D. kunayalae, and D. microtus). Dactyloa insignis can readily be distinguished from these six species by its unique coloration of fine reticulate lines forming transverse crossbands on dorsum and flanks, and a preaxillary dark blotch (Fig. 14). Moreover, D. insignis has all dorsal and lateral body scales smooth (vs. conical, keeled, rugose or wrinkled in the other species), and the highest number of scales around midbody (190 or more vs. 178 or fewer). In addition, D. insignis differs from D. casildae, D. frenata, and D. ibanezi in having short legs (tip of fourth toe of adpressed hind limb reaching at most to a point between shoulder and tympanum in D. insignis), and from D. ginaelisae, D. ibanezi, D. kunayalae, and D. microtus in having more subdigital lamellae under the fourth toe (52 or more in D. insignis vs. 50 or fewer) as well as under the fourth finger (40 in D. insignis vs. 37 or fewer). Description. Total length to 464 mm; SVL to 160 mm in males, to 135 mm in females; tail long, about two times SVL, compressed, without dorsal crest; legs short, tip of fourth toe of adpressed hind limb reaching at most to a point between shoulder and tympanum; internasals, canthals, and loreals keeled; scales of frontal and prefrontal area mostly flat, smooth; IP usually distinct, surrounded by not much smaller scales (indistinct from surrounding scales in our juvenile); parietal eye usually distinct (barely visible in our juvenile); scales of SS barely enlarged, smooth; scales of supraorbital disk not conspicuously enlarged, smooth; two or three only slightly elongate, keeled anterior superciliary scales, none reaching a length of one-fourth of horizontal eye diameter; anterior sublabials distinctly enlarged, about as high as INL, keeled; scales of temporal arch conspicuously larger than those above and below; ear opening moderate to large, higher than SPL and INL together, slightly lower to higher than eye, much larger than IP; nuchal and dorsal crests present; 0���2 rows of smooth middorsal scales slightly enlarged; other dorsal scales as well as lateral scales flat, smooth; ventrals much larger than largest dorsals, smooth; scales on anterodorsal surface of thigh mostly smooth, only on anterior edge unicarinate with a few bi- or tricarinate; scales on dorsal surface of forearm smooth, becoming unicarinate towards wrist and anterior edge; fourth toe with well-developed dilated pad, about three times width of distal phalanx; female dewlap moderate, extending posteriorly to slightly beyond axilla, with broad, diffuse gorgetal-sternal rows leaving very narrow interspaces with less densely packed scales. The hemipenis of Dactyloa insignis remains undescribed. Coloration in life. Dorsal and lateral surfaces white to brown; dorsum, flanks, limbs, and tail with fine reticulate lines conglomerating to form light-centered crossbands that become solid only on posterior portion of tail; lips with dark vertical bars; a pronounced, light-centered blotch between tympanum and axilla, and a dark postorbital stripe extending to above tympanum; ventral surfaces largely white, except for dark crossbands forming complete rings under tail and dark flank markings and lip bars extending onto ventral surfaces; iris brown; female dewlap brown, with dark and light blotches and lines (Figs. 2; 14). Dactyloa insignis exhibits considerable metachrosis with ground color ranging from white to dark brown, and the dark markings being green, brown, or almost black. Most notably, in certain situations numerous beige to orange blotches appear on body and head, and even the color of the iris can range from almost white to dark reddish brown (compare Figures 14 B, C, and F). Color photographs of D. insignis have been published by Savage (2002), K��hler (2003, 2008), Fl��schendr��ger and Wijffels (2009), Stadler (2010), and Uetz (2013). The coloration in life of the adult female (SMF 89482, Figs. 2; 14 B���D, F, G) was recorded as follows: Ground color Sulphur Yellow (57) with Sepia (219) broken crossbands on dorsum which contain Pale Pinkish Buff (121 D) blotches; vertical series of Salmon Color (106) blotches between crossbands; several Spectrum Orange (17) blotches on neck; a Raw Umber (23) postorbital stripe; a Burnt Umber (22) blotch with Raw Umber (23) center anterior to shoulder; tail with Sepia (119) crossbands, anterior ones suffused with Natal Brown (219 A); ventral surfaces of body and tail dirty white; Dorsal surfaces of limbs with Dark Brownish Olive (129) and Lime Green (59) transverse lines as well as transverse series of Pale Pinkish Buff (121 D) flecks; iris Cinnamon-Rufous (40); dewlap Sayal Brown (223 C) with Indigo (73) horizontal lines and horizontal series of dirty white flecks. The coloration in life of the juvenile female (SMF 91477, Figs. 2; 14 A, E, H���L) was recorded as follows: Dorsal ground color Opaline Green (162 D), grading into dirty white with a suggestion of Pale Horn Color (92) ventrolaterally; head and body with a reticulum of fine Sepia (119) lines that are suffused with, and partly turn into, Emerald Green (163), reach onto venter and accumulate along middorsum to give the impression of four blotches; limbs, toes, and fingers with pairs of transverse Sepia (119) lines; tail with six Sepia (119) transverse bands, the anterior three composed of dense reticula of lines similar to the middorsal blotches; head, body, and limbs with irregularly distributed Orange-Rufous (132 C) and Cream Color (54) blotches; ventral ground color dirty white with a suggestion of Pale Horn Color (92); iris Pale Pinkish Buff (121 D); tongue Olive-Gray (42); dewlap dirty white with a suggestion of Pale Horn Color (92) at base, grading into Ground Cinnamon (239) mottled with Buff (24) towards apex, with Sepia (119) longitudinal lines, longitudinal series of dirty white spots, and a Pale Horn Color (92) margin. Coloration in preservative. After 23���48 months of preservation in 70 % ethanol, the ground color has lightened to dirty white, and all markings have assumed grayish or brownish tonalities. Only slight suggestions of blue where green was present in life in our juvenile are still discernable after 23 months (Figs. 18 K���M). Geographic distribution. Dactyloa insignis is distributed throughout Costa Rica and Panama at low and premontane elevations of 0���1600 m asl. In Panama, the species has been recorded from the provinces of Bocas del Toro, Chiriqu��, Cocl��, Col��n, Dari��n, Panam��, and Veraguas, and from the Comarca Ng��be-Bugl�� (Fig. 21). According to Young et al. (1999), it is also present in the Comarca Kuna Yala. Natural history notes. Both specimens were encountered at night while they were sleeping on branches 5���6 m above ground. Around our collection sites, Dactyloa insignis co-occurs with D. frenata, D. ibanezi, and D. kunayalae. Our automatized temperature recordings around our collection sites of Dactyloa insignis (640 and 880 m asl) range between 18.2���26.2 ��C. According to our combined dataset of 36 georeferenced occurrences, the species inhabits LMF, LWF, PMMF, and PMWF, with temperatures between 12.4���33.7 ��C, mean annual temperatures of 18.0��� 26.4 ��C and a total annual precipitation of 1696���4373 mm. Conservation. Jaramillo et al. (2010) calculated an EVS of 11 for Dactyloa insignis, and assigned the species to the IUCN category LC. We calculated the species��� EVS as 4 (range) + 3 (persecution) + 4 (ecological distribution) = 11. Its extent of occurrence of more than 63000 km 2 does not qualify D. insignis for any of the ���Threatened��� IUCN categories. Considering the continuing deforestation we observed in the region, we place the species in the category ���Near Threatened��� (NT). Remarks. Our record from Willie Maz�� (Fig. 1: loc. 9) constitutes the first record for the Comarca Ng��be- Bugl��. In the maps of K��hler (2003, 2008), the distribution area of Dactyloa insignis extends into Colombia. Although it is very likely that the species ranges at least into the Colombian Dari��n, no specimens from this country are known to date. The occurrence of D. insignis in the Comarca Kuna Yala as stated by Young et al. (1999) seems plausible considering that the specimen FMNH 170087 was collected at Paradise Camp, which lies less than 4 km west of Camp Summit (Fig. 1: loc. 20) that is situated on the border between Dari��n province and the Comarca Kuna Yala. Unfortunately, our sample is composed of just two females. Savage (2002) pictured male individuals from Monteverde in Costa Rica (plates 256 and 257), and described the male dewlap as dark red (key on p. 451) or as ���primarily orange-red, with or without several horizontal green or white bars and dark spotting��� (p. 455) with a white to greenish free margin. Both Savage (2002) and K��hler (2008) stated that Dactyloa insignis has 51 or more lamellae under the fourth toe, and used this as a key characteristic to distinguish this species from D. microtus, which has at most 49 according to these authors. Nevertheless, Savage and Talbot (1978) counted just 50 lamellae on the fourth toe of MCZ 16297, a value that is also reached by one of our specimens of D. ginaelisae (which formerly was included in D. microtus). All previous authors have reported the presence of a distinct IP and parietal eye as a diagnostic character to distinguish D. insignis from D. microtus. However, the IP is indistinct and the parietal eye barely visible in one of our two specimens (Fig. 14 H), suggesting that strict adherence to this key characteristic might not always be recommendable (also see remarks concerning this matter for D. ginaelisae and D. microtus). Savage and Talbot (1978) pointed out differences in anterior head scale surface and coloration between individuals from Costa Rica and Pacific west-central Panama on the one hand and examples from northwestern, central, and eastern Panama on the other hand, concluding that additional material was required to finally evaluate whether or not two species are involved. Our specimens have smooth anterior dorsal head scales and, just as the individuals photographed by Michael Castillo in Donoso district in Col��n province, agree in exhibiting dark lip bars, a dark postorbital marking (in form of a short stripe rather than a spot or blotch), a dark preaxillary blotch, and in having their transverse bands formed by reticula. Thus, our specimens would fall somewhere between Savage and Talbot���s (1978) specimens from Costa Rica and their specimen from Valle de Ant��n (Cocl�� province). Since we have no male from our study area available, we cannot contribute further to the issue of geographical variation in male dewlap coloration., Published as part of Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix & K��hler, Gunther, 2013, Distribution and variation of the giant alpha anoles (Squamata: Dactyloidae) of the genus Dactyloa in the highlands of western Panama, with the description of a new species formerly referred to as D. microtus, pp. 1-54 in Zootaxa 3626 (1) on pages 32-34, DOI: 10.11646/zootaxa.3626.1.1, http://zenodo.org/record/283424

Dactyloa ginaelisae sp. nov. Figures 2; 8; 9; 17 G���I; 18 R���S. Anolis microtus: Dunn (1937: in part.); Slevin (1942); Taylor (1956: in part.); Peters and Donoso-Barros (1970: in part.); Savage and Talbot (1978: in part.); Arosemena et al. (1992: in part.); Auth (1994: in part); Young et al. (1999); Ib����ez et al. (2001); K��hler et al. (2008); Fl��schendr��ger and Wijffels (2009: in part.); Hamad (2009: in part.); Lotzkat et al. (2010 a); Jaramillo et al. (2010); Stadler (2010); Casta��eda & de Queiroz (2011: in part.). Dactyloa microtus: Savage and Guyer (1989: in part.); Savage (2002: in part.); K��hler (2003, 2008: in part.); Uetz (2013: in part.). Holotype. SMF 91504 (Figs. 2; 8; 9 S���T), adult male, from the banks of Quebrada Jugl�� (Fig. 19 F) on the southeastern slope of Cerro Sagu�� (also known as Cerro Rat��n; locality 11 in Fig. 1) at Finca Alto Cedro, about 2 km north-northeast of the village Rat��n, 8.5576 ��N, 81.8262 ��W, 1710 m asl, Corregimiento de Piedra Roja, Distrito de Kankint��, Comarca Ng��be-Bugl��, Panama; collected by Andreas Hertz and Sebastian Lotzkat on 0 8 July 2010; original field number SL 660. Paratypes. Collected by Sebastian Lotzkat and Andreas Hertz, if not indicated otherwise. All from the Comarca Ng��be-Bugl��, Panama. Southeastern slope of Cerro Sagu�� (Fig. 1: loc. 11): MHCH 2240, juvenile female, same collecting data as holotype; MHCH 2239 and SMF 91503, females, near type locality, 8.5561 ��N, 81.8252 ��W, 1700 m asl, 0 7 July 2010; SMF 91502, juvenile female, about 850 m NNE of type locality, 8.5636 ��N, 81.8217 ��W, 1960 m asl, 0 8 July 2010. Western slope of Cerro Santiago, La Nevera (Fig. 1: loc. 14): SMF 89496 and 89497, juvenile female and adult female, 8.4997 ��N, 81.7724 ��W, 1700 m asl, 11 May and 17 August 2008; SMF 85069, juvenile female, 8.5000��N, 81.7722 ��W, 1600 m asl, collected by Abel Batista, Gunther K��hler, Marcos Ponce, and Javier Sunyer on 22 January 2006; SMF 89498, male, 8.5011 ��N, 81.7694 ��W, 1580 m asl, 14 August 2008; MHCH 2235, juvenile female, 8.5018 ��N, 81.7689 ��W, 1560 m asl, 10 August 2008; MHCH 2238, female, 8.4989 ��N, 81.7682 ��W, 1620 m asl, 11 November 2009; MHCH 2234, female, 8.5032 ��N, 81.7675 ��W, 1530 m asl, 10 August 2008; SMF 90133, female, 8.4954 ��N, 81.7673 ��W, 1810 m asl, 11 November 2009. Eastern slope of Cerro Santiago, Quebrada Ardilla (Fig. 1: loc. 15): MHCH 1338, female, 8.4974 ��N, 81.7228 ��W, 1600 m asl, 24 March 2009. Referred specimens. Apart from the material collected by ourselves and listed in the Appendix, all specimens reported as Dactyloa microtus that have been collected in Panama to our knowledge are referrable to D. ginaelisae in view of their collection localities in the vicinities of Boquete, where we also collected three specimens of this species: ANSP 22418 and 22419 from above Boquete (Dunn 1937; Savage & Talbot 1978); CAS 79598 from the north slope of Volc��n Bar�� (Slevin 1942); MVUP 966 from Cerro Horqueta and MVUP 969 from Bajo Mono near Boquete (MVUP catalogue); CHP 1038 from Cerro Horqueta (CHP catalogue). Diagnosis. A large species (maximum SVL 112 mm) of the genus Dactyloa (sensu Nicholson et al. 2012) that is most similar in external morphology to the other members of this genus found in western Panama (D. casildae, D. frenata, D. ibanezi, D. insignis, D. kunayalae, and D. microtus). These species share a moderate to large adult size (SVL> 70 mm), a large dewlap and enlarged postcloacal scales in males, and smooth or faintly keeled ventrals. Dactyloa ginaelisae can readily be distinguished from these six species by its color pattern described below and shown in Figs. 2, 8, 9, and 18 R���S. It further differs from all mentioned species except D. microtus by its low numbers of horizontal loreal rows (4 or fewer in D. ginaelisae vs. 5 or more) and total loreal scales (25 or fewer in D. ginaelisae vs. 39 or more), and by its low number of scales around midbody (100 or usually much fewer in D. ginaelisae vs. 110 or more). Moreover, D. ginaelisae differs from D. casildae, D. frenata, and D. ibanezi in having short legs (tip of fourth toe of adpressed hind limb reaching to a point between tympanum and eye, very rarely to posterior border of eye, in D. ginaelisae vs. beyond eye; shank length/SVL = 0.22 or less in D. ginaelisae vs. 0.25 or more). Among the short-legged species of Dactyloa in western Panama, D. ginaelisae further differs from D. insignis in having fewer subdigital lamellae under the fourth toe (50 or fewer in D. ginaelisae vs. 52 or more) as well as under the fourth finger (36 or fewer in D. ginaelisae vs. 40), and from D. kunayalae in having more subdigital lamellae under the fourth toe (41 or more in D. ginaelisae vs. 35 or fewer) as well as under the fourth finger (29 or more in D. ginaelisae vs. 25 or fewer). Dactyloa ginaelisae is very similar to D. microtus, from which it differs in having longer legs (tip of fourth toe of adpressed hind limb reaching to a point between tympanum and eye in D. ginaelisae vs. to a point between shoulder and tympanum in D. microtus; shank length/ SVL = 0.19 or more in D. ginaelisae vs. 0.183 or less) and by its conspicuous and clear-cut coloration pattern between eye and shoulder (a prominent light stripe extending from supralabials posteriorly above or a across the ear before bending down towards shoulder, delineating a dark preaxillary blotch above and posteriorly, and paralleled above by a dark postorbital stripe with darker borders that extends at least to a level above the preaxillary blotch in D. ginaelisae vs. light postsupralabial and dark postorbital stripe oriented more ventrally and losing their conspicuousness around ear). Description of the holotype. Adult male as indicated by everted hemipenes (Fig. 8 G), a pair of enlarged postcloacal scales (Fig. 8 E), and presence of large dewlap (Fig. 8 D); snout-vent length 107 mm; tail complete; tail length 244 mm, tail length/SVL ratio 2.28; tail compressed in cross section, tail height 7.0 mm, tail width 4.3 mm; axilla to groin distance 45.0 mm; head length 29.1 mm, HL/SVL ratio 0.27; snout length 14.2 mm; head width 16.7 mm; longest toe of adpressed hind limb reaching to a point between tympanum and eye; shank length 22.3 mm, shank length/SVL ratio 0.21, shank length/HL ratio 0.77; longest finger of extended forelimb reaching well beyond tip of snout; longest finger of adpressed forelimb reaching to anterior insertion of hind limbs; prefrontal ridges distinct, parietal ridges conspicuously protruding; scales on snout mostly rugose to wrinkled; 5 postrostrals; 6 scales between nasals; scales in distinct prefrontal depression wrinkled; supraorbital semicircles differentiated, composed of very rugose and partly wrinkled scales, separated by a minimum of 2 scales; supraorbital disc composed of 15 enlarged rugose scales; one slightly elongated, keeled anterior superciliary, followed posteriorly by a much smaller, keeled, elongate scale; about 2 rows of small keeled scales extending between enlarged supraorbitals and superciliary; interparietal plate not distinct, no parietal eye visible; canthal ridge distinct, composed of 5 large (posterior) and 1 small (anterior) canthal scales; 7 scales present between second canthals; 8 scales present between posterior canthals; 19 / 20 (right side/left side) wrinkled loreal scales in a maximum of 3 horizontal rows; subocular scales flat, wrinkled, subocular row well-defined; 7 supralabials to level below center of eye; ear opening 1.1 x 2.2 mm (length x height); mental distinctly wider than long, almost completely divided medially, bordered posteriorly by 6 postmentals; 7 infralabials to level below center of eye; sublabials enlarged, about as high as INL anterior to level of orbit, first two sublabials posterior to mental greatly enlarged, higher than INL; keeled granular to elongate scales present on chin and throat; dewlap large, extending well onto body, anterior insertion at a level halfway between orbit and tip of snout, posterior insertion at a level between one-fourth and one-third the distance between axilla and groin, with about 5 gorgetal-sternal rows 2���4 scales wide, becoming more diffuse posteriorly; low nuchal and dorsal crests present, dorsum of body with elevated, wrinkled scales, 2 middorsal rows of prominently protruding, keeled, but not otherwise enlarged scales, largest dorsal scales about 0.8 x 0.8 mm (length x width); about 42 medial dorsal scales in one HL; about 72 medial dorsal scales between axilla and groin; lateral scales raised, rugose to wrinkled, average size 0.7 mm in diameter, with minute granules occupying varying portions of the interspaces between them; ventrals at midbody smooth, subimbricate, about 0.5 x 0.5 mm (length x width); about 57 ventral scales in one HL; about 88 ventral scales between axilla and groin; about 94 scales around midbody; caudal scales strongly keeled, without whorls of enlarged scales, subcaudal scales with a single prominent keel; a pair of greatly enlarged postcloacal scales, larger one about 1.8 x 3.1 mm (length x width); tube-like axillary pocket not developed; scales on anterodorsal surface of thigh and on dorsal surface of forearm multicarinate, imbricate; digital pads dilated, dilated pad about 3 times width of non-dilated scales under distal phalanx; distal phalanx narrower than and raised from dilated pad; 33 / 35 lamellae under phalanges ii to iv of 4 th toe; 12 / 13 scales under distal phalanx of 4 th toe; 24 lamellae under phalanges ii to iv of 4 th finger; 12 / 11 scales under distal phalanx of 4 th finger. The completely everted hemipenis is a medium-sized, bilobate organ; sulcus spermaticus bordered by welldeveloped sulcal lips, opening at base of apex into two broad concave areas, one on each lobe; large asulcate processus and ridge present; a knob-like processus present on each lateral side of truncus below base of apex; lobes finely calyculate, truncus with transverse folds. Coloration in life. The coloration in life, including two stages of metachrosis, is shown in Figs. 8 A���E. The holotype represents the most contrastingly colored morph known for the species, with the dark crossbands on the dorsal and lateral surfaces of body, limbs, and anterior portion of tail well-delineated against the very light ground color by darker bands. Equally well-demarcated are the dark postorbital stripe extending posteriorly to above shoulder, and the large, elongate preaxillary blotch. The dirty to bright white supralabial stripe extends below the dark postorbital one, over and across the ear, to above the posterior portion of the preaxillary blotch, where it curves down towards the shoulder. Otherwise, no detailed notes of the holotype were taken. Coloration after approximately two years of preservation in 70 % ethanol (Figs. 8 F���M) is similar to that in life, apart from that all reddish, greenish, and bluish tonalities have faded. Variation. The paratypes and referred specimens agree well with the holotype in terms of general morphometrics and pholidosis (see Tables 2 and 3), but are very variable in coloration (Fig. 9). The following account of variation among our sample of Dactyloa ginaelisae is congruent with the standardized description sections in the subsequent species accounts for the other six species of Dactyloa found in western Panama. Total length to 362 mm; SVL to 112 mm in males, to 108 mm in females; tail long, about 1.7���2.4 times SVL, compressed, with a low dorsal crest on the anterior portion; legs short, tip of fourth toe of adpressed hind limb reaching to a point between anterior border of tympanum and posterior border of eye; dorsal and lateral head scales generally large; internasals, canthals, and loreals rugose to wrinkled; scales of frontal and prefrontal area mostly rugose to wrinkled; IP indistinct in most specimens, if discernable, then usually surrounded by scales of both smaller and equal size; scales of parietal area generally rugose to wrinkled; parietal eye indistinct in most specimens; scales of SS distinctly enlarged, rugose; scales of supraorbital disk conspicuously enlarged, rugose; one or sometimes two usually only slightly elongate, keeled anterior superciliary scale(s), one-fourth to half as long as horizontal eye diameter, usually followed by several similarly keeled, but much shorter scales; one, two, or more anterior sublabials greatly enlarged, higher than INL; scales of temporal arch usually larger than those above and below; ear opening small, by far not as high as eye, less high than SPL and INL together, usually about as high as SPL; low nuchal and dorsal crests present; two rows of middorsal scales strongly keeled and projecting upwards, but not laterally enlarged; other dorsal scales as well as lateral scales elevated and rugose to wrinkled in adults, smooth in very small juveniles, with very small granules more or less densely filling the interspaces between them; ventrals not or only slightly larger than largest dorsals, smooth; scales on anterodorsal surface of thigh multicarinate; scales on dorsal surface of forearm multicarinate; fourth toe with well-developed dilated pad, about three times width of distal phalanx; male dewlap large, extending posteriorly to between one-fourth and one-third of the distance between axilla and groin in large specimens, with well-demarcated gorgetal-sternal scale rows of densely arranged scales and widely spaced scales in the broad interspaces between the rows; female dewlap moderate, extending posteriorly to slightly beyond axilla, with more diffuse gorgetal-sternal rows owing to the less widely spaced scales in their interspaces. The completely everted hemipenis of SMF 89498 (Figs. 17 G���I) is a medium-sized, bilobate organ; sulcus spermaticus bordered by well-developed sulcal lips, opening at base of apex into two broad concave areas, one on each lobe; large asulcate processus and ridge present; a knob-like processus present on each lateral side of truncus just below base of apex; lobes finely calyculate, truncus with transverse folds. Coloration in life. Dactyloa ginaelisae exhibits a very variable coloration. Among the typical pattern elements are dark crossbands around the tail and, mostly with light centers, on dorsal surfaces of limbs and digits; a light stripe extending posteriorly from below the eye over the ear opening before bending down towards shoulder, paralleled above by a dark postorbital stripe with darker borders; a dark preaxillary blotch between tympanum and shoulder. Otherwise, dorsal and lateral surfaces with spots, blotches, reticula, or solid bars or chevrons that are lighter and/or darker than ground color; ventral surfaces usually comparably unicolor white or yellow; ground and marking colors of dorsal and lateral surfaces ranging from bright white over different shades of yellow, green, blue, and brown to black; iris gray to blue; male dewlap light salmon color; female dewlap pinkish to salmon color, in some individuals with dark blotches (Figs. 2; 8; 9). Apart from its highly variable pattern, this species is capable of overwhelming metachrosis (compare Figs. 9 B and C, D and E) as already noticed by Dunn (1937). The green phase, usually shown while the animal is sleeping, can culminate in restricting all colors to white, green, and blue, lightening up to invisibility some or all of the otherwise contrasting markings. Similarly, the dark or brown phase, often assumed when the animal is handled, can cause the whole animal to appear solid dark brown at its climax. In between, most coloration pattern elements of a given individual can apparently assume almost every color from the palette of this species. The only individual spotted at daytime was in the brown phase and showed no green at all. Color photographs of D. ginaelisae have been published by K��hler (2008), K��hler et al. (2008); Fl��schendr��ger and Wijffels (2009), Hamad (2009), and Uetz (2013). The coloration in life of an adult female (SMF 89501, Figs. 9 B���C, M���P) was recorded in the brown phase as follows: Dorsal and lateral ground color of body and limbs Straw Yellow (56) suffused with Tawny Olive (223 D); dorsal and lateral surfaces of body and limbs with Jet Black (89) irregular, sometimes broken, lines forming a reticulum suggesting transverse bands that enclose Robin Rufous (340) blotches, are suffused with Emerald Green (163) spots and disintegrate into mottles lateroventrally; dorsal surface of head Light Russet Vinaceous (221 D) with a reticulum of Sepia (219) bordered by shadings of Brick Red (132 A) and Leaf Green (146); ventral surface of head Cream Color (54) with Tawny (38) shadings and Maroon (31) mottling posteriorly; a Buff (124) supralabial stripe originating anterior to eye with shadings of Paris Green (63) (also present on borders of infralabials) and grading to dirty white posterior to eye continues bordered by Sepia (119) above ear to level above shoulder; a Brick Red (132 A) postorbital stripe bordered by Sepia (119) almost reaching to above shoulder; an elongate blotch of the same color behind ear opening; tail Drab-Gray (119 D) with Walnut Brown (221 B) transverse bands and various Vandyke Brown (221) scales; iris Sky blue (66); dewlap Pink (7) with dirty white scales and longitudinal series of Olive-Brown (28) flecks that fade away posteriorly. The coloration in life of another female (MHCH 2234, Figs. 9 G, Q) was recorded as follows: Dorsal ground color Lime Green (159), grading into Chartreuse (158) laterally; dorsal surfaces of body and limbs with partly broken Burnt Sienna (132) transverse bands, Spectrum Yellow (55) transverse stripes and Burnt Sienna (132), Sepia (119) as well as Shamrock Green (162 B) mottling; dorsal surface of head Tawny Olive (223 D) with a Sepia (219) V-shaped marking and some scale margins shaded in the same color; a Burnt Sienna (132) postocular stripe and blotch anterior to shoulder bordered by Sepia (119); a Spectrum Yellow (55) supralabial stripe extending above ear to above shoulder; supralabial Region Spectrum Yellow (55) with a suggestion of Yellow-Green (58), grading into Opaline Green (162 D) towards ventral surface of head; ventral surfaces of body, limbs and base of tail Straw Yellow (56) with sparse Cinnamon (123 A) mottling; tail grading into Olive-Yellow (52) with broad Sepia (219) transverse bands grading into Army Brown (219 B) ventrally; iris Sky Blue (66); dewlap Pink (7) with a suggestion of Salmon Color (6), with Opaline Green (162 D) scales aggregating to form longitudinal rows, and a suggestion of Yellowish Olive-Green (50) mottling. Coloration in preservative. After 22���76 months of preservation in 70 % ethanol, colors are largely reduced to white, cream, different shades of brown, and black. Some individuals have retained bluish-gray tonalities in certain places, often between eye and shoulder (Figs. 8 F���M; 18 R���S). Geographic distribution. As far as we know, Dactyloa ginaelisae is endemic to the Cordillera Central of western Panama, where it occurs at premontane and lower montane elevations on both Caribbean and Pacific versants. The species has been recorded, Published as part of Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix & K��hler, Gunther, 2013, Distribution and variation of the giant alpha anoles (Squamata: Dactyloidae) of the genus Dactyloa in the highlands of western Panama, with the description of a new species formerly referred to as D. microtus, pp. 1-54 in Zootaxa 3626 (1) on pages 14-20, DOI: 10.11646/zootaxa.3626.1.1, http://zenodo.org/record/283424

Dactyloa casildae (Arosemena, Ib����ez and de Sousa 1992) Figures 2; 10���11; 17 A���C; 18 A���D. Anolis casildae: Arosemena et al. (1992); Auth (1994); Young et al. (1999); Hofer and Bersier (2001); Ib����ez et al. (2001); Nicholson et al. (2001, 2005); Hamad (2009); Fl��schendr��ger and Wijffels (2009); Carrizo (2010); Chun (2010); Jaramillo et al. (2010); Stadler (2010); Casta��eda & de Queiroz (2011). Holotype. MVUP 755, male, from Quebrada Frank, 1100 m, 8 �� 44 ���N, 82 �� 13 ���W, Reserva Forestal de Fortuna, Chiriqu�� province, Panama (Fig. 1: loc. 29; Fig. 20). Diagnosis. A large species (maximum SVL 114 mm) of the genus Dactyloa (sensu Nicholson et al. 2012) that is most similar in external morphology to the other members of this clade found in western Panama (D. frenata, D. ginaelisae, D. ibanezi, D. insignis, D. kunayalae, and D. microtus). Dactyloa casildae can readily be distinguished from these six species by its coloration (described below, shown in Figs. 2, 10, and 11), and from all except D. ibanezi by its very long tail (more than 2.5 times SVL). Moreover, D. casildae differs from D. ginaelisae, D. insignis, D. kunayalae, and D. microtus in having long legs (tip of fourth toe of adpressed hind limb reaching to eye or beyond, usually to nostril or beyond, in D. casildae vs. at most to posterior border of eye), and from D. ginaelisae, D. insignis and D. microtus in having more horizontal loreal rows (6 or more in D. casildae vs. 5 or fewer). Additionally, D. casildae has more subdigital lamellae under the fourth toe (42 or more in D. casildae vs. 41 or fewer) as well as under the fourth finger (29 or more in D. casildae vs. 28 or fewer) than D. ibanezi and D. kunayalae. Description. Total length to 446 mm; SVL to 114 mm in males, to 99 mm in females; tail very long, about 2.5���3 times SVL, compressed (only very slightly in many individuals), without dorsal crest; legs long, tip of fourth toe of adpressed hind limb reaching at least to eye, in most individuals well beyond eye, and in some even beyond snout; internasals, canthals, and loreals keeled; scales of frontal and prefrontal area mostly keeled, some rugose or almost smooth (especially in juveniles); IP distinct, surrounded by smaller, keeled scales; parietal eye distinct; scales of SS enlarged, keeled; scales of supraorbital disk distinctly enlarged, keeled; a very elongate (in several specimens two, rarely three, accordingly shorter scales), keeled anterior superciliary scale, more than half as long (in some specimens almost as long) as horizontal eye diameter, usually followed by up to three similarly keeled but much shorter scales; anterior sublabials slightly enlarged, but never as high as INL, keeled; temporal arch weakly defined, with enlarged scales usually only in its anterior portion; ear opening large, higher than SPL and INL together, at least as high as eye, much larger than IP; nuchal and dorsal crests present in males; 2 rows of keeled middorsal scales slightly enlarged only in few specimens; other dorsal scales as well as lateral scales granular, conical to keeled; ventrals larger than largest dorsals, smooth; scales on anterodorsal surface of thigh mostly unicarinate, some bi- or tricarinate; scales on dorsal surface of forearm unicarinate, becoming multicarinate towards wrist; fourth toe with well-developed dilated pad, about three times width of distal phalanx; male dewlap very large, extending posteriorly to halfway between axilla and groin in large specimens, with well-demarcated gorgetal-sternal scale rows of densely arranged scales and widely spaced scales in the broad interspaces between the rows; female dewlap moderate, extending posteriorly to slightly beyond axilla, with more diffuse gorgetalsternal rows owing to the less widely spaced scales in their interspaces. The completely everted hemipenis of SMF 91454 (Figs. 17 A���C) is a small, bilobate organ; sulcus spermaticus bordered by well-developed sulcal lips, opening at base of apex into two broad concave areas, one on each lobe; an asulcate ridge present; a knob-like processus present on asulcate side of truncus; lobes finely calyculate, truncus with transverse folds. Coloration in life. Dorsal and lateral surfaces green to bluish green, either almost unicolor, or with brown, blue, and yellow spots or blotches, or additionally with oblique transverse bands that may appear solid or composed of blotches; some females of the unicolor and spotted variants with a dark-bordered light middorsal stripe; blue and yellow mottling usually most contrasting around chin, gular area, throat, and shoulders; light and dark lip bars often present; usually a postorbital stripe with dark borders; postorbital stripes of both sides often vaguely connected by a diffuse nuchal band; a less prominent preorbital stripe extending to snout in many specimens; tail with dark mottling suggesting diffuse crossbands; ventral surfaces unicolor yellow or yellowish green, in some individuals with dark mottling; iris brown; male dewlap cream-colored with gorgetal-sternal rows of yellow scales and interspersed blue or green scales, scales in interspaces between gorgetal-sternal rows widely spaced; female dewlap with contrasting yellow and green or blue mottling suggesting longitudinal stripes or a reticulate pattern, with only weakly demarcated gorgetal-sternal rows (Figs. 2; 10���11). Apart from the extremely variable color pattern, Dactyloa casildae is capable of considerable metachrosis (compare Figs. 10 A and I, or 10 E and F). While the description above refers to the colorful or green phase usually shown while the animal is sleeping, the dark or brown phase, typically assumed when the animal is handled, can cause all colors to turn dark brown, making most or all elements of a possible spotted or banded pattern disappear. Specimens spotted at daytime showed either of the color phases. Color photographs of D. casildae have been published by K��hler (2003, 2008), Hamad (2009), Fl��schendr��ger and Wijffels (2009), and Chun (2010). The coloration in life of a subadult male (MHCH 2121, Figs. 9 B, J) was recorded as follows: Dorsal and lateral ground color Apple Green (61); body with a series of broad transverse Greenish Olive (49) bands, interrupted by a Buff-Yellow (52) reticulate mottling covering dorsal and lateral surfaces of body, and mottled with Dark Grayish Brown (20); dorsal surface of head densely mottled with Buff (124) and Sepia (219); lateral and ventral surfaces of head Robin���s Egg Blue (93) mottled with Spectrum Yellow (55); a Sulphur Yellow (57) postorbital stripe bordered by Sepia (219) extending from eye to above ear opening; chin region with a Sulfur Yellow (157) reticulate mottling bordered by Sepia (119); dorsal surfaces of limbs and base of tail with Greenish Olive (49) mottling forming transverse bands; posterior portion of tail Citrine (51) with mostly ill-defined Sepia (219) transverse bands; ventral surface of body Cream Color (54) mottled with Citrine (51) and a suggestion of Lime Green (59); ventral surfaces of limbs and base of tail Citrine (51) mottled with Cream Color (54) and Lime Green (59); iris Clay Color (26); dewlap with a continuum of the chin coloration in anterior portion, grading into Pale Horn Color (92) longitudinal lines of scales suffused with Leaf Green (146); dewlap skin between these lines Lime Green (59) with solitary Leaf Green (146) scales; dewlap margin Cream Color (54). The coloration in life of an adult male (SMF 89455) was recorded as follows (translated from Hamad 2009): Dorsal surfaces with alternating crossbands from anterior head to tip of tail; transverse bands Yellowish Olive- Green (50), alternating with Olive-Yellow (52) bands with Straw Yellow (56) mottling, which grade into Turquoise Blue (65) and Straw Yellow (56) laterally; dorsal surface of head Spectrum Yellow (55), Greenish Olive (49), and Paris Green (63); lateral surfaces of head Spectrum Yellow (55), Turquoise Blue (65), and Sulphur Yellow (57), extending over ventral surfaces of head and body posteriorly to midventer; then grading into Beige (219 D) that continues to ventral surface of base of tail; tail thereafter with Sepia (219) and Raw Umber (23) crossbands; dewlap Cream (84) with longitudinal Orange-Yellow (18) stripes and Cerulean Blue (67) scales. Coloration in preservative. After 21���48 months of preservation in 70 % ethanol, all green and blue elements have turned to brown or, especially around head, to bluish gray; yellow throat markings are dirty white to yellowish brown (Figs. 18 A���D). After 10 years (SMF 85370), also the bluish shades have vanished almost completely. Geographic distribution. As currently understood, Dactyloa casildae is an endemic of the Panamanian Cordillera Central in Chiriqu�� province and the Comarca Ng��be-Bugl��. The species ranges from the eastern extreme of the Serran��a de Talamanca at Cerro Guayabo (RFLF, Chiriqu��) and nearby BPPS (Comarca Ng��be- Bugl��) over 60 airline km east to Cerro Santiago. It occurs on both Caribbean and Pacific slopes, at premontane and lower montane elevations of 990���1720 m asl (Fig. 20). Natural history notes. All collected specimens were encountered at night while they were sleeping on branches, lianas, or leaves 0.3���4 m above ground. Hamad (2009) spotted individuals of this species sleeping as high as 8 m above the R��o Hornito at RFLF. The only two individuals seen at daytime were perched head down on treetrunks 1.5 and 5 m above ground, respectively. During some surveys, Dactyloa casildae appeared very abundant at certain localities both at RFLF and Cerro Santiago in the Comarca Ng��be-Bugl��, with many sleeping individuals encountered very close to each other in some nights. In these surroundings, we also found this species to occur syntopically with D. ginaelisae, and in one case even to share a sleeping branch with that species. At the lowest collection site at R��o Hacha, we found D. casildae to occur in syntopy with D. kunayalae. Our automatized temperature recordings at collection localities of Dactyloa casildae (990���1560 m asl) range between 12.6���23.1 ��C. According to our combined dataset of 41 georeferenced occurrences, the species inhabits PMWF and LMWF, with temperatures between 11.8���27.4 ��C, mean annual temperatures of 17.8���21.2 ��C and a total annual precipitation of 2385���3376 mm. Conservation. Jaramillo et al. (2010) calculated an EVS of 13 for Dactyloa casildae, and assigned the species to the IUCN category NT. We calculated the species��� EVS as 5 (range) + 3 (persecution) + 4 (ecological distribution) = 12. Its extent of occurrence of just 426 km 2 and the continuing deforestation we observed in the region qualify D. casilde for the IUCN category ���Endangered��� (EN). Remarks. Our records expand the known range about 56 km southeastwards and 320 m uphill from previously reported localities (Nicholson et al. 2001; Chun 2010). Hofer and Bersier (2001) and consequently Nicholson et al. (2001) stated the westernmost specimens from BPPS to come from Bocas del Toro province. However, the region in question (Fig. 1: loc. 27) is presently situated within the Comarca Ng��be-Bugl��. As far as we know, Dactyloa casildae has not been reported from Bocas del Toro yet. Based on a sample of eight specimens, Nicholson et al. (2001) restricted the color morphs without transverse bands to females. Among our sample, there are also male specimens which exhibit a non-banded pattern (Figs. 10 G; 11 E), just like the specimen LACM FS 1084 pictured by Chun (2010: Fig. 4). Likewise, the middorsal stripe (Fig. 10 C) is not present in all specimens, not even in all females, that lack transverse bands (Fig. 10 D). One of our adult male specimens (SMF 89673, Figs. 10 A, I, Q) has no enlarged postcloacal scales. The same is true for the young male MHCH 2121 (Figs. 10 B, J; 18 A, B), that was erroneously pictured as a female by K��hler (2008: p. 99: Fig. 197) and could only be confidently sexed by verifying the presence of testes through an incision in the flanks. Considering the lack of postcloacal scales in some males, we assume that those allegedly female specimens having a chiefly white dewlap similar to the male dewlap (Nicholson et al. 2001) are actually males lacking enlarged postcloacal scales, whereas the female dewlap in reality always is contrastingly mottled., Published as part of Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix & K��hler, Gunther, 2013, Distribution and variation of the giant alpha anoles (Squamata: Dactyloidae) of the genus Dactyloa in the highlands of western Panama, with the description of a new species formerly referred to as D. microtus, pp. 1-54 in Zootaxa 3626 (1) on pages 23-27, DOI: 10.11646/zootaxa.3626.1.1, http://zenodo.org/record/283424

Dactyloa kunayalae (Hulebak, Poe, Ib����ez and Williams 2007) Figures 2; 15; 17 M��� O; 18 N���Q. Anolis kunayalae: Hulebak et al. (2007); Poe et al. (2009); Jaramillo et al. (2010); Casta��eda & de Queiroz (2011). Anolis new species 1: Nicholson et al. (2005). Holotype. MSB 72605, male, from along the trails of Parque National General de Divisi��n Omar Torrijos Herrera, 5 km North of El Cope, Cocl�� Province, Panama (8 �� 40.315 'N, 80 �� 35.518 'W) (Fig. 1: loc. 31; Fig. 22). Diagnosis. A large species (maximum SVL 109 mm) of the genus Dactyloa (sensu Nicholson et al. 2012) that is most similar in external morphology to the other members of this clade found in western Panama (D. casildae, D. frenata, D. ginaelisae, D. ibanezi, D. insignis, and D. microtus). Dactyloa kunayalae can readily be distinguished from these six species by its coloration (described below, shown in Figs. 2 and 15), and by the morphology of its toes: In contrast to the other species, D. kunayalae has narrow lamellar pads that are barely dilated under all toes and fingers, a distal phalanx (including claw) of the fourth toe that is longer than phalanges ii and iii combined, and the lowest number of subdigital lamellae under the fourth toe (35 or fewer in D. kunayalae vs. 38 or more). Moreover, it has fewer subdigital lamellae under the fourth finger than the other species except D. ibanezi (25 or fewer in D. kunayalae vs. 29 or more). In addition, D. kunayalae differs from D. casildae, D. frenata, and D. ibanezi in having short legs (tip of fourth toe of adpressed hind limb reaching to tympanum in D. kunayalae). Description. Total length to 354 mm (in our material); SVL to 109 mm in males, to 81 mm in females; tail long, about 2���2.5 times SVL, compressed, without dorsal crest; legs short, tip of fourth toe of adpressed hind limb reaching to tympanum; internasals, canthals, and loreals keeled; scales of frontal and prefrontal area mostly rugose to keeled; IP distinct, surrounded by smaller, rugose scales; parietal eye distinct; scales of SS slightly but distinctly enlarged, rugose to keeled; scales of supraorbital disk less conspicuously enlarged, rugose or usually keeled; an elongate, keeled anterior superciliary scale (or two or three accordingly shorter ones, then together) at least half as long as horizontal eye diameter, usually followed by 1���3 similarly keeled, but much shorter scales; anterior sublabials slightly enlarged, but less high than INL; scales of temporal arch much larger than those above and below; ear opening moderate to large, higher than SPL and INL together, slightly lower to higher than eye, much larger than IP; nuchal and dorsal crests present; females without, males with two rows of keeled middorsal scales slightly enlarged; other dorsal scales as well as lateral scales granular, conical to keeled; ventrals larger than largest dorsals, smooth to slightly keeled; scales on anterodorsal surface of thigh unicarinate; scales on dorsal surface of forearm unicarinate, becoming multicarinate towards wrist; all fingers and toes with very weakly developed dilated pad, not even twice the width of distal phalanx; distal phalanx including claw of fourth toe longer than phalanges ii and iii combined; male dewlap large, extending posteriorly to one-third of the distance between axilla and groin; female dewlap moderate to large, extending posteriorly to up to one-fourth of the distance between axilla and groin in large specimens; interspaces between well-demarcated gorgetal-sternal scale rows almost free of scales in both sexes. The completely everted hemipenis of SMF 91484 (Figs. 17 M��� O) is a small, slightly bilobate organ; sulcus spermaticus bordered by well-developed sulcal lips, opening at base of apex into two broad concave areas, one on each lobe; no asulcate processus or ridge present; lobes finely calyculate, truncus with transverse folds. Coloration in life. Dactyloa kunayalae presents a pronounced sexual dichromatism and apparently also ontogenetic changes in coloration. Dorsal and lateral surfaces in males largely green, turquoise, and blue, with darker green or blue as well as brown and yellow blotches that tend to form transverse and reticulate bands and lines (Figs. 15 A, F); females chiefly green with dark green to brown blotches usually suggesting transverse bands, and interspersed light markings (Figs. 15 B���D, G���L); juvenile females with conspicuous dark vertical lip bars that extend onto ventral surfaces of head; larger females and the adult male with rather immaculate, yellowish green lips; similarly, lateral pattern extending further onto venter in juveniles, whereas adults have a largely unicolor venter. Both sexes with crossbanded limbs, a diffuse postorbital blotch or stripe, more or less pronounced dark crossbands throughout the tail, and a reddish brown iris; male dewlap white with an orange-yellow margin; female dewlap white with a narrower margin of lighter yellow (Fig. 15). In the course of metachrosis, the ground color may assume a reddish brown similar to iris color, approximate the tonality of the dark markings, or the dark markings lighten up to become diffuse or barely visible. Color photographs of D. kunayalae have been published by Hulebak et al. (2007) and K��hler (2008). The coloration in life of the adult male (SMF 91484, Figs. 15 A, F, M) was recorded as follows: Dorsal ground color Bunting Green (150) with a suggestion of Lime Green (159), suffused with Cyan (164) on the flanks and bearing a few scattered Natal Brown (219 A) areas; body with mottling partly fusing to form a reticulum of Cream Color (54), grading into Raw Sienna (136) anteriorly and on the head; dorsal surfaces of body and legs mottled with Sepia (119); tail Light Drab (119 C), grading into Drab-Gray (119 D) ventrally, with Sepia (119) crossbands; lips and ventral surface of head Chartreuse (158); ventral surfaces of limbs, body and base of tail Sulphur Yellow (57); iris Mahogany Red (132 B); dewlap Sulphur Yellow (57) on basal portions, grading into dirty white towards the center, and a broad Spectrum Orange (17) with a suggestion of Orange Yellow (18) margin; dewlap scales dirty white and Chartreuse (158). The coloration in life of a young female (SMF 91485, Figs. 2; 15 B, H���L, N, O) was recorded as follows: Dorsal and lateral ground color of body, limbs, base of tail, and head Lime Green (159), grading into Pistachio (161) laterally, with Sepia (119) mottling except on snout that partly fuses to form broken crossbands on body, limbs, and base of tail; head, body, and limbs with scattered, diffuse blotches of Mahogany Red (132 B); tail Ground Cinnamon (239) with Warm Sepia (221 A) crossbands that grade into Sepia (119) dorsally, their Sepia (119) portions ventrally bordered by dirty white mottling; ventral surfaces of body, head, and limbs Tawny Olive (223 D), mottled with Warm Sepia (221 A) and dirty white, giving the impression of transverse stripes especially laterally and under head; iris Mahogany Red (132 B); dewlap Yellowish Olive-Green (50) with a Buff Yellow (53) margin and broad, partly broken gorgetal-sternal rows of small Buff-Yellow (53) scales. Coloration in preservative. After 22���74 months of preservation in 70 % ethanol all yellow, green, and blue colors have given way to brown, cream, and white. Only a very faint suggestion of yellow is retained on the margin of the male dewlap after 23 months (Figs. 18 N���Q). Geographic distribution. Dactyloa kunayalae is distributed from eastern Panama to western Panama, mostly along the Caribbean versant. This Panamanian endemic has been recorded from the Comarcas Kuna Yala and Ng��be-Bugl�� as well as from the provinces of Chiriqu��, Cocl��, Col��n, Veraguas, and probably also Dari��n (see remarks). It is a denizen of low and premontane elevations at 320���1050 m asl (Fig. 22). Natural history notes. Our specimens were encountered at night while they were sleeping on branches and lianas 2���8 m above ground. We found this species to occur in syntopy with Dactyloa frenata, D. ibanezi, and D. insignis at two sites (Fig. 1: locs. 9 and 16), and with D. casildae at one locality (Fig. 1: loc. 13). Our automatized temperature recordings at localities where we encountered Dactyloa kunayalae (480���1050 m asl) range between 17.6���27.6 ��C. According to our combined dataset of 26 georeferenced occurrences, the species inhabits LMF, PMMF, and PMWF, with temperatures between 14.7���30.1 ��C, mean annual temperatures of 20.6���25.4 ��C and a total annual precipitation of 1735���3452 mm. Conservation. Jaramillo et al. (2010) calculated an EVS of 12 for Dactyloa kunayalae, and assigned the species to the IUCN category LC. We calculated the species��� EVS as 5 (range) + 3 (persecution) + 4 (ecological distribution) = 12. Its extent of occurrence of 19441 km 2 and the continuing deforestation we observed in the region qualify D. kunayalae for the IUCN category ���Vulnerable��� (VU). Remarks. Our findings of Dactyloa kunayalae raise the altitudinal maximum by 250 m, originate from up to 180 km west of the documented range, and constitute the first records for Chiriqu��, Veraguas, and the Comarca Ng��be-Bugl�� (Hulebak et al. 2007). Moreover, GK had the privilege to examine the specimen FMNH 170034 (collected in August 1967 by Michael Duever at "border of Darien, Summit site, 8 �� 55 'N 77 �� 51 'W, 320m ") and identified it as an adult male D. kunayalae, probably the first specimen of this species ever collected. This specimen further extends the documented range some 135 km east-southeast from the nearest record in Nusagand�� (Hulebak et al. 2007), may be regarded as the first record for Dari��n province, and sets the new lower vertical limit for the species. Hulebak et al. (2007) described Dactyloa kunayalae to be long-legged, with the fourth toe reaching ���to eye when hindleg pressed against body.��� This trait was implied as diagnostic in the key of K��hler (2008). Strictly adhering to that key characteristic, all our specimens would have been identified as D. insignis or D. microtus, since their hind legs are short with the fourth toe merely reaching to tympanum. Although we do not know if there is really so much geographical variation in relative hind limb length in D. kunayalae, we consider it rather unlikely and prefer to attribute this discrepancy to different ways of extending or adpressing hind limbs, or bending the specimens��� bodies while doing so., Published as part of Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix & K��hler, Gunther, 2013, Distribution and variation of the giant alpha anoles (Squamata: Dactyloidae) of the genus Dactyloa in the highlands of western Panama, with the description of a new species formerly referred to as D. microtus, pp. 1-54 in Zootaxa 3626 (1) on pages 36-38, DOI: 10.11646/zootaxa.3626.1.1, http://zenodo.org/record/283424

Dactyloa frenata (Cope 1899) Figures 2; 12; 17 D���F; 18 E���G. Anolis frenatus: Cope (1899); Dunn (1937); Breder (1946); Peters and Donoso-Barros (1970); Myers (1971); Scott et al. (1976); Myers (1977); Savage and Talbot (1978); Rand and Myers (1990); Arosemena et al. (1992); Auth (1994); Mart��nez and Rodr��guez (1994); Mart��nez et al. (1995); Ib����ez et al. (1995, 1996, 1997); Young et al. (1999); Ib����ez et al. (2001); Poe (2004); Nicholson et al. (2005); Hamad (2009); Fl��schendr��ger and Wijffels (2009); Poe et al. (2009); Ugueto et al. (2009); Carrizo (2010); Jaramillo et al. (2010); Lotzkat et al. (2010 b); Stadler (2010); Casta��eda & de Queiroz (2011). Anolis purpurescens: Taylor (1956) and references therein. Holotype. Lost; type locality in Colombia, but exact location uncertain according to Savage and Talbot (1978). Diagnosis. A large species (maximum SVL 143 mm) of the genus Dactyloa (sensu Nicholson et al. 2012) that is most similar in external morphology to the other members of this clade found in western Panama (D. casildae, D. ginaelisae, D. ibanezi, D. insignis, D. kunayalae, and D. microtus). Dactyloa frenata can readily be distinguished from these six species by the presence of a pronounced light interorbital bar with dark anterior and posterior borders, and by its unique color pattern on dorsum and flanks consisting of dark blotches or ocelli arranged to form oblique bands (Fig. 12). Moreover, D. frenata has 9 or more SPL (vs. usually 8 or fewer) to the level below center of eye. In addition, D. frenata differs from D.ginaelisae, D. insignis, D. kunayalae, and D. microtus in having long legs (tip of fourth toe of adpressed hind limb reaching to eye or beyond in D. frenata), and from D. ginaelisae, D. ibanezi, D. insignis, D. kunayalae, and D. microtus in having the suboculars separated from the SPL by at least one scale row (vs. suboculars and SPL in contact). Description. Total length to 430 mm; SVL to 143 mm in males, to 118 mm in females; tail long, about two times SVL, compressed, without dorsal crest; legs long, tip of fourth toe of adpressed hind limb reaching to eye or beyond; internasals, canthals, and loreals keeled; scales of frontal and prefrontal area mostly smooth, some weakly keeled; IP distinct, surrounded by smaller, mostly smooth, scales; parietal eye distinct; scales of SS not distinctly enlarged, keeled; scales of supraorbital disk only slightly enlarged, keeled; an elongate, keeled anterior superciliary scale, at least half as long as horizontal eye diameter, often followed by one or two similarly keeled, but much shorter scales; anterior sublabials not distinctly enlarged, keeled; scales of temporal arch slightly larger than those above and below; ear opening large, higher than SPL and INL together, at least as high as eye, much larger than IP; nuchal and dorsal crests pronounced in males, weakly developed in female; 2���4 rows of keeled middorsal scales slightly enlarged; other dorsal scales as well as lateral scales granular, smooth to slightly keeled; ventrals slightly larger than largest dorsals, smooth to very slightly keeled; scales on anterodorsal surface of thigh mostly unicarinate, some bi- or tricarinate; scales on dorsal surface of forearm unicarinate, becoming multicarinate towards wrist; fourth toe with well-developed dilated pad, about three times width of distal phalanx; male dewlap very large, extending posteriorly to almost one-half of the distance between axilla and groin in large specimens; female dewlap moderate, extending posteriorly to slightly beyond axilla; dewlap in both sexes with welldemarcated gorgetal-sternal scale rows and scattered scales in their interspaces. The completely everted hemipenis of SMF 89467 (Figs. 17 D���F) is a small, slightly bilobate organ; sulcus spermaticus bordered by well-developed sulcal lips, opening at base of apex into two broad concave areas, one on each lobe; a small asulcate ridge present; lobes finely calyculate, truncus with transverse folds. Coloration in life. Dorsal and lateral surfaces green, usually involving brownish, yellowish, or bluish tonalities; series of darker green to dark brown blotches or ocelli form oblique transverse bands on dorsum, flanks, limbs, and anterior portion of tail; posterior portion of tail with dark crossbands; alternating yellow and dark stripes radiate more or less distinctly from eye, the upper ones meeting on dorsal surface of head to form a pronounced light-colored interorbital bar usually with dark anterior and posterior borders; ventral surfaces largely unicolor cream or gray, becoming lighter towards head; iris brown; male dewlap white, female dewlap brown (Figs. 2; 12). We did not observe metachrosis in Dactyloa frenata. Color photographs of D. frenata have been published by Savage (2002), K��hler (2003, 2008), Fl��schendr��ger and Wijffels (2009), Stadler (2010), Carrizo (2010), and Uetz (2013), among others. The coloration in life of an adult male (SMF 89467, Figs. 12 A, E) was recorded as follows (translated from Stadler 2010): Dorsal surfaces of body, limbs, and anterior portion of tail Opaline Green (162 D) with numerous Parrot Green (60) ocelli, partly bordered by Straw Yellow (56) and Spectrum Yellow (55), which are arranged in a pattern of transverse bands; middle portion of tail Sepia (119), posterior portion Olive-Yellow (52); anterior dorsal surface of head with a heterogeneous pattern of Salmon Color (6), Olive-Gray (42), and Cyan (164); a Salmon Color (6) interorbital bar bordered anteriorly and posteriorly by Greenish Olive (59) stripes; posterior dorsal surface of head with a random pattern of Flesh Color (5), Paris Green (63), and Salmon Color (6) scales; lateral surfaces of head from snout to posterior border of orbit Olive Gray (42), those posterior to orbit Cyan (164); Spectrum Yellow (55) and Grayish Olive (43) stripes extending from eyes radially in all directions; ventral surfaces largely dirty white, grading into Spectrum Yellow (53) laterally; ventral surfaces of hind limbs and anterior portion of tail Straw Yellow (53), those of middle and posterior portions of tail Sepia (119) and Olive-Yellow (52), respectively; dewlap Cream Color (54) with Sulphur Yellow (157) scales on the upper and Glaucous (80) scales in the lower half; margin dirty white anteriorly and Cream Color (54) posteriorly; iris Antique Brown (37). Coloration in preservative. After 21���45 months of preservation in 70 % ethanol, the greenish tonalities have turned to brown, gray, or bluish gray (the latter especially on head and flanks), venters are cream-colored (Figs. 18 E���G). Geographic distribution. Dactyloa frenata is distributed from the Caribbean versant of Costa Rica to Colombia, at low and premontane elevations of 2���900 m asl. In Costa Rica and extreme western Panama, it is restricted to the Caribbean versant of the Talamancan highlands. From the Fortuna depression eastwards, D. frenata is also found in the Pacific drainage. In Panama, the species has been recorded from the provinces of Bocas del Toro, Chiriqu��, Cocl��, Col��n, Dari��n, Panam��, and Veraguas, and from the Comarcas Kuna Yala and Ng��be-Bugl�� (Fig. 20). Natural history notes. All specimens were encountered at night while they were sleeping on branches or lianas 1.5���4 m above ground. Around our collection sites of Dactyloa frenata, this species shares its habitat with D. ibanezi, D. insignis, and D. kunayalae. Our automized temperature recordings at collection localities of Dactyloa frenata (700���900 m asl) range between 16.1���26.2 ��C. According to our combined dataset of 75 georeferenced occurrences, the species inhabits LMF, LWF, PMMF, and PMWF, with temperatures between 15.2���32.4 ��C, mean annual temperatures of 21.2���26.8 ��C and a total annual precipitation of 1735���4417 mm. Conservation. Jaramillo et al. (2010) calculated an EVS of 8 for Dactyloa frenata, and assigned the species to the IUCN category LC. We calculated the species��� EVS as 1 (range) + 3 (persecution) + 4 (ecological distribution) = 8. Its extent of occurrence of more than 49000 km 2 does not qualify D. frenata for any of the ���Threatened��� IUCN categories. Considering the continuing deforestation we observed in the region, we place the species in the category ���Near Threatened��� (NT). Remarks. The adult male in our sample raised the upper vertical limit for this species to 900 m asl (Stadler 2010). As confirmed by Ugueto et al. (2009), Dactyloa frenata does definitely not occur in Venezuela., Published as part of Lotzkat, Sebastian, Hertz, Andreas, Bienentreu, Joe-Felix & K��hler, Gunther, 2013, Distribution and variation of the giant alpha anoles (Squamata: Dactyloidae) of the genus Dactyloa in the highlands of western Panama, with the description of a new species formerly referred to as D. microtus, pp. 1-54 in Zootaxa 3626 (1) on pages 27-29, DOI: 10.11646/zootaxa.3626.1.1, http://zenodo.org/record/283424

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 R��o Chiriqu�� Mal��, approximately 5 km N of the La Fortuna dam reservoir (8.7890 ��N, 82.2154 ��W, 1060 m), Bosque Protector Palo Seco, Comarca Ng��be-Bugl�� (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 Iba��ez, 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 Bola��os 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 Bola��os 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, Garc��a-Rodr��guez, Artavia, Bola��os, 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, Garc��a-Rodr��guez, Artavia, Bola��os, 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 R��o Chiriqu�� Mal��, 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 K��hler. 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., Published as part of Hertz, Andreas, Lotzkat, Sebastian & K��hler, Gunther, 2013, A new species of Bolitoglossa (Caudata, Plethodontidae) from the continental divide of western Panama, pp. 463-475 in Zootaxa 3636 (3) on pages 465-471, DOI: 10.11646/zootaxa.3636.3.5, http://zenodo.org/record/221943

Anolis guentherii Bocourt Anolis guentherii Bocourt 1873: 61; holotype (MNHN 712) from ��� Mexique.��� Troschel (1874: 160), Boulenger (1885: 54), Cope (1885: 391), Cope (1887: 28), Anolis guentheri G��nther (1885: 46), Barbour (1934: 133), Guib�� (1954: 37), Stuart (1955: 4) Anolis guntheri Stuart (1955: 19), Peters and Donoso-Barros (1970: 59), Fitch and Henderson (1973: 126) Anolis guntherii. Smith and Taylor (1950 a: 63), Smith and Taylor (1950 b: 320), Brygoo (1989: 52) Description of the holotype of Anolis guentherii (MNHN 712; Fig. 2). Adult female as indicated by shape of base of tail and absence of dewlap; SVL 45.5 mm; tail incomplete, regenerated; tail slightly compressed in cross section, tail height 1.7 mm, tail width 1.6 mm; axilla to groin distance 18.5 mm; head length 12.8 mm, head length/ SVL ratio 0.28; snout length 6.0 mm; head width 7.7 mm; longest toe of adpressed hind limb reaching to a point between tympanum and eye; shank length 10.6 mm, shank length/head length ratio 0.83; longest finger of extended forelimb reaching to tip of snout; longest finger of adpressed forelimb failing to reach anterior insertion of hind limbs by 2.5 mm. Scales on snout keeled; 6 postrostrals; 8 scales between nasals; 1 elongate prenasal scale in contact with rostral; scales in shallow prefrontal depression keeled; supraorbital semicircles well developed, separated by a minimum of 2 scale rows; supraorbital disc composed of 5���7 distinctly enlarged keeled scales; circumorbital row of small scales complete; a single elongate superciliary; 3���4 rows of small scales extending between enlarged supraorbitals and superciliary; interparietal scale well developed, 1.7 x 1.2 mm (length x width), surrounded by scales of moderate size; 4 scales present between interparietal and supraorbital semicircles; canthal ridge distinct, composed of 3 large (anteriormost one largest) and 3 small anterior canthal scales; 9 scales present between second canthals; 11 scales present between posterior canthals; 28 (right)��� 29 (left) mostly obtusely keeled loreal scales in a maximum of 5 horizontal rows; subocular scales in a single series, 3 scales broadly contacting supralabials; 6 (right)��� 7 (left) supralabials to level below center of eye; ear opening 0.6 x 1.1 mm (length x height), ratio tympanum height/parietal scale length 0.65; mental distinctly wider than long, almost completely divided medially, bordered posteriorly by 5 postmentals (outer pair largest); 6 infralabials to level below center of eye; 2 anterior sublabials greatly enlarged, in contact with infralabials; slightly keeled granular scales present on chin and throat; dewlap absent; dorsal trunk scales weakly keeled, juxtaposed to subimbricate with rounded posterior margins, 2 medial rows slightly enlarged, largest dorsal scales about 0.24 x 0.23 mm (length x width); about 60 medial dorsal scales in one head length; about 94 medial dorsal scales between axilla and groin; lateral scales keeled, granular, homogeneous, average size 0.15 mm in diameter; ventrals at midbody smooth, slightly bulging, subimbricate, about 0.29 x 0.34 mm (length x width); about 48 ventral scales in one head length; about 64 ventral scales between axilla and groin; about 94 scales around midbody; caudal scales keeled; caudal middorsal scales enlarged; lateral caudal scales keeled, homogeneous, although indistinct division in segments discernible; postcloacal scales not enlarged; no tube-like axillary pocket present; scales on dorsal surface of forelimb weakly keeled, imbricate; digital pads greatly dilated, dilated pad (width 1.3 mm) several times the width of non-dilated scales on distal phalanx (0.3 mm); 36 (right)��� 35 (left) lamellae under phalanges II���IV of fourth toe; 6 scales under distal phalanx of fourth toe. Snout and frontal region cream color with a suffusion of grayish brown; dorsal surfaces of posterior portion of head, neck, and anterior body with a faint pattern of pale spots; ventral surface of head pale grayish brown with brown wavy longitudinal lines, forming a reticulum on sides of head; venter pale grayish brown; dorsal and ventral surfaces of limbs and tail pale cream color. Taxonomic conclusions: A comparison of the holotype of Anolis guentherii with all other known species of anoles leads me to the conclusion that it is conspecific with A. grahami Gray 1845. Diagnostic characters include: (1) two anterior sublabials greatly enlarged, in contact with infralabials; (2) digital pads greatly dilated, dilated pad several times the width of non-dilated scales on distal phalanx; (3) supraorbital semicircles well developed, separated by a minimum of 2 scale rows, 4 scales present between interparietal and supraorbital semicircles; (4) a single elongate superciliary; (5) longest toe of adpressed hind limb reaching to a point between tympanum and eye, shank length/head length ratio 0.83; (6) smooth ventral scales; (7) dorsal trunk scales weakly keeled, juxtaposed to subimbricate with rounded posterior margins, 2 medial rows slightly enlarged; (8) ventral surface of head pale with brown wavy longitudinal lines, forming a reticulum on sides of head, dorsal surfaces of posterior portion of head, neck, and anterior body with a faint pattern of pale spots. Therefore, I herewith place Anolis guentherii Bocourt 1973 in the synonymy of A. grahami Gray 1845. This species occurs natively on Jamaica, and has been introduced on Bermuda (Russell and Bauer 1991, Schwartz and Henderson 1991). Because of its smooth ventral scales and its large interparietal plate, MNHN 712 is assigned to the nominal subspecies. The type locality ��� Mexique ��� as stated in the original description of A. guentherii is clearly erroneous., Published as part of K��hler, Gunther, 2012, Taxonomic status of two enigmatic Mexican anoles: Anolis cumingii Peters 1863 and Anolis guentherii Bocourt 1873 (Reptilia, Squamata, Dactyloidae), pp. 82-88 in Zootaxa 3551 on pages 85-86, DOI: 10.5281/zenodo.282852, {"references":["Bocourt, M. F. (1873) In A. Dumeril, Bocourt, M. - F. & Mocquard, F. (eds.), Etudes sur les reptiles. In: Recherches zoologiques pour sevir a l´histoire de la faune de l´Amerique Centrale et du Mexique. Mission Scientifique au Mexique et dans l´Amerique Centrale, recherches Zool., part 3, sect. 1. Imprimerie au Nat.; Paris.","Troschel, F. H. (1874) Bericht uber die Leistungen in der Herpetologie wahrend des Jahres 1873. Archiv fur Naturgeschichte, 40 (2), 146 - 170.","Boulenger, G. A. (1885) Catalogue of the Lizards in the British Museum (Natural History). 2 nd ed., Vol. II. London (Trustees of the British Museum).","Cope, E. D. (1885) A contribution to the herpetology of Mexico. V. Notes on anoles. Proceedings of the American Philosophical Society, 22, 390 - 393.","Cope, E. D. (1887) Catalogue of Batrachians and reptiles of Central America and Mexico. Bulletin of the United States National Museum, 32, 1 - 98.","Gunther, A. C. L. G. (1885 - 1902) Biologia Centrali - Americana. Reptila and Batrachia. Porter, London.","Barbour, T. (1934) The anoles II. The mainland species from Mexico southward. Bulletin of the Museum of Comparative Zoology, 77, 119 - 155.","Guibe, J. (1954) Catalogue des types de lezards du Museum National d'Histoire Naturelle. Paris.","Stuart, L. C. (1955) A brief review of the Guatemalan lizards of the genus Anolis. Miscellaneous Publication Museum of Zoology University of Michigan, 91, 1 - 31.","Peters, J. A. & Donoso-Barros, R. (1970) Catalogue of the Neotropical Squamata. Part II. Lizards and Amphisbaenians. Smithsonian Institution United Stated National Museum Bulletin, 297, 1 - 293.","Fitch, H. S. & Henderson, R. W. (1973) A new anole (Reptilia: Iguanidae) from southern Veracruz, Mexico. Journal of Herpetology, 7 (2), 125 - 128.","Smith, H. M., & Taylor, E. H. (1950 a) An annotated checklist and key to the reptiles of Mexico exclusive of the snakes. Bulletin of the United States National Museum, 199, 1 - 253.","Smith, H. M., & Taylor, E. H. (1950 b) Type localities of Mexican reptiles and amphibians. Kansas University Science Bulletin, 33 (8), 313 - 380.","Brygoo, E. R. (1989) Les types d'Iguanides (Reptiles, Sauriens) du Museum national d'Histoire naturelle Catalogue critique. Bulletin Museum National d'Histoire Naturelle, 4 Ser. 11 section A no. 3, supplement, 1 - 112.","Russell, A. P. & Bauer, A. M. (1991) Anolis grahami. Catalogue of American Amphibians and Reptiles, 514, 1 - 4.","Schwartz, A. & Henderson, R. W. (1991) Amphibians and reptiles of the West Indies. University of Florida Press, Gainesville."]}

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"]}

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

Anolis tenorioensis sp. nov. Figs. 1, 2 a, 3, 4, 5, 6 Holotype. SMF 91985, an adult male from near Laguna Danta, Volc��n Tenorio (10.70521 ��N, 85.03068 ��W, WGS 84), 1160 m, Alajuela Province, Costa Rica; collected 5 March 2011 by Gunther K��hler, Donald Varela Soto, and Jos��ph Vargas. Field tag number GK 3324. Paratypes. SMF 91986, UCR 21182, same collecting data as holotype. Diagnosis. A medium-sized species (SVL in largest specimen 45.0 mm) of the genus Anolis (sensu Poe 2004) that differs from all Lower Central American anoles except Anolis altae, A. carpenteri, A. fortunensis, A. fuscoauratus, A. gruuo, A. kemptoni, A. monteverde and A. pseudokemptoni in that it is short-legged (longest toe of adpressed hind leg reaches only to ear opening), has smooth to slightly keeled ventral scales, and a slender habitus, often delicate. Anolis tenorioensis differs from all the species in the cluster listed above by having the ventral surfaces of body and limbs distinctly reticulated with dark brown pigment (versus immaculate dirty white or cream colored); and a male dewlap that is dark red with brown blotches (versus more or less uniformly reddish orange in A. altae and A. monteverde, uniform orange in A. carpenteri, posterior portion of male dewlap greenish beige, anterior portion reddish orange in A. fortunensis, uniform pinkish brown to red in A. fuscoauratus, more or less uniform dull orange in A. gruuo, posterior portion rose pink, anterior portion burnt orange, small cream colored basal blotch in some populations in A. kemptoni, posterior portion vinaceous, anterior portion orange, anterodorsal corner cream color in A. pseudokemptoni); and by having a divided prenasal scale (versus usually a single elongated prenasal scale). It also differs from A. fortunensis, A. kemptoni, and A. monteverde by having a bilobate hemipenis (versus unilobate). Additionally, it can be differentiated from A. carpenteri by having a brownish dorsal ground color (versus greenish). Anolis tenorioensis is most similar to A. altae and A. monteverde, from which it further differs in several morphometric and pholidotic characteristics, such as relative head length (ratio head length / SVL) and number of scales around midbody (see Table 1). Description of the holotype. Adult male as indicated by everted hemipenes and presence of large dewlap (Fig. 3); SVL 42.0 mm; tail length 36.0 mm, tail incomplete; tail round in cross section, tail height and width 1.9 mm; axilla to groin distance 18.7 mm; head length 10.7 mm, head length/SVL ratio 0.255; snout length 4.9 mm; head width 6.1 mm; longest toe of adpressed hind limb reaching to tympanum; shank length 9.9 mm, shank length/head length ratio 0.93; longest finger of extended forelimb reaching nostril; longest finger of adpressed forelimb reaches a point 3.6 mm in front of anterior insertion of hind limbs. Most scales on snout keeled (Fig. 4); 6 postrostrals; 7 scales between nasals; 2 prenasal scales, the lower one in contact with both rostral and first supralabial; scales in distinct prefrontal depression slightly wrinkled; supraorbital semicircles well developed, separated by a minimum of 1 scale; supraorbital disc composed of 6���7 distinctly enlarged keeled scales; circumorbital row incomplete, therefore, one enlarged supraorbital in contact with supraorbital semicircles; two elongated, strongly overlapping superciliaries, the anterior one much larger than the posterior one; about 3 or 4 rows of small keeled scales extending between enlarged supraorbitals and superciliaries; parietal depression absent; interparietal scale well developed, 2.0 x 1.2 mm (length x width), surrounded by scales of moderate size; 3 scales present between interparietal and supraorbital semicircles; canthal ridge distinct, composed of 3 large (posterior two largest) and 5 small anterior canthal scales; 7 scales present between second canthals; 7 scales present between posterior canthals; 30 (right)��� 28 (left) mostly keeled loreal scales in a maximum of 5 horizontal rows; 6 keeled subocular scales arranged in a single row; 7 supralabials to level below center of eye; 5 suboculars broadly in contact with supralabials; ear opening 0.7 x 0.9 mm (length x height); mental distinctly wider than long, almost completely divided medially, bordered posteriorly by 6 postmentals; 7 (right)��� 9 (left) infralabials to level below center of eye; sublabials undifferentiated; keeled granular scales present on chin and throat; dewlap extending from level below posterior margin of eye to level of axilla; dorsum of body with weakly keeled imbricate scales with rounded posterior margins, medial rows not enlarged, largest dorsal scales about 0.34 x 0.31 mm (length x width); about 47 medial dorsal scales in one head length; about 72 medial dorsal scales between axilla and groin; lateral scales keeled, granular and homogeneous, average size 0.15 mm in diameter; ventrals at midbody smooth, slightly bulging, subimbricate, about 0.34 x 0.29 mm (length x width); about 48 ventral scales in one head length; about 74 ventral scales between axilla and groin; 101 scales around midbody; caudal scales keeled except ventrally at base of tail; caudal middorsal scales not enlarged, without whorls of enlarged scales, although an indistinct division in segments is discernible; a pair of slightly enlarged postcloacal scales present, about 0.60 mm wide; no tube-like axillary pocket present; scales on dorsal surface of forelimb keeled, imbricate, about 0.21 x 0.31 mm (length x width); digital pads dilated, dilated pad about 3 times width of non-dilated scales on distal phalanx; distal phalanx narrower than and raised from dilated pad; 24 (right)��� 26 (left) lamellae under phalanges II���IV of fourth toe; 8 scales under distal phalanx of fourth toe. The completely everted hemipenis is a large bilobate organ (Fig. 5); sulcus spermaticus bordered by well developed sulcal lips and bifurcating at base of apex into branches that continue to tips of lobes; a weak asulcate ridge present; apex strongly calyculate, truncus with transverse folds. The extended dewlap in life (Fig. 2 a) has six horizontal gorgetal-sternal rows with a declining number of scales per row from base to tip (17���19 scales per row on basal portion, 5���9 towards tip); about 50 scales in marginal series. Coloration in life was recorded as follows: Dorsal surface of head Burnt Umber (22); dorsal ground color of body and limbs Grayish Horn Color (91), with shades of Buff (124) and some Dark Brownish Olive (129) pigmentation along vertebral line and indistinct Dark Brownish Olive (129) crossbands on limbs; tail Straw Yellow (56) with Ground Cinnamon (239) bands; venter Buff (124) with Dark Brownish Olive (129) speckling; chin Buff (124) laterally and dirty white with Dark Brownish Olive (129) reticulum medially; iris Mikado Brown (121 C); dewlap Pratt���s Ruby (210) with Vandyke Brown (121) blotches. Coloration after 4 months preservation in 70 % ethanol was recorded as follows: Dorsal ground color grayish brown with indistinct paler shadings, a cream colored temporal blotch, a dark brown interorbital bar, and faint crossbands on limbs and tail; ventral surfaces of head, body, limbs, and tail dirty pale grayish cream with a distinct dark brown reticulum; dewlap reddish (except damaged basal area) with dark brown and dirty white gorgetals, and with dark brown blotches, mostly around gorgetals. Variation. The paratypes agree well with the holotype in general appearance, morphometrics and scalation (see Table 1). Coloration in life of an adult female paratype (GK 3326) was recorded as follows: Dorsal ground color Raw Umber (123) with shades of Straw Yellow (56), a Mars Brown (223 A) interorbital bar; snout Verona Brown (223 B), and a few Dark Brownish Olive (129) markings along vertebral line; tail with Verona Brown (223 B) crossbands; venter Cream Color (54) with a paramedian zone of Sepia (219) reticulation; iris Verona Brown (223 B); dewlap Pale Horn Color (92) with a few Grayish Horn Color (91) gorgetals. Number of scales between 2 nd canthals 5���11 (8.63 �� 1.51) 6���9 (7.81 �� 0.98) 7 Number of scales between posterior canthals 7���15 (9.80 �� 1.70) 7���11 (9.63 �� 1.26) 7���10 Etymology. The name tenorioensis is derived from Volc��n Tenorio where the type series of the species was collected and to where it is probably restricted, and the Latin suffix - ensis (denoting place or locality). Natural history notes. All type specimens were collected at night while the lizards were sleeping on low vegetation (mostly on fern leaves) within the forest (Premontane Rainforest fide Savage 2002) near an open area that is flooded most of the year and known locally as Laguna Danta (see also Figs. 7 and 8). At the type locality, a recently fallen tree had created a hole in the canopy allowing sun to reach the ground. No daytime searches were done. Geographic distribution and conservation. As currently known, Anolis tenorioensis is restricted to the highlands of Volc��n Tenorio in the Cordillera de Guanacaste, Costa Rica. It occurs in a protected area (Parque Nacional Volc��n Tenorio). Only three specimens of this species are known to science. However, given its presumably small geographic range, it seems justified to classify A. tenorioensis as Critically Endangered based on criterion B. 1.a. of the IUCN Red List Categories and Criteria (IUCN 2011)., Published as part of K��hler, Gunther, 2011, A new species of anole related to Anolis altae from Volc��n Tenorio, Costa Rica (Reptilia, Squamata, Polychrotidae), pp. 29-42 in Zootaxa 3120 on pages 30-39, DOI: 10.5281/zenodo.205375, {"references":["Poe, S. (2004) Phylogeny of anoles. Herpetological Monographs, 18, 37 - 89.","Savage, J. M. (2002) The amphibians and reptiles of Costa Rica. A herpetofauna between two continents, between two seas. University of Chicago Press, Chicago, Illinois, U. S. A., and London, U. K."]}

Anolis charlesmyersi sp. nov. Figs. 4, 5, and 6 Anolis pentaprion: Wettstein (1934), Taylor (1956; in part), Williams and Smith (1968 ��� 1966 ���), Myers (1971; in part), Anolis pentaprion pentaprion: Etheridge (1959; in part.), Holotype. SMF 89688, adult male, from trail to R��o Majagua, Los Algarrobos, 8.48927 ��N, 82.43333 ��W, ca. 130 m a.s.l., Province Chiriqu��, Panama; collected by Sebastian Lotzkat and Andreas Hertz on 23 April 2009. Paratypes. SMF 89508, 90053, adult males, from trail to R��o Majagua, Los Algarrobos, 8.48456 ��N, 82.43417 ��W, ca. 110 m a.s.l., Province Chiriqu��, Panama; collected by Sebastian Lotzkat and Andreas Hertz on 7 June 2008. Diagnosis. A medium-sized species (SVL in largest specimen examined 78.0 mm in males, 76.7 mm in females) of the genus Anolis (sensu Poe, 2004) that differs from all Central American anoles except A. beckeri, A. cristifer, A. fungosus, A. pentaprion, A. salvini, A. utilensis by (1) having (1) extremely short hind legs (fourth toe of adpressed hindlimb usually reaching to level of tympanum; ratio shank length / head length 0.49���0.84); (2) smooth, juxtaposed ventral scales; (3) a relatively short tail (ratio tail length / SVL 1.0��� 1.5). Anolis charlesmyersi differs from A. beckeri by having a pronounced serrated caudal crest (versus lacking a pronounced serrated caudal crest). Anolis charlesmyersi differs from A. cristifer by having a double row of slightly enlarged middorsal scales, not forming a serrated crest (versus a serrated middorsal crest, at least on posterior portion of dorsum, present in A. cristifer). Anolis charlesmyersi differs from A. fungosus by lacking bony parietal protuberances (versus having a pair of small bony parietal protuberances posterior and lateral to interparietal plate). Anolis charlesmyersi differs from A. pentaprion by having a brick red (in life) dewlap with few large widely spaced gorgetal scales scattered across the dewlap or in loosely arranged rows with 4���9 widely spaced scales (versus a pink dewlap with purple pigment between rows of gorgetal scales and with regular rows of 17���25 small narrowly spaced gorgetal scales). Anolis charlesmyersi differs from A. salvini by having smooth, obliquely conical ventral scales (versus keeled), lacking a dark interorbital bar (versus present), and lacking enlarged postcloacal scales (versus males with a pair of greatly enlarged postcloacal scales). Anolis charlesmyersi differs from A. utilensis by having the proximal subdigital scales of toes differentiated as slightly broadened lamellae (versus those scales granular) and by having a pronounced serrated caudal crest (versus lacking a pronounced serrated caudal crest). Description of holotype. Adult male as indicated by everted hemipenes; SVL 65.0 mm; tail length 93.0 mm, tail complete; tail length/SVL ratio 1.43; tail compressed in cross section, tail height 4.4 mm, tail width 3.1 mm; axilla to groin distance 24.9 mm; head length 17.5 mm, head length/SVL ratio 0.27; snout length 8.5 mm; head width 10.4 mm; longest toe of adpressed hind limb reaching to a point between shoulder and tympanum; shank length 13.6 mm, shank length/head length ratio 0.78; longest finger of extended forelimb reaching to a point between eye and nostril; longest finger of adpressed forelimb reaches to a point 2 mm anterior to insertion of hind limbs. Scales on snout tuberculate to weakly keeled; 10 postrostrals; 7 scales between nasals; scales in distinct prefrontal depression slightly wrinkled to tuberculate; supraorbital semicircles well developed, 2 (right)��� 3 (left) scales broadly in contact; supraorbital disc composed of 11���14 distinctly enlarged smooth scales; three short superciliaries, posterior one shortest; about 4 rows of small keeled scales extending between enlarged supraorbitals and superciliaries; interparietal scale well developed, 3.1 x 1.3 mm (length x width), surrounded by scales of moderate size; interparietal and supraorbital semicircles in broad contact; canthal ridge distinct, composed of three large and four small anterior canthal scales; 6 scales present between second canthals; 8 scales present between posterior canthals; 19 (right)��� 20 (left) loreal scales in a maximum of 3 (right)��� 4 (left) horizontal rows, with the scales of lower rows mostly smooth, and those of upper rows mostly weakly keeled; subocular scales smooth to wrinkled, five subocular scales broadly in contact with five supralabials; 8 supralabials to level below center of eye; ear opening 0.8 x 0.9 mm (length x height); mental 2.0 x 0.9 mm (width x length), almost completely divided medially, bordered posteriorly by 6 postmentals (outer pair largest); 8 infralabials to level below center of eye; sublabials undifferentiated; smooth granular scales present on chin and throat; dewlap extending well onto chest, from level below mideye to 3.5 mm beyond level of axilla; dorsum of body with smooth conical nonimbricate scales, 2 medial rows of slightly enlarged scales, largest dorsal scales about 0.35 x 0.29 mm (length x width); about 51 medial dorsal scales in one head length; about 75 medial dorsal scales between axilla and groin; lateral scales smooth, granular and homogeneous, average size 0.24 mm in diameter; ventrals at midbody smooth, bulging, nonimbricate, about 0.34 x 0.31 mm (length x width); about 52 ventral scales in one head length; about 77 ventral scales between axilla and groin; 154 scales around midbody; ventral caudal scales strongly keeled, forming longitudinal ridges; caudal middorsal scales enlarged, forming a serrated crest; lateral caudal scales mostly smooth, without whorls of enlarged scales, although an indistinct division in segments is discernible; precloacal scales pointed and granular; postcloacal scales not enlarged; no tube-like axillary pocket present; scales on dorsal surface of forelimb smooth, conical, subimbricate, about 0.25 x 0.24 mm (length x width); digital pads dilated, dilated pad about three times width of non-dilated scales on distal phalanx; distal phalanx narrower than and raised from dilated pad; 33 (right)��� 34 (left) lamellae under phalanges II���IV of fourth toe; seven scales under distal phalanx of fourth toe. The completely everted left hemipenis is a large bilobate organ; sulcus spermaticus bordered by well developed sulcal lips and bifurcates at base of apex, branches continue to tip of lobes; a small asulcate processus present; apex covered with fine calyces, truncus with transverse folds. The extended dewlap has four horizontal gorgetal-sternal rows with 8���10 large scales per row; about 75 scales in marginal series, modal number of anterior marginal pairs 4; about 35 apicogorgetals; gorgetals widely spaced, no scales between gorgetal rows; about 20 apicosternals. In life, dorsal and lateral surfaces Dark Drab (119 B), with diffuse spots and blotches of Vandyke Brown (221) and circular dirty white blotches with a suggestion of Pearl Gray (81); a dirty white middorsal stripe with a suggestion of Pearl Gray (81) interrupted by Sepia (119) blotches; a diffuse dirty white lateral stripe with a suggestion of Pearl Gray (81) extending from shoulder to groin; tail Beige (219 D), except for its base, with broad Brussels Brown (121 B) transverse bands and fine Sepia (119) mottling; a diffuse Vandyke Brown (221) pre- and postorbital stripe extending about 0.5 cm each anteriorly and posteriorly of orbit; ventral surfaces of body, limbs, and base of tail dirty white with a suggestion of Pearl Gray (81), with reticulate Dark Drab (119 B) mottling; chin and gular region Flaxflower Blue (170 C) with a suggestion of Light Sky Blue (168 D), with reticulate Hair Brown (119 A) mottling; iris Cinnamon (123 A); throat Blackish Neutral Gray (82); tongue Cream Color (54); dewlap Poppy Red (108 A) with circular Crimson (108) blotches between gorgetal rows; dewlap scales dirty white with a suggestion of Lavender Blue (170 D). Color in preservative (70 % ethanol) similar to color in life except for chin and gular region is dirty white. Variation. The paratypes agree well with the holotype in general appearance, morphometrics, and scalation (see Table 1). The coloration in life of SMF 89508 was recorded as follows: Dorsal and lateral surfaces of body, limbs and anterior half of tail Smoke Gray (45) suffused with Cinnamon-Drab (219 C), sparse Sepia (219) mottling and scattered flecks of dirty white with a suggestion of Pearl Gray (81); posterior half of tail Drab-Gray (119 D) with broad Raw Sienna (136) transverse bands; ventral surfaces of body, limbs, and anterior half of tail dirty white with a suggestion of Opaline Green (162 D); dorsal and lateral surfaces of head Drab-Gray (119 D) with Sepia (219) mottling on supraorbital semicircles, snout, and parietal region; ventral surface of head Light Sky Blue (168 D) with Hair Brown (119 A) spots on anterior half; dewlap Spectrum Red (11) with Carmine (8) blotches and dirty white scales. Natural history notes. In the area of Hacienda Bar�� (near Dominical, Costa Rica) Anolis charlesmyersi appears to be fairly common, especially along the beach and flat areas where there are many Terminalia catappa trees (Almendro de playa) (R. Mason pers. communication 23 January 2009). This statement is also supported by the relative large number of specimens collected from this site (see Appendix I). However, I failed to detect this species during a two days stay at Hacienda Bar�� during the dry season in March 2009. Whether my lack of spotting this species indicates some kind of seasonality remains to be investigated. Barbour (1934: 145) stated that Anolis pentaprion was ���not uncommon in the Canal Zone during the rainy season, excessively rare during the dry months.��� At the type locality near Los Algarrobos, Chiriqu��, Panama, a similar seasonality seems to occur with several observations during a single search in the rainy season and none but the holotype during numerous searches at the peak of the dry season (Lotzkat pers. comm. Oct. 2009). The holotype was collected at 22: 40 hours sleeping on a vertical twig of a small tree, about 2.5 m above the ground in relatively open secondary vegetation. Geographic distribution. As currently understood, Anolis charlesmyersi is distributed along the lowlands of the Pacific versant in Costa Rica and western Panama (Fig. 3). Etymology. The specific name is a patronym for my colleague Charles W. Myers in recognition of his important contributions to the herpetology of Lower Central America in general and to the understanding of the taxonomy and sytematics of the anoles of the pentaprion group in particular., Published as part of K��hler, Gunther, 2010, A revision of the Central American species related to Anolis pentaprion with the resurrection of A. beckeri and the description of a new species (Squamata: Polychrotidae), pp. 1-18 in Zootaxa 2354 on pages 6-9, DOI: 10.5281/zenodo.193523, {"references":["Wettstein, O. (1934) Ergebnisse der osterreichischen biologischen Costa Rica-Expedition 1930. Die Amphibien und Reptilien. Aus den Sitzungsberichten der Akademie der Wissenschaften in Wien Mathematischnaturwissenschaftliche Klasse, Abteilung I, 143, 1 - 39.","Taylor, E. H. (1956) A review of the lizards of Costa Rica. - University of Kansas Science Bulletin, 38, 1 - 322.","Williams, K. L. & Smith, H. M. (1968) Noteworthy lizards of the genus Anolis from Costa Rica. Caribbean Journal of Science, 6, 163 - 166. (\" 1966 \").","Myers, C. W. (1971) Central American lizards related to Anolis pentaprion: Two new species from the Cordillera de Talamanca. American Museum Novitates, 2471, 1 - 40.","Etheridge, R. (1959) The Relationships of the Anoles (Reptilia: Sauria: Iguanidae). An Interpretation Based on Skeletal Morphology. Ph. D. Dissertation, University of Michigan, Ann Arbor, Michigan, U. S. A.","Poe, S. (2004) Phylogeny of anoles. Herpetological Monographs, 18, 37 - 89.","Barbour, T. (1934) The anoles. II. The mainland species from Mexico southward. Bulletin of the Museum of Comparative Zoology, Harvard University, 77, 121 - 155."]}

Oedipina nica sp. nov. Nicaraguan Highland Worm Salamander Figure 1 Oedipina cyclocauda (in part). K��hler 1999, 2001. Holotype. MVZ 263774, an adult male, from El Gobiado, Reserva Natural Cerro Datanl��-El Diablo, 13 ��09���N, 85 �� 52 ���W, 1420 m above sea level (a.s.l.), Dept. Jinotega, Nicaragua, collected 30 August 2009 by S. M. Rovito, T. J. Papenfuss, J. Sunyer, and L. A. Obando (original field number SMR 842). Paratypes (5). UF 156445, from Reserva Natural Cerro Kilamb��, 13 �� 34 ���N, 85 �� 42 ���W, 1625 m a.s.l., Dept. Jinotega, Nicaragua; UF 156446 ��� 47, from Reserva Natural Cerro Kilamb��, 13 �� 35 ���N, 85 �� 43 ���W, 1660 m a.s.l., Dept. Jinotega, Nicaragua; UF 156453 ��� 54, from Reserva Natural Macizos de Pe��as Blancas, 13 �� 17 ���N, 85 �� 43 ���W, 1515 m a.s.l., Dept. Jinotega, Nicaragua. Referred specimens (12). SMF 78736, from Camp El Hielo, Reserva Natural Cerro Kilamb��, 1490 m a.s.l., Dept. Jinotega, Nicaragua; SMF 78737, 78739��� 40, from Camp 2, Reserva Natural Cerro Kilamb��, 13 �� 35.25 'N, 85 �� 41.50 'W, 1360 m a.s.l., Dept. Jinotega, Nicaragua; UF 156443 ��� 44, from Reserva Natural Cerro Kilamb��, 13 �� 34 ���N, 85 �� 42 ���W, 1625 m a.s.l., Dept. Jinotega, Nicaragua; UF 156448 ��� 50, from Reserva Natural Cerro Kilamb��, 13 �� 35 ���N, 85 �� 43 ���W, 1660 m a.s.l., Dept. Jinotega, Nicaragua; UF 156451 ��� 52, two tails dropped by escaped adults from Reserva Natural Cerro Kilamb��, 13 �� 34 ���N, 85 �� 42 ���W, 1600 m a.s.l., Dept. Jinotega, Nicaragua; UF 156455 from Reserva Natural Macizos de Pe��as Blancas, 13 �� 17 ���N, 85 �� 43 ���W, 1515 m a.s.l., Dept. Jinotega, Nicaragua. Diagnosis. A slender, small-sized (largest specimen 48.5 mm SL) species assigned to the genus Oedipina based on the presence of a high number (19���20) of costal grooves between the short limbs, very narrow manus and pes, long tail (tail length ca 2 x SL), overall morphological similarity to other Oedipina species, and molecular phylogenetic evidence. This species is a member of the subgenus Oeditriton (based on molecular data; Fig. 2) and is distinguished from the only other members of that clade (O. kasios and O. quadra) by its more slender habitus and fewer vomerine teeth. Further distinguished from other Nicaraguan members of the genus Oedipina as follows: from O. collaris by being much smaller and more slender, having short limbs and narrow manus and pes, and having a short, rather bluntly tipped snout; from O. cyclocauda by being smaller and more slender without nearly pad-like feet; from O. pseudouniformis by being more slender with shorter, less robust limbs and narrower manus and pes, and by having almost uniformly dark brownishblack coloration with no distinct light pigmentation on the upper parts of the proximal segment of the limbs. Description of the holotype. The male holotype is 39.2 mm SL and is judged to be an adult by the presence of small papillae in the anterior margins of the vent and a small mental gland behind the anterior tip of the mandible. The small head is rounded in dorsal profile, and somewhat flattened in lateral profile with a relatively blunt snout and distinct nasolabial protuberances. Snout to posterior angle of vent 9.8 times head width and 6.5 times head length. Nostrils are small but conspicuous under magnification. The small eyes are inconspicuous and do not extend beyond the lateral margins of the head. The suborbital groove does not intercept the lip line. The skin of the dorsal surface of the head behind the eyes is depressed in a pattern that suggests the presence of a frontoparietal fenestra. There is a single slightly enlarged premaxillary tooth that is located distinctly anterior to the maxillary teeth. Maxillary teeth total 41. The small vomerine teeth are in a short, fairly straight series and total 11. There are 20 costal grooves between the small limbs with a limb interval of 13 intercostal folds. Hands and feet are tiny, narrow, and elongated. Digit I is fused with digit II and digit IV is fused with digit III on the forelimbs, with only a part of a phalangeal segment of digit III free between digits II���III on the forelimbs. Digit I is fused with digit II and digit V is fused with digit IV on the hind limbs; both outer digits are extremely short on the hind limbs. The triangular tip of digit III of the hind limb is distinct from the tiny rounded tips of digits II and IV, but less than a full phalangeal segment of that digit is free of the others. Subdigital pads are nearly imperceptible. Digits on forelimbs in order of decreasing length are III���II���IV���I; digits on hind limbs are III���II���IV���(V���I). The tail is round in cross section and tapers gradually to a narrow tip; the tail is almost twice SL. Measurements (in millimeters) of the holotype. Head width 4.0; snout to gular fold (head length) 6.0; head depth at posterior angle of jaw 1.8; eyelid width 1.1; eyelid length 1.5; eye to nostril 0.6; anterior rim of eye to snout 1.2; horizontal orbital diameter 1.0; interorbital distance 1.7; distance separating eyelids 0.7; nostril diameter 0.2; snout projecting beyond mandible 0.4; distance from eye to distal end of postorbital groove 1.9; snout to posterior angle of vent (SL) 39.2; snout to anterior angle of vent 37.0; snout to forelimb 9.4; axilla to groin 26.0; limb interval 13; shoulder width 2.7; tail length 77.0; tail width at base 2.4; tail depth at base 2.4; forelimb length (to tip of longest digit) 3.7; hind limb length (to tip of longest digit) 4.5; forelimb foot width 0.6; hind limb foot width 0.9; free length of longest digit 0.2. Coloration of the holotype in alcohol. Dorsal and lateral surfaces of the head, body, and tail are black. Costal grooves on the body and tail are the same color as the body or slightly lighter dark gray, but at the deepest part of the groove pigment is lacking so the grooves are prominent. Ventral surfaces of the head, body, and tail are slightly paler black than those dorsal surfaces. Dorsal and ventral surfaces of the limbs are the same as for the body. Tiny white speckles are most visible behind the eye and over the shoulder, then in an irregular, narrow line along the dorsolateral part of the body to the hind limbs, but they are few in number and obscure. White spots are a little larger in the gular and chest areas ventrally. The nasolabial protuberances are unpigmented and contrast with the surrounding tissue. Variation. Variation based on study of the holotype and three male (UF 156445, 156447, 156453), one female (UF 156446), and one juvenile (UF 156454) paratypes: the four males, judged to be mature based on having inconspicuous mental glands and anterior vent margins bearing fine pinnae, range from 38.5���44.5 SL; the female is 48.5 SL. Tails are present in the holotype, three male paratypes, and the juvenile and all taper gradually to slender tips. SL is from 0.86 (adult) to 0.52 (holotype) times tail length. SL in males is 9.8���11.8 times head width and 9.9 in the single female. Limbs are relatively short; number of costal folds between adpressed limbs is 13���14 in males and 13 in the female. There are 19���20 costal grooves between the limbs in the males and 20 in the female. Feet are narrow; 0.8���1.1. The digits are joined to their neighbors, with only the tips of the longest toes free. Males have 1���2 premaxillary teeth, the female has 4. Maxillary teeth range from 41���47 in males; 48 in the female. Males have 9���16 vomerine teeth; the female 18. If one adds the referred specimens, the size range of the adult males increases to 35.3���48.5 SL, and that of females from 37.5���48.5 SL. These specimens are consistent in measurements, tooth and costal groove counts, and coloration with the type series. SMF 78738, from Reserva Natural Cerro Kilamb�� (44.0 SL), fits within the range of the new species in all measurements (see discussion). Variation of coloration based on color photographs taken in life during daytime. Coloration is in general uniformly dark brownish-black with no distinguishing features. The upper side of the proximal segment of the limbs is the same color. The costal grooves are very evident because of the lack of pigment at their centers. Details for particular individuals follow. Adult male paratype (UF 156447; Fig. 1 b): Dorsum of body and tail blackish brown becoming a faded lighter brown on the dorsal and lateral surfaces of head extending to the chin. A lighter brown speckling is faintly noticeable dorsally from the head becoming imperceptible towards the base of the tail. Ventral surfaces of the body are blackish brown, similar in coloration to the tail. Limbs faded brown similar to color of head. Skin creases such as costal grooves, gular fold, and those around the paratoid region lack pigmentation and appear light grey. Iris dark rust brown. Adult male (UF 156450): Dorsum of body and tail blackish brown becoming faded lighter brown on the anterior and lateral surfaces of head. Gold speckling mixed with some white is located on the dorsal surface of the head and body. Speckling is most prominent on the top of the head becoming sparser posteriad, not extending beyond base of tail. Dorsolaterally above the anterior limbs a slight concentration of tiny white speckles forms a short, irregular line. Below the extent of the speckling ventrolaterally coloration is same as that of tail. Limb coloration is same as that of anterior and lateral surface of head. Iris dark rust brown. Juvenile paratype (UF 156454; Fig. 1 d): Dorsal and lateral surfaces of body, head, tail, and limbs are a uniform jet black. There is a faint overlay of tiny dirty white flecks along the dorsum of head, body, and tail giving a somewhat ashy appearance. This flecking is most concentrated dorsolaterally giving the impression of an irregular line extending from behind the jaw to the hind limb. Flecking is also present around the mouth and chest area. Larger white speckles are mixed sparsely and intermittently along the head, body, tail, and limbs. Habitat and distribution. This species is known to occur from 1360���1660 m elevation at three isolated localities in the Lower Montane Moist Forest formation (Holdridge 1967) in north-central Nicaragua (Fig. 3). The holotype was collected in a small patch of secondary cloud forest (Fig. 4 a���b) mostly surrounded by cattle pastures and agricultural land. It was found while raking during the daytime inside a moderately large rotten log. The paratypes and UF referred specimens from Reserva Natural Cerro Kilamb�� were collected in undisturbed, primary cloud forest (Fig. 4 c) during both diurnal and nocturnal surveys. All were found underneath logs, rocks, and other debris in areas along the mountain ridge where the substrate comprised a spongy mass of plant roots and mosses. The SMF referred specimens from Reserva Natural Cerro Kilamb�� were also collected in undisturbed, primary cloud forest during diurnal surveys. Most of them were found inside dead basal ramifications of giant ferns. The paratypes from Reserva Natural Macizos de Pe��as Blancas were collected in undisturbed, primary cloud forest (Fig. 4 d). One was uncovered during the daytime beneath a fallen log, and two others were found active on the forest floor around the campsite at 2130 h. Those found active within the camp were likely dislodged or disturbed by raking and uprooting logs that occurred during set-up of the campsite earlier the same day. Phylogenetic relationships. Phylogenetic analysis of sequence data from two mitochondrial genes (16 S and cyt b) confirms the monophyly of this taxon (ML bootstrap support = 99, Bayesian posterior probability = 1.0) and its position as sister species to Oedipina kasios in the Oeditriton clade (Fig. 2). A sister-species relationship between O. kasio s and O. nica is strongly supported (ML bootstrap support = 93, Bayesian posterior probability = 0.99) as is the monophyly of an Oeditriton clade containing O. kasio s, O. nica, and O. quadra (ML bootstrap support = 96, Bayesian posterior probability = 1.0). Oeditriton is the sister clade to the remaining Oedipina, which are divided into the subgenera Oedipina and Oedopinola (Fig. 2), which supports previous phylogenetic hypotheses for Oedipina (Garc��a-Par��s & Wake 2000; McCranie et al. 2008). All wellsupported phylogenetic relationships are congruent between the ML and Bayesian results. Etymology. The specific name nica is a short name for Nicaraguan (Nicarag��ense in Spanish) and is a colloquial word used in Central America to denote Nicaraguan people. The name alludes to the country of origin of the type specimens. Conservation status. Using the criteria established by IUCN for evaluating threatened species, Oedipina nica should be classified as Endangered (EN B 2 ab[iii]), due to its limited distribution (known only from three isolated highland forest areas with a total extent of less than 500 km 2) and the continued loss of habitat at these localities., Published as part of Sunyer, Javier, Wake, David B., Townsend, Josiah H., Travers, Scott L., Rovito, Sean M., Papenfuss, Theodore J., Obando, Lenin A. & K��hler, Gunther, 2010, A new species of worm salamander (Caudata: Plethodontidae: Oedipina) in the subgenus Oeditriton from the highlands of northern Nicaragua, pp. 29-39 in Zootaxa 2613 on pages 32-35, DOI: 10.5281/zenodo.197858, {"references":["Kohler, G. (1999) The amphibians and reptiles of Nicaragua - a distributional checklist with keys. Courier Forschungsinstut Senckenberg, 213, 1 - 212.","Kohler, G. (2001) Anfibios y Reptiles de Nicaragua. Herpeton. Offenbach, Germany.","Holdridge, L. R. (1967) Life Zone Ecology. Tropical Science Center. San Jose, Costa Rica.","McCranie, J. R., Vieites, D. R. & Wake, D. B. (2008) Description of a new divergent lineage and three new species of Honduran salamanders of the genus Oedipina (Caudata, Plethodontidae). Zootaxa, 1930, 1 - 17."]}

Anolis pentaprion COPE 1862 Anolis pentaprion: Dunn (1930), Schmidt (1933), Barbour (1934), Gaige et al. (1937), Taylor (1956; in part.), Etheridge (1959; in part), Myers (1971), Poe (2004; in part.). Anolis panamensis BOULENGER, 1890: 81; two syntypes BMNH 1946.8.13.7��� 8 from ��� Panama.��� Norops pentaprion: K��hler (2001; in part.). Neotype designation. The holotype of Anolis pentaprion, supposedly designated to be deposited at USNM, is lost (Barbour 1934, Myers 1971). K.A. Tighe (pers. comm. 2 September 2009) wrote that ���the holotype [of Anolis pentaprion] was apparently never returned to the USNM after Cope described it. There is an old lost type card in Cochran's handwriting that has the information from the original description, but no catalogue number.��� To avoid future uncertainties I herewith designate SMF 83608 as the neotype of Anolis pentaprion Cope. SMF 83608 was collected on 12 August 2009 by Johannes K��hler and Arne Schulze at San Rafael, ca. 15 km S Los Chiles, 10.73719 ��N, 84.49378 ��W, 60 m elevation, Alajuela Province, Costa Rica. Original field number JJK 182. SMF 83608 is an adult male as indicated by its partially everted hemipenes; SVL 61.0 mm; tail length 79.0 mm (tail complete); head length 15.4 mm; head width 9.5 mm; snout length 7.4 mm; ear length 0.8 mm; ear height 1.5 mm; interparietal length 1.3 mm; interparietal width 1.1 mm; axilla���groin���distance 26.0 mm; shank length 12.5 mm. Scalation characteristics of SMF 83608 are given in parentheses in the description below. Diagnosis. A medium-sized species (SVL in largest specimen examined 77.0 mm in males, 74 mm in females) of the genus Anolis (sensu Poe, 2004) that differs from all Central American anoles except A. beckeri, A. charlesmyersi, A. cristifer, A. fungosus, A. salvini, A. utilensis by (1) having extremely short hind legs (fourth toe of adpressed hindlimb usually reaching to level of tympanum; ratio shank length / head length 0.49���0.84); (2) smooth, juxtaposed ventral scales; (3) a relatively short tail (ratio tail length / SVL 1.0��� 1.5). Anolis pentaprion differs from A. beckeri by having a pronounced serrated caudal crest (versus lacking a pronounced serrated caudal crest) and by having gorgetal rows with 17���25 narrowly spaced scales (versus gorgetal rows with 4���10 widely spaced scales). Anolis pentaprion differs from A. charlesmyersi by having a pink dewlap in life with purple pigment between rows of gorgetal scales and regular rows with 17���25 small narrowly spaced gorgetal scales (versus a brick red dewlap with few large widely spaced gorgetal scales scattered across the dewlap or in loosely arranged rows with 4���9 widely spaced scales). Anolis pentaprion differs from A. cristifer by having a double row of slightly enlarged middorsal scales, not forming a serrated crest (versus a serrated middorsal crest, at least on posterior portion of dorsum, present in A. cristifer). Anolis pentaprion differs from A. fungosus by lacking bony parietal protuberances (versus having a pair of small bony parietal protuberances posterior and lateral to interparietal plate). Anolis pentaprion differs from A. salvini by having smooth, obliquely conical ventral scales (versus keeled), lacking a dark interorbital bar (versus present), and lacking enlarged postcloacal scales (versus males with a pair of greatly enlarged postcloacal scales). Anolis pentaprion differs from A. utilensis by having the proximal subdigital scales of toes differentiated as slightly broadened lamellae (versus those scales granular) and by having a pronounced serrated caudal crest (versus lacking a pronounced serrated caudal crest). Description. SVL 52.0���77.0 (62.6 �� 5.7) mm in males, 46.0���64.0 (56.4 �� 5.4) mm in females; for other morphometric data see Table 1; dorsal head scales in internasal, prefrontal, and frontal areas rugose; scales in parietal area rugose to smooth; shallow frontal and parietal depressions present; 5���8 postrostrals (7); anterior nasal divided, lower scale contacting rostral and first supralabial; 5���9 internasals (6); canthal ridge sharply defined; scales comprising supraorbital semicircles ridged, especially anterior ones, largest scale in semicircles larger than largest supraocular scale; supraorbital semicircles well defined, broadly in contact with each other or separated by one scale at narrowest point (in contact); 1���4 scales separating supraorbital semicircles and interparietal at narrowest point (2); interparietal well defined, irregular in outline, longer than wide, larger than size of ear opening; about 6���16 enlarged, smooth to faintly keeled supraocular scales on each side (9), in 2���3 rows; enlarged supraoculars completely separated from supraorbital semicircles; three short superciliaries, posterior one shortest; usually 3 enlarged canthals; 5���9 scales between second canthals (8); 6���15 scales present between posterior canthals (6); loreal region slightly concave, 13���29 mostly smooth or rugose (some keeled) loreal scales (16) in a maximum of 3���6 horizontal rows (3); 7���10 supralabials (9) and 7���10 infralabials (9) to level below center of eye; suboculars smooth to rugose, in broad contact with supralabials; ear opening vertically oval; scales anterior to ear opening slightly larger than those posterior to ear opening; usually 6, occasionally 5 postmentals (6), outer pair largest; gular scales not keeled; male dewlap moderately large, extending to beyond level of axillae onto chest; male dewlap with 5���7 horizontal gorgetalsternal scale rows, about 16���18 scales per row; female dewlap well developed, extending to level of axillae; female dewlap with 4���7 horizontal gorgetal-sternal scale rows, with about 9���15 scales per row; about 2 middorsal scale rows slightly enlarged, mostly smooth, dorsal scales lateral to middorsal series grading into granular lateral scales; no enlarged scales among granular laterals; lateral scales granular, slightly smaller than largest dorsal scales; about 73���108 dorsal scales along vertebral midline between levels of axillae and groin in males (73), about 88���110 in females; about 56���80 dorsal scales along vertebral midline contained in one head length in males (52), about 45���70 in females; ventral scales on midsection slightly larger than largest dorsal scales; ventral body scales smooth, obliquely conical, juxtaposed; about 67���86 ventral scales along midventral line between levels of axilla and groin in males (84), about 65���93 in females; about 34���62 ventral scales contained in one head length in males (44), about 36���52 in females; about 134���190 scales around midbody in males (154), about 136���162 in females; tubelike axillary pocket absent; precloacal scales not keeled; enlarged postcloacal scales absent; tail oval at about midlength; basal subcaudal scales smooth to faintly keeled; lateral caudal scales keeled, homogeneous, although an indistinct division in segments is discernible; dorsal medial caudal scale row enlarged, keeled, not forming a crest; 2 median subcaudal scale rows distinctly enlarged, keeled; most scales on lateral surface of antebrachium smooth; 29���33 subdigital lamellae on Phalanges II���IV of Toe IV of hind limbs (29); 6���9 subdigital scales on Phalanx I of Toe IV of hind limbs (6); longest toe of adpressed hind limb usually reaching to shoulder region. The coloration in life of an adult male (SMF 80961) from Bartola, Department R��o San Juan, Nicaragua, was recorded as follows: Dorsal and lateral surfaces of body, limbs and anterior half of tail Clay Color (123 B) with Raw Umber (223) and Beige (219 D) blotches; dorsal surface of head Cinnamon Drab (119 C); dewlap Vinaceous (4) with rows of Magenta (2) blotches and dirty white scales. The coloration of the dewlap in life of another adult male (SMF 82100) from the same locality was recorded as Pink (7) with rows of Magenta (2) blotches and dirty white scales. Geographic distribution: Along the Caribbean versant of Lower Central America from southern Nicaragua to northwestern Colombia; also on Pacific versant in central and eastern Panama., Published as part of K��hler, Gunther, 2010, A revision of the Central American species related to Anolis pentaprion with the resurrection of A. beckeri and the description of a new species (Squamata: Polychrotidae), pp. 1-18 in Zootaxa 2354 on pages 12-13, DOI: 10.5281/zenodo.193523, {"references":["Cope, E. D. (1862) Contributions to Neotropical saurology. Proceedings of the Academy of Natural Sciences of Philadelphia, 14, 176 - 188.","Dunn, E. R. (1930) Notes on Central American Anolis. Proceedings of the New England Zoological Club, 12, 15 - 24.","Schmidt, K. P. (1933) Amphibians and reptiles collected by the Smithsonian Biological Survey of the Panama Canal Zone. Smithsonian Miscellaneous Collections, 89, 1 - 20.","Barbour, T. (1934) The anoles. II. The mainland species from Mexico southward. Bulletin of the Museum of Comparative Zoology, Harvard University, 77, 121 - 155.","Gaige, H. T., Hartweg, N. & Stuart, L. C. (1937) Notes on a collection of amphibians and reptiles from eastern Nicaragua. Occasional Papers Museum of Zoology University of Michigan, 357, 1 - 18.","Taylor, E. H. (1956) A review of the lizards of Costa Rica. - University of Kansas Science Bulletin, 38, 1 - 322.","Etheridge, R. (1959) The Relationships of the Anoles (Reptilia: Sauria: Iguanidae). An Interpretation Based on Skeletal Morphology. Ph. D. Dissertation, University of Michigan, Ann Arbor, Michigan, U. S. A.","Myers, C. W. (1971) Central American lizards related to Anolis pentaprion: Two new species from the Cordillera de Talamanca. American Museum Novitates, 2471, 1 - 40.","Poe, S. (2004) Phylogeny of anoles. Herpetological Monographs, 18, 37 - 89.","Boulenger, G. A. (1890) First report on additions to the lizard collection in the British Museum (Natural History). Proceedings of the Zoological Society London, 1890, 77 - 86.","Kohler, G. (2001) Anfibios y Reptiles de Nicaragua. Herpeton, Offenbach, Germany, 208 pp."]}

Anolis osa sp. nov. Figs. 9���11 Anolis polylepis: Poe (2004, in part.). Holotype. SMF 80645, an adult male from about 6.3 km WSW Rinc��n de Osa, 8 �� 40 ��� 36.7 ������N, 83 �� 32 ��� 7.9 ������W, about 150 m elevation, Puntarenas Province, Costa Rica, collected 22 February 2008 by Gunther K��hler and Johannes K��hler. Field tag number GK 2092. Paratypes. SMF 89227 ��� 32, same collecting data as holotype. SMF 89215 ��� 21, UCR 20733 ���35, 2 km W Rinc��n de Osa, about 10 m elevation, Puntarenas Province, Costa Rica, collected 22 February 2008 by Gunther K��hler and Johannes K��hler. SMF 89260 ���61, 89427��� 29, Parque Nacional Corcovado, Estaci��n Sirena, 8.48 ��N, 83.58 ��W, Puntarenas Province, Costa Rica, collected 10���11 August 2005 by Matthias Dehling. SMF 89227 ��� 29, SMF 89260 ���61, 89427��� 29, UCR 20733 ��� 35 are adult males with everted hemipenes. SMF 89230 ��� 32 are adult females. Referred Specimen: See Appendix I. Diagnosis. A medium-sized species (SVL in largest specimen 57 mm) of the genus Anolis (sensu Poe 2004) that differs from all other Lower Central American beta anoles (sensu Etheridge 1967) except Anolis apletophallus, A. cryptolimifrons, and A. limifrons in that it is long-legged (longest toe of adpressed hind leg reaches to at least center of eye), has smooth ventral scales, and a slender habitus, often delicate. Anolis osa differs from the species in the cluster listed above, except A. polylepis, by having two elongate, overlapping superciliaries (one such scale in A. apletophallus, A. cryptolimifrons, and A. limifrons), usually two anterior prenasal scales (versus one elongate prenasal scale) and dewlap coloration (usually orange with a darker orange basal portion in A. osa versus dirty white with a basal orange-yellow blotch in A. cryptolimifrons and A. limifrons, almost uniformly orange-yellow in A. apletophallus). Anolis osa differs from A. polylepis by having a unilobed hemipenis (bilobed in A. polylepis). Other anole species from Lower Central America that are somewhat similar in appearance to both A. osa and A. polylepis are A. tropidogaster and A. cupreus, the latter of which occurs syntopically at some localities with A. polylepis. However, both A. tropidogaster and A. cupreus have strongly keeled ventral scales (smooth in A. osa and A. polylepis). Also, the majority of males in A. tropidogaster have a pair of greatly enlarged postcloacal scales (these scales usually not enlarged in A. osa and A. polylepis). In the field, most individuals of A. osa and A. polylepis have a pale longitudinal lateral stripe (absent in Costa Rican populations of A. cupreus). For variation in selected morphometric and scalation characters of A. osa see Table 3. Description of the holotype. Adult male as indicated by everted hemipenes and presence of large dewlap; SVL 50.5 mm; tail length 106 mm, tail complete; tail compressed in cross section, tail height 2.0 mm, tail width 1.4 mm; axilla to groin distance 20.8 mm; head length 13.3 mm, head length/SVL ratio 0.26; snout length 6.6 mm; head width 8.2 mm; longest toe of adpressed hind limb reaching to a point between eye and snout; shank length 14.6 mm, shank length/head length ratio 1.1; longest finger of extended forelimb reaching tip of snout; longest finger of adpressed forelimb reaches to anterior insertion of hind limbs. Scales on snout keeled (Fig. 10); 9 postrostrals; 10 scales between nasals; scales in distinct prefrontal depression slightly wrinkled; supraorbital semicircles well developed, separated by a minimum of 2 scales; supraorbital disc composed of 6 distinctly enlarged keeled scales; two elongated, strongly overlapping superciliaries, the anterior one much larger than the posterior scale; about 3���4 rows of small keeled scales extending between enlarged supraorbitals and superciliaries; interparietal scale well developed, 1.8 x 1.0 mm (length x width), surrounded by scales of moderate size; 2 scales present between interparietal and supraorbital semicircles; canthal ridge distinct, composed of 3 large (posterior two largest) and 6 small anterior canthal scales; 11 scales present between second canthals; 12 scales present between posterior canthals; 70 (right)��� 63 (left) mostly keeled loreal scales in a maximum of 8 (right)��� 9 (left) horizontal rows, with the scales of lower rows mostly keeled, and those of upper rows mostly tuberculated; subocular scales keeled, narrowly in contact with supralabials; 8 supralabials to level below center of eye; ear opening 0.7 x 1.4 mm (length x height); mental distinctly wider than long, almost completely divided medially, bordered posteriorly by 8 postmentals (outer larger than inner); 7 infralabials to level below center of eye; sublabials undifferentiated; slightly keeled granular scales present on chin and throat; dewlap extending well onto chest, anterior insertion at level of posterior canthals, posterior insertion about 6.0 mm beyond level of axilla; dorsum of body with keeled granular scales, 2 medial rows of slightly enlarged scales, largest dorsal scales about 0.28 x 0.22 mm (length x width); about 54 medial dorsal scales in one head length; about 85 medial dorsal scales between axilla and groin; lateral scales keeled, granular and homogeneous, average size 0.15 mm in diameter; ventrals at midbody smooth, bulging, subimbricate, about 0.23 x 0.23 mm (length x width); about 54 ventral scales in one head length; about 94 ventral scales between axilla and groin; about 176 scales around midbody; caudal scales strongly keeled; caudal middorsal scales slightly enlarged, without whorls of enlarged scales; precloacal scales pointed and granular; postcloacal scales not enlarged; no tube-like axillary pocket present; scales on dorsal surface of forelimb slightly keeled, imbricate, about 0.25 x 0.27 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; 25 (right)��� 25 (left) lamellae under phalanges II���IV of fourth toe; 9 scales under distal phalanx of fourth toe. The completely everted hemipenis is a small unilobed organ; sulcus spermaticus bordered by well developed sulcal lips and opens at base of apex into a concave field without ornamentation; a small asulcate processus present; asulcal side of apex covered with fine calyces, truncus with transverse folds. The extended dewlap in life has 6���7 horizontal gorgetal-sternal rows with 26���42 scales per row, rows somewhat irregular, some of them with pairs or triplets of scales, apical portion of dewlap between marginal pairs and last gorgetal-sternal row free of scales; modal number of anterior marginal pairs 4���5. Coloration in life was recorded as follows: Dorsal ground color Dark Drab (119 B) with a Pale Horn Color (92) broken longitudinal lateral stripe, edged diffusely by Mars Brown (223 A) pigment; a Sepie (119) shoulder spot, edged posteriorly by a Yellow Ocher (123 C) blotch; dorsum of head Dark Drab (119 B) with a Hair Brown (119 A) interorbital bar; dorsal surfaces of limbs Army Brown (219 B) with indistinct Tawny Olive (223 D) markings; tail Brick Red (132 A) with indistinct Tawny Olive (223 D) banding; tip of tail Tawny Olive (223 D); ventral surfaces Glaucus (79); dewlap Orange Yellow (18), basal portion suffused with Raw Umber (23), gorgetals dirty white; iris Mars Brown (219 A). Coloration after one year preservation in 70 % ethanol agrees well with this description. Variation. The paratypes agree well with the holotype in general appearance, morphometrics and scalation (see Table 1). We documented variation in relative hind leg length in this species. Of 66 specimens examined, the longest toe of the adpressed hind leg reached to mid-eye in 5 individuals (7.6 %), to anterior margin of eye in 33 individuals (50.0%), and to a point between eye and snout in 28 individuals (42.4 %). For variation in nasal region see Table 2 and Figure 1. For variation in dorsal pattern see Table 2 and Figure 2. Etymology. The name osa is used as a noun in apposition and in reference to the Peninsula de Osa where the type series of the species was collected and where it is probably restricted. A. polylepis A. osa 3 47 3 22 �� 24 �� 19 SVL 3 42.2���59.4 (51.3 �� 3.76) 49.9���55.4 (50.2 �� 2.28) �� 39.6���53.5 (47.3 �� 3.52) 41.9���50.7 (46.4 �� 2.60) Tail length 3 80.1 ���117.0 (102.1 �� 10.28) 84.9���107.2 (97.3 �� 6.88) �� 70.0��� 101.5 (86.3 �� 8.29) 73.1 ���93.0 (82.0 �� 4.59) HL 3 10.8���14.4 (12.8 �� 0.88) 11.3���13.9 (12.4 �� 0.62) �� 10.1���13.7 (11.9 �� 0.78) 10.4���12.8 (11.5 �� 0.60) HW 3 6.7���8.5 (7.7 �� 0.48) 7.0��� 8.2 (7.5 �� 0.35) �� 6.3���8.5 (7.3 �� 0.56) 6.7���7.9 (7.1 �� 0.36) Shank length 3 11.6 ���16.0 (13.9 �� 1.12) 12.2 ���15.0 (13.6 �� 0.66) �� 10.5 ���14.0 (12.4 �� 0.98) 11.0��� 13.2 (12.3 �� 0.65) Axilla���groin distance 3 16.3���26.1 (21.2 �� 2.16) 18.5���24.4 (21.3 �� 1.32) �� 15.5���24.1 (19.87 �� 2.14) 17.8���29.4 (20.7 �� 2.60) Natural history notes. Anolis osa is an extremely common anole that reaches high population densities. At most sites on the Peninsula de Osa it was the most abundant species of anole. During daytime, most individuals were observed perching head down on the lower parts (30���120 cm above the ground) of trunks of small trees and other plants such as heliconias (Heliconiaceae) and banana plants (Musaceae). Males of A. osa are very territorial and will show aggressive display behavior whenever another adult male is placed in reaching distance (less than about 150 cm). The species was studied in detail by Andrews (1971, 1983), and Savage (2002) summarized much of the information provided by her. Geographic distribution. As currently known, Anolis osa is restricted to the Peninsula de Osa, Costa Rica. It occurs both in protected areas (Corcovado National Park) and in disturbed areas such as at the type locality. The documented vertical range of the species is from sea level to about 550 m elevation., Published as part of K��hler, Gunther, Dehling, Matthias & K��hler, Johannes, 2010, Cryptic species and hybridization in the Anolis polylepis complex, with the description of a new species from the Osa Peninsula, Costa Rica (Squamata: Polychrotidae), pp. 23-38 in Zootaxa 2718 on pages 28-33, DOI: 10.5281/zenodo.199772, {"references":["Poe, S. (2004) Phylogeny of anoles. Herpetological Monographs, 18, 37 - 89.","Etheridge, R. (1967) Lizard caudal vertebrae. Copeia, 1967, 699 - 721.","Andrews, R. M. (1971) Structural habitat and time budget of a tropical Anolis lizard. Ecology, 52, 262 - 279.","Andrews, R. M. (1983) Norops polylepis (lagartija, anole, anolis lizard.). Pp. 409 - 410. In Janzen, D. H. (Ed.), Costa Rican Natural History. University of Chicago Press, Chicago, Illinois, USA.","Savage, J. M. (2002) The amphibians and Reptiles of Costa Rica. A Herpetofauna between two Continents, between two Seas. University of Chicago Press, Chicago, Illinois, USA."]}

Anolis beckeri Boulenger 1881 Anolis beckeri Boulenger, 1881: 921; two syntypes, IRSNB 2010 (1���2), from ���Yucat��n.��� Barbour (1934). Anolis pentaprion: Stuart (1937; in part), Stuart (1948; in part), Myers (1971; in part), Meyer and Wilson (1973; in part); Fitch and Seigel (1984; in part), Lee (1996), Poe (2004; in part.). Anolis pentaprion beckeri: Etheridge (1959), Stuart (1963). Norops pentaprion: K��hler (2001; in part), McCranie et al. (2006; in part). Diagnosis. A medium-sized species (SVL in largest specimen examined 60.6 mm in males, 55.4 mm in females) of the genus Anolis (sensu Poe, 2004) that differs from all Central American anoles except A. charlesmyersi, A. cristifer, A. fungosus, A. pentaprion, A. salvini, and A. utilensis by having (1) extremely short hind legs (fourth toe of adpressed hindlimb usually reaching to level of tympanum; ratio shank length / head length 0.49���0.84); (2) smooth, juxtaposed ventral scales; (3) a relatively short tail (ratio tail length / SVL 1.0��� 1.5). Anolis beckeri differs from A. charlesmyersi by having a pink dewlap in life with purple pigment between rows of gorgetal scales and regular rows of numerous small narrowly spaced gorgetal scales (versus a brick red dewlap with few large widely spaced gorgetal scales scattered across the dewlap). Anolis beckeri differs from A. cristifer by having a double row of slightly enlarged middorsal scales, not forming a serrated crest (versus a serrated middorsal crest, at least on posterior portion of dorsum, present in A. cristifer). Anolis beckeri differs from A. fungosus by lacking bony parietal protuberances (versus having a pair of small bony parietal protuberances posterior and lateral to interparietal plate). Anolis beckeri differs from A. pentaprion by lacking a pronounced serrated caudal crest (versus having a pronounced serrated caudal crest) and by having gorgetal rows with 4���10 widely spaced scales (versus gorgetal rows with 17���25 narrowly spaced scales). Anolis beckeri differs from A. salvini by having smooth, obliquely conical ventral scales (versus keeled), lacking a dark interorbital bar (versus present), and lacking enlarged postcloacal scales (versus males with a pair of greatly enlarged postcloacal scales). Anolis beckeri differs from A. utilensis by having the proximal subdigital scales of toes differentiated as slightly broadened lamellae (versus those scales granular). Description. SVL 46.5���60.6 (51.0��5.0) mm in males, 38.0��� 55.4 (58.3 �� 9.6) mm in females; for other morphometric data see Table 1; dorsal head scales in internasal, prefrontal, and frontal areas rugose; scales in parietal area rugose to smooth; shallow frontal and parietal depressions present; 5���8 postrostrals; anterior nasal divided, lower scale contacting rostral and first supralabial; 5���9 internasals; canthal ridge sharply defined; scales comprising supraorbital semicircles ridged, especially anterior ones, largest scale in semicircles larger than largest supraocular scale; supraorbital semicircles well defined; supraorbital semicircles broadly in contact with each other or separated by one scale at narrowest point; 1���3 scales separating supraorbital semicircles and interparietal at narrowest point; interparietal well defined, irregular in outline, longer than wide, larger than size of ear opening; about 4���11 enlarged, smooth to faintly keeled supraocular scales on each side, in 2���3 rows; enlarged supraoculars completely separated from supraorbital semicircles; three short superciliaries, posterior one shortest; usually 3 enlarged canthals; 6���14 scales between second canthals; 7���9 scales present between posterior canthals; loreal region slightly concave, 18���30 mostly smooth or rugose (some keeled) loreal scales in a maximum of 3���4 horizontal rows; 7���10 supralabials and 7��� 10 infralabials to level below center of eye; suboculars smooth to rugose, in broad contact with supralabials; ear opening vertically oval; scales anterior to ear opening slightly larger than those posterior to ear opening; 5���7 postmentals, outer pair largest; gular scales not keeled; male dewlap moderately large, extending to beyond level of axillae onto chest; male dewlap with 5���6 horizontal gorgetal-sternal scale rows, about 16���18 scales per row; female dewlap well developed, extending to level of axillae; female dewlap with 4���7 horizontal gorgetal-sternal scale rows, with about 9���15 scales per row; about 2 middorsal scale rows slightly enlarged, mostly smooth, dorsal scales lateral to middorsal series grading into granular lateral scales; no enlarged scales among granular laterals; lateral scales granular, slightly smaller than largest dorsal scales; about 95���104 dorsal scales along vertebral midline between levels of axillae and groin in males, about 84���123 in females; about 56���80 dorsal scales along vertebral midline contained in one head length in males, about 45���70 in females; ventral scales on midsection slightly larger than largest dorsal scales; ventral body scales smooth, obliquely conical, juxtaposed; about 67���86 ventral scales along midventral line between levels of axilla and groin in males, about 67���92 in females; about 34���60 ventral scales contained in one head length in males, about 34���60 in females; about 146���178 scales around midbody in males, about 130���178 in females; tubelike axillary pocket absent; precloacal scales not keeled; enlarged postcloacal scales absent; tail oval at about midlength; tail height/tail width 0.80���1.25; basal subcaudal scales smooth to faintly keeled; lateral caudal scales keeled, homogeneous, although an indistinct division in segments is discernible; dorsal medial caudal scale row enlarged, keeled, not forming a crest; 2 median subcaudal scale rows distinctly enlarged, keeled; most scales on lateral surface of antebrachium smooth; 18���32 subdigital lamellae on Phalanges II���IV of Toe IV of hind limbs; 7���10 subdigital scales on Phalanx I of Toe IV of hind limbs; longest toe of adpressed hind limb usually reaching to shoulder region. Geographic distribution: Along the Caribbean versant of Nuclear Central America from Tabasco, Mexico, to northern Nicaragua, including the southern portion of the Yucat��n Peninsula., Published as part of K��hler, Gunther, 2010, A revision of the Central American species related to Anolis pentaprion with the resurrection of A. beckeri and the description of a new species (Squamata: Polychrotidae), pp. 1-18 in Zootaxa 2354 on pages 9-12, DOI: 10.5281/zenodo.193523, {"references":["Boulenger, G. A. (1881) Description of a new species of Anolis from Yucatan. Proceedings of the Zoological Society London, 1881, 921 - 922.","Barbour, T. (1934) The anoles. II. The mainland species from Mexico southward. Bulletin of the Museum of Comparative Zoology, Harvard University, 77, 121 - 155.","Stuart, L. C. (1948) The amphibians and reptiles of Alta Verapaz, Guatemala. Miscellaneous Publications Museum of Zoology University of Michigan, 69, 1 - 109.","Myers, C. W. (1971) Central American lizards related to Anolis pentaprion: Two new species from the Cordillera de Talamanca. American Museum Novitates, 2471, 1 - 40.","Meyer, J. R. & Wilson, L. D. (1973) A distributional checklist of the turtles, crocodilians, and lizards of Honduras. Contributions in Science, Natural History Museum, Los Angeles County, 244, 1 - 39.","Fitch, H. S. & Seigel, R. A. (1984) Ecological and taxonomic notes on Nicaraguan anoles. Milwaukee Public Museum Contributions in Biology and Geology, 57, 1 - 13.","Lee, J. C. (1996) The amphibians and reptiles of the Yucatan Peninsula. Cornell University Press, Ithaca, New York, U. S. A., 500 pp.","Poe, S. (2004) Phylogeny of anoles. Herpetological Monographs, 18, 37 - 89.","Etheridge, R. (1959) The Relationships of the Anoles (Reptilia: Sauria: Iguanidae). An Interpretation Based on Skeletal Morphology. Ph. D. Dissertation, University of Michigan, Ann Arbor, Michigan, U. S. A.","Stuart, L. C. (1963) A checklist of the herpetofauna of Guatemala. Miscellaneous Publications Museum of Zoology University of Michigan, 122, 1 - 150.","Kohler, G. (2001) Anfibios y Reptiles de Nicaragua. Herpeton, Offenbach, Germany, 208 pp.","McCranie, J. R., Townsend, J. H. & Wilson, L. D. (2006) The amphibians and reptiles of the Honduran Mosquitia. - Krieger Publishing Company, Malabar, Florida, U. S. A., 291 pp."]}

Anolis polylepis Peters Anolis polylepis Peters, 1873: 738; syntypes (ZMB 7825 -26, 7830, 58002 -09) from ���Chiriqui���. Barbour (1934), Dunn (1930), Slevin (1942), Taylor (1956), Etheridge (1959), Poe (2004; in part.), K��hler et al. (2008). Diagnosis. A medium-sized species (SVL in largest specimen 59.4 mm) of the genus Anolis (sensu Poe 2004) that differs from all other Lower Central American beta anoles (sensu Etheridge 1967) except Anolis apletophallus, A. cryptolimifrons, and A. limifrons in that it is long-legged (longest toe of adpressed hind leg reaches to at least center of eye), has smooth ventral scales, and a slender habitus, often delicate. Anolis polylepis differs from the species in the cluster listed above by having two elongate, overlapping superciliaries (one such scale in A. apletophallus, A. cryptolimifrons, and A. limifrons), usually two anterior prenasal scales (versus one elongate prenasal scale) and dewlap coloration (usually orange with a darker orange basal portion in A. polylepis versus dirty white with a basal orange-yellow blotch in A. cryptolimifrons and A. limifrons, almost uniformly orange-yellow in A. apletophallus). Additionally, A. polylepis differs from A. limifrons, the only species of this cluster with which it occurs sympatrically, by having a bilobed hemipenis (unilobed in A. limifrons). For a comparison with the somewhat similar species A. tropidogaster and A. cupreus see Diagnosis section of A. osa. For variation in selected morphometric and scalation characters of A. polylepis see Table 3. Description. Norops polylepis is a medium sized anole (maximum recorded SVL 59.4 mm in males, 53.5 mm in females); dorsal head scales in internasal region keeled (Fig. 8), in prefrontal, parietal, and frontal areas rugose to tuberculate; deep frontal depression present, parietal depression absent; 7���11 (7.90 �� 0.87) postrostrals; anterior nasal divided in most individuals, usually in contact with rostral and first supralabial, for variation in nasal region see also Table 2 and Figure 1; 7���11 (8.52 �� 1.19) internasals; canthal ridge sharply defined; scales comprising supraorbital semicircles weakly keeled, largest scale in semicircles about same size as largest supraocular scale; supraorbital semicircles well defined; 1���5 (2.69 �� 0.83) scales separating supraorbital semicircles at narrowest point; 2���7 (3.80 �� 1.03) scales separating supraorbital semicircles and interparietal at narrowest point; interparietal well defined, greatly enlarged relative to adjacent scales, surrounded by scales of moderate size, longer than wide, usually larger than ear opening; 5���22 enlarged, keeled supraocular scales on each side, in 2���4 rows; enlarged supraoculars not in contact with supraorbital semicircles; 2 elongate supraciliaries, posterior one much shorter than anterior one; 3���6 enlarged canthals; 7���16 (1.46��1.66) scales between second canthals; 12���18 (14.4 �� 1.9) scales present between posterior canthals; loreal region slightly concave, 37���92 (57.69 �� 11.38) mostly keeled (some smooth or rugose) loreal scales in a maximum of 6���10 (8.06 �� 1.05) horizontal rows; 6���9 (7.17 �� 0.65) supralabials to level below center of eye; suboculars keeled, suboculars usually in broad contact with supralabials; ear opening vertically oval; scales anterior to ear opening granular, similar in size to those posterior to ear opening; 6���9 (6.82 �� 0.96) postmentals, outer pair usually largest; keeled granular scales present on chin and throat; male dewlap large, extending onto chest; 6���8 horizontal gorgetal-sternal rows with 26���38 scales per row, rows somewhat irregular, some of them with pairs or triplets of scales, apical portion of dewlap between marginal pairs and last gorgetal-sternal row free of scales; modal number of marginal pairs 4���5; female dewlap absent; no nuchal crest or dorsal ridge; 2 middorsal scale rows slightly enlarged, weakly keeled, dorsal scales lateral to middorsal series gradually larger than granular lateral scales; no enlarged scales scattered among granular laterals; 62���118 (87.3 �� 13.18) dorsal scales along vertebral midline between levels of axillae and groin in males, 66���111 (95.1 �� 11.2) in females; 35���68 (51.7 �� 6.9) dorsal scales along vertebral midline contained in one head length in males, 39���76 (55.8 �� 7.99) in females; ventral scales on midsection about the same size as largest dorsal scales; ventral body scales smooth, subimbricate; 56���85 (69.9 �� 8.3) ventral scales along midventral line between levels of axilla and groin in males, 57���76 (66.92 �� 5.30) in females; 36���62 (48.9 �� 5.72) ventral scales contained in one head length in males, 36���56 (44.0�� 5.12) in females; 134���212 (164.6 �� 14.5) scales around midbody in males, 140���212 (178.5 �� 17.2) in females; tubelike axillary pocket absent; preanal scales not keeled; postcloacal scales usually not enlarged; tail laterally compressed in cross section, tail height/tail width 1.20���1.85 (1.43 �� 0.13); basal subcaudal scales smooth; lateral caudal scales keeled, homogeneous; dorsal medial caudal scale row slightly enlarged, keeled, not forming a crest; most scales on lateral surface of antebrachium weakly keeled, unicarinate; 21���29 (24.66 �� 1.52) subdigital lamellae on Phalanges II���IV of Toe IV of hind limbs; 7���12 (9.4 ��1.0) subdigital scales on Phalanx I of Toe IV of hind limbs; SVL 42.2���59.4 (51.3 �� 3.76) mm in males, 39.6���53.5 (47.3 �� 3.52) mm in females; HL 10.8��� 14.4 (12.8 �� 0.88) mm in males, 10.1���13.7 (11.9 �� 0.78) mm in females; TAL 80.1 ���117.0 (102.1 �� 10.28) mm in males, 70.0��� 101.5 (86.3 �� 8.29) mm in females; SHL 11.6 ���16.0 (13.9 �� 1.12) mm in males, 10.5 ���14.0 (12.4 �� 0.98) mm in females; TAL/SVL 1.53���2.22 (2.01 �� 0.23) in males, 1.39���2.50 (1.92 �� 0.3) in females; HL/SVL 0.24���0.27 (0.26 �� 0.01) in males, 0.24���0.29 (0.26 ��0.0) in females; SHL/SVL 0.25���0.32 (0.29 �� 0.02) in males, 0.25���0.34 (0.25 ��0.0) in females; SHL/HL 0.98���1.22 (1.14 �� 0.08) in males, 0.97���1.17 (1.06 �� 0.1) in females; longest toe of adpressed hind limb usually reaching to a point between anterior border of eye and tip of snout. Of 45 specimens examined, the longest toe of the adpressed hind leg reached to mid-eye in 3 individuals (6.7 %), to anterior margin of eye in 29 individuals (64.4 %), and to a point between eye and snout in 13 individuals (28.9 %). Coloration in life of an adult male specimen (SMF 89509) of Anolis polylepis from Hacienda Caf�� de Eleta, 8.87157 ��N, 82.77099 ��W, 1380 m elevation, Chiriqu��, Panama, was recorded as follows: Dorsal ground color Citrine (51) suffused with Dark Brownish Olive (129); a narrow Dark Brownish Olive (129) interorbital bar present; lateral longitudinal stripe Cream Color (54); venter Cream Color (54) with a suggestion of Lime Green (59); ventral surfaces of limbs Dark Drab (119 B); dewlap Spectrum Orange (17) with the apical portion more yellowish; gorgetals Raw Umber (23); iris Raw Sienna (136). Coloration in life of the dewlap of an adult male specimen (SMF 89606) of Anolis polylepis from near Uvita, Province Puntarenas, Costa Rica, was recorded as follows: Orange Yellow (18) except for dirty white basal portion and a Raw Umber (23) central area; iris Raw Umber (23). For variation in dorsal pattern see also Table 2 and Figure 2. Geographic distribution. Anolis polylepis is distributed along the Pacific versant of central Costa Rica to western Panama, exclusive of the Penisula de Osa, from near sea level to 1615 m elevation. Anolis osa A. polylepis, Published as part of K��hler, Gunther, Dehling, Matthias & K��hler, Johannes, 2010, Cryptic species and hybridization in the Anolis polylepis complex, with the description of a new species from the Osa Peninsula, Costa Rica (Squamata: Polychrotidae), pp. 23-38 in Zootaxa 2718 on pages 27-28, DOI: 10.5281/zenodo.199772, {"references":["Peters, W. (1873) Uber neue Saurier (Spaeriodactylus, Anolis, Phrynosoma, Tropidolepisma, Lygosoma, Ophioscincus) aus Centralamerica, Mexico und Australien. - Monatsberichte der koniglichen Akademie der Wissenschaften zu Berlin, 1873, 738 - 747.","Barbour, T. (1934) The anoles II. The mainland species from Mexico southward. Bulletin of the Museum of Comparative Zoology, 77, 119 - 155.","Dunn, E. R. (1930) Notes on Central American Anolis. Proceedings of the New England Zoological Club, 12, 15 - 24.","Slevin, J. R. (1942) Notes on a collection of reptiles from Boquete, Panama, with the description of a new species of Hydromorphus. Proceedings of the California Academy of Sciences, 23, 463 - 480.","Taylor, E. H. (1956) A review of the lizards of Costa Rica. University of Kansas Science Bulletin, 38, 1 - 322.","Etheridge, R. (1959) The Relationships of the Anoles (Reptilia: Sauria: Iguanidae). An Interpretation Based on Skeletal Morphology. Ph. D. Dissertation, University of Michigan, Ann Arbor, Michigan, USA.","Poe, S. (2004) Phylogeny of anoles. Herpetological Monographs, 18, 37 - 89.","Etheridge, R. (1967) Lizard caudal vertebrae. Copeia, 1967, 699 - 721."]}

Xenopholis werdingorum sp. nov. (Figs. 2���5) Holotype: MNKR 4700, adult female; Hacienda San Sebasti��n (S 16 �� 21.676 ���, W 62 ��00.017���, 550 m; Fig. 6), Province of ��uflo de Ch��vez, Department of Santa Cruz, Bolivia; collected about 2300 hours on 25 October 2006 by M. Jansen and A. Schulze. Paratypes: ZSM 5113 / 2005, adult female; old cattle post San Ignacio near San Jos�� de Chiquitos (������verlassene Ranchogruppe San Ignacio���,���, Krieg 1931: 71), Province of Chiquitos, Department of Santa Cruz, Bolivia; collected by the German Chaco Expedition, which was between 20 and 28 September 1926 in that area according to the narration of the expedition by Krieg (1931); MNKR 4701, adult female, same locality data as holotype, collected about 2055 hours on 3 December 2008 by A. Schulze. Etymology: The name werdingorum is a noun in the genitive case, proposed to honor the family Werding, owners of type locality Hacienda San Sebasti��n. Lutz Werding provided great hospitality and encouragement for our inventories of the flora and fauna on their property. Furthermore, the family Werding generously provided broad logistic and financial support for the building of a biological research station (Centro de Investigaci��nes Ecol��gicas Chiquitos) on their grounds. Diagnosis: The new species can be identified as a member of the genus Xenopholis by its characteristic vertebral morphology (see Boulenger (1896): Externally, the new species differs from all other species of Xenopholis and all members of Clelia by having a unique dorsal color pattern (dorsal and dorsolateral surfaces of head and body dark brown with an iridescent sheen, grading to yellowish orange laterally and onto venter). The new species can be distinguished from all other species of Xenopholis and all members of Clelia by the following combination of characters: (1) 19 - 19 - 17 dorsal scale rows; (2) 171���179 ventrals (mean 176, n = 3); (3) 38���40 subcaudals (mean 39, n = 3); (4) no apical pits on dorsal scales; (5) two to three supralabials in contact with the eye; (6) eye diameter 0.35 times the distance between the eye and the tip of the snout; (7) one anterior and two posterior temporals; (8) two prefrontals; (9) a narrow septum within the neural spine and perpendicular to its long axis, as seen in x-ray images (Fig. 7). Xenopholis werdingorum differs from X. scalaris (features of X. scalaris in parentheses) in color pattern (dirty brick red above, narrow black spots on the sides; Fig. 8); number of prefrontals (one prefrontal); shape of loreal (loreal as long as wide or only slightly longer); number of supralabials in contact with the eye (two); number of ventrals (127���146); dorsal scale rows at midbody (17 - 17 - 17); number of subcaudals (28���36); and vertebral morphology (no narrow septum within the neural spine visible in x-ray images, Fig. 7). From X. undulatus, the new species is distinguished (features of X. undulatus in parentheses) in body color pattern (dorsal ground color yellowish, with a brown or black undulating band and lateral series of black spots; Fig. 9); coloration of head (dark streak behind the eye present; upper lip yellowish); and number of supralabials entering the eye (usually two, rarely three). From the superficially similar species of the genus Clelia (C. bicolor, C. clelia, C. equatoriana, C. errabunda, C. hussami, C. langeri, C. montana, C. plumbea, C. rustica, C. scytalina, and C. quimi) the new species of Xenopholis differs (characters of Clelia in parentheses) by number of subcaudals (more than 43 in all species of Clelia); number of ventrals in females (more than 192, except C. bicolor); supralabials in contact with the eye (two); anterior temporals (usually two, rarely one in C. rustica and C. bicolor; Fig. 10); and eye diameter (0.5 times the distance between the eye and the tip of the snout). Description of holotype: Adult female (Figs. 2���4), preserved in 70 % alcohol. Head little wider than neck; body long and diameter equal in all extensions; tail short relative to body, with conical tip; iris black and pupil round; nostril at anterior part of nasal scale; maxillary teeth aproximately 12, the last two grooved and separated from the others by a large diastema. Measurements: SVL 338 mm; TL 52 mm; TTL 390 mm; head length 13.9 mm; height of head at level of parietals 4.9 mm; height of head at level of nostrils 3.0 mm; largest width at level of parietals 7.9 mm; width at level of nostrils 3.6 mm; diameter of orbit 1.6 mm, approximately 0.35 times the distance from eye to snout; diameter of body at level of neck 6.2 mm; diameter of body at level of midbody 10.0 mm; diameter of body one head length before cloaca 6.4 mm; diameter of base of tail 5.3 mm; diameter of middle of tail 4.0 mm; diameter of final part at level of last middorsocaudal scale 1.5 mm; length of conical tip of tail 1.8 mm; length of ultimate dorsal 0.9 mm; length of ultimate subcaudal 0.8 mm; proportions: TL/SVL: 0.15; head length/total length 0.36. Pholidosis: Rostral wider than high; portion of tip of rostral visible from above less than length of suture between internasals; internasals wider than long (right internasal: width 1.8 mm, length 1.4 mm); prefrontals about as long as wide (right prefrontal: width 2.3 mm, length 2.2 mm); supraocular small (width 0.9 mm, length 1.3 mm); frontal irregularly heptagonal (width 3.7 mm, length 3.5 mm); nasal scales long (length 2.5 mm), in contact with loreal; preoculars 1 / 1, (right preocular: height 1.7 mm, length 1.2 mm); one loreal; 2 / 2 postoculars, lower postocular pentagonal and smaller than upper; supralabials 8 / 8; third to fifth supralabials in contact with eye; first supralabial triangular and smaller than others, third supralabial elongated posterodorsally at tip and reaching to eye; fourth supralabial below eye, fifth below eye and lower postocular; sixth and seventh supralabial below anterior temporal; sixth largest, irregular pentagonal; 1 / 1 anterior and 2 / 2 posterior temporals; anterior temporal narrow and long (right anterior temporal: height 1.4 mm, length 2.9 mm); upper and lower posterior temporals about same size (right lower posterior temporal: height 1.1 mm, length 2.0 mm; right upper posterior temporal: height 1.2 mm, length 1.7 mm); mental triangular (width 1.8 mm, length 1.4 mm); infralabials 9 / 9; first four infralabials in contact with anterior chin shields; fourth and fifth infralabials in contact with posterior chin shields; ninth infralabial smallest, fifth infralabial largest; anterior chin shields longer than posterior chin shields; anterior chin shields wider anteriorly than posteriorly; seven gular scales; one preventral; 178 ventrals; cloacal scute entire; 38 subcaudals; subcaudals/ventrals 0.21; dorsal scales smooth, in 19 - 19 - 17 rows, without apical pits; 12 dorsal scale rows at base of tail, eight at middle, and three before terminal scale. Color in life: Dorsal and dorsolateral surfaces of head and body dark brown with an iridescent sheen, grading to yellowish orange laterally and onto venter, scales of pale lateral region with dark edges; throat and venter creamy white; supralabials creamy white to pale yellow, except upper tips of supralabials below eye and posterior to eye, which are the same color as dorsal portion of head. Color in alcohol: Very similar as described for specimen in life, except that lateral brownish orange and orange-yellow portions have faded to creamy white. Variation: The paratypes are very similar in scalation and pholidosis to the holotype (see Table 1). ZSM 5113 / 2005 (adult female; SVL 353mm; TL 58mm): 179 ventrals and 40 subcaudals; fourth to fifth supralabials contact eye; posterior upper edge of third supralabial elongated, reaching eye on both sides; only right parietal in contact with sixth supralabial, maxillary teeth aproximately 11. MNKR 4701 (adult female; SVL 312mm; TL 55mm): 171 ventrals and 40 subcaudals; 8 supralabials (3 rd to 5 th in contact with the eye); left and right parietals in contact with sixth supralabial. length; TL: Tail length. Coloration of paratypes in alcohol very similar to that of holotype: uniform dorsal and dorsolateral coloration on head and body (chocolate brown in ZSM 5113 / 2005, dark chocolate brown in MNKR 4701); in ZSM 5113 / 2005 dorsal coloration bordered by a longitudinal line of scattered, similarly dark dots, extending to level of cloaca (Fig. 5). Distribution and natural history: The new species is known only from the type locality and from close to San Jos��, Department of Santa Cruz (Fig. 6). Photographs and descriptions of specimens reported by Str��ssman and Sazima (1993) and Marques et al. (2005) suggest this taxon might also be distributed in the Pantanal (see Str��ssmann and Sazima 1993; Marques et al. 2005). Its occurrence in other neighboring open areas is possible, such as the savanna and Cerrado habitats of the Beni in Bolivia. The holotype was active on the ground at 2300 hours in Cerrado savanna used for extensive cattle farming. The paratype MNKR 4701 was found at about 2100 hours at the edge of a wetland. Upon being captured, it was calm, exhibiting no aggressive behavior, but produced a strong-smelling secretion from the cloacal glands (A. Schulze, pers. comm.). The x-ray image of the paratype ZSM 5113 / 2005 showed a frog in the stomach. The preference of anurans as prey for snakes of the genus Xenopholis was mentioned by Cunha and Nascimento (1978), Cunha et al. (1985), Duellman (1990), and Fran��a and Ara��jo (2007)., Published as part of Jansen, Martin, ��lvarez, Lucindo Gonzales & K��hler, Gunther, 2009, Description of a new species of Xenopholis (Serpentes: Colubridae) from the Cerrado of Bolivia, with comments on Xenopholis scalaris in Bolivia, pp. 31-45 in Zootaxa 2222 on pages 33-41, DOI: 10.5281/zenodo.190108, {"references":["Krieg, H. (1931) Wissenschaftliche Ergebnisse der Deutschen Gran Chaco-Expedition. Geographische Ubersicht und illustrierter Routenbericht. Strecker und Schroder Verlag, Stuttgart. 95 pp.","Boulenger, G. A. (1896) Catalogue of the snakes in the British Museum (Natural History). Vol. 3. Containing the Colubridae (Opistoglyphae and Proteroglyphae), Amblycephalidae, and Viperidae. London, Trustees of the British Museum. 727 pp.","Strussmann, C. & C. & Sazima, I. (1993) The snake assemblage of the Pantanal at Pocone, Western Brazil: Faunal Composition and Ecological Summary. Studies on Neotropical Fauna and Environment 28, 157 - 168.","Marques, O. A. V., Eterovic, A., Strussmann, C. & I. Sazima, (2005) Serpentes do Pantanal. Guia ilustrado. Holos Editora, Ribeirao Preto, 184 pp.","Cunha, O. R. da & Nascimento, F. P do (1978) Ofidios da Amazonia. X - As cobras da regiao leste do Para. Museu Paranense Emilio Goeldi Publicacoes Avulsas, 31, 1 - 218.","Cunha, O. R. da, Nascimento, F. P. do & Avila-Pires, T. C. Sauer de (1985) Os repteis da area de Carajas, Para, Brasil (Testudines e Squamata). Museu Paranense Emilio Goeldi Publicacoes Avulsas, 40, 9 - 92.","Duellman, W. E. (1990) Herpetofaunas in Neotropical rainforests: comparative composition, history, and resource use. In: Gentry A. H. (ed.), Four Neotropical rain forests. New Haven: Yale University Press, pp. 455 - 505.","Franca, F. G. R & Araujo, A. F. B. (2007) Are there co-occurrence patterns that structure snake communities in Central Brazil? Brazilian Journal of Biology, 67, 33 - 40."]}

Pristimantis ventriguttatus new species Holotype: SMF 86413, an adult female (Fig. 1), collected in amplexus with male SMF 86414, from Hacienda Taulis (approx. 6 �� 50 ' S, 79 �� 10 ' W, 1800 m), Provincia de Santa Cruz, Departamento de Cajamarca, Peru, obtained by H.-W. Koepcke on 2 February 1952. Paratypes: Three adult males: SMF 86414, collected with the holotype; SMF 86416, SMF 86415, partially desiccated, from Hacienda Taulis (approx. 6 �� 50 'S, 79 �� 10 ' W, 1700 m), Provincia de Santa Cruz, Departamento de Cajamarca, Peru, obtained by H.-W. Koepcke on January 1952. Diagnosis. A member of the Pristimantis orestes Group having the following combination of characters: (1) skin on dorsum shagreen with small tubercles coalesced into ridges dorsolaterally, forming discontinuous dorsolateral folds; venter areolate; discoidal fold present; (2) tympanic membrane and annulus present, upper margin slightly concealed by supratympanic fold; (3) snout long, acutely rounded in dorsal view, rounded in lateral view; (4) upper eyelid bearing small tubercles; upper eyelid width narrower than IOD; cranial crests absent; (5) dentigerous processes of vomers small, oblique; (6) males with vocal sac and vocal slits, nuptial pads absent; (7) Finger I shorter than Finger II; discs on outer fingers broadly expanded, slightly truncate; (8) fingers with lateral fringes; (9) ulnar and tarsal tubercles present; (10) heel with small tubercles; inner tarsal fold present; (11) inner metatarsal tubercle oval, twice the size of subconical outer metatarsal tubercle, elevated, slightly conical in lateral view; few, low supernumerary plantar tubercles present; (12) toes with lateral fringes; toes webbed basally; Toe V slightly longer than Toe III; toe discs about same size as fingers, slightly truncate; (13) in ethanol, dorsum pale brown with dark brown flecks and narrow, tan middorsal stripe; venter brown with tan blotches and spots; axilla, groin, flanks, anterior and posterior surfaces of thighs, and inner surfaces of shanks, dark brown with tan blotches and spots; iris dark gray; (14) SVL in single female 29.4 mm, in males 17.8���21.8 mm (n = 3). Pristimantis ventriguttatus is tentatively assigned to the P. o re s t e s Group by having a contrasting pattern of tan blotches and spots on dark brown in the groin, a small SVL of no more than 33 mm (Duellman et al. 2006), and a robust body. Pristimantis ventriguttatus is readily distinguished from the other 11 species (see Table 1) currently assigned to the P. o re s t e s Group (Duellman et al. 2006, Lehr & Duellman 2007 a) by having tan blotches and spots on a brown venter, anterior and posterior surfaces of the thighs, and groin, discs on outer fingers broadly expanded, and a long snout with a small terminal tubercle. Pristimantis ventriguttatus shares with P. cordovae and P. vidua dorsolateral folds, dentigerous processes of the vomers, and vocal slits, but male P. v e n - triguttatus and P. v i d u a lack nuptial pads (present in P. cordovae). Furthermore the digits are broadly expanded and mostly truncate in P. ventriguttatus, whereas narrowly expanded and marginate in P. cordovae. Pristimantis vidua has a short snout without terminal tubercle (snout long with terminal tubercle in P. v e n - triguttatus), and lacks lateral fringes (present in P. ventriguttatus). Furthermore, dorsolateral folds are usually tan in P. vidua (not distinct from surrounding color or blackish brown in P. ventriguttatus) and the venter is tan with fine, small brown spots (brown with tan blotches and spots in P. ventriguttatus). Pristimantis ventriguttatus and P. pinguis occur syntopically at Hacienda Taulis. Both species are of similar size, have dorsolateral folds, tympanum present and males that have vocal slits present and nuptial pads absent. However, fingers and toes are short in P. pinguis (long in P. ventriguttatus) without lateral fringes (present in P. ventriguttatus), with rounded discs (truncate) that are barely wider than digit proximal to it (discs broadly expanded in P. ventriguttatus), snout short (long), and venter uniformly tan (contrasting pattern consisting of tan blotches and spots on brown). Both Pristimantis cajamarcensis (Barbour and Noble) and P. lymani (Barbour and Noble) are widespread in the western Andes of northern Peru and can easily be distinguished from P. ventriguttatus by having males with nuptial pads (absent in P. ventriguttatus). Furthermore, P. cajamarcensis has dorsolateral folds and lateral fringes absent (both present in P. ventriguttatus), and P. l y m a n i are much larger (male SVL to 45.3 mm in P. lymani vs. 21.8 mm in P. ventriguttatus) and have Finger I> II (Finger I P. ventriguttatus). Pristimantis ventriguttatus and P. s i m o n s i i (Boulenger) may occur in sympatry in Departamento de Cajamarca, but P. simonsii lacks a tympanum (present in P. ventriguttatus) and lacks marginal grooves (present in P. ventriguttatus). Other Pristimantis having a distinctive color pattern consisting of tan or white blotches and spots on a brown venter are P. douglasi (Lynch) belonging to the galdi Group from eastern Colombia and Pristimantis phalaroinguinis (Duellman and Lehr) belonging to the conspicillatus Group from the Cordillera Occidental in northern Peru (Departamento de Cajamarca). Both P. ventriguttatus and P. phalaroinguinis have discs on outer fingers broadly expanded and truncate, but P. ventriguttatus has fingers and toes with lateral fringes (absent in P. phalaroinguinis), is much smaller (SVL in males to 21.8 mm vs. 31.4 mm in P. phalaroinguinis: Duellman & Lehr 2007 a), has areolate skin on the venter (smooth in P. phalaroinguinis), and has vocal slits (absent in P. phalaroinguinis). Pristimantis ventriguttatus and P. douglasi are similar in size; both have vocal slits, fingers and toes with lateral fringes, and areolate skin on the venter. Pristimantis ventriguttatus has long dorsolateral folds (extending only to shoulder in P. douglasi), posterior border of tongue notched (not notched in P. douglasi), and no nuptial pads in males (present in P. douglasi). Pristimantis muscosus (Duellman & Pramuk) a member of the unistrigatus Group from the eastern slopes of the Cordillera Central in northern Peru also has an areolate venter with tan spots and fingers and toes with lateral fringes, but P. ventriguttatus has dorsolateral folds (absent in P. muscosus), tarsal folds (absent in P. muscosus), and lacks conical heel tubercles (present in P. muscosus). Furthermore, P. ventriguttatus is much smaller (female SVL 29.4 mm in E. ventriguttatus vs. 46.1 mm in P. m u s c o s u s: Duellman & Pramuk 1999). Description of the holotype. Head as wide as body and nearly as wide as long; head width 39.1 % of SVL; head length 40.1 % of SVL; snout long with small terminal tubercle, acutely rounded in dorsal view, rounded in lateral view (Figs. 2 A, B); eye diameter 91.7 % of eye���nostril distance; nostrils slightly protuberant, directed dorsolaterally; canthus rostralis nearly straight in dorsal view, angular in profile; loreal region concave; lips rounded; upper eyelid width 93.3 % of interorbital distance, with small tubercles; tympanic annulus round, its upper and posterodorsal part obscured by supratympanic fold, tympanum diameter 48.5 % of eye diameter; tympanum-eye distance about equal to tympanum diameter; enlarged postrictal tubercles fused into a ridge on each side of head. Choanae small, oval, not concealed by palatal shelf of maxilla; dentigerous processes of vomers small, narrowly separated medially, situated posteromedial to choanae; tongue 3 �� as long as wide, slightly notched posteriorly, posterior half free. Skin on dorsum shagreen with small tubercles; dorsolateral tubercles forming slender, discontinuous dorsolateral folds, two on each side; skin on flanks tuberculate with tubercles coalescing into ridges; skin on ventral thighs, belly, chest, and throat areolate, skin on other ventral surfaces smooth; discoidal and thoracic folds present; cloacal sheath short; cloacal region without large tubercles. Three small ulnar tubercles on each forearm, round, low; palmar tubercles slightly elevated, outer palmar tubercle bifid, approximately 2 �� size of oval, inner palmar tubercle; subarticular tubercles well defined, elongate in ventral view, slightly subconical in lateral view; one supernumerary tubercle at base of each finger except Finger I, round, low, approximately one-fourth size of subarticular tubercles; fingers with lateral fringes; Finger I shorter than Finger II; discs on Fingers I���II slightly expanded, most prominent on Fingers III���IV; discs on Fingers III���IV nearly truncate, discs on Fingers I���II rounded; ventral pads of fingers well defined by circumferential grooves (Fig. 2 C). Hind limbs slender, tibia length 53.1 % of SVL; foot length 50.3 % of SVL; upper surfaces of hind limbs tuberculate; posterior and ventral surfaces of thighs areolate; no distinct tubercles on heel or on outer surface of tarsus; inner tarsal fold extending along distal half of tarsus; inner metatarsal tubercle elevated, ovoid, twice size of subconical outer metatarsal tubercle; one, low plantar supernumerary tubercle at base of Toes II, IV, and V, two supernumerary tubercles at base of Toe III; subarticular tubercles well defined, round in ventral view, subconical in lateral view; toes with lateral fringes; basal webbing between Toes IV and V; discs about equal to those on fingers, most prominent on Toe IV; discs nearly truncate; toes having ventral pads well defined by circumferential grooves; relative lengths of toes: 1 Variation. Measurements (in mm) for the three males (SMF 86414, 86416, 86415): SVL 21.3, 21.8, 17.8; tibia length 13.1, 11.1, 7.8; foot length 12.8, 9.4, 7.1; head length 9.3, 8.2, 5.6; head width 9.1, 7.7, 7.9; eye diameter 2.4, 2.5, 2.7; tympanum diameter 1.4, 1.2, 1.1; IOD 2.8, 2.9, 2.8; upper eyelid width 2.8, 1.9, 1.9; inter narial distance 2.3, 1.8, 1.5; eye-nostril distance 2.5, 2.5, 2.3. In one male (SMF 86414), the dorsum and flanks are uniformly dark brown. This individual has fewer tan spots and blotches in the axilla and groin and on the anterior and posterior surfaces of the thighs and the inner surfaces of shanks than the holotype; the venter is tan and brown with few, barely distinct tan blotches and spots. In the other males (SMF 86415, 86416), the dorsum and flanks are brown and tan with blackish-brown flecks dorsolaterally. The groin is pale brown with large tan flecks; anterior surfaces of thighs are dark brown with tan spots, and the posterior surfaces of the thighs are dark brown with elongate, tan blotches. The concealed surfaces of the shanks are pale brown with tan blotches and spots, and the venter is tan with small dark brown spots. All males have nearly straight vocal slits located at posterior half of mouth floor closer to margin of jaw than to tongue. Distribution and ecology. Pristimantis ventriguttatus is known only from the type locality described as montane forest by H.-W. Koepcke. Sympatric Pristimantis include P. cajamarcensis, P. lymani, and P. pinguis. Etymology. The specific name ventriguttatus is derived from the Latin noun venter meaning belly and the Latin adjective guttatus meaning spotted. The name refers to the distinct ventral pattern consisting of tan spots and flecks on dark brown coloration. Remarks. The Pristimantis orestes Group was defined by Lynch and Duellman (1997) and modified by Duellman et al. (2006) and Lehr and Duellman (2007 a) as three new species were assigned to the group. With P. ventriguttatus the P. o re s t e s Group contains 12 species and occurs in p��ramo, puna, and cloud forest at elevations of 1700���4010 m above sea level in the Andes from southern Ecuador to northern Peru. With more intensive exploration of the Cordillera Occidental, the discovery of new species of Pristimantis is likely. Phylogeny of the orestes Group is unknown. Both members of the orestes Group and Phrynopus are similar in several morphological characters: Phrynopus have narrow digits that lack pads and marginal grooves, whereas members of the Pristimantis orestes Group have narrow digits with pads that have reduced or absent (only in Pristimantis simonsii) marginal grooves. As in Phrynopus, most of the members of the orestes Group have limited distributional ranges which are restricted to the type locality or its close surroundings. Sexual male characters such as nuptial pads or vocal slits are useful to distinguish species from each other. Unfortunately these important diagnostic characters are frequently ignored or simply overseen in species descriptions, even if adult males are present. This led to wrong conclusions that Phrynopus lack nuptial pads (see comments by Lehr 2006) or that nuptial pads are intraspecific variable (present or absent) in males. Unlike in bufonids or Telmatobius where males usually develop seasonally dark brown, spinous nuptial pads, nuptial pads in Pristimantis (and Phrynopus) are more secretive, cream colored usually non spinous and seem to be present permanently, indicating that reproduction may take place throughout the year (Lehr et al. 2006). In none of the thousands observed specimens from different museum collections did the senior author find nuptial pads in adult male Pristimantis (or Phrynopus) to be seasonally present or intraspecific variable (absent or present). Only slight differences in size and shape of nuptial pads were observed. Thanks to the detailed field notes by H.-W. Koepcke, we know that female holotype and male paratype (SMF 86414) of P. ventriguttatus were found in amplexus, indicating that both were reproductive adults. Despite the long time in preservative, the type series is well preserved., Published as part of Lehr, Edgar & K��hler, Gunther, 2007, A new species of the Pristimantis orestes Group (Anura: Leptodactylidae) from the Cordillera Occidental in northern Peru, pp. 45-54 in Zootaxa 1621 on pages 46-52, DOI: 10.5281/zenodo.179186, {"references":["Duellman, W. E., Lehr, E. & Venegas, P. J. (2006) Two new species of Eleutherodactylus (Anura: Leptodactylidae) from northern Peru. Zootaxa, 1285, 51 - 64.","Lehr, E. & Duellman, W. E. (2007 a) Two new species of Eleutherodactylus (Anura: Leptodactylidae) from the Cordillera Occidental in Peru. Copeia, 2007, 140 - 149.","Duellman, W. E. & Pramuk, J. B. (1999) Frogs of the genus Eleutherodactylus (Anura: Leptodactylidae) in the Andes of northern Peru. Scientific Papers Natural History Museum University of Kansas, 13, 1 - 78.","Lynch, J. D. & Duellman, W. E. (1997) Frogs of the genus Eleutherodactylus in western Ecuador. Systematics, ecology, and biogeography. The University of Kansas, Natural History Museum Special Publication, 23, 1 - 236."]}