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Stenocercus qalaywasi sp. nov. Figures 8���9 Holotype. CORBIDI 20567, adult male from Lampa Pariahuanca, Pariahuanca district, Huancayo province, Jun��n Department, Peru (11��58���51.9���S, 74��53���47.6���O, 2,587 m), collected by Luis A. Garc��a-Ayachi on 19 June 2019. Paratype. CORBIDI 20568, an adult female collected with the holotype. Diagnosis. From all currently known species of Stenocercus (including the new species described herein), S. qalaywasi sp. nov. is only similar to thirteen species (e.g., S. asenlignus sp. nov., S. arndti, S. flagracanthus, S. leybachi sp. nov., S. bolivarensis, S. carrioni, S. chlorostictus, S. crassicaudatus, S. empetrus, S. eunetopsis, S. nigrocaudatus sp. nov., S. torquatus and S. simonsii) in sharing the following characters: granular scales on posterior surface of thighs, relatively short tail, caudals spinose, and two caudal whorls per autotomic segment. From these species, S. qalaywasi possesses a smaller number of vertebrals than S. nigrocaudatus and S. torquatus (77���79 in S. qalaywasi versus 96���108 in S. nigrocaudatus and 83���115 in S. torquatus), scales on dorsal surface of neck keeled (granular in S. asenlignus, S. nigrocaudatus and S. torquatus), a longer tail than S. asenlignus, S. nigrocaudatus and S. torquatus (tail length 57���60% of total length in S. qalaywasi, 51���55% in S. asenlignus, 50���55% in S. nigrocaudatus, and 47���54% in S. torquatus). Stenocercus leybachi is easily distinguished from S. qalaywasi by having a distinct, serrate, low crest on neck (absent in S. qalaywasi) and by lacking a distinct, black antehumeral collar (complete middorsally in S. qalaywasi). Stenocercus arndti, S. crassicaudatus, S. empetrus, S. flagracanthus, and S. simonsii can also be easily distinguished from S. qalaywasi by having the scales on dorsal surface of neck granular, while in the new species they are keeled. Furthermore, S. crassicaudatus and S. flagracanthus possess more vertebrals (83���97) than S. qalaywasi (77���79). In the case of S. flagracanthus, the tail is shorter (50���54% of total length) than S. qalaywasi (57���60%). Species such as S. bolivarensis, S. carrioni, S. chlorostictus, and S. eunetopsis possess keeled scales on the dorsal surface of neck, like S. qalaywasi, and preserved specimens can be confused. However, these species can be distinguished by the following features: from S. arndti, S. carrioni, S. chlorostictus, S. empetrus, and S. simonsii by having a distinct black, middorsally complete, antehumeral collar and two nuchal bands (antehumeral collar incomplete middorsally and nuchal bands absent in the aforementioned species); from S. carrioni and S. chlorostictus by having more vertebral scales (55���72 in S. carrioni, 63���73 in S. chlorostictus and 77���79 in S. qalaywasi); from S. bolivarensis by having the lateral scales of body granular (strongly keeled in S. bolivarensis); and from S. eunetopsis by having more scales around midbody (60���80 in S. eunetopsis and 87���92 in S. qalaywasi) and a shorter tail (57���60% of total length vs. 62���66% in S. eunetopsis). Characterization. (1) Maximum SVL in males 88 mm (n = 1); (2) maximum SVL in females 95 mm (n = 1); (3) vertebrals 77���79; (4) paravertebrals 104���108; (5) scales around midbody 87���92; (6) supraoculars six; (7) internasals four; (8) postrostrals six; (9) loreals five; (10) gulars 44; (11) subdigitals on Finger IV 26���29; (12) subdigitals on Toe IV 33���34; (13) posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; (14) postfemoral mite pocket distinct with slit-like opening [Type 2 of Torres-Carvajal (2007b)]; (15) parietal eye not visible; (16) scales on occipitoparietal region small, rugose, juxtaposed; (17) projecting angulate temporal absent; (18) row of enlarged supraoculars occupying most of supraocular region absent; (19) scales on frontonasal region juxtaposed, rugose; (20) preauricular fringe present; (21) antegular, antehumeral, gular, longitudinal, and postauricular neck folds present; (22) lateral and dorsal nuchals similar in size; (23) posterior gulars cycloid, smooth, slightly imbricate, not notched; (24) lateral body scales smaller than dorsals, reduced in size, approximately one third of the size of dorsal body scales closer to vertebral line; (25) vertebrals larger than adjacent paravertebrals forming a distinct vertebral row; (26) dorsolateral crest absent; (27) ventrals smooth, imbricate; (28) scales on posterior surface of thighs granular; (29) prefemoral fold present; (30) inguinal groove present; (31) preanals not projected; (32) tail not compressed laterally in adult males; (33) tail relatively short (tail length 57���60% of total length); (34) two caudal whorls per autotomic segment; (35) caudals spinose; (36) dark stripe extending anterodorsally from subocular region to supraciliaries absent; (37) dark patch extensively covering gular region of females absent; (38) dark patch extensively covering gular region of adult males absent; (39) black patch on ventral surface of neck in adult males absent; (40) dark midventral longitudinal mark, such as faint line, conspicuous stripe, or extensive patch in adult males absent; (41) black patches on ventral surfaces of thighs in adult males absent; (42) background color of dorsum green or dark gray, with distinct black bands in adults individuals of both sexes; (43) postxiphisternal inscriptional ribs not examined. Description of the holotype. Male (Fig. 8); SVL 88 mm; TL 136 mm; maximum head width 16.8 mm; head length 21.4 mm; head height 13.2 mm; posterior dorsal head scales small, smooth, juxtaposed; parietal eye not visible; supraoculars in six rows, smooth, juxtaposed, with the most lateral two rows less than half the size of the medial adjacent rows; distinct circumorbitals absent; canthals two; internasals four; postrostrals five, two lateral ones larger; supralabials six; infralabials six; loreals four; lorilabials in one row; preoculars two, the dorsal-most in contact with posterior canthal; lateral temporals granular; gulars in 44 rows between tympanic openings; all gulars cycloid, smooth, imbricate, not notched; second infralabial in contact with first two sublabials; first pair of postmentals partially in contact medially; mental in contact with first pair of infralabials but not with the first pair of postmentals; dorsal scales of neck small, weakly keeled, slightly imbricate, and slightly larger than granular laterals; dorsal scales of body imbricate, keeled, becoming smaller, slightly keeled or smooth towards flanks; scales around midbody 87; vertebrals 77, enlarged, keeled, imbricate, forming a distinct vertebral row; paravertebrals adjacent to vertebral row 104, same size or slightly smaller than vertebrals; ventrals smooth, imbricate, larger than dorsals; preauricular fringe short, composed by two enlarged, projected scales; antegular, antehumeral, gular, longitudinal, and postauricular neck folds present; ventrolateral fold present; humeral dorsal scales imbricate, weakly keeled; dorsal scales of forelimbs imbricate, keeled; dorsal scales of hindlimbs imbricate, keeled, mucronate; ventral humeral scales imbricate; ventral scales of forearms keeled, imbricate, and ventral scales of hindlimbs imbricate, smooth; palmars imbricate, keeled; plantars imbricate, keeled, mucronate; lamellae on Finger IV 26; lamellae on Toe IV 33; tail rounded; caudals strongly keeled, mucronate, imbricate dorsally; two caudal whorls per autotomic segment; basal subcaudals strongly keeled, imbricate; posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; postfemoral mite pocket distinct with slit-like opening; [Type 2 of Torres-Carvajal (2007b)]. Color in life of holotype (Fig. 9 A-C): Head bright green with a narrow gray postocular stripe and two gray stripes below it; dorsal surface of neck with two pale gray transverse bands and a distinct, middorsally complete black antehumeral collar; dorsum, limbs and tail with a green hue and scattered dark gray flecks; dorsum and forelimbs also covered by bright green specks; gular region dark gray, spotted with light gray blotches, and bearing a pale gray medial patch posteriorly; ventral surface of neck pale gray; chest greenish gray; belly light pink; ventral surface of hips and hindlimbs cream; cloacal region and ventral surface of base of tail cream with a light pink hue; ventral surface of tail gray becoming dark gray towards tip. Iris brown. Color in preservative (Fig. 8): dorsal surface dark gray; the postorbital gray stripe black; bands on neck and antehumeral collar black; dorsum with short irregular black bands along the vertebral region and covered with light gray specks; forelimbs covered with light gray specks and hindlimbs with darker bands than the gray background; fingers and toes light gray with black spots. Ventral surface similar as in life, but without the pink and green hues. Intraspecific variation. Scale counts and measurements for Stenocercus qalaywasi are presented in Table 1. Loreals 4���5; postrostrals 4���5; second infralabial not in contact with third sublabial in all specimens; first pair of postmentals in contact in both specimens. Sexual dimorphism is not evident in S. qalaywasi; the single female paratype (CORBIDI 20568) differs from the male holotype only in having the antehumeral black collar incomplete (Fig. 9E). Ventrally, the gular coloration is greenish cream with cream spots laterally in the female paratype (Fig. 9F), and dark gray with light gray spots in the male holotype. With a low sample of two specimens, the female is larger (95 mm) than the male (88 mm SVL). Distribution and natural history. Stenocercus qalaywasi is known only from the village of Lampa Pariahuanca in the inter-Andean valley of the Mantaro River in central Peru (Fig. 3). It occurs at an elevation of 2,587 m in the department of Jun��n. The type locality lies within the Peruvian Yungas ecoregion following Olson et al. (2001) and Yungas ecoregion according to Brack-Egg (1986) and Pe��aherrera del ��guila (1989). Six individuals of S. qalaywasi were observed basking on sunny days at 0900 hours on the rooftops of Lampa Pariahuanca village houses. When disturbed, they hid under the roof tiles, in holes between bricks or in the slit between roof and wall. The surrounded areas of the village are croplands of corn (Zea mays), potato (Solanum tuberosum), carrot (Daucus carota), and barley (Hordeum vulgare), with a few scattered patches of secondary montane forest on the steepest slopes. No other species of lizards or snakes were recorded at this locality. When the adult individuals of S. qalaywasi are basking, they possess the head bright green and the dorsal surface of body and tail with a green hue bearing conspicuous light green specks on back (Fig 9G). At the moment of capture the single male collected changed dramatically in coloration to dark gray with the dorsum containing light gray specks (Fig. 9A). The single collected female specimen did not show a dramatic change of coloration becoming simply dull green (Fig. 9D). Etymology. The specific epithet ��� qalaywasi ��� is a noun in apposition derived from two words in Quechua, ��� qalaywa ��� that means lizard and ��� wasi ��� that means home or house. The specific name refers to the habitus of this species of living in the houses of Lampa Pariahuanca village., Published as part of Venegas, Pablo J., Garc��a-Ayachi, Luis A., Ch��vez-Arribasplata, Juan C. & Garc��a-Bravo, Antonio, 2022, Four new species of polychromatic spiny-tailed iguanian lizards, genus Stenocercus (Iguania: Tropiduridae), from Peru, pp. 1-28 in Zootaxa 5115 (1) on pages 19-23, DOI: 10.11646/zootaxa.5115.1.1, http://zenodo.org/record/6345982, {"references":["Torres-Carvajal, O. (2007 b) A taxonomic revision of south American Stenocercus (Squamata: Iguania) lizards. Herpetological monographs, 21, 76 - 178. https: // doi. org / 10.1655 / 06 - 001.1","Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N. D., Powell, G. V. N., Underwood, E. C., D'Amico, J. A., Itoua, I., Strand, H. E., Morrison, J. C., Loucks, C. J., Allnutt, T. F., Ricketts, T. H., Kura, Y., Lamoreux, J. F., Wettengel, W. W., Hedao, P. & Kassem, K. R. (2001) Terrestrial ecoregions of the world: A new map of life on earth. BioScience, 51, 933 - 938. https: // doi. org / 10.1641 / 0006 - 3568 (2001) 051 [0933: TEOTWA] 2.0. CO; 2","Penaherrera del Aguila, C. (1989) Atlas del Peru. Instituto Geografico Nacional, Lima, 400 pp."]}

Stenocercus leybachi sp. nov. Figures 4���5 Holotype. CORBIDI 16397, an adult male, from Comunidad Saria, Chaglla (9��43���27.7���S, 75��48���35.9���W, 1,082 m), Hu��nuco Province, Hu��nuco Department, Peru, collected by D. V��squez on 11 October 2015. Paratypes (22). PERU: HU��NUCO DEPARTMENT: Hu��nuco Province: CORBIDI 9323, adult female, from Chinchavito (9�� 33��� 27.2���S, 75�� 55��� 07.1���W, 824 m), collected by P.J. Venegas on 16 June 2011; CORBIDI 13659, juvenile male, from Miraflores (9��41���0.11���S, 75��50���33.44���W, 1,270 m), collected by V. Duran and L. Lujan on 11 October 2013; CORBIDI 16390, a juvenile, CORBIDI 16394���95 and 16398, males, CORBIDI 16399���16400, females, from Saria, Chaglla (9��42���55.4���S, 75��49���3���W, 1,151 m), collected by D. V��squez on 24 August 2015 and 10 October 2015; CORBIDI 16412 juvenile, CORBIDI 16428���29, 16431, females, CORBIDI 16432, juvenile, CORBIDI 16437, female, from Agua Nueva, Chinchao (9��41���59.82���S, 75��49���55.39���W, 1,115 m), collected by J. C. Ch��vez-Arribasplata within 24 to 28 September 2015 and 1 to 4 October 2015; CORBIDI 16525���26, a juvenile and female, CORBIDI 16532���33, male and juvenile, CORBIDI 16539, female, and CORBIDI 16594, juvenile, also from Agua Nueva, Chinchao (9��43���6.67���S, 75��48���56.1���W, 1,185 m), collected by A.C. Barboza within 10 to 15 November 2015; Pachitea Province: CORBIDI 13333, male, from Casa de M��quinas, Chaglla (9��42���17.81���S, 75��49���47.96���W, 928 m), collected by V. Duran and L. Luj��n on August 2013; CORBIDI 13334, male, from El Waro, Chaglla (9��41���55.87���S, 75��49���46.56���W, 927 m), collected by V. Duran and L. Luj��n on August 2013. Diagnosis. Stenocercus leybachi differs from all other Stenocercus species, except S. arndti, S. asenlignus sp. nov., S. bolivarensis, S. carrioni, S. chlorostictus, S. crassicaudatus, S. empetrus, S. eunetopsis, S. flagracanthus, S. nigrocaudatus sp. nov., S. qalaywasi sp. nov., S. torquatus, and S. simonsii, by having granular scales on posterior surface of thighs, relatively short tail, caudals spinose and two caudal whorls per autotomic segment. Nevertheless, S. leybachi can be easily distinguished from all aforementioned species by having a distinct serrate low crest on neck, that in some specimens reach to the middle of the dorsum. We found the arboreal S. chinchaoensis, a species known from the Upper Huallaga Basin at Hu��nuco Department (Venegas et al. 2013), sympatric with S. leybachi in the locality of Agua Nueva, Chinchao district, at elevations of 1,115 to 1,200 m. This species also has the ability to change color between green and gray, however this has a longer tail (tail length 61���64% of total length versus 52���58% of total length, respectively) and caudal scales not spinose. Stenocercus boettgeri is another green species distributed in the Amazon slope of central Peru from Hu��nuco, Pasco and Jun��n departments, at elevations between 2900 and 3250 m (Torres-Carvajal 2007b). Although this species can be confused with S. leybachi, the former species can be readily distinguished by having the lateral neck scales less than half the size of dorsal neck scales (dorsal and lateral neck scales similar in size in S. leybachi) and lacking a spinose tail. Characterization. (1) Maximum SVL in males 84mm (n=6); (2) maximum SVL in females 82mm (n=7); (3) vertebrals 63���73; (4) paravertebrals 100���112; (5) scales around midbody 74���97; (6) supraoculars 7���9; (7) internasals 4; (8) postrostrals 4���6; (9) loreals 3���5; (10) gulars 42���61; (11) subdigitals on Finger IV 21���28; (12) subdigitals on Toe IV 28���32; (13) posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; (14) postfemoral mite pocket distinct with slit-like opening [Type 2 of Torres-Carvajal (2007b)]; (15) parietal eye not visible; (16) scales on occipitoparietal region small, some rugose and other smooth, juxtaposed; (17) projecting angulate temporal absent; (18) row of enlarged supraoculars occupying most of supraocular region absent; (19) scales on frontonasal region juxtaposed, smooth, slightly imbricate; (20) preauricular fringe present; (21) antegular, antehumeral, gular, longitudinal, and postauricular neck folds present; (22) lateral and dorsal nuchals similar in size; (23) posterior gulars cycloid, smooth, slightly imbricate, not notched; (24) lateral body scales smaller than dorsals, reduced in size, approximately one third of the size of dorsal body scales closer to vertebral line; (25) vertebrals larger than adjacent paravertebrals forming a conspicuous vertebral row and a distinct low serrate crest on neck that in some specimens comes to the middle of dorsum; (26) dorsolateral crest absent; (27) ventrals smooth, imbricate; (28) scales on posterior surface of thighs granular; (29) prefemoral fold present; (30) inguinal groove present; (31) preanals not projected; (32) tail not compressed laterally in adult males; (33) tail relatively short (tail length 52���58 % of total length); (34) two caudal whorls per autotomic segment; (35) caudals spinose; (36) dark stripe extending anterodorsally from subocular region to supraciliaries absent; (37) dark patch extensively covering gular region of females absent; (38) dark patch extensively covering gular region of adult males absent; (39) black patch on ventral surface of neck in adult males absent; (40) dark midventral longitudinal mark, such as faint line, conspicuous stripe, or extensive patch in adult males absent; (41) black patches on ventral surfaces of thighs in adult males absent; (42) background color of dorsum green, dark brown or gray and without distinct black bands in adult individuals of both sexes; (43) postxiphisternal inscriptional ribs not in contact midventrally [Pattern 4A of Torres-Carvajal 2004)]. Description of the holotype. Male (Fig. 4); SVL 83 mm; TL 117 mm; maximum head width 16.5 mm; head length 20.1 mm; head height 12.7 mm; posterior dorsal head scales small, smooth, slightly imbricate, juxtaposed; parietal eye not visible; supraoculars in seven rows, smooth, slightly imbricate, with the most lateral two rows less than half the size of the medial adjacent rows; distinct circumorbitals absent; canthals two; internasals four; postrostrals four, two median ones larger; supralabials six; infralabials five; loreals four; lorilabials in two rows; preoculars three, the middle one tiny, the dorsal-most in contact with posterior canthal; lateral temporals granular; gulars in 47 rows between tympanic openings; all gulars cycloid, smooth, slightly imbricate, not notched; second infralabial in contact with first three sublabials; first pair of postmentals not in contact medially; mental in contact with first pair of infralabials and first pair of postmentals; dorsal scales of neck keeled and lateral scales of neck granular; dorsal scales of body imbricate, keeled, feebly mucronate becoming smaller and slightly keeled to smooth towards flanks; scales around midbody 86; vertebrals enlarged, keeled, imbricate, in 73 rows, forming distinct vertebral row and a low serrate crest from the neck to the middle of dorsum; paravertebrals adjacent to vertebral row same size as dorsals, keeled, and imbricate; paravertebrals 107; ventrals smooth, imbricate, one third larger than dorsals; preauricular fringe short, composed of two enlarged, posteriorly projected scales; antegular, antehumeral, gular, longitudinal, and postauricular neck folds present; ventrolateral fold present; dorsal humeral scales imbricate, weakly keeled; scales of forearms imbricate, keeled; dorsal scales of hindlimbs imbricate, strongly keeled, mucronate; ventral humeral scales imbricate, weakly keeled; ventral scales of forearms and hindlimbs imbricate, smooth; palmars imbricate, weakly keeled; plantars imbricate, smooth; lamellae on Finger IV 24; lamellae on Toe IV 29; tail rounded (tail length 58% of total length); caudals strongly keeled, mucronate, imbricate dorsally; two caudal whorls per autotomic segment; basal subcaudals slightly keeled, imbricate; posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; postfemoral mite pocket distinct with slit-like opening; [Type 2 of Torres-Carvajal (2007b)]. Color in life of holotype (Fig. 5 A-C): dorsum emerald green with whitish and turquoise spots on head and neck, and pale greenish yellow spots on dorsum; pelvic region, hindlimbs and tail pale brown covered by dark gray reticulations and pale greenish yellow spots; throat brownish gray with pale gray spots; ventral surface including base of tail dirty cream and rest of tail darker than the base. Once captured it quickly changes its dorsal color from green to dark brown, sprinkled with dirty cream spots on head and neck, and yellowish cream spots on dorsum; forelimbs, hindlimbs and tail dark grayish brown, darker on tail and with a greenish tone on forelimbs; some dirty cream spots are present on hindlimbs. Iris pale brown. Color in preservative (Fig. 4 D-E): head, forelimbs and dorsum to pelvic region greenish gray, sprinkled with cream spots; anterior surface of thighs greenish gray and the rest of hindlimbs, pelvic region and tail grayish brown with scattered dark gray flecks and reticulations. The last one third of tail is black. The throat is greenish gray without spots, the rest of ventral surface of body grayish cream. Intraspecific variation. Measurements,scutellation, and other morphological characters of Stenocercus leybachi are presented in Table 1. Supralabials 4���6; infralabials 5���6; second infralabials not in contact with third sublabials in sixteen specimens (69.5%); first pair of postmentals in contact medially only in four specimens (17.3%). In two dissected specimens, the rib pattern was three xiphisternal and two long postxiphisternal pairs of inscriptional ribs not in contact midventrally, [Pattern 4A of Torres-Carvajal (2004)]. Females are smaller than males (Table 1) and both sexes are able to change color from green to dark brown or grayish brown (see Fig. 5). The seven adult males in the type series lack a black antehumeral collar and of the seven adult females, one (CORBIDI 16429) possesses a middorsally complete collar. Complete black antehumeral collar is present in eight of nine juveniles, only one specimen (CORBIDI 16431) has an incomplete black collar. Adult specimens of both sexes and juveniles possess pale spots on dorsum and some specimens also have scattered black flecks. The throat in adult females can be brownish green with pale green spots, chest with a greenish tone and the belly is dirty cream with or without a pinkish hue on the belly and base of tail. Juveniles have throat and venter yellowish green. Distribution and natural history. Stenocercus leybachi is known from five close localities in the Amazon foothills of the R��o Huallaga basin, at elevations between 824���1,270 m, in central Peru (Fig. 3). The known localities of this species lie in the Hu��nuco Department within the Yungas (500���2,300 m) ecoregion according to Brack���Egg (1986) and Pe��aherrera del Aguila (1989), and Peruvian Yungas following Olson et al. (2001). The general habitat where the new species was collected is composed of croplands of coca (Erythroxylum coca), coffee (Coffea sp.), cacao (Theobroma cacao), corn (Zea mays), orange (Citrus sp.) and pastures for cattle ranching with scattered patches of secondary montane forest, especially on the steepest slopes. Individuals of S. leybachi were observed active during sunny days basking on tree trunks at 1.5���6 m above ground. When these individuals felt disturbed, they climbed the tree trunk finding refuge up high. Some individuals were observed and collected with a fishing rod while basking on fallen trees. The individuals found basking were light green and at the moment of capture suddenly changed to brown or brownish gray. Most specimens were collected in the forest canopy, during a flood for the construction of a hydroelectric dam. The specimens were collected by hand on the branches of high trees (5 to 10 m) from a boat, and some individuals tried to escape by hiding in tree holes (Fig. 5 K-L) and cracks in the bark of the trees. Stenocercus leybachi was syntopic in the area of the flood with other arboreal lizards such as Anolis punctatus, Euspondylus excelsum, and S. chinchaoensis. Other sympatric species of squamate reptiles collected with S. leybachi were: A. ortonii, Bothrops chloromelas, Dipsas catesbyi, D. indica, D. schunkii, Drymoluber dichrous, Enyalioides feiruzae, Micrurus annelatus, Oxyrhopus leucomelas, Phrynonax polylepis, Proctoporus sp., and S. prionotus. Etymology. The specific name leybachi is a patronym for Achim Leybach of Marktsteft, Germany, in recognition of his financial support for the taxonomic work and biodiversity research through the BIOPAT-Programme., Published as part of Venegas, Pablo J., Garc��a-Ayachi, Luis A., Ch��vez-Arribasplata, Juan C. & Garc��a-Bravo, Antonio, 2022, Four new species of polychromatic spiny-tailed iguanian lizards, genus Stenocercus (Iguania: Tropiduridae), from Peru, pp. 1-28 in Zootaxa 5115 (1) on pages 11-15, DOI: 10.11646/zootaxa.5115.1.1, http://zenodo.org/record/6345982, {"references":["Torres-Carvajal, O. (2007 b) A taxonomic revision of south American Stenocercus (Squamata: Iguania) lizards. Herpetological monographs, 21, 76 - 178. https: // doi. org / 10.1655 / 06 - 001.1","Torres-Carvajal, O. (2004) The abdominal skeleton of tropidurid lizards (Squamata: Tropiduridae). Herpetologica, 60, 75 - 83. https: // doi. org / 10.1655 / 03 - 15","Penaherrera del Aguila, C. (1989) Atlas del Peru. Instituto Geografico Nacional, Lima, 400 pp.","Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N. D., Powell, G. V. N., Underwood, E. C., D'Amico, J. A., Itoua, I., Strand, H. E., Morrison, J. C., Loucks, C. J., Allnutt, T. F., Ricketts, T. H., Kura, Y., Lamoreux, J. F., Wettengel, W. W., Hedao, P. & Kassem, K. R. (2001) Terrestrial ecoregions of the world: A new map of life on earth. BioScience, 51, 933 - 938. https: // doi. org / 10.1641 / 0006 - 3568 (2001) 051 [0933: TEOTWA] 2.0. CO; 2"]}

Stenocercus asenlignus sp. nov. Figures 1���2 & 10 Holotype. CORBIDI 20219, an adult male, from Omia (6��28���12.702���S, 77��23���45.187���W, 1,381 m), Rodr��guez de Mendoza Province, Amazonas Department, Peru, collected by P.J. Venegas and L.A. Garc��a-Ayachi on 5 December 2018. Paratypes (24): PERU: SAN MART��N DEPARTMENT: Mariscal C��ceres Province: MUSM 23034, adult male, MUSM 23035���38, adult females, and MUSM 23039, a juvenile, from El Dorado (06��46���00���S, 77��32���42���W, 1,600 m), collected by P.J. Venegas between 6 and 8 December 2003; CORBIDI 00622 adult female from Los Chilchos (06��42���28.7���S, 77��34���0.9���W, 1,800 m), collected by P.J. Venegas on 28 January 2008; CORBIDI 14780, adult male from R��o Lej��a (06�� 50���11.6���S, 77��29���09.7���W, 1,500 m), collected by M. Salas on 14 June 2012; AMAZONAS DEPARTMENT: Rodr��guez de Mendoza Province: CORBIDI 15658, juvenile, from Omia village (6��27���58.5���S, 77��23���50.9���W, 1,390 m), collected by A. Garc��a-Bravo on 11 November 2014; CORBIDI 20215���16, 20218, and 20221, adult males, CORBIDI 20214, 20217, and 20220, adult females, and CORBIDI 20222, a juvenile male, from Omia (6��28���12.702���S, 77��23���45.187���W, 1,381 m), collected by P.J. Venegas and L.A. Garc��a-Ayachi on 5 December 2018; CORBIDI 20135, adult female, from Santo Toribio village (6��1���55.179���S, 77��19���45.308���W, 1,622 m), collected by P.J. Venegas and A. Garc��a-Ayachi on 11 November 2018; CORBIDI 20198, juvenile female, CORBIDI 20199, juvenile male, and CORBIDI 20200���01, adult females, from Fundo Playas del Inca, Santo Toribio, Vista Alegre (6��1���56.356���S, 77��21���6.385���W, 1,682 m), collected by P.J. Venegas and L.A. Garc��a-Ayachi on 17 November 2018; CORBIDI 23003, a juvenile, from Limabamba (6��37���53.98���S, 77��26���22.33���W, 2,036 m), collected by Ivan Wong on 30 July 2019; HU��NUCO DEPARTMENT: Mara����n Province: CORBIDI 19549, adult female, from Nuevo Cajh��n village, Chol��n (8��52���8.076���S, 76��28���44.688���W, 1,625 m), collected by L.A. Garc��a-Ayachi on 23 June 2017. Diagnosis. From the 76 previously known species Stenocercus and the new species described herein, only S. arndti, S. leybachi sp. nov., S. bolivarensis Castro & Ayala, 1982, S. carrioni Parker, 1934, S. chlorostictus Cadle, 1991, S. crassicaudatus, S. empetrus, S. eunetopsis, S. flagracanthus, S. nigrocaudatus sp. nov., S. qalaywasi sp. nov., S. torquatus and S. simonsii Boulenger, 1901, share with S. asenlignus granular scales on posterior surface of thighs, relatively short tail, caudals spinose, and two caudal whorls per autotomic segment. However, S. torquatus and S. qalaywasi, are easily distinguished from S. asenlignus (state of character between parentheses) by having transverse black nuchal bands (absent), a middorsally complete antehumeral black collar (incomplete) and, only in the case of S. torquatus (Fig. 10A), the dorsum is emerald green without bands (green with black bands in males). Moreover, S. qalaywasi possesses a longer tail with 57���60 % of the total length versus 49���57% in S. asenlignus. Live male individuals of S. nigrocaudatus can be readily distinguished from S. asenlignus by having the tail black with scattered turquoise spots (tail light brown or grayish brown with dark gray spots or bands) and the body without dorsal black bands (present). In addition, in S. asenlignus the dorsal scales of the body are keeled, becoming gradually strongly keeled and mucronate toward the hindlimb insertion and in S. nigrocaudatus dorsal scales are smooth and feebly keeled becoming strongly keeled but not mucronate close to the hindlimbs insertion. Stenocercus leybachi can be readily distinguished from S. asenlignus by having a distinct, low, serrate crest on the neck that can reach or not to the middle of dorsum, while in S. asenlignus, vertebrals form an indistinct row of enlarged scales along the body. Moreover, S. asenlignus has more vertebrals than S. leybachi (71���106, = 89.8 versus 63���73, = 68.6, respectively). Stenocercus torquatus has more scales around midbody than S. asenlignus (102 to 137, = 116.9 versus 78 to 111, = 96.4, respectively). The recently described S. flagracanthus differs from S. asenlignus in having a tail strongly armed with projected mucronate scales that are less developed in S. asenlignus. In the distal half of the tail of S. flagracanthus the spines are conspicuously alternating in size, with large spines followed by small spines per each caudal whorl, and in S. asenlignus, the alternation in size of spines is undistinguishable. Dorsal scales between the posterior half of body and hindlimbs insertion become enlarged and strongly keeled with strongly projected mucronate scales in S. flagracanthus, while in S. asenlignus, they are keeled and, in some specimens, slightly mucronate. Furthermore, the dorsal and gular pattern is different in both species (state of characters in S. flagracanthus between parentheses): usually adult males of S. asenlignus possess a ���zig-zag��� pattern of black dorsal bars on dorsum (bold black bars; see Fig. 11 in Venegas et al. 2020a), antehumeral collar bordered by cream spots (cream line), and preserved specimens have the gular region gray with scattered black flecks (gray with cream dots). The other species such as S. carrioni, S. chlorostictus and S. eunetopsis differ from S. anselignus by having dorsal scales of neck keeled and imbricate (granular in S. asenlignus). Stenocercus eunetopsis has a considerably longer tail than S. asenlignus (64���66% of total length vs. 49���57%, respectively), and S. chlorostictus has a strong sexual dichromatism, with the dorsal background color bright green in males and brown in females (males and females gray, brown or green in S. asenlignus). Stenocercus bolivarensis differs from S. asenlignus by having strongly keeled and imbricate lateral body scales (Torres-Carvajal 2007b), which are granular or smooth in S. asenlignus. Stenocercus asenlignus differs from S. crassicaudatus by having fewer paravertebrals (92 to 118, = 105.8 versus 107 to 166, = 126.6), and a shorter tail (49 to 57% of total length versus 57 to 62%). Males of S. arndti are easily distinguished from S. asenlignus by having a bold black transverse band at midbody that extends ventrolaterally (absent in S. asenlignus) and dorsal scales of neck slightly keeled and subimbricate (granular in S. asenlignus). Stenocercus simonsii differs from S. asenlignus (character states in parentheses) by having dorsals imbricate, moderately keeled, but not mucronate (dorsals granular in the anterior half of body, getting gradually enlarged and keeled from the posterior half of body to strongly keeled and mucronate near to the hindlimbs insertion). Stenocercus empetrus differs from S. asenlignus by having the venter yellowish-orange with black reticulations, or completely black, whereas in the new species the venter is gray without reticulations. In addition, S. simonsii and S. empetrus are larger than S. asenlignus (maximum SVL = 88 mm in males and 79 mm in females of S. simonsii, SVL = 103 mm in males and 90 mm in females of S. empetrus, and maximum SVL = 73 mm in males and 70 mm in females of S. asenlignus). Characterization. (1) Maximum SVL in males 73 mm (n = 9); (2) maximum SVL in females 70 mm (n = 14); (3) vertebrals 71���106; (4) paravertebrals 92���118; (5) scales around midbody 78���111; (6) supraoculars 5���8; (7) internasals 4���5; (8) postrostrals 4���6; (9) loreals 2���6; (10) gulars 47���60; (11) lamellae on Finger IV 23���30; (12) lamellae on Toe IV 28���34; (13) posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; (14) postfemoral mite pocket distinct with slit-like opening [Type 2 of Torres-Carvajal (2007b)]; (15) parietal eye absent; (16) occipital scales small, smooth, juxtaposed; (17) projecting angulate temporal absent; (18) row of enlarged supraoculars occupying most of supraocular region absent; (19) scales on frontonasal region juxtaposed, smooth, not imbricate; (20) preauricular fringe short; (21) antehumeral, gular, longitudinal, oblique, postauricular, supra-auricular, antegular neck folds present; (22) lateral nuchals and dorsals similar in size; (23) lateral body scales smaller than dorsals; (24) vertebrals slightly enlarged, forming a distinct row of scales from forelimbs to hindlimbs; (25) dorsolateral crest absent; (26) paravertebrals and adjacent scales from midbody keeled, becoming strongly keeled and mucronate at the posterior half of body; (27) ventral scales smooth, imbricate; (28) scales on posterior surface of thighs granular; (29) prefemoral fold present; (30) inguinal fold present, weakly defined; (31) preanals not projected; (32) tail not compressed laterally; (33) tail relatively short (tail length 49���57% of total length); (34) caudal whorls per autotomic segment two; (35) tail spinose; (36) postocular stripe absent or present; (37) gular region in males dark gray with black dots; (38) gular region in females grayish yellow with light green dots; (39) black patch on ventral surface of neck in adult males absent; (40) dark midventral stripe in adult males absent; (41) dark patch on ventral surface of thighs, vent and tail in adult males absent; (42) background color of dorsum green, brown or grayish in both males and females, with a ���zig-zag��� pattern of black transversal bands in adult males; (43) two long postxiphisternal pairs of inscriptional ribs not in contact midventrally (pattern 4A of Torres-Carvajal, 2004). Description of holotype. Male (Fig. 1); SVL 69 mm; TL 88 mm; maximum head width 14.61 mm; head length 17.93 mm; head height 11.20 mm; scales on parietal and occipital regions small, smooth, juxtaposed, subequal in size; parietal eye not visible: supraoculars seven, smooth, juxtaposed; circumorbitals absent; canthals two; loreals three; postrostrals four; internasals four; supralabials five; infralabials five; lorilabials in one row; preocular divided into three scales, most dorsal in contact with posterior canthal; lateral temporals granular; gulars in 57 rows between tympanic openings; all gulars cycloid, smooth, imbricate; second infralabial in contact with first, second and third sublabials; first pair of postmentals in contact; mental not separated from the infralabials by the first pair of postmentals; dorsal and lateral scales of neck and body granular; scales around midbody 97; vertebrals 90 enlarged, keeled, imbricate, forming distinct vertebral row; paravertebrals and adjacent scales slightly smaller than vertebral row, keeled, imbricate, becoming mucronate from midbody to hindlimb insertion; paravertebrals 100; ventrals smooth, imbricate, more than twice the size of dorsals, except for paravertebrals at the second half of body, which are nearly equal in size or slightly longer than ventrals; preauricular fringe short, composed of four enlarged, granular scales; suprauricular, antehumeral, gular, longitudinal, oblique, antegular, postauricular and rictal neck folds present; dorsolateral, ventrolateral and prefemoral folds present; dorsal scales of forelimbs imbricate, keeled; dorsal scales of hindlimbs imbricate, strongly keeled and mucronate; ventral humeral scales granular; ventral scales of fore and hindlimbs smooth, imbricate; palmar scales imbricate, keeled; plantar scales imbricate, strongly keeled, mucronate; lamellae on Finger IV 23; lamellae on Toe IV 29; tail rounded (tail length 55% of total length); caudal scales strongly keeled, mucronate, imbricate; basal subcaudal scales smooth, imbricate; posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; postfemoral mite pocket distinct with slit-like opening [Type 2 of Torres-Carvajal (2007b)]. Color in life of holotype: dorsally green (from head to the anterior half of body, including forelimbs) speckled with dark gray flecks and with transverse rows of light green dots; posterior half of body and hindlimbs pale brown with the same pattern of flecks and dots scattered of the green half but the light green dots are faint on hindlimbs; tail light brown with black speckles and black bands on the distal third; incomplete middorsal black collar at antehumeral fold and a dorsal ���zig-zag��� pattern of black bands on vertebral line. Ventrally, the gular region is dark gray with scattered black flecks, chest pale green and venter grayish white. Once captured, it quickly changes its dorsal color from green to brownish gray, with scattered black flecks and cream dots (Fig. 2A). Limbs change from green with black flecks to gray with scattered black flecks and pale cream dots. Distal portion of tail remains light brown with black bands. Iris dark brown. Color in preservative (Fig. 1 D-E): dorsal surface dark gray with scattered black and pale dots, dorsal ���zig-zag��� bands pattern remains evident. Gular region dark gray with black dots and venter gray. Intraspecific variation. Measurements, scutellation, and other morphological characters of Stenocercus asenlignus are presented in Table 1. Supralabials 4���7; infralabials 4���7; second infralabials in contact with third sublabials in all specimens, except in MUSM 23035; first pair of postmentals in contact medially only in the specimen MUSM 23039. In two dissected specimens, the pattern was three xiphisternal and two long postxiphisternal pairs of inscriptional ribs not in contact midventrally (Pattern 4A; Torres-Carvajal 2004). Sexual dimorphism in size is slightly noticeable (Table 1). The head of adult males looks more robust than the females, and both sexes are able to change color from green to brownish gray or bluish gray in males (Fig. 2) and green to dark gray or pale brown in females. Black collar and ���zig-zag��� bands dorsal pattern are distinct only in males (Fig. 2A, C & L) and absent or faint in females (Fig. 2D, F & H) and juveniles (Fig. 2I). In adult females throat can be cream with gray flecks and the chest dirty cream (Fig. 2E) or throat grayish green with light green dots and chest green (Fig. 2G), and the venter is gray with a pink tone (Fig. 2E). Juveniles have the throat yellowish with light yellow flecks (Fig. 2J) or green with light green flecks, and the chest and ventral surface of forelimbs pale green. Some males possess transverse rows of yellow or light green dots on body surface and the throat and chest yellowish when basking (Figs. 2K & 10B). Distribution and natural history observations. Stenocercus asenlignus is known from five localities along the Amazon slope of the northern portion of the Central Andes, along the Huallabamba River basin in the departments of Amazonas and San Mart��n, at elevations between 1500 and 2036 m (Fig. 3). According to Olson et al. (2001), the distribution of S. asenlignus lies inside the Peruvian Yungas ecoregion. At the localities of El Dorado and R��o Lej��a the habitat is characterized by dense primary forests with some coffee and yuca plots. In Los Chilchos the habitat was secondary forest with open areas for pasture. The general landscape in Playa Los Incas was open areas for cattle ranching near the rivers and slopes covered by primary montane forest. Omia is a small village surrounded by crops of coffee (Coffea sp.), cacao (Theobroma cacao) and mango (Mangifera indica). Most specimens were found on tree trunks inside the forest or at the edge of crop areas, between 2 and 3 m above ground; however, several uncollected individuals were observed at heights between 5 and 10 m. A couple of adult individuals were observed on the rocky slope of a cliff at 5 m height. One adult male (MUSM 23034) was collected on a wall of a ���tambo,��� a rural house made of logs. When disturbed it hid in a carpenter bee hole. Hatchlings and juveniles were observed basking on tambo walls and fallen trees around crops and near the base of large trees. In Omia village Stenocercus asenlignus is a common species and several individuals were observed basking on house walls and using the holes and crevices between bricks as retreats (Fig. 2L). Individuals observed basking on trees and house walls were found with green dorsum and at the moment of capture this changed abruptly to dark gray or brownish gray color. However, individuals observed on trees and rock walls in the shadow were also found with a grayish brown dorsum. Female CORBIDI 0622 (SVL = 61) collected in the rainy season (January 2008) had 2 and 4 follicles, in the left and right ovary, respectively. Female MUSM 23038 (SVL = 62 mm) had two underdeveloped eggs (pale yellow) that were 9.6���10.3 mm long, 8��� 7.8 mm wide, and 156.82���172.57 mm 3 in volume. Female MUSM 23036 (SVL = 70 mm) had two fully developed eggs with white, hard shell (22���22.1 mm long, 8.5���8.6 mm wide and 393.42���396.25 mm 3 of volume); it was found digging a hole in a cavity between roots, this behavior suggests that it was building a nest to deposit the eggs. Also, the presence of hatchlings and juveniles around buttress roots suggests that S. asenlignus deposits their eggs between them. Both gravid females (MUSM 23036, 23038) and a juvenile (MUSM 23039, SVL = 37) were collected at the beginning of the rainy season (December, 2003). Stenocercus asenlignus is not sympatric with other Stenocercus or Tropiduridae species. The only known lizard species sympatric with S. asenlignus are Alopoglossus buckleyi, Anolis fuscoauratus, Cercosaura doanae, and Enyalioides sophiarothschildae. Etymology. The specific epithet asenlignus is a noun in apposition and derives from the Latin words ��� asen ��� (= climb or ascend) and ��� lignus ��� (= trunk). It refers to the arboreal habitus of the new species. Remarks. Stenocercus crassicaudatus was reported by Fritts (1974) for San Mart��n and Loreto departments, based on a neonate (AMNH 57176) and a subadult male (AMNH 57173), respectively, that were examined later by Torres-Carvajal et al. (2005). According to Torres-Carvajal et al. (2005), both specimens are more similar morphologically to S. torquatus; they concluded that S. crassicaudatus is restricted to Cusco Department. While in the absence of a major sample from Loreto and San Mart��n, they also concluded that S. torquatus is restricted to the Chanchamayo Valley and adjacent areas in Jun��n and Pasco departments in Central Peru (Torres-Carvajal et al. 2005). Although we did not review the specimens reported by Fritts (1974), we cons, Published as part of Venegas, Pablo J., Garc��a-Ayachi, Luis A., Ch��vez-Arribasplata, Juan C. & Garc��a-Bravo, Antonio, 2022, Four new species of polychromatic spiny-tailed iguanian lizards, genus Stenocercus (Iguania: Tropiduridae), from Peru, pp. 1-28 in Zootaxa 5115 (1) on pages 5-10, DOI: 10.11646/zootaxa.5115.1.1, http://zenodo.org/record/6345982, {"references":["Cadle, J. E. (1991) Systematics of lizards of the genus Stenocercus (Iguania: Tropiduridae) from northern Peru: New species and comments on relationships and distribution patterns. Proceedings of the Academy of Natural Sciences of Philadelphia, 143, 1 - 96.","Venegas, P. J., Garcia-Ayachi, L. A., Chavez-Arribasplata, J. C., Chavez, G., Wong, I. & Garcia-Bravo, A. (2020 a) Four new species of Stenocercus Dumeril & Bibron, 1837 (Squamata, Iguania) from the Department of Amazonas in northeastern Peru. Evolutionary Systematics, 4, 79 - 108. https: // doi. org / 10.3897 / evolsyst. 4.57578","Torres-Carvajal, O. (2007 b) A taxonomic revision of south American Stenocercus (Squamata: Iguania) lizards. Herpetological monographs, 21, 76 - 178. https: // doi. org / 10.1655 / 06 - 001.1","Torres-Carvajal, O. (2004) The abdominal skeleton of tropidurid lizards (Squamata: Tropiduridae). Herpetologica, 60, 75 - 83. https: // doi. org / 10.1655 / 03 - 15","Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N. D., Powell, G. V. N., Underwood, E. C., D'Amico, J. A., Itoua, I., Strand, H. E., Morrison, J. C., Loucks, C. J., Allnutt, T. F., Ricketts, T. H., Kura, Y., Lamoreux, J. F., Wettengel, W. W., Hedao, P. & Kassem, K. R. (2001) Terrestrial ecoregions of the world: A new map of life on earth. BioScience, 51, 933 - 938. https: // doi. org / 10.1641 / 0006 - 3568 (2001) 051 [0933: TEOTWA] 2.0. CO; 2","Fritts, T. H. (1974) A multivariate evolutionary analysis of the Andean iguanid lizards of the genus Stenocercus. San Diego Society of Natural History, 7, 1 - 86.","Torres-Carvajal, O., Lehr, E. & Lundberg, M. (2005) Resurrection of Stenocercus torquatus Boulenger, a spiny-tailed iguanid lizard (Squamata: Iguania) from Peru. Herpetologica, 61, 440 - 448. https: // doi. org / 10.1655 / 05 - 15.1"]}

Stenocercus nigrocaudatus sp. nov. Figures 6–7 Holotype. CORBIDI 19768, an adult male from San Lorenzo, San José de Lourdes (5º4’33.16”S, 78º52’53.749”W, 1,892 m), San Ignacio Province, Cajamarca Department, Peru, collected by P.J. Venegas on 5 September 2018. Paratypes (8): CAJAMARCA DEPARTMENT: San Ignacio Province: CORBIDI 4387, an adult male from Tabaconas (5°14’8.16”S, 79°5’56.58”W, 1,716 m) collected by Maik Dobiey in August 2008; CORBIDI 4388-90, adult females, from Namballe (5°11’40.139”S, 79°4’49.738”W, 1,716 m) collected by M. Dobiey in August 2008; CORBIDI 19765-66, adult females, and CORBIDI 19767 and 19769, adult males, from San Lorenzo, San José de Lourdes (5°4’33.16”S, 78°52’53.749”W, 1,892 m), collected by P.J. Venegas on 5 September 2018. Diagnosis. Among the 80 species of Stenocercus (including the four species described herein), S. nigrocaudatus sp. nov. is only similar to species with granular scales on posterior surface of thighs, relatively short tail, caudal scales spinose, and two caudal whorls per autotomic segment; such as S. asenlignus sp. nov., S. arndti, S. qalaywasi sp. nov., S. flagracanthus, S. leybachi sp. nov., S. bolivarensis, S. carrioni, S. chlorostictus, S. crassicaudatus, S. empetrus, S. eunetopsis, S. torquatus and S. simonsii. Adult males of S. nigrocaudatus can be easily distinguished from the aforementioned species by a contrasting black tail with scattered turquoise flecks (Fig. 7C). Moreover, S. nigrocaudatus can be readily distinguished from S. qalaywasi, S. leybachi, S. bolivarensis, S. carrioni, S. chlorostictus, and S. eunetopsis by having granular scales on dorsal surface of neck (keeled in the compared species).Furthermore, S. leybachi possesses a distinct serrate low crest on neck (absent in S. nigrocaudatus), S. bolivarensis strongly keeled scales on flanks (granular), S. qalaywasi a distinct complete middorsally black antehumeral collar and two black nuchal bands (incomplete and absent, respectively), S. carrioni and S. chlorostictus by having strongly keeled dorsal scales on body (smooth and feebly keeled), and S. eunetopsis by having a longer tail with 62–66% of total length (50–55%). In addition, S. nigrocaudatus possesses more vertebrals (96–108) than S. carrioni (55–72), S. chlorostictus (63–73), and S. eunetopsis (59–80). Stenocercus nigrocaudatus shares with S. asenlignus, S. arndti, S. flagracanthus, S. crassicaudatus, S. empetrus, S. torquatus, and S. simonsii the presence of granular scales on neck. Nevertheless, the new species can be differentiated from these species by the coloration pattern, number of scales, or the length of tail. Adult males of S. arndti are easily distinguished from S. nigrocaudatus (state of character in parentheses) by having a bold black transverse band at midbody that extends ventrolaterally (bands on body absent); adult males of S. empetrus have the belly black (cream); S. simonsii has distinct black bands on body in both sexes (absent). Additionally, S. nigrocaudatus has a greater number of vertebrals (96–108) than S. arndti (74–94), more subdigitals (26–31) than S. empetrus (17–24), and a shorter tail in relation to the total length (50–55%) than S. simonsii (57–63%). Stenocercus asenlignus and S. flagracanthus differ from S. nigrocaudatus in having dorsal scales on the posterior half of body strongly keeled and mucronate (feebly keeled in S. nigrocaudatus). Moreover, adult males of S. asenlignus and S. flagracanthus have black bands on the body (absent in S. nigrocaudatus). Stenocercus crassicaudatus has a longer tail (57–62% of total length) than S. nigrocaudatus (50–55%). Adult males of S. torquatus differ from S. nigrocaudatus in having the antehumeral black collar complete middorsally and two distinct black nuchal transverse bands, whereas the new species has the antehumeral collar incomplete middorsally and lacks nuchal bands. Characterization. (1) Maximum SVL in males 72 mm (n = 4); (2) maximum SVL in females 76 mm (n = 5); (3) vertebrals 96–108; (4) paravertebrals 110–128; (5) scales around midbody 99–117; (6) supraoculars 5–7; (7) internasals 4–5; (8) postrostrals 5–6; (9) loreals 3–7; (10) gulars 48–59; (11) lamellae on Finger IV 26–31; (12) lamellae on Toe IV 30–36; (13) posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; (14) postfemoral mite pocket distinct with slit-like opening [Type 2 of Torres-Carvajal (2007b)]; (15) parietal eye absent; (16) occipital scales small, smooth, juxtaposed; (17) projecting angulate temporal absent; (18) row of enlarged supraoculars occupying most of supraocular region absent; (19) scales on frontonasal region juxtaposed and smooth, not imbricate; (20) preauricular fringe short; (21) antegular, antehumeral, gular, longitudinal, oblique, postauricular, and supra-auricular neck folds present; (22) lateral nuchals and dorsals similar in size; (23) lateral body scales smaller than dorsals; (24) vertebrals enlarged, forming a distinct row of scales from forelimbs to hindlimbs; (25) dorsolateral crest absent; (26) paravertebrals and adjacent scales from midbody, smooth or feebly keeled, becoming keeled on the second half of body and mucronate between hindlimbs; (27) ventral scales smooth, imbricate; (28) scales on posterior surface of thighs granular; (29) prefemoral fold present; (30) inguinal fold present; (31) preanals not projected; (32) tail not compressed laterally; (33) tail relatively short (tail length 50–55% of total length); (34) caudal whorls per autotomic segment two; (35) tail spinose; (36) postocular stripe present; (37) dark patch on gular region in males absent; (38) dark patch on gular region in females absent; (39) black patch on ventral surface of neck in adult males absent; (40) dark midventral stripe in adult males absent; (41) dark patch on ventral surface of thighs, vent and tail in adult males absent; (42) background color of dorsum green, dark brown or olive green in both sexes; (43) postxiphisternal inscriptional ribs not examined. Description of holotype. Male (Fig. 6); SVL 57 mm; TL 114 mm; maximum head width 12.3 mm; head length 16.3 mm; head height 10.1 mm; scales on parietal and occipital regions small, smooth, juxtaposed, subequal in size; parietal eye not visible: supraoculars six, smooth, juxtaposed; circumorbitals absent; canthals two; loreals one; postrostrals four; internasals four; supralabials five; infralabials five; lorilabials in one row; preocular divided into two scales, most dorsal in contact with posterior canthal; lateral temporals granular; gulars in 55 rows between tympanic openings; all gulars cycloid, smooth, imbricate; second infralabial in contact with first, second and third sublabials; first pair of postmentals in contact; mental not separated from the infralabials by the first pair of postmentals; dorsal and lateral scales of neck and body granular; scales around midbody 107; vertebrals 104 moderately enlarged, keeled, imbricate, forming distinct vertebral row; paravertebrals and adjacent scales slightly smaller than vertebrals, smooth, subimbricate, becoming feebly keeled from the posterior half of dorsum and strongly keeled near hindlimbs insertion; paravertebrals 127; scales on flanks granulars; ventrals smooth, imbricate, more than twice the size of dorsals, only paravertebrals on the second half of body, nearly equal in size or slightly longer than ventrals; preauricular fringe short composed of three enlarged scales with round edge; suprauricular, antehumeral, gular, longitudinal, oblique, antegular, postauricular and rictal neck folds present; dorsolateral, ventrolateral and prefemoral folds present; dorsal scales of forelimbs imbricate, keeled; dorsal scales of hindlimbs imbricate, strongly keeled and mucronate; ventral humeral scales granular; ventral scales of fore- and hindlimbs smooth, imbricate; palmar scales imbricate, keeled and minutely mucronate; plantar scales imbricate, smooth or strongly keeled and mucronate; lamellae on Finger IV 28; lamellae on Toe IV 35; tail rounded (tail length 50% of total length); caudal scales strongly keeled, mucronate, imbricate; basal subcaudal scales smooth, imbricate; posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; postfemoral mite pocket distinct with slit-like opening [Type 2 of Torres-Carvajal (2007b)]. Color in life (Fig. 7 A-C): head and neck brownish gray with white flecks on head and white dots on neck; labials white with dark brown bars; postocular stripes dark brown, extending through the temporal and edge of occipital region to a black rhomboid blotch in the nuchal region; antehumeral collar black, distinct; dorsal surface of body green covered by bright green dots; forelimbs green with scattered bright green dots; hindlimbs brown with scattered white flecks and dots on posterior surface of thighs; tail black with scattered turquoise flecks on the basal half. Ventrally, gular region grayish white with white dots and black flecks; neck gray with few scattered black flecks; chest and belly cream with an orange hue on the posterior half; thighs, cloacal region, and tail dull orange. Iris dark brown. Color in ethanol 70% (Fig. 6 D-E): the dorsal surface turns dark brownish gray; postocular stripes and the nuchal blotch remain black; the white dots on the neck and the scattered white flecks on hindlimbs are dirty white; the bright green dots on the body and forelimbs are pale gray; tail turns dark brown. Ventrally, the gular region and neck is pale gray with dirty white dots and scattered black flecks, and chest, belly, limbs and base of tail are grayish white; the two distal thirds of the tail are dark brown with grayish white bars. Intraspecific variation. Measurements and scutellation of Stenocercus nigrocaudatus are presented in Table 1. Supralabials 4–5; infralabials 5–6; no specimens showed the second infralabials in contact with third sublabial; first pair of postmentals in contact medially in eight specimens (88.8%). Females larger than males (see Table 1), and both sexes are able to change color from green to dark brown or grayish brown. Sexual dimorphism in coloration is evident in the tail, which is black with scattered turquoise speckles in males (Fig. 7C) and brown with black and gray speckles in the proximal half and dark brown with gray rings in the distal half in females (Fig. 7J). The antehumeral collar is bold black in males and faint black or dark gray in females. Ventrally, the gular region in females is gray with scattered black flecks and speckles, the chest has a greenish hue, the belly is grayish white, and the ventral surface of hindlimbs, cloacal region and tail are tannish cream (Fig. 7I). The single female juvenile specimen (CORBIDI 19766) is dorsally identical to the adult female, but the gular region and chest possess a green hue. Distribution and natural history. Stenocercus nigrocaudatus is known from both sides of the inter-Andean valley of the Chinchipe River in the Cajamarca Department of northeastern Peru, between elevations of 1,700 and 1,892 m (Fig. 3). The two known localities of this species lie at the Yungas (500–2,300 m) ecoregion according to Brack-Egg (1986) and Peñaherrera del Águila (1989), and Eastern Cordillera Real montane forest following Olson et al. (2001). The general landscape in Alto Ihuamaca is composed of humid montane forest covering the steepest slopes and open areas with croplands for corn (Zea mays), coffee (Coffea sp.), and pastures for cattle ranching. In San José de Lourdes, the general landscape includes pastures for cattle ranching and small croplands for coffee, mango (Mangifera indica), and oranges (Citrus sp.), with some scattered patches of secondary forest and fringes of forest bordering ravines and along streams. Several individuals of S. nigrocaudatus were observed basking during sunny days between 900 and 1,500 hours. In small villages and single houses in the field, we observed individuals of S. nigrocaudatus basking on the walls at heights between 2 and 3 m (Fig. 7L), and one individual was observed at 7 m in the wall of a small building. This species also was common in open areas for cattle ranching basking on big tree trunks between 1 and 8 m above ground (Fig. 7K). Up to five individuals, a pair of adults and three juveniles, were observed in the same tree. All individuals observed basking showed a green dorsal coloration that changed to brownish gray to the moment of be captured. Sympatric squamate reptiles collected with S. nigrocaudatus were Alopoglossus buckleyi, Atractus gigas, Enyalioides anisolepis, and Potamites strangulatus; and in San José de Lourdes only Varzea altamazonica. Etymology. The specific epithet nigrocaudatus is a noun in apposition that derives from the Latin word “ nigrum ” that means black and “cauda” that means tail, with the suffix “ tus ” that means provided with. It refers to the black tail that is characteristic of the adult males of this new species.

Stenocercus asenlignus sp. nov. Figures 1–2 & 10 Holotype. CORBIDI 20219, an adult male, from Omia (6°28’12.702”S, 77°23’45.187”W, 1,381 m), Rodríguez de Mendoza Province, Amazonas Department, Peru, collected by P.J. Venegas and L.A. García-Ayachi on 5 December 2018. Paratypes (24): PERU: SAN MARTÍN DEPARTMENT: Mariscal Cáceres Province: MUSM 23034, adult male, MUSM 23035–38, adult females, and MUSM 23039, a juvenile, from El Dorado (06°46’00”S, 77°32’42”W, 1,600 m), collected by P.J. Venegas between 6 and 8 December 2003; CORBIDI 00622 adult female from Los Chilchos (06°42’28.7”S, 77°34’0.9”W, 1,800 m), collected by P.J. Venegas on 28 January 2008; CORBIDI 14780, adult male from Río Lejía (06° 50’11.6”S, 77°29’09.7”W, 1,500 m), collected by M. Salas on 14 June 2012; AMAZONAS DEPARTMENT: Rodríguez de Mendoza Province: CORBIDI 15658, juvenile, from Omia village (6°27’58.5”S, 77°23’50.9”W, 1,390 m), collected by A. García-Bravo on 11 November 2014; CORBIDI 20215–16, 20218, and 20221, adult males, CORBIDI 20214, 20217, and 20220, adult females, and CORBIDI 20222, a juvenile male, from Omia (6°28’12.702”S, 77°23’45.187”W, 1,381 m), collected by P.J. Venegas and L.A. García-Ayachi on 5 December 2018; CORBIDI 20135, adult female, from Santo Toribio village (6°1’55.179”S, 77°19’45.308”W, 1,622 m), collected by P.J. Venegas and A. García-Ayachi on 11 November 2018; CORBIDI 20198, juvenile female, CORBIDI 20199, juvenile male, and CORBIDI 20200–01, adult females, from Fundo Playas del Inca, Santo Toribio, Vista Alegre (6°1’56.356”S, 77°21’6.385”W, 1,682 m), collected by P.J. Venegas and L.A. García-Ayachi on 17 November 2018; CORBIDI 23003, a juvenile, from Limabamba (6°37’53.98”S, 77°26’22.33”W, 2,036 m), collected by Ivan Wong on 30 July 2019; HUÁNUCO DEPARTMENT: Marañón Province: CORBIDI 19549, adult female, from Nuevo Cajhán village, Cholón (8°52’8.076”S, 76°28’44.688”W, 1,625 m), collected by L.A. García-Ayachi on 23 June 2017. Diagnosis. From the 76 previously known species Stenocercus and the new species described herein, only S. arndti, S. leybachi sp. nov., S. bolivarensis Castro & Ayala, 1982, S. carrioni Parker, 1934, S. chlorostictus Cadle, 1991, S. crassicaudatus, S. empetrus, S. eunetopsis, S. flagracanthus, S. nigrocaudatus sp. nov., S. qalaywasi sp. nov., S. torquatus and S. simonsii Boulenger, 1901, share with S. asenlignus granular scales on posterior surface of thighs, relatively short tail, caudals spinose, and two caudal whorls per autotomic segment. However, S. torquatus and S. qalaywasi, are easily distinguished from S. asenlignus (state of character between parentheses) by having transverse black nuchal bands (absent), a middorsally complete antehumeral black collar (incomplete) and, only in the case of S. torquatus (Fig. 10A), the dorsum is emerald green without bands (green with black bands in males). Moreover, S. qalaywasi possesses a longer tail with 57–60 % of the total length versus 49–57% in S. asenlignus. Live male individuals of S. nigrocaudatus can be readily distinguished from S. asenlignus by having the tail black with scattered turquoise spots (tail light brown or grayish brown with dark gray spots or bands) and the body without dorsal black bands (present). In addition, in S. asenlignus the dorsal scales of the body are keeled, becoming gradually strongly keeled and mucronate toward the hindlimb insertion and in S. nigrocaudatus dorsal scales are smooth and feebly keeled becoming strongly keeled but not mucronate close to the hindlimbs insertion. Stenocercus leybachi can be readily distinguished from S. asenlignus by having a distinct, low, serrate crest on the neck that can reach or not to the middle of dorsum, while in S. asenlignus, vertebrals form an indistinct row of enlarged scales along the body. Moreover, S. asenlignus has more vertebrals than S. leybachi (71–106, = 89.8 versus 63–73, = 68.6, respectively). Stenocercus torquatus has more scales around midbody than S. asenlignus (102 to 137, = 116.9 versus 78 to 111, = 96.4, respectively). The recently described S. flagracanthus differs from S. asenlignus in having a tail strongly armed with projected mucronate scales that are less developed in S. asenlignus. In the distal half of the tail of S. flagracanthus the spines are conspicuously alternating in size, with large spines followed by small spines per each caudal whorl, and in S. asenlignus, the alternation in size of spines is undistinguishable. Dorsal scales between the posterior half of body and hindlimbs insertion become enlarged and strongly keeled with strongly projected mucronate scales in S. flagracanthus, while in S. asenlignus, they are keeled and, in some specimens, slightly mucronate. Furthermore, the dorsal and gular pattern is different in both species (state of characters in S. flagracanthus between parentheses): usually adult males of S. asenlignus possess a “zig-zag” pattern of black dorsal bars on dorsum (bold black bars; see Fig. 11 in Venegas et al. 2020a), antehumeral collar bordered by cream spots (cream line), and preserved specimens have the gular region gray with scattered black flecks (gray with cream dots). The other species such as S. carrioni, S. chlorostictus and S. eunetopsis differ from S. anselignus by having dorsal scales of neck keeled and imbricate (granular in S. asenlignus). Stenocercus eunetopsis has a considerably longer tail than S. asenlignus (64–66% of total length vs. 49–57%, respectively), and S. chlorostictus has a strong sexual dichromatism, with the dorsal background color bright green in males and brown in females (males and females gray, brown or green in S. asenlignus). Stenocercus bolivarensis differs from S. asenlignus by having strongly keeled and imbricate lateral body scales (Torres-Carvajal 2007b), which are granular or smooth in S. asenlignus. Stenocercus asenlignus differs from S. crassicaudatus by having fewer paravertebrals (92 to 118, = 105.8 versus 107 to 166, = 126.6), and a shorter tail (49 to 57% of total length versus 57 to 62%). Males of S. arndti are easily distinguished from S. asenlignus by having a bold black transverse band at midbody that extends ventrolaterally (absent in S. asenlignus) and dorsal scales of neck slightly keeled and subimbricate (granular in S. asenlignus). Stenocercus simonsii differs from S. asenlignus (character states in parentheses) by having dorsals imbricate, moderately keeled, but not mucronate (dorsals granular in the anterior half of body, getting gradually enlarged and keeled from the posterior half of body to strongly keeled and mucronate near to the hindlimbs insertion). Stenocercus empetrus differs from S. asenlignus by having the venter yellowish-orange with black reticulations, or completely black, whereas in the new species the venter is gray without reticulations. In addition, S. simonsii and S. empetrus are larger than S. asenlignus (maximum SVL = 88 mm in males and 79 mm in females of S. simonsii, SVL = 103 mm in males and 90 mm in females of S. empetrus, and maximum SVL = 73 mm in males and 70 mm in females of S. asenlignus). Characterization. (1) Maximum SVL in males 73 mm (n = 9); (2) maximum SVL in females 70 mm (n = 14); (3) vertebrals 71–106; (4) paravertebrals 92–118; (5) scales around midbody 78–111; (6) supraoculars 5–8; (7) internasals 4–5; (8) postrostrals 4–6; (9) loreals 2–6; (10) gulars 47–60; (11) lamellae on Finger IV 23–30; (12) lamellae on Toe IV 28–34; (13) posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; (14) postfemoral mite pocket distinct with slit-like opening [Type 2 of Torres-Carvajal (2007b)]; (15) parietal eye absent; (16) occipital scales small, smooth, juxtaposed; (17) projecting angulate temporal absent; (18) row of enlarged supraoculars occupying most of supraocular region absent; (19) scales on frontonasal region juxtaposed, smooth, not imbricate; (20) preauricular fringe short; (21) antehumeral, gular, longitudinal, oblique, postauricular, supra-auricular, antegular neck folds present; (22) lateral nuchals and dorsals similar in size; (23) lateral body scales smaller than dorsals; (24) vertebrals slightly enlarged, forming a distinct row of scales from forelimbs to hindlimbs; (25) dorsolateral crest absent; (26) paravertebrals and adjacent scales from midbody keeled, becoming strongly keeled and mucronate at the posterior half of body; (27) ventral scales smooth, imbricate; (28) scales on posterior surface of thighs granular; (29) prefemoral fold present; (30) inguinal fold present, weakly defined; (31) preanals not projected; (32) tail not compressed laterally; (33) tail relatively short (tail length 49–57% of total length); (34) caudal whorls per autotomic segment two; (35) tail spinose; (36) postocular stripe absent or present; (37) gular region in males dark gray with black dots; (38) gular region in females grayish yellow with light green dots; (39) black patch on ventral surface of neck in adult males absent; (40) dark midventral stripe in adult males absent; (41) dark patch on ventral surface of thighs, vent and tail in adult males absent; (42) background color of dorsum green, brown or grayish in both males and females, with a “zig-zag” pattern of black transversal bands in adult males; (43) two long postxiphisternal pairs of inscriptional ribs not in contact midventrally (pattern 4A of Torres-Carvajal, 2004). Description of holotype. Male (Fig. 1); SVL 69 mm; TL 88 mm; maximum head width 14.61 mm; head length 17.93 mm; head height 11.20 mm; scales on parietal and occipital regions small, smooth, juxtaposed, subequal in size; parietal eye not visible: supraoculars seven, smooth, juxtaposed; circumorbitals absent; canthals two; loreals three; postrostrals four; internasals four; supralabials five; infralabials five; lorilabials in one row; preocular divided into three scales, most dorsal in contact with posterior canthal; lateral temporals granular; gulars in 57 rows between tympanic openings; all gulars cycloid, smooth, imbricate; second infralabial in contact with first, second and third sublabials; first pair of postmentals in contact; mental not separated from the infralabials by the first pair of postmentals; dorsal and lateral scales of neck and body granular; scales around midbody 97; vertebrals 90 enlarged, keeled, imbricate, forming distinct vertebral row; paravertebrals and adjacent scales slightly smaller than vertebral row, keeled, imbricate, becoming mucronate from midbody to hindlimb insertion; paravertebrals 100; ventrals smooth, imbricate, more than twice the size of dorsals, except for paravertebrals at the second half of body, which are nearly equal in size or slightly longer than ventrals; preauricular fringe short, composed of four enlarged, granular scales; suprauricular, antehumeral, gular, longitudinal, oblique, antegular, postauricular and rictal neck folds present; dorsolateral, ventrolateral and prefemoral folds present; dorsal scales of forelimbs imbricate, keeled; dorsal scales of hindlimbs imbricate, strongly keeled and mucronate; ventral humeral scales granular; ventral scales of fore and hindlimbs smooth, imbricate; palmar scales imbricate, keeled; plantar scales imbricate, strongly keeled, mucronate; lamellae on Finger IV 23; lamellae on Toe IV 29; tail rounded (tail length 55% of total length); caudal scales strongly keeled, mucronate, imbricate; basal subcaudal scales smooth, imbricate; posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; postfemoral mite pocket distinct with slit-like opening [Type 2 of Torres-Carvajal (2007b)]. Color in life of holotype: dorsally green (from head to the anterior half of body, including forelimbs) speckled with dark gray flecks and with transverse rows of light green dots; posterior half of body and hindlimbs pale brown with the same pattern of flecks and dots scattered of the green half but the light green dots are faint on hindlimbs; tail light brown with black speckles and black bands on the distal third; incomplete middorsal black collar at antehumeral fold and a dorsal “zig-zag” pattern of black bands on vertebral line. Ventrally, the gular region is dark gray with scattered black flecks, chest pale green and venter grayish white. Once captured, it quickly changes its dorsal color from green to brownish gray, with scattered black flecks and cream dots (Fig. 2A). Limbs change from green with black flecks to gray with scattered black flecks and pale cream dots. Distal portion of tail remains light brown with black bands. Iris dark brown. Color in preservative (Fig. 1 D-E): dorsal surface dark gray with scattered black and pale dots, dorsal “zig-zag” bands pattern remains evident. Gular region dark gray with black dots and venter gray. Intraspecific variation. Measurements, scutellation, and other morphological characters of Stenocercus asenlignus are presented in Table 1. Supralabials 4–7; infralabials 4–7; second infralabials in contact with third sublabials in all specimens, except in MUSM 23035; first pair of postmentals in contact medially only in the specimen MUSM 23039. In two dissected specimens, the pattern was three xiphisternal and two long postxiphisternal pairs of inscriptional ribs not in contact midventrally (Pattern 4A; Torres-Carvajal 2004). Sexual dimorphism in size is slightly noticeable (Table 1). The head of adult males looks more robust than the females, and both sexes are able to change color from green to brownish gray or bluish gray in males (Fig. 2) and green to dark gray or pale brown in females. Black collar and “zig-zag” bands dorsal pattern are distinct only in males (Fig. 2A, C & L) and absent or faint in females (Fig. 2D, F & H) and juveniles (Fig. 2I). In adult females throat can be cream with gray flecks and the chest dirty cream (Fig. 2E) or throat grayish green with light green dots and chest green (Fig. 2G), and the venter is gray with a pink tone (Fig. 2E). Juveniles have the throat yellowish with light yellow flecks (Fig. 2J) or green with light green flecks, and the chest and ventral surface of forelimbs pale green. Some males possess transverse rows of yellow or light green dots on body surface and the throat and chest yellowish when basking (Figs. 2K & 10B). Distribution and natural history observations. Stenocercus asenlignus is known from five localities along the Amazon slope of the northern portion of the Central Andes, along the Huallabamba River basin in the departments of Amazonas and San Martín, at elevations between 1500 and 2036 m (Fig. 3). According to Olson et al. (2001), the distribution of S. asenlignus lies inside the Peruvian Yungas ecoregion. At the localities of El Dorado and Río Lejía the habitat is characterized by dense primary forests with some coffee and yuca plots. In Los Chilchos the habitat was secondary forest with open areas for pasture. The general landscape in Playa Los Incas was open areas for cattle ranching near the rivers and slopes covered by primary montane forest. Omia is a small village surrounded by crops of coffee (Coffea sp.), cacao (Theobroma cacao) and mango (Mangifera indica). Most specimens were found on tree trunks inside the forest or at the edge of crop areas, between 2 and 3 m above ground; however, several uncollected individuals were observed at heights between 5 and 10 m. A couple of adult individuals were observed on the rocky slope of a cliff at 5 m height. One adult male (MUSM 23034) was collected on a wall of a “tambo,” a rural house made of logs. When disturbed it hid in a carpenter bee hole. Hatchlings and juveniles were observed basking on tambo walls and fallen trees around crops and near the base of large trees. In Omia village Stenocercus asenlignus is a common species and several individuals were observed basking on house walls and using the holes and crevices between bricks as retreats (Fig. 2L). Individuals observed basking on trees and house walls were found with green dorsum and at the moment of capture this changed abruptly to dark gray or brownish gray color. However, individuals observed on trees and rock walls in the shadow were also found with a grayish brown dorsum. Female CORBIDI 0622 (SVL = 61) collected in the rainy season (January 2008) had 2 and 4 follicles, in the left and right ovary, respectively. Female MUSM 23038 (SVL = 62 mm) had two underdeveloped eggs (pale yellow) that were 9.6–10.3 mm long, 8– 7.8 mm wide, and 156.82–172.57 mm 3 in volume. Female MUSM 23036 (SVL = 70 mm) had two fully developed eggs with white, hard shell (22–22.1 mm long, 8.5–8.6 mm wide and 393.42–396.25 mm 3 of volume); it was found digging a hole in a cavity between roots, this behavior suggests that it was building a nest to deposit the eggs. Also, the presence of hatchlings and juveniles around buttress roots suggests that S. asenlignus deposits their eggs between them. Both gravid females (MUSM 23036, 23038) and a juvenile (MUSM 23039, SVL = 37) were collected at the beginning of the rainy season (December, 2003). Stenocercus asenlignus is not sympatric with other Stenocercus or Tropiduridae species. The only known lizard species sympatric with S. asenlignus are Alopoglossus buckleyi, Anolis fuscoauratus, Cercosaura doanae, and Enyalioides sophiarothschildae. Etymology. The specific epithet asenlignus is a noun in apposition and derives from the Latin words “ asen ” (= climb or ascend) and “ lignus ” (= trunk). It refers to the arboreal habitus of the new species. Remarks. Stenocercus crassicaudatus was reported by Fritts (1974) for San Martín and Loreto departments, based on a neonate (AMNH 57176) and a subadult male (AMNH 57173), respectively, that were examined later by Torres-Carvajal et al. (2005). According to Torres-Carvajal et al. (2005), both specimens are more similar morphologically to S. torquatus; they concluded that S. crassicaudatus is restricted to Cusco Department. While in the absence of a major sample from Loreto and San Martín, they also concluded that S. torquatus is restricted to the Chanchamayo Valley and adjacent areas in Junín and Pasco departments in Central Peru (Torres-Carvajal et al. 2005). Although we did not review the specimens reported by Fritts (1974), we cons

Stenocercus qalaywasi sp. nov. Figures 8–9 Holotype. CORBIDI 20567, adult male from Lampa Pariahuanca, Pariahuanca district, Huancayo province, Junín Department, Peru (11°58’51.9”S, 74°53’47.6”O, 2,587 m), collected by Luis A. García-Ayachi on 19 June 2019. Paratype. CORBIDI 20568, an adult female collected with the holotype. Diagnosis. From all currently known species of Stenocercus (including the new species described herein), S. qalaywasi sp. nov. is only similar to thirteen species (e.g., S. asenlignus sp. nov., S. arndti, S. flagracanthus, S. leybachi sp. nov., S. bolivarensis, S. carrioni, S. chlorostictus, S. crassicaudatus, S. empetrus, S. eunetopsis, S. nigrocaudatus sp. nov., S. torquatus and S. simonsii) in sharing the following characters: granular scales on posterior surface of thighs, relatively short tail, caudals spinose, and two caudal whorls per autotomic segment. From these species, S. qalaywasi possesses a smaller number of vertebrals than S. nigrocaudatus and S. torquatus (77–79 in S. qalaywasi versus 96–108 in S. nigrocaudatus and 83–115 in S. torquatus), scales on dorsal surface of neck keeled (granular in S. asenlignus, S. nigrocaudatus and S. torquatus), a longer tail than S. asenlignus, S. nigrocaudatus and S. torquatus (tail length 57–60% of total length in S. qalaywasi, 51–55% in S. asenlignus, 50–55% in S. nigrocaudatus, and 47–54% in S. torquatus). Stenocercus leybachi is easily distinguished from S. qalaywasi by having a distinct, serrate, low crest on neck (absent in S. qalaywasi) and by lacking a distinct, black antehumeral collar (complete middorsally in S. qalaywasi). Stenocercus arndti, S. crassicaudatus, S. empetrus, S. flagracanthus, and S. simonsii can also be easily distinguished from S. qalaywasi by having the scales on dorsal surface of neck granular, while in the new species they are keeled. Furthermore, S. crassicaudatus and S. flagracanthus possess more vertebrals (83–97) than S. qalaywasi (77–79). In the case of S. flagracanthus, the tail is shorter (50–54% of total length) than S. qalaywasi (57–60%). Species such as S. bolivarensis, S. carrioni, S. chlorostictus, and S. eunetopsis possess keeled scales on the dorsal surface of neck, like S. qalaywasi, and preserved specimens can be confused. However, these species can be distinguished by the following features: from S. arndti, S. carrioni, S. chlorostictus, S. empetrus, and S. simonsii by having a distinct black, middorsally complete, antehumeral collar and two nuchal bands (antehumeral collar incomplete middorsally and nuchal bands absent in the aforementioned species); from S. carrioni and S. chlorostictus by having more vertebral scales (55–72 in S. carrioni, 63–73 in S. chlorostictus and 77–79 in S. qalaywasi); from S. bolivarensis by having the lateral scales of body granular (strongly keeled in S. bolivarensis); and from S. eunetopsis by having more scales around midbody (60–80 in S. eunetopsis and 87–92 in S. qalaywasi) and a shorter tail (57–60% of total length vs. 62–66% in S. eunetopsis). Characterization. (1) Maximum SVL in males 88 mm (n = 1); (2) maximum SVL in females 95 mm (n = 1); (3) vertebrals 77–79; (4) paravertebrals 104–108; (5) scales around midbody 87–92; (6) supraoculars six; (7) internasals four; (8) postrostrals six; (9) loreals five; (10) gulars 44; (11) subdigitals on Finger IV 26–29; (12) subdigitals on Toe IV 33–34; (13) posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; (14) postfemoral mite pocket distinct with slit-like opening [Type 2 of Torres-Carvajal (2007b)]; (15) parietal eye not visible; (16) scales on occipitoparietal region small, rugose, juxtaposed; (17) projecting angulate temporal absent; (18) row of enlarged supraoculars occupying most of supraocular region absent; (19) scales on frontonasal region juxtaposed, rugose; (20) preauricular fringe present; (21) antegular, antehumeral, gular, longitudinal, and postauricular neck folds present; (22) lateral and dorsal nuchals similar in size; (23) posterior gulars cycloid, smooth, slightly imbricate, not notched; (24) lateral body scales smaller than dorsals, reduced in size, approximately one third of the size of dorsal body scales closer to vertebral line; (25) vertebrals larger than adjacent paravertebrals forming a distinct vertebral row; (26) dorsolateral crest absent; (27) ventrals smooth, imbricate; (28) scales on posterior surface of thighs granular; (29) prefemoral fold present; (30) inguinal groove present; (31) preanals not projected; (32) tail not compressed laterally in adult males; (33) tail relatively short (tail length 57–60% of total length); (34) two caudal whorls per autotomic segment; (35) caudals spinose; (36) dark stripe extending anterodorsally from subocular region to supraciliaries absent; (37) dark patch extensively covering gular region of females absent; (38) dark patch extensively covering gular region of adult males absent; (39) black patch on ventral surface of neck in adult males absent; (40) dark midventral longitudinal mark, such as faint line, conspicuous stripe, or extensive patch in adult males absent; (41) black patches on ventral surfaces of thighs in adult males absent; (42) background color of dorsum green or dark gray, with distinct black bands in adults individuals of both sexes; (43) postxiphisternal inscriptional ribs not examined. Description of the holotype. Male (Fig. 8); SVL 88 mm; TL 136 mm; maximum head width 16.8 mm; head length 21.4 mm; head height 13.2 mm; posterior dorsal head scales small, smooth, juxtaposed; parietal eye not visible; supraoculars in six rows, smooth, juxtaposed, with the most lateral two rows less than half the size of the medial adjacent rows; distinct circumorbitals absent; canthals two; internasals four; postrostrals five, two lateral ones larger; supralabials six; infralabials six; loreals four; lorilabials in one row; preoculars two, the dorsal-most in contact with posterior canthal; lateral temporals granular; gulars in 44 rows between tympanic openings; all gulars cycloid, smooth, imbricate, not notched; second infralabial in contact with first two sublabials; first pair of postmentals partially in contact medially; mental in contact with first pair of infralabials but not with the first pair of postmentals; dorsal scales of neck small, weakly keeled, slightly imbricate, and slightly larger than granular laterals; dorsal scales of body imbricate, keeled, becoming smaller, slightly keeled or smooth towards flanks; scales around midbody 87; vertebrals 77, enlarged, keeled, imbricate, forming a distinct vertebral row; paravertebrals adjacent to vertebral row 104, same size or slightly smaller than vertebrals; ventrals smooth, imbricate, larger than dorsals; preauricular fringe short, composed by two enlarged, projected scales; antegular, antehumeral, gular, longitudinal, and postauricular neck folds present; ventrolateral fold present; humeral dorsal scales imbricate, weakly keeled; dorsal scales of forelimbs imbricate, keeled; dorsal scales of hindlimbs imbricate, keeled, mucronate; ventral humeral scales imbricate; ventral scales of forearms keeled, imbricate, and ventral scales of hindlimbs imbricate, smooth; palmars imbricate, keeled; plantars imbricate, keeled, mucronate; lamellae on Finger IV 26; lamellae on Toe IV 33; tail rounded; caudals strongly keeled, mucronate, imbricate dorsally; two caudal whorls per autotomic segment; basal subcaudals strongly keeled, imbricate; posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; postfemoral mite pocket distinct with slit-like opening; [Type 2 of Torres-Carvajal (2007b)]. Color in life of holotype (Fig. 9 A-C): Head bright green with a narrow gray postocular stripe and two gray stripes below it; dorsal surface of neck with two pale gray transverse bands and a distinct, middorsally complete black antehumeral collar; dorsum, limbs and tail with a green hue and scattered dark gray flecks; dorsum and forelimbs also covered by bright green specks; gular region dark gray, spotted with light gray blotches, and bearing a pale gray medial patch posteriorly; ventral surface of neck pale gray; chest greenish gray; belly light pink; ventral surface of hips and hindlimbs cream; cloacal region and ventral surface of base of tail cream with a light pink hue; ventral surface of tail gray becoming dark gray towards tip. Iris brown. Color in preservative (Fig. 8): dorsal surface dark gray; the postorbital gray stripe black; bands on neck and antehumeral collar black; dorsum with short irregular black bands along the vertebral region and covered with light gray specks; forelimbs covered with light gray specks and hindlimbs with darker bands than the gray background; fingers and toes light gray with black spots. Ventral surface similar as in life, but without the pink and green hues. Intraspecific variation. Scale counts and measurements for Stenocercus qalaywasi are presented in Table 1. Loreals 4–5; postrostrals 4–5; second infralabial not in contact with third sublabial in all specimens; first pair of postmentals in contact in both specimens. Sexual dimorphism is not evident in S. qalaywasi; the single female paratype (CORBIDI 20568) differs from the male holotype only in having the antehumeral black collar incomplete (Fig. 9E). Ventrally, the gular coloration is greenish cream with cream spots laterally in the female paratype (Fig. 9F), and dark gray with light gray spots in the male holotype. With a low sample of two specimens, the female is larger (95 mm) than the male (88 mm SVL). Distribution and natural history. Stenocercus qalaywasi is known only from the village of Lampa Pariahuanca in the inter-Andean valley of the Mantaro River in central Peru (Fig. 3). It occurs at an elevation of 2,587 m in the department of Junín. The type locality lies within the Peruvian Yungas ecoregion following Olson et al. (2001) and Yungas ecoregion according to Brack-Egg (1986) and Peñaherrera del Águila (1989). Six individuals of S. qalaywasi were observed basking on sunny days at 0900 hours on the rooftops of Lampa Pariahuanca village houses. When disturbed, they hid under the roof tiles, in holes between bricks or in the slit between roof and wall. The surrounded areas of the village are croplands of corn (Zea mays), potato (Solanum tuberosum), carrot (Daucus carota), and barley (Hordeum vulgare), with a few scattered patches of secondary montane forest on the steepest slopes. No other species of lizards or snakes were recorded at this locality. When the adult individuals of S. qalaywasi are basking, they possess the head bright green and the dorsal surface of body and tail with a green hue bearing conspicuous light green specks on back (Fig 9G). At the moment of capture the single male collected changed dramatically in coloration to dark gray with the dorsum containing light gray specks (Fig. 9A). The single collected female specimen did not show a dramatic change of coloration becoming simply dull green (Fig. 9D). Etymology. The specific epithet “ qalaywasi ” is a noun in apposition derived from two words in Quechua, “ qalaywa ” that means lizard and “ wasi ” that means home or house. The specific name refers to the habitus of this species of living in the houses of Lampa Pariahuanca village.

Stenocercus leybachi sp. nov. Figures 4–5 Holotype. CORBIDI 16397, an adult male, from Comunidad Saria, Chaglla (9°43’27.7”S, 75°48’35.9”W, 1,082 m), Huánuco Province, Huánuco Department, Peru, collected by D. Vásquez on 11 October 2015. Paratypes (22). PERU: HUÁNUCO DEPARTMENT: Huánuco Province: CORBIDI 9323, adult female, from Chinchavito (9° 33’ 27.2”S, 75° 55’ 07.1”W, 824 m), collected by P.J. Venegas on 16 June 2011; CORBIDI 13659, juvenile male, from Miraflores (9°41’0.11”S, 75°50’33.44”W, 1,270 m), collected by V. Duran and L. Lujan on 11 October 2013; CORBIDI 16390, a juvenile, CORBIDI 16394–95 and 16398, males, CORBIDI 16399–16400, females, from Saria, Chaglla (9°42’55.4”S, 75°49’3”W, 1,151 m), collected by D. Vásquez on 24 August 2015 and 10 October 2015; CORBIDI 16412 juvenile, CORBIDI 16428–29, 16431, females, CORBIDI 16432, juvenile, CORBIDI 16437, female, from Agua Nueva, Chinchao (9°41’59.82”S, 75°49’55.39”W, 1,115 m), collected by J. C. Chávez-Arribasplata within 24 to 28 September 2015 and 1 to 4 October 2015; CORBIDI 16525–26, a juvenile and female, CORBIDI 16532–33, male and juvenile, CORBIDI 16539, female, and CORBIDI 16594, juvenile, also from Agua Nueva, Chinchao (9°43’6.67”S, 75°48’56.1”W, 1,185 m), collected by A.C. Barboza within 10 to 15 November 2015; Pachitea Province: CORBIDI 13333, male, from Casa de Máquinas, Chaglla (9°42’17.81”S, 75°49’47.96”W, 928 m), collected by V. Duran and L. Luján on August 2013; CORBIDI 13334, male, from El Waro, Chaglla (9°41’55.87”S, 75°49’46.56”W, 927 m), collected by V. Duran and L. Luján on August 2013. Diagnosis. Stenocercus leybachi differs from all other Stenocercus species, except S. arndti, S. asenlignus sp. nov., S. bolivarensis, S. carrioni, S. chlorostictus, S. crassicaudatus, S. empetrus, S. eunetopsis, S. flagracanthus, S. nigrocaudatus sp. nov., S. qalaywasi sp. nov., S. torquatus, and S. simonsii, by having granular scales on posterior surface of thighs, relatively short tail, caudals spinose and two caudal whorls per autotomic segment. Nevertheless, S. leybachi can be easily distinguished from all aforementioned species by having a distinct serrate low crest on neck, that in some specimens reach to the middle of the dorsum. We found the arboreal S. chinchaoensis, a species known from the Upper Huallaga Basin at Huánuco Department (Venegas et al. 2013), sympatric with S. leybachi in the locality of Agua Nueva, Chinchao district, at elevations of 1,115 to 1,200 m. This species also has the ability to change color between green and gray, however this has a longer tail (tail length 61–64% of total length versus 52–58% of total length, respectively) and caudal scales not spinose. Stenocercus boettgeri is another green species distributed in the Amazon slope of central Peru from Huánuco, Pasco and Junín departments, at elevations between 2900 and 3250 m (Torres-Carvajal 2007b). Although this species can be confused with S. leybachi, the former species can be readily distinguished by having the lateral neck scales less than half the size of dorsal neck scales (dorsal and lateral neck scales similar in size in S. leybachi) and lacking a spinose tail. Characterization. (1) Maximum SVL in males 84mm (n=6); (2) maximum SVL in females 82mm (n=7); (3) vertebrals 63–73; (4) paravertebrals 100–112; (5) scales around midbody 74–97; (6) supraoculars 7–9; (7) internasals 4; (8) postrostrals 4–6; (9) loreals 3–5; (10) gulars 42–61; (11) subdigitals on Finger IV 21–28; (12) subdigitals on Toe IV 28–32; (13) posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; (14) postfemoral mite pocket distinct with slit-like opening [Type 2 of Torres-Carvajal (2007b)]; (15) parietal eye not visible; (16) scales on occipitoparietal region small, some rugose and other smooth, juxtaposed; (17) projecting angulate temporal absent; (18) row of enlarged supraoculars occupying most of supraocular region absent; (19) scales on frontonasal region juxtaposed, smooth, slightly imbricate; (20) preauricular fringe present; (21) antegular, antehumeral, gular, longitudinal, and postauricular neck folds present; (22) lateral and dorsal nuchals similar in size; (23) posterior gulars cycloid, smooth, slightly imbricate, not notched; (24) lateral body scales smaller than dorsals, reduced in size, approximately one third of the size of dorsal body scales closer to vertebral line; (25) vertebrals larger than adjacent paravertebrals forming a conspicuous vertebral row and a distinct low serrate crest on neck that in some specimens comes to the middle of dorsum; (26) dorsolateral crest absent; (27) ventrals smooth, imbricate; (28) scales on posterior surface of thighs granular; (29) prefemoral fold present; (30) inguinal groove present; (31) preanals not projected; (32) tail not compressed laterally in adult males; (33) tail relatively short (tail length 52–58 % of total length); (34) two caudal whorls per autotomic segment; (35) caudals spinose; (36) dark stripe extending anterodorsally from subocular region to supraciliaries absent; (37) dark patch extensively covering gular region of females absent; (38) dark patch extensively covering gular region of adult males absent; (39) black patch on ventral surface of neck in adult males absent; (40) dark midventral longitudinal mark, such as faint line, conspicuous stripe, or extensive patch in adult males absent; (41) black patches on ventral surfaces of thighs in adult males absent; (42) background color of dorsum green, dark brown or gray and without distinct black bands in adult individuals of both sexes; (43) postxiphisternal inscriptional ribs not in contact midventrally [Pattern 4A of Torres-Carvajal 2004)]. Description of the holotype. Male (Fig. 4); SVL 83 mm; TL 117 mm; maximum head width 16.5 mm; head length 20.1 mm; head height 12.7 mm; posterior dorsal head scales small, smooth, slightly imbricate, juxtaposed; parietal eye not visible; supraoculars in seven rows, smooth, slightly imbricate, with the most lateral two rows less than half the size of the medial adjacent rows; distinct circumorbitals absent; canthals two; internasals four; postrostrals four, two median ones larger; supralabials six; infralabials five; loreals four; lorilabials in two rows; preoculars three, the middle one tiny, the dorsal-most in contact with posterior canthal; lateral temporals granular; gulars in 47 rows between tympanic openings; all gulars cycloid, smooth, slightly imbricate, not notched; second infralabial in contact with first three sublabials; first pair of postmentals not in contact medially; mental in contact with first pair of infralabials and first pair of postmentals; dorsal scales of neck keeled and lateral scales of neck granular; dorsal scales of body imbricate, keeled, feebly mucronate becoming smaller and slightly keeled to smooth towards flanks; scales around midbody 86; vertebrals enlarged, keeled, imbricate, in 73 rows, forming distinct vertebral row and a low serrate crest from the neck to the middle of dorsum; paravertebrals adjacent to vertebral row same size as dorsals, keeled, and imbricate; paravertebrals 107; ventrals smooth, imbricate, one third larger than dorsals; preauricular fringe short, composed of two enlarged, posteriorly projected scales; antegular, antehumeral, gular, longitudinal, and postauricular neck folds present; ventrolateral fold present; dorsal humeral scales imbricate, weakly keeled; scales of forearms imbricate, keeled; dorsal scales of hindlimbs imbricate, strongly keeled, mucronate; ventral humeral scales imbricate, weakly keeled; ventral scales of forearms and hindlimbs imbricate, smooth; palmars imbricate, weakly keeled; plantars imbricate, smooth; lamellae on Finger IV 24; lamellae on Toe IV 29; tail rounded (tail length 58% of total length); caudals strongly keeled, mucronate, imbricate dorsally; two caudal whorls per autotomic segment; basal subcaudals slightly keeled, imbricate; posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; postfemoral mite pocket distinct with slit-like opening; [Type 2 of Torres-Carvajal (2007b)]. Color in life of holotype (Fig. 5 A-C): dorsum emerald green with whitish and turquoise spots on head and neck, and pale greenish yellow spots on dorsum; pelvic region, hindlimbs and tail pale brown covered by dark gray reticulations and pale greenish yellow spots; throat brownish gray with pale gray spots; ventral surface including base of tail dirty cream and rest of tail darker than the base. Once captured it quickly changes its dorsal color from green to dark brown, sprinkled with dirty cream spots on head and neck, and yellowish cream spots on dorsum; forelimbs, hindlimbs and tail dark grayish brown, darker on tail and with a greenish tone on forelimbs; some dirty cream spots are present on hindlimbs. Iris pale brown. Color in preservative (Fig. 4 D-E): head, forelimbs and dorsum to pelvic region greenish gray, sprinkled with cream spots; anterior surface of thighs greenish gray and the rest of hindlimbs, pelvic region and tail grayish brown with scattered dark gray flecks and reticulations. The last one third of tail is black. The throat is greenish gray without spots, the rest of ventral surface of body grayish cream. Intraspecific variation. Measurements,scutellation, and other morphological characters of Stenocercus leybachi are presented in Table 1. Supralabials 4–6; infralabials 5–6; second infralabials not in contact with third sublabials in sixteen specimens (69.5%); first pair of postmentals in contact medially only in four specimens (17.3%). In two dissected specimens, the rib pattern was three xiphisternal and two long postxiphisternal pairs of inscriptional ribs not in contact midventrally, [Pattern 4A of Torres-Carvajal (2004)]. Females are smaller than males (Table 1) and both sexes are able to change color from green to dark brown or grayish brown (see Fig. 5). The seven adult males in the type series lack a black antehumeral collar and of the seven adult females, one (CORBIDI 16429) possesses a middorsally complete collar. Complete black antehumeral collar is present in eight of nine juveniles, only one specimen (CORBIDI 16431) has an incomplete black collar. Adult specimens of both sexes and juveniles possess pale spots on dorsum and some specimens also have scattered black flecks. The throat in adult females can be brownish green with pale green spots, chest with a greenish tone and the belly is dirty cream with or without a pinkish hue on the belly and base of tail. Juveniles have throat and venter yellowish green. Distribution and natural history. Stenocercus leybachi is known from five close localities in the Amazon foothills of the Río Huallaga basin, at elevations between 824–1,270 m, in central Peru (Fig. 3). The known localities of this species lie in the Huánuco Department within the Yungas (500–2,300 m) ecoregion according to Brack–Egg (1986) and Peñaherrera del Aguila (1989), and Peruvian Yungas following Olson et al. (2001). The general habitat where the new species was collected is composed of croplands of coca (Erythroxylum coca), coffee (Coffea sp.), cacao (Theobroma cacao), corn (Zea mays), orange (Citrus sp.) and pastures for cattle ranching with scattered patches of secondary montane forest, especially on the steepest slopes. Individuals of S. leybachi were observed active during sunny days basking on tree trunks at 1.5–6 m above ground. When these individuals felt disturbed, they climbed the tree trunk finding refuge up high. Some individuals were observed and collected with a fishing rod while basking on fallen trees. The individuals found basking were light green and at the moment of capture suddenly changed to brown or brownish gray. Most specimens were collected in the forest canopy, during a flood for the construction of a hydroelectric dam. The specimens were collected by hand on the branches of high trees (5 to 10 m) from a boat, and some individuals tried to escape by hiding in tree holes (Fig. 5 K-L) and cracks in the bark of the trees. Stenocercus leybachi was syntopic in the area of the flood with other arboreal lizards such as Anolis punctatus, Euspondylus excelsum, and S. chinchaoensis. Other sympatric species of squamate reptiles collected with S. leybachi were: A. ortonii, Bothrops chloromelas, Dipsas catesbyi, D. indica, D. schunkii, Drymoluber dichrous, Enyalioides feiruzae, Micrurus annelatus, Oxyrhopus leucomelas, Phrynonax polylepis, Proctoporus sp., and S. prionotus. Etymology. The specific name leybachi is a patronym for Achim Leybach of Marktsteft, Germany, in recognition of his financial support for the taxonomic work and biodiversity research through the BIOPAT-Programme.

Stenocercus nigrocaudatus sp. nov. Figures 6���7 Holotype. CORBIDI 19768, an adult male from San Lorenzo, San Jos�� de Lourdes (5��4���33.16���S, 78��52���53.749���W, 1,892 m), San Ignacio Province, Cajamarca Department, Peru, collected by P.J. Venegas on 5 September 2018. Paratypes (8): CAJAMARCA DEPARTMENT: San Ignacio Province: CORBIDI 4387, an adult male from Tabaconas (5��14���8.16���S, 79��5���56.58���W, 1,716 m) collected by Maik Dobiey in August 2008; CORBIDI 4388-90, adult females, from Namballe (5��11���40.139���S, 79��4���49.738���W, 1,716 m) collected by M. Dobiey in August 2008; CORBIDI 19765-66, adult females, and CORBIDI 19767 and 19769, adult males, from San Lorenzo, San Jos�� de Lourdes (5��4���33.16���S, 78��52���53.749���W, 1,892 m), collected by P.J. Venegas on 5 September 2018. Diagnosis. Among the 80 species of Stenocercus (including the four species described herein), S. nigrocaudatus sp. nov. is only similar to species with granular scales on posterior surface of thighs, relatively short tail, caudal scales spinose, and two caudal whorls per autotomic segment; such as S. asenlignus sp. nov., S. arndti, S. qalaywasi sp. nov., S. flagracanthus, S. leybachi sp. nov., S. bolivarensis, S. carrioni, S. chlorostictus, S. crassicaudatus, S. empetrus, S. eunetopsis, S. torquatus and S. simonsii. Adult males of S. nigrocaudatus can be easily distinguished from the aforementioned species by a contrasting black tail with scattered turquoise flecks (Fig. 7C). Moreover, S. nigrocaudatus can be readily distinguished from S. qalaywasi, S. leybachi, S. bolivarensis, S. carrioni, S. chlorostictus, and S. eunetopsis by having granular scales on dorsal surface of neck (keeled in the compared species).Furthermore, S. leybachi possesses a distinct serrate low crest on neck (absent in S. nigrocaudatus), S. bolivarensis strongly keeled scales on flanks (granular), S. qalaywasi a distinct complete middorsally black antehumeral collar and two black nuchal bands (incomplete and absent, respectively), S. carrioni and S. chlorostictus by having strongly keeled dorsal scales on body (smooth and feebly keeled), and S. eunetopsis by having a longer tail with 62���66% of total length (50���55%). In addition, S. nigrocaudatus possesses more vertebrals (96���108) than S. carrioni (55���72), S. chlorostictus (63���73), and S. eunetopsis (59���80). Stenocercus nigrocaudatus shares with S. asenlignus, S. arndti, S. flagracanthus, S. crassicaudatus, S. empetrus, S. torquatus, and S. simonsii the presence of granular scales on neck. Nevertheless, the new species can be differentiated from these species by the coloration pattern, number of scales, or the length of tail. Adult males of S. arndti are easily distinguished from S. nigrocaudatus (state of character in parentheses) by having a bold black transverse band at midbody that extends ventrolaterally (bands on body absent); adult males of S. empetrus have the belly black (cream); S. simonsii has distinct black bands on body in both sexes (absent). Additionally, S. nigrocaudatus has a greater number of vertebrals (96���108) than S. arndti (74���94), more subdigitals (26���31) than S. empetrus (17���24), and a shorter tail in relation to the total length (50���55%) than S. simonsii (57���63%). Stenocercus asenlignus and S. flagracanthus differ from S. nigrocaudatus in having dorsal scales on the posterior half of body strongly keeled and mucronate (feebly keeled in S. nigrocaudatus). Moreover, adult males of S. asenlignus and S. flagracanthus have black bands on the body (absent in S. nigrocaudatus). Stenocercus crassicaudatus has a longer tail (57���62% of total length) than S. nigrocaudatus (50���55%). Adult males of S. torquatus differ from S. nigrocaudatus in having the antehumeral black collar complete middorsally and two distinct black nuchal transverse bands, whereas the new species has the antehumeral collar incomplete middorsally and lacks nuchal bands. Characterization. (1) Maximum SVL in males 72 mm (n = 4); (2) maximum SVL in females 76 mm (n = 5); (3) vertebrals 96���108; (4) paravertebrals 110���128; (5) scales around midbody 99���117; (6) supraoculars 5���7; (7) internasals 4���5; (8) postrostrals 5���6; (9) loreals 3���7; (10) gulars 48���59; (11) lamellae on Finger IV 26���31; (12) lamellae on Toe IV 30���36; (13) posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; (14) postfemoral mite pocket distinct with slit-like opening [Type 2 of Torres-Carvajal (2007b)]; (15) parietal eye absent; (16) occipital scales small, smooth, juxtaposed; (17) projecting angulate temporal absent; (18) row of enlarged supraoculars occupying most of supraocular region absent; (19) scales on frontonasal region juxtaposed and smooth, not imbricate; (20) preauricular fringe short; (21) antegular, antehumeral, gular, longitudinal, oblique, postauricular, and supra-auricular neck folds present; (22) lateral nuchals and dorsals similar in size; (23) lateral body scales smaller than dorsals; (24) vertebrals enlarged, forming a distinct row of scales from forelimbs to hindlimbs; (25) dorsolateral crest absent; (26) paravertebrals and adjacent scales from midbody, smooth or feebly keeled, becoming keeled on the second half of body and mucronate between hindlimbs; (27) ventral scales smooth, imbricate; (28) scales on posterior surface of thighs granular; (29) prefemoral fold present; (30) inguinal fold present; (31) preanals not projected; (32) tail not compressed laterally; (33) tail relatively short (tail length 50���55% of total length); (34) caudal whorls per autotomic segment two; (35) tail spinose; (36) postocular stripe present; (37) dark patch on gular region in males absent; (38) dark patch on gular region in females absent; (39) black patch on ventral surface of neck in adult males absent; (40) dark midventral stripe in adult males absent; (41) dark patch on ventral surface of thighs, vent and tail in adult males absent; (42) background color of dorsum green, dark brown or olive green in both sexes; (43) postxiphisternal inscriptional ribs not examined. Description of holotype. Male (Fig. 6); SVL 57 mm; TL 114 mm; maximum head width 12.3 mm; head length 16.3 mm; head height 10.1 mm; scales on parietal and occipital regions small, smooth, juxtaposed, subequal in size; parietal eye not visible: supraoculars six, smooth, juxtaposed; circumorbitals absent; canthals two; loreals one; postrostrals four; internasals four; supralabials five; infralabials five; lorilabials in one row; preocular divided into two scales, most dorsal in contact with posterior canthal; lateral temporals granular; gulars in 55 rows between tympanic openings; all gulars cycloid, smooth, imbricate; second infralabial in contact with first, second and third sublabials; first pair of postmentals in contact; mental not separated from the infralabials by the first pair of postmentals; dorsal and lateral scales of neck and body granular; scales around midbody 107; vertebrals 104 moderately enlarged, keeled, imbricate, forming distinct vertebral row; paravertebrals and adjacent scales slightly smaller than vertebrals, smooth, subimbricate, becoming feebly keeled from the posterior half of dorsum and strongly keeled near hindlimbs insertion; paravertebrals 127; scales on flanks granulars; ventrals smooth, imbricate, more than twice the size of dorsals, only paravertebrals on the second half of body, nearly equal in size or slightly longer than ventrals; preauricular fringe short composed of three enlarged scales with round edge; suprauricular, antehumeral, gular, longitudinal, oblique, antegular, postauricular and rictal neck folds present; dorsolateral, ventrolateral and prefemoral folds present; dorsal scales of forelimbs imbricate, keeled; dorsal scales of hindlimbs imbricate, strongly keeled and mucronate; ventral humeral scales granular; ventral scales of fore- and hindlimbs smooth, imbricate; palmar scales imbricate, keeled and minutely mucronate; plantar scales imbricate, smooth or strongly keeled and mucronate; lamellae on Finger IV 28; lamellae on Toe IV 35; tail rounded (tail length 50% of total length); caudal scales strongly keeled, mucronate, imbricate; basal subcaudal scales smooth, imbricate; posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007b)]; postfemoral mite pocket distinct with slit-like opening [Type 2 of Torres-Carvajal (2007b)]. Color in life (Fig. 7 A-C): head and neck brownish gray with white flecks on head and white dots on neck; labials white with dark brown bars; postocular stripes dark brown, extending through the temporal and edge of occipital region to a black rhomboid blotch in the nuchal region; antehumeral collar black, distinct; dorsal surface of body green covered by bright green dots; forelimbs green with scattered bright green dots; hindlimbs brown with scattered white flecks and dots on posterior surface of thighs; tail black with scattered turquoise flecks on the basal half. Ventrally, gular region grayish white with white dots and black flecks; neck gray with few scattered black flecks; chest and belly cream with an orange hue on the posterior half; thighs, cloacal region, and tail dull orange. Iris dark brown. Color in ethanol 70% (Fig. 6 D-E): the dorsal surface turns dark brownish gray; postocular stripes and the nuchal blotch remain black; the white dots on the neck and the scattered white flecks on hindlimbs are dirty white; the bright green dots on the body and forelimbs are pale gray; tail turns dark brown. Ventrally, the gular region and neck is pale gray with dirty white dots and scattered black flecks, and chest, belly, limbs and base of tail are grayish white; the two distal thirds of the tail are dark brown with grayish white bars. Intraspecific variation. Measurements and scutellation of Stenocercus nigrocaudatus are presented in Table 1. Supralabials 4���5; infralabials 5���6; no specimens showed the second infralabials in contact with third sublabial; first pair of postmentals in contact medially in eight specimens (88.8%). Females larger than males (see Table 1), and both sexes are able to change color from green to dark brown or grayish brown. Sexual dimorphism in coloration is evident in the tail, which is black with scattered turquoise speckles in males (Fig. 7C) and brown with black and gray speckles in the proximal half and dark brown with gray rings in the distal half in females (Fig. 7J). The antehumeral collar is bold black in males and faint black or dark gray in females. Ventrally, the gular region in females is gray with scattered black flecks and speckles, the chest has a greenish hue, the belly is grayish white, and the ventral surface of hindlimbs, cloacal region and tail are tannish cream (Fig. 7I). The single female juvenile specimen (CORBIDI 19766) is dorsally identical to the adult female, but the gular region and chest possess a green hue. Distribution and natural history. Stenocercus nigrocaudatus is known from both sides of the inter-Andean valley of the Chinchipe River in the Cajamarca Department of northeastern Peru, between elevations of 1,700 and 1,892 m (Fig. 3). The two known localities of this species lie at the Yungas (500���2,300 m) ecoregion according to Brack-Egg (1986) and Pe��aherrera del ��guila (1989), and Eastern Cordillera Real montane forest following Olson et al. (2001). The general landscape in Alto Ihuamaca is composed of humid montane forest covering the steepest slopes and open areas with croplands for corn (Zea mays), coffee (Coffea sp.), and pastures for cattle ranching. In San Jos�� de Lourdes, the general landscape includes pastures for cattle ranching and small croplands for coffee, mango (Mangifera indica), and oranges (Citrus sp.), with some scattered patches of secondary forest and fringes of forest bordering ravines and along streams. Several individuals of S. nigrocaudatus were observed basking during sunny days between 900 and 1,500 hours. In small villages and single houses in the field, we observed individuals of S. nigrocaudatus basking on the walls at heights between 2 and 3 m (Fig. 7L), and one individual was observed at 7 m in the wall of a small building. This species also was common in open areas for cattle ranching basking on big tree trunks between 1 and 8 m above ground (Fig. 7K). Up to five individuals, a pair of adults and three juveniles, were observed in the same tree. All individuals observed basking showed a green dorsal coloration that changed to brownish gray to the moment of be captured. Sympatric squamate reptiles collected with S. nigrocaudatus were Alopoglossus buckleyi, Atractus gigas, Enyalioides anisolepis, and Potamites strangulatus; and in San Jos�� de Lourdes only Varzea altamazonica. Etymology. The specific epithet nigrocaudatus is a noun in apposition that derives from the Latin word ��� nigrum ��� that means black and ���cauda��� that means tail, with the suffix ��� tus ��� that means provided with. It refers to the black tail that is characteristic of the adult males of this new species., Published as part of Venegas, Pablo J., Garc��a-Ayachi, Luis A., Ch��vez-Arribasplata, Juan C. & Garc��a-Bravo, Antonio, 2022, Four new species of polychromatic spiny-tailed iguanian lizards, genus Stenocercus (Iguania: Tropiduridae), from Peru, pp. 1-28 in Zootaxa 5115 (1) on pages 15-18, DOI: 10.11646/zootaxa.5115.1.1, http://zenodo.org/record/6345982, {"references":["Torres-Carvajal, O. (2007 b) A taxonomic revision of south American Stenocercus (Squamata: Iguania) lizards. Herpetological monographs, 21, 76 - 178. https: // doi. org / 10.1655 / 06 - 001.1","Penaherrera del Aguila, C. (1989) Atlas del Peru. Instituto Geografico Nacional, Lima, 400 pp.","Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N. D., Powell, G. V. N., Underwood, E. C., D'Amico, J. A., Itoua, I., Strand, H. E., Morrison, J. C., Loucks, C. J., Allnutt, T. F., Ricketts, T. H., Kura, Y., Lamoreux, J. F., Wettengel, W. W., Hedao, P. & Kassem, K. R. (2001) Terrestrial ecoregions of the world: A new map of life on earth. BioScience, 51, 933 - 938. https: // doi. org / 10.1641 / 0006 - 3568 (2001) 051 [0933: TEOTWA] 2.0. CO; 2"]}

Venegas, Pablo J., Echevarría, Lourdes Y., García-Ayachi, Luis A., Landauro, Caroll Z. (2020): Two new sympatric species of Stenocercus (Squamata: Iguania) from the inter- Andean valley of the Mantaro River, Peru. Zootaxa 4858 (4): 555-575, DOI: https://doi.org/10.11646/zootaxa.4858.4.5

Stenocercus nigrobarbatus sp. nov. Figures 6–10, Table 1. Holotype. CORBIDI 13725, adult male from Fundición (12°17’36.41”S, 74°39’39.67” W) at 2,158 m asl, Tayacaja Province, Huancavelica Region, Peru, collected on January 31, 2014 by L. Y. Echevarría. Paratypes. Peru: Huancavelica Department: Tayacaja Province: CORBIDI 13637 a juvenile female from Jatuspata (12°15’14.617’’ S, 74°41’ 16.93’’ W) at 2,972 m asl, collected on September 3, 2013 by C. Landauro; CORBIDI 13638 a juvenile female from Pichiu (12°19’11.815’’ S, 74°39’3.588’’ W) at 2,036 m asl, collected on September 13, 2013 by C. Landauro; CORBIDI 13640 juvenile female from Pichiu, collected on September 12, 2013 by C. Landauro; CORBIDI 13641 adult female from Barropata (12°17’21.943’’ S, 74°41’19.035’’ W) at 1,725 m asl, collected on October 2, 2013 by C. Landauro; CORBIDI 13639 and CORBIDI 13642 female juveniles from Fundición (12°17’32.347’’ S, 74°39’45.433’’ W), at 2,169 m asl, collected on September 28, 2013 by C. Landauro; CORBIDI 13718 adult male from Chupto (12°19’52.96” S, 74°39’13” W) at 2,058 m asl, collected on January 23, 2014 by L. Y. Echevarría; CORBIDI 13719 adult male from Chupto (12°18’22.19” S, 74°39’18.42” W) at 2,314 m asl, on January 24, 2014 by L. Y. Echevarría; CORBIDI 13722 adult female from Chupto (12°19’52.96” S, 74°39’13” W) at 2,058 m asl, collected on January 21, 2014 by L. Y. Echevarría; CORBIDI 13728 adult female from Barropata (12°17’34.95” S, 74°41’34.08” W) at 1,704 m asl, collected on February 2, 2014 by L. Y. Echevarría; CORBIDI 13729 adult female from Barropata (12°17’22.11” S, 74°41’19.07” W) at 1,783 m asl, collected on February 3, 2014 by L. Y. Echevarría; CORBIDI 14669 adult female from Chupto (12°18’48.18” S, 74°39’1.83” W) at 2,345 m asl, collected on July, 1 2014 by L. Y. Echevarría; CORBIDI 14674 adult female from type locality, collected on July 6, 2014 by L. Y. Echevarría; CORBIDI 14676 juvenile female and CORBIDI 14677 adult male from Barropata (12°17’37.66” S, 74°41’39.18” W) at 1,693 m asl, collected on July 7, 2014 by L. Y. Echevarría; CORBIDI 15765–66 (adult male and female, respectively) and CORBIDI 15767 (juvenile male) from Pichiu (12°15’31.57” S, 75°6’3.6” W) at 2,206 m asl, collected on January 14, 2015 by L. Lujan; CORBIDI 16033 adult male from Pichiu (12°19’50” S, 74°39’13.07” W) at 2,090 m asl, collected on July 9, 2015 by J. Malqui; CORBIDI 20546–47 two adult males from San Luis de Estanque (12º19’8.62” S, 74º44’24.19” W) at 2,636 m asl., collected on Jun 17, 2019 by L.A. García-Ayachi. Diagnosis. Adult males of Stenocercus nigrobarbatus can be easily distinguished from all species of Stenocercus by having a continuous black patch covering the infralabials, throat, chest, ventral surfaces of forelimbs, belly (as a black midventral line), ventral surfaces of hind limbs, and pelvic region (Figs. 6B, 8B, 9B and 9C). However, among the 69 currently described species of Stenocercus, S. nigrobarbatus resembles S. frittsi Torres-Carvajal, 2005, and S. variabilis Boulenger, 1901, by the following combinations of characters: (1) granular scales on the posterior surface of thighs, (2) imbricate and keeled lateral body scales, (3) a distinct row of enlarged vertebral scales, (4) unnotched gular scales, (5) three caudal whorls per autotomic segment, (6) brown dorsal ground color, and (7) distinct oblique neck fold, and antegular fold not continuous medially. Stenocercus nigrobarbatus differs from the geographically close S. frittsi (character state of latter in parenthesis) by having a postfemoral mite pocket composed by one or more vertical folds or ridges, Type 1 of Torres-Carvajal (2007a) (postfemoral pocket absent in S. frittsi), 2–4 postrostrals (5–7), and two pairs of postxiphisternal inscriptional ribs, long, not in contact midventrally, pattern 1A of Torres-Carvajal (2004) (three pairs of postxiphisternal inscriptional ribs, two pairs long and the last pair short, not in contact midventrally; pattern 2B of Torres-Carvajal 2004). Stenocercus nigrobarbatus can be distinguished from S. variabilis (character state in parenthesis) by having 2–4 postrostrals (6), postfemoral pocket composed by one or more vertical folds or ridges, Type 1 of Torres-Carvajal (2007a) (slit-like opening, Type 2 of Torres-Carvajal 2007a), and inguinal groove absent (present). In addition, lateral body scales in S. nigrobarbatus are slightly smaller than dorsals, while S. frittsi and S. variabilis have lateral body scales approximately half the size of dorsals. Characterization. (1) Maximum SVL in males 67.73 mm (n = 4); (2) maximum SVL in females 62.29 mm (n = 5); (3) vertebrals 47–73; (4) paravertebrals 69–87; (5) scales around midbody 57–74; (6) supraoculars 5–7; (7) internasals 3–5; (8) postrostrals 2–4; (9) loreals 3–7; (10) gulars 19–30; (11) lamellae on Finger IV 18–23; (12) lamellae on Toe IV 27–34; (13) posthumeral pocket as one or more vertical folds or ridges, Type 1 of Torres-Carvajal (2007a); (14) postfemoral pocket composed by one or more vertical folds or ridges, Type 1 of Torres-Carvajal (2007a); (15) parietal eye not visible; (16) occipital scales small, smooth, juxtaposed; (17) no projecting angulate temporals; (18) supraoculars of varying sizes; (19) scales in frontonasal region slightly imbricate anteriorly; (20) short preauricular fringe present; (21) antehumeral, antegular, gular, oblique, suprauricular and postauricular neck folds present; (22) nuchal mite pocket absent; (23) lateral nuchals less than half size of dorsal nuchals; (24) posterior gulars in adults smooth, imbricate, not mucronate; (25) lateral scales slightly smaller than dorsal body scales; (26) vertebrals larger than adjacent paravertebrals; (27) dorsolateral crest absent; (28) ventrals in adults smooth, imbricate, not mucronate; (29) scales on posterior surfaces of thighs granular; (30) inguinal granular pocket absent; (31) inguinal groove absent; (32) preanals not projecting; (33) tail not compressed laterally in adult males; (34) tail length 62–70% of total length; (35) three caudal whorls per autotomic segment; (36) caudals not spinose; (37) dark stripe extending anterodorsally from subocular region to supraciliaries present; (38) gular region of adult females dark or densely pigmented in all specimens; (39) gular region of adult males dark or densely pigmented in all specimens; (40) black blotch on ventral surface of neck in adult males absent; (41) bold black midventral line present in males; (42) black patch on ventral surface of thighs present in males; (43) dorsal ground color brown, in females and males; and (44) postxiphisternal inscriptional ribs not articulating midventrally, both pairs long, Pattern 1A of Torres-Carvajal (2004). Description of holotype. Male (Fig. 6-8); SVL= 66.02 mm; TL = 130.0 mm; maximum head width = 12.24; head length = 15.87; head height = 8.63; scales on parietal and occipital regions small, smooth, juxtaposed; parietal eye not visible; supraoculars in six rows, smooth, slightly imbricate, of varying sizes; canthals two; anterior most canthal separated from nasal by one minute scale; scales in frontonasal region slightly imbricate; internasals four; postrostrals three, subequal in size, wider than long; supralabials seven; infralabials seven; loreals six; lorilabials in one row; preocular not divided, in contact with posterior canthal; lateral temporals imbricate, weakly keeled; gulars in 29 rows between tympanic openings; all gulars smooth, imbricate; second infralabial in contact with first two sublabials; mental in contact with first pair of infralabials and first pair of postmentals; lateral and dorsal scales of body and dorsal scales of neck keeled, imbricate; lateral scales of neck granular; scales around midbody 63; vertebrals large, in 68 rows, not forming a prominent serrate vertebral crest; paravertebrals 86; ventrals smooth, imbricate; preauricular fringe short, composed of two enlarged scales, both same size; antehumeral, antegular, gular, oblique, suprauricular, postauricular neck folds and ventrolateral body fold present; limb scales keeled, imbricate; ventral scales of hind limbs and upper arms smooth; lamellae on Finger IV 23; lamellae on Toe IV 31; tail not compressed laterally; caudals keeled, imbricate; basal subcaudals smooth, imbricate; vertebral crest extending more than half length of tail; tail length 1.9 times SVL; posthumeral pocket shallow (Type 1 Torres-Carvajal 2007a) with vertically oriented fold approximately 2.57 mm long; postfemoral pocket composed by one vertical fold (type 1 of Torres-Carvajal 2007a); postfemoral region composed of imbricate, smooth scales, becoming smaller toward insertion of hind limbs (Fig. 10A). Color in life of the holotype. Dorsal surfaces of head and neck cinnamon brown with yellow spots, denser on neck; lorilabials and supralabials yellow; infralabials black; ventrolateral regions of head and neck black; dorsal surfaces of body, limbs and tail dusty brown with pale yellow spots; flanks light blue, densely covered with cream spots; limbs and tail with thin dark brown marks on dorsal surfaces (Fig. 8A). Ventral surface of head, throat, chest black; ventral surfaces of forelimbs black, although pigmentation is faint at elbow joint; a bold black midventral line, continuous with chest pigmentation, flanked by pink; ventral surfaces of hind limbs, and pelvic region black, except by the cloaca; posterior surfaces of thighs pink; pre-cloacal region dark yellow and post post-cloacal region pink; ventral surface of tail pink, with a yellow blotch and blackish speckles at the base (Fig. 8B). All black pigmentation on ventral surfaces constitutes a continuous patch, except black pigmentation on tail. Coloration of holotype in ethanol 70%. Dorsally, all yellow spots turned to pale yellow. Dorsal surfaces of head and neck turned brown. Ventrally, pink and yellow pigmentation turned to a less vivid (pale) shade, almost vanished on cloaca. Variation. Scale counts and measurements for Stenocercus nigrobarbatus are presented in Table 1. Loreals 3–7; supralabials 6–8; infralabials 5–8; postrostrals 2–4; second infralabial not in contact third sublabial in all specimens; first pair of postmentals in contact in all specimens. Dorsal coloration in adult males of Stenocercus nigrobarbatus is the same in all adult male paratypes (n = 5) (Fig. 9A). The black coloration on ventral surfaces as a continuous patch is exclusive to adult males (Fig. 9 B– C). The juvenile specimen CORBIDI 13718 has a faint black midventral line on the belly (Fig. 9 D–E). Another juvenile specimen (CORBIDI 13719) has the gular region and ventral surface of neck covered by a dark gray patch and the midventral region of the belly has scattered dark gray dots (Fig. 9 F–G). These immature males (CORBIDI 13718 and 13719) have the pelvic region and the base of tail yellow and the ventral surface of forelimbs cream (Fig. 9 D–G). In both specimens the pink coloration on the flanks of the belly is also less intense than in the holotype. In the adult male paratype (CORBIDI 20547) the base of the tail has a black blotch and the pre-cloacal region is yellow with black speckles (Fig. 9C). Sexual dimorphism is evident in coloration and size (maximum SVL in males 67.0 mm versus 62.0 mm in females). Dorsal coloration in adult females can be gray or brown with irregular dark brown blotches over the vertebral line and a light brown dorsolateral stripe from the temporal region to the base of tail (Fig. 9H); dorsal surface of limbs has dark brown or gray transversal bars; sides of head brown or gray with a thin postocular stripe and some scattered dark brown dots; sides of neck yellowish brown with light yellow flecks; jaw and ventrolateral region of neck black or dark gray; body flanks covered by a dark brown reticulation. Ventrally all adult females have the gular region and neck dark gray, belly and limbs are light cream, and the tail is light orange (Fig. 9I). Juvenile females, between 43.0–51.0 mm of SVL, have the same dorsal and ventral pattern than adult females, only two specimens (CORBIDI 13637 and 13642) have the gular region gray with dark gray speckles, and chest and belly light cream with gray speckles. The smallest juvenile female (CORBIDI 14676) has the gular region covered by gray flecks. The juvenile male CORBIDI 13640 (SVL = 43.0 mm) has the dorsum with dark brown dorsal marks as adult and juvenile females, however, the gular region and chest are gray and the belly is grayish cream. The juvenile male CORBIDI 15767 (SVL = 53.0 mm) has the same dorsal coloration as adult males but ventrally the gular region and chest are gray, and has a faint gray midventral line on belly flanked by a weak pink hue; pelvic region, ventral surface of hindlimbs and tail are light cream. Distribution and natural history observations. Stenocercus nigrobarbatus is known from four localities in SDF of the Mantaro River Valley at elevations between 1,693 and 2,345 m asl, Huancavelica Department, Peru (Fig. 5). Stenocercus nigrobarbatus inhabits a seasonal dry forest, according to classification of Linares-Palomino (2004), and Central Andean Yungas according to Olson et al. (2001). In most localities the habitat has been modified by crops of Zea mays, Opuntia, and Persea sp., roads and the construction of a hydroelectric plant. However, this species is very conspicuous in most localities. Specimens of Stenocercus nigrobarbatus were found inactive under rocks during cloudy days and active, basking or running, during warm days, among rocks, cacti or shrubs. Other species of Squamate reptiles collected at the same localities of S. nigrobarbatus include Ameiva reticulata, Epictia sp., Mastigodryas boddaerti, Micrurus sp., and Oxyrhopus cf. erdisii, and Stenocercus diploauris. Etymology. The specific epithet “ nigrobarbatus ” is a noun derived from the Latin words “ nigro ” (= black) and “ barbatus ” (= bearded). It refers to the black patch covering the gular region and ventral surface of neck, including the jaw angle, of adult males which resembles a black beard.

Stenocercus diploauris sp. nov. Figures 1���4, Table 1. Holotype. CORBIDI 13643, adult male from Limonal (12��13���55.097������ S, 74��41���27.487������ W), at 1,678 m asl, Surcubamba District, Tayacaja Province, Huancavelica Department, Peru, collected on September 1, 2013 by A. Escobar. Paratypes. Peru: Huancavelica Department: Tayacaja Province: CORBIDI 9913 (adult female), CORBIDI 9914 (juvenile male), CORBIDI 9915 (adult female), CORBIDI 9916 (adult male) from Jatuspata (12��15���1.2������ S, 74��41���33.6������ W), at 2,609 m asl, collected on April 7, 2011 by D. Amaya; CORBIDI 14672 adult female from Chupto (12��18���42.56������ S, 74��39���13.23������ W) at 2,328 m asl, collected on July 2, 2014 by L. Y. Echevarr��a; COR- BIDI 14901 adult female from Jatuspata (12��15���28.83������ S, 74��41���28.74������ W) at 2,816 m asl, collected on July 2, 2014 by C. Landauro; CORBIDI 14902 and CORBIDI 14903 adult females from Jatuspata (12��15���35.31������ S, 74��41���31.74������ W) at 2,835 m asl, collected on July 3, 2014 by C. Landauro; CORBIDI 14904 adult male from Jatuspata (12��15���34.23������ S, 74��41���25.12������ W) at 2,920 m asl, collected on July 3, 2014 by C. Landauro; CORBIDI 14906 adult male from Limonal (12��13���58.82������ S, 74��41���23.09������ W) at 1,753 m asl, collected on July 8, 2014 by C. Landauro; CORBIDI 16036 adult female from Pichiu (12��19���50������ S, 74��39���13.07������ W) at 2090 m asl, collected on July 12, 2015 by J. Malqui. Diagnosis. Adult specimens of Stenocercus diploauris can be easily distinguished from all known species of Stenocercus by having a C-shaped nuchal mite pocket around the oblique fold, posteriorly limited by the antehumeral fold (Fig. 4). However, among the 69 currently known species of Stenocercus, S. diploauris resembles S. formosus Tschudi, 1845, and S. ochoai Fritts, 1972, by the combination of the following characters: (1) imbricate scales on the posterior surface of thighs, (2) well developed postfemoral mite pocket, (3) antehumeral and oblique neck folds, and (4) absence of posthumeral mite pocket. The new species can be distinguished from S. ochoai and S. formosus by having dorsal and lateral nuchals similar in size and the presence of longitudinal neck fold, in the former species lateral nuchals are smaller than dorsal nuchals and the longitudinal neck fold is absent. Furthermore, S. diploauris differs from S. formosus (character state of latter in parenthesis) by having three whorls per autotomic segment (four), smooth dorsal head scales (keeled), and fewer scales, 50���61, around midbody (74���82). Characterization. (1) Maximum SVL in males 94.82 mm (n = 5); (2) maximum SVL in females 76.06 mm (n = 5); (3) vertebrals 40���50; (4) paravertebrals 63���78; (5) scales around midbody 50���61; (6) supraoculars 5���7; (7) internasals 3���4; (8) postrostrals 2���4; 9) loreals 4���8; (10) gulars 17���24; (11) lamellae on Finger IV 17���21; (12) lamellae on Toe IV 24���30; (13) posthumeral pocket absent; (14) postfemoral pocket distinct with slit-like opening, Type 2 of Torres-Carvajal (2007a); (15) parietal eye not visible; (16) occipital scales large, smooth, imbricate; (17) projecting angulate temporals absent; (18) enlarged supraoculars occupying most of supraocular region in one row; (19) scales on frontonasal region weakly imbricate anteriorly; (20) preauricular fringe present; (21) antehumeral, longitudinal and oblique neck folds present; (22) C-shaped nuchal mite pocket around the oblique neck fold, posteriorly limited by the antehumeral neck fold; (23) lateral nuchals slightly smaller than dorsal nuchals; (24) posterior gulars in adults smooth, imbricate, not mucronate, not notched; (25) lateral and dorsal body scales similar in size; (26) vertebral crest prominent; (27) dorsolateral crest absent; (28) ventrals in adults smooth, imbricate, mucronate; (29) scales on posterior surfaces of thighs keeled, imbricate, mucronate; (30) prefemoral fold absent; (31) inguinal groove absent; (32) preanals projected; (33) tail not strongly compressed laterally in adult males; (34) tail length 50���70% of total length; (35) three caudal whorls per autotomic segment; (36) caudals not spinose; (37) dark stripe extending anterodorsally from subocular region to supraciliaries present in most specimens; (38) gular region of adult females black or densely pigmented; (39) gular region of adult males gray or reddish; (40) black blotch on ventral surface of neck in adult males absent; (41) light gray or cream midventral line present; (42) black patch on ventral surface of thighs absent; (43) background color of dorsum, in females and males, brown; (44) two xiphisternal, and two long postxiphisternal pairs of inscriptional ribs that do not articulate midventrally (Pattern 1A of Torres-Carvajal 2004). Description of holotype. Male; SVL 94.82 mm; TL 217.5 mm; maximum head width 19.18 mm; head length 21.99 mm; head height 13.69 mm; occipitals, parietals, interparietal, and postparietals large, smooth, slightly imbricate (Fig. 2); parietal eye not visible; supraoculars in six rows, smooth, slightly imbricate, with one row two times larger than adjacent rows; anterior and posterior circumorbitals smooth, imbricate; canthals two; anteriormost canthal in contact with nasal; scales in frontonasal region not imbricate; internasals four; postrostrals four, wider than long, median ones smaller; supralabials six; infralabials six; loreals eight; lorilabials in one row; preocular not divid- ed, in contact with posterior canthal; lateral temporals imbricate, keeled; gulars in 22 rows between tympanic openings; all gulars smooth, imbricate; second infralabial in contact with first three sublabials; mental in contact with first pair of infralabials and first pair of postmentals; dorsal and lateral scales of body and neck keeled, imbricate; scales around midbody 53; vertebrals large, in 48 rows, forming a distinct serrate vertebral crest; paravertebrals 77; ventrals smooth, imbricate; preauricular fringe short, composed of two enlarged scales, of similar size; antehumeral, longitudinal and oblique neck folds present; a C-shaped nuchal mite pocket around the oblique fold, posteriorly limited by the antehumeral fold (Fig. 4); limb scales keeled, imbricate; ventral scales of hind limbs smooth; lamellae on Finger IV 20; lamellae on Toe IV 30; tail slightly compressed laterally; caudals keeled, imbricate; basal subcaudals keeled, imbricate; vertebral crest extending two thirds of tail length; tail length 2.3 times SVL; posthumeral pocket absent; postfemoral pocket shallow with slit-like opening (Type 2 of Torres-Carvajal 2007a). Color in life of the holotype. Dorsal surface dusty brown with light cream chevrons over the vertebral line, irregular light cream transversal bars on limbs and a reddish hue on tail; flanks cinnamon, dotted with yellowish cream scales; dorsal surface of head darker than dorsum and with some scattered cream dots; temporal surface of head dark brown, with scattered reddish pigmentation and white dots; loreal region and canthus rostralis dark brown and subocular region, supralabials and infralabials bright cream; sublabials forming a thin black stripe along the lower jaw; superciliaries cream with transversal brown bars; dark brown stripe from postocular region to supratemporals; sides of neck dusty brown with reddish pigmentation; a black blotch covers the nuchal mite pocket; black line below antehumeral fold. Gular region and throat cream with reddish pigmentation on the sides and a faint pink blotch over the throat; chest and ventral surfaces of forelimbs dirty cream; first portion of venter, immediately after forelimb insertion, yellow; belly dirty cream flanked by pink pigmentation; pelvic region and ventral surfaces of thighs yellow; tail pale pink, with yellow pigmentation on the base (Fig. 11B). Irises light brown. Color of the holotype in ethanol 70%. Similar to coloration in life. However, flanks turned dusty brown as dorsum and the reddish coloration to the sides of gular region and ventral surface of neck turned brown (Fig. 1). Variation. Scale counts and measurements for Stenocercus diploauris are presented in Table 1. Loreals 4���8; supralabials 5���6; infralabials 5���6; postrostrals 2���4; second infralabial not in contact with third sublabial in 91% of specimens; first pair of postmentals in contact in 91% of specimens (n = 11). The nuchal mite pocket is conspicuous only in adult specimens, after preservation it becomes less defined. The type series of S. diploauris contains three adult males (CORBIDI 9916, 14904 and 14906) of 68.0 mm to 78.0 mm of SVL, and one juvenile male (CORBIDI 9914) with 61.0 mm of SVL. Two adult males (CORBIDI 9916 and 14906) are identical in dorsal coloration to the holotype. However, ventrally both specimens differ from the holotype by having the posterior portion of the gular region and ventral surface of neck gray and a gray midventral line on the belly. The dorsal coloration of one adult male (CORBIDI 14904) is similar to the holotype differing only by having the flanks dusty brown as the dorsum and not cinnamon as the holotype (Fig. 3A). Ventrally this specimen differs from the holotype by having the gular region cream and the ventral surface of neck gray, the chest and belly are cream with an indistinct light gray midventral line and a dark cream patch covering the pelvic region and the ventral surface of thighs (Fig. 3B). A juvenile male (CORBIDI 9914), has the dorsum grayish and the flanks dusty brown (Fig. 3C). Ventrally this specimen has the gular region, ventral surfaces of neck and chest cream with gray reticulations, and the belly cream with a faint gray midventral line (Fig. 3D). Sexual dimorphism is conspicuous with respect to size (maximum SVL in males 94.82 mm versus 76.0 mm in females) and coloration. Females have brown dorsal surfaces with a gray dorsolateral stripe, and pairs of dark brown triangular blotches along the vertebral line becoming chevrons on the tail (Fig. 3E); flanks are brown as dorsum or cinnamon (CORBIDI 14901) without marks; sides of head brown with a dark brown subocular stripe; jaw, ventrolateral region of head and sides of neck black (Fig. 3E). Ventral surfaces in adult females are brownish cream with a black patch covering the gular region and ventral surface of neck (Fig. 3F). One specimen (CORBIDI 14902) has the gular region and ventral surface of neck black with a cream blotch on the middle. One of the five female paratypes (CORBIDI 14672) has a dusty brown dorsum with dark brown chevrons along the vertebral line and the loreal and subocular regions bright cream like adult males. One juvenile female (CORBIDI 9913) differs from the adult females by having a light cream dorsolateral stripe (gray in adult females). Distribution and natural history observations. Stenocercus diploauris is known from four localities in the SDF of the Mantaro River Valley at elevations between 1,678 to 2,920 m asl, Huancavelica Department, Peru (Fig. 5). Stenocercus diploauris inhabits a seasonal dry forest, according to Linares-Palomino (2004) classification, and Central Andean Yungas according to Olson et al. (2001). Limonal (1,678 m asl), corresponds to the locality with the lowest elevation record of S. diploauris, its general landscape is that of a typical dry forest but with scattered crop lands with plantations of corn Zea mays, avocado pear Persea sp., citrus fruit trees and several other species of fruit trees. The agriculture is more intensive in the surrounding areas of Pichiu village (2,090 m asl); however, the remnants of native vegetation are dominated by shrubs, Opuntia and other cacti species. Chupto, at 2,328 m asl, is a very steep area dominated by shrubs and grasses. The locality of Jatuspata is located within an evergreen forest at elevations between 2,609 m and 2,920 m asl. The holotype of Stenocercus diploauris was collected by chance, it was found dead but not decomposed in a Sherman trap. Most paratypes were collected inactive under rocks, among cacti and shrubs, during cloudy days. Only CORBIDI 14672 was collected while active and moving among shrubs. Stenocercus diploauris and S. nigrobarbatus are known to co-occur at Chupto and Pichiu. Other species of Squamate reptiles collected at the same localities of S. diploauris include Ameiva reticulata, Mastigodryas boddaerti, Micrurus sp., and Wilsonosaura josyi. Etymology. The specific epithet ��� diploauris ��� is a noun (in apposition) in the nominative singular and derives from the Greek word diploos (= double) and the Latin word auris (= ear). It refers to the depression on both sides of the neck of the new species, appearing like a second tympanic opening., Published as part of Venegas, Pablo J., Echevarr��a, Lourdes Y., Garc��a-Ayachi, Luis A. & Landauro, Caroll Z., 2020, Two new sympatric species of Stenocercus (Squamata: Iguania) from the inter- Andean valley of the Mantaro River, Peru, pp. 555-575 in Zootaxa 4858 (4) on pages 557-562, DOI: 10.11646/zootaxa.4858.4.5, http://zenodo.org/record/4412698, {"references":["Torres-Carvajal, O. (2007 a) A taxonomic revision of South American Stenocercus (Squamata: Iguania) lizards. Herpetological Monographs, 21, 76 - 178. https: // doi. org / 10.1655 / 06 - 001.1","Torres-Carvajal, O. (2004) The abdominal skeleton of tropidurid lizards (Squamata: Tropiduridae). Herpetologica, 60, 75 - 83. https: // doi. org / 10.1655 / 03 - 15","Linares-Palomino, R. (2004) Los Bosques Tropicales Estacionalmente Secos: II. Fitogeografia y Composicion Floristica. Arnaldoa, 11 (1), 103 - 138.","Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N. D., Powell, G. V. N., Underwood, E. C., D'amico, J. A., Itoua, I., Strand, H. E., Morrison, J. C., Loucks, C. J., Allnutt, T. F., Ricketts, T. H., Kura, Y., Lamoreux, J. F., Wettengel, W. W., Hedao, P. & Kassem, K. R. (2001) Terrestrial Ecoregions of the World: A New Map of Life on Earth. BioScience, 51 (11), 933 - 938. https: // doi. org / 10.1641 / 0006 - 3568 (2001) 051 [0933: teotwa] 2.0. co; 2"]}

Stenocercus diploauris sp. nov. Figures 1–4, Table 1. Holotype. CORBIDI 13643, adult male from Limonal (12°13’55.097’’ S, 74°41’27.487’’ W), at 1,678 m asl, Surcubamba District, Tayacaja Province, Huancavelica Department, Peru, collected on September 1, 2013 by A. Escobar. Paratypes. Peru: Huancavelica Department: Tayacaja Province: CORBIDI 9913 (adult female), CORBIDI 9914 (juvenile male), CORBIDI 9915 (adult female), CORBIDI 9916 (adult male) from Jatuspata (12°15’1.2’’ S, 74°41’33.6’’ W), at 2,609 m asl, collected on April 7, 2011 by D. Amaya; CORBIDI 14672 adult female from Chupto (12°18’42.56’’ S, 74°39’13.23’’ W) at 2,328 m asl, collected on July 2, 2014 by L. Y. Echevarría; COR- BIDI 14901 adult female from Jatuspata (12°15’28.83’’ S, 74°41’28.74’’ W) at 2,816 m asl, collected on July 2, 2014 by C. Landauro; CORBIDI 14902 and CORBIDI 14903 adult females from Jatuspata (12°15’35.31’’ S, 74°41’31.74’’ W) at 2,835 m asl, collected on July 3, 2014 by C. Landauro; CORBIDI 14904 adult male from Jatuspata (12°15’34.23’’ S, 74°41’25.12’’ W) at 2,920 m asl, collected on July 3, 2014 by C. Landauro; CORBIDI 14906 adult male from Limonal (12°13’58.82’’ S, 74°41’23.09’’ W) at 1,753 m asl, collected on July 8, 2014 by C. Landauro; CORBIDI 16036 adult female from Pichiu (12°19’50’’ S, 74°39’13.07’’ W) at 2090 m asl, collected on July 12, 2015 by J. Malqui. Diagnosis. Adult specimens of Stenocercus diploauris can be easily distinguished from all known species of Stenocercus by having a C-shaped nuchal mite pocket around the oblique fold, posteriorly limited by the antehumeral fold (Fig. 4). However, among the 69 currently known species of Stenocercus, S. diploauris resembles S. formosus Tschudi, 1845, and S. ochoai Fritts, 1972, by the combination of the following characters: (1) imbricate scales on the posterior surface of thighs, (2) well developed postfemoral mite pocket, (3) antehumeral and oblique neck folds, and (4) absence of posthumeral mite pocket. The new species can be distinguished from S. ochoai and S. formosus by having dorsal and lateral nuchals similar in size and the presence of longitudinal neck fold, in the former species lateral nuchals are smaller than dorsal nuchals and the longitudinal neck fold is absent. Furthermore, S. diploauris differs from S. formosus (character state of latter in parenthesis) by having three whorls per autotomic segment (four), smooth dorsal head scales (keeled), and fewer scales, 50–61, around midbody (74–82). Characterization. (1) Maximum SVL in males 94.82 mm (n = 5); (2) maximum SVL in females 76.06 mm (n = 5); (3) vertebrals 40–50; (4) paravertebrals 63–78; (5) scales around midbody 50–61; (6) supraoculars 5–7; (7) internasals 3–4; (8) postrostrals 2–4; 9) loreals 4–8; (10) gulars 17–24; (11) lamellae on Finger IV 17–21; (12) lamellae on Toe IV 24–30; (13) posthumeral pocket absent; (14) postfemoral pocket distinct with slit-like opening, Type 2 of Torres-Carvajal (2007a); (15) parietal eye not visible; (16) occipital scales large, smooth, imbricate; (17) projecting angulate temporals absent; (18) enlarged supraoculars occupying most of supraocular region in one row; (19) scales on frontonasal region weakly imbricate anteriorly; (20) preauricular fringe present; (21) antehumeral, longitudinal and oblique neck folds present; (22) C-shaped nuchal mite pocket around the oblique neck fold, posteriorly limited by the antehumeral neck fold; (23) lateral nuchals slightly smaller than dorsal nuchals; (24) posterior gulars in adults smooth, imbricate, not mucronate, not notched; (25) lateral and dorsal body scales similar in size; (26) vertebral crest prominent; (27) dorsolateral crest absent; (28) ventrals in adults smooth, imbricate, mucronate; (29) scales on posterior surfaces of thighs keeled, imbricate, mucronate; (30) prefemoral fold absent; (31) inguinal groove absent; (32) preanals projected; (33) tail not strongly compressed laterally in adult males; (34) tail length 50–70% of total length; (35) three caudal whorls per autotomic segment; (36) caudals not spinose; (37) dark stripe extending anterodorsally from subocular region to supraciliaries present in most specimens; (38) gular region of adult females black or densely pigmented; (39) gular region of adult males gray or reddish; (40) black blotch on ventral surface of neck in adult males absent; (41) light gray or cream midventral line present; (42) black patch on ventral surface of thighs absent; (43) background color of dorsum, in females and males, brown; (44) two xiphisternal, and two long postxiphisternal pairs of inscriptional ribs that do not articulate midventrally (Pattern 1A of Torres-Carvajal 2004). Description of holotype. Male; SVL 94.82 mm; TL 217.5 mm; maximum head width 19.18 mm; head length 21.99 mm; head height 13.69 mm; occipitals, parietals, interparietal, and postparietals large, smooth, slightly imbricate (Fig. 2); parietal eye not visible; supraoculars in six rows, smooth, slightly imbricate, with one row two times larger than adjacent rows; anterior and posterior circumorbitals smooth, imbricate; canthals two; anteriormost canthal in contact with nasal; scales in frontonasal region not imbricate; internasals four; postrostrals four, wider than long, median ones smaller; supralabials six; infralabials six; loreals eight; lorilabials in one row; preocular not divid- ed, in contact with posterior canthal; lateral temporals imbricate, keeled; gulars in 22 rows between tympanic openings; all gulars smooth, imbricate; second infralabial in contact with first three sublabials; mental in contact with first pair of infralabials and first pair of postmentals; dorsal and lateral scales of body and neck keeled, imbricate; scales around midbody 53; vertebrals large, in 48 rows, forming a distinct serrate vertebral crest; paravertebrals 77; ventrals smooth, imbricate; preauricular fringe short, composed of two enlarged scales, of similar size; antehumeral, longitudinal and oblique neck folds present; a C-shaped nuchal mite pocket around the oblique fold, posteriorly limited by the antehumeral fold (Fig. 4); limb scales keeled, imbricate; ventral scales of hind limbs smooth; lamellae on Finger IV 20; lamellae on Toe IV 30; tail slightly compressed laterally; caudals keeled, imbricate; basal subcaudals keeled, imbricate; vertebral crest extending two thirds of tail length; tail length 2.3 times SVL; posthumeral pocket absent; postfemoral pocket shallow with slit-like opening (Type 2 of Torres-Carvajal 2007a). Color in life of the holotype. Dorsal surface dusty brown with light cream chevrons over the vertebral line, irregular light cream transversal bars on limbs and a reddish hue on tail; flanks cinnamon, dotted with yellowish cream scales; dorsal surface of head darker than dorsum and with some scattered cream dots; temporal surface of head dark brown, with scattered reddish pigmentation and white dots; loreal region and canthus rostralis dark brown and subocular region, supralabials and infralabials bright cream; sublabials forming a thin black stripe along the lower jaw; superciliaries cream with transversal brown bars; dark brown stripe from postocular region to supratemporals; sides of neck dusty brown with reddish pigmentation; a black blotch covers the nuchal mite pocket; black line below antehumeral fold. Gular region and throat cream with reddish pigmentation on the sides and a faint pink blotch over the throat; chest and ventral surfaces of forelimbs dirty cream; first portion of venter, immediately after forelimb insertion, yellow; belly dirty cream flanked by pink pigmentation; pelvic region and ventral surfaces of thighs yellow; tail pale pink, with yellow pigmentation on the base (Fig. 11B). Irises light brown. Color of the holotype in ethanol 70%. Similar to coloration in life. However, flanks turned dusty brown as dorsum and the reddish coloration to the sides of gular region and ventral surface of neck turned brown (Fig. 1). Variation. Scale counts and measurements for Stenocercus diploauris are presented in Table 1. Loreals 4–8; supralabials 5–6; infralabials 5–6; postrostrals 2–4; second infralabial not in contact with third sublabial in 91% of specimens; first pair of postmentals in contact in 91% of specimens (n = 11). The nuchal mite pocket is conspicuous only in adult specimens, after preservation it becomes less defined. The type series of S. diploauris contains three adult males (CORBIDI 9916, 14904 and 14906) of 68.0 mm to 78.0 mm of SVL, and one juvenile male (CORBIDI 9914) with 61.0 mm of SVL. Two adult males (CORBIDI 9916 and 14906) are identical in dorsal coloration to the holotype. However, ventrally both specimens differ from the holotype by having the posterior portion of the gular region and ventral surface of neck gray and a gray midventral line on the belly. The dorsal coloration of one adult male (CORBIDI 14904) is similar to the holotype differing only by having the flanks dusty brown as the dorsum and not cinnamon as the holotype (Fig. 3A). Ventrally this specimen differs from the holotype by having the gular region cream and the ventral surface of neck gray, the chest and belly are cream with an indistinct light gray midventral line and a dark cream patch covering the pelvic region and the ventral surface of thighs (Fig. 3B). A juvenile male (CORBIDI 9914), has the dorsum grayish and the flanks dusty brown (Fig. 3C). Ventrally this specimen has the gular region, ventral surfaces of neck and chest cream with gray reticulations, and the belly cream with a faint gray midventral line (Fig. 3D). Sexual dimorphism is conspicuous with respect to size (maximum SVL in males 94.82 mm versus 76.0 mm in females) and coloration. Females have brown dorsal surfaces with a gray dorsolateral stripe, and pairs of dark brown triangular blotches along the vertebral line becoming chevrons on the tail (Fig. 3E); flanks are brown as dorsum or cinnamon (CORBIDI 14901) without marks; sides of head brown with a dark brown subocular stripe; jaw, ventrolateral region of head and sides of neck black (Fig. 3E). Ventral surfaces in adult females are brownish cream with a black patch covering the gular region and ventral surface of neck (Fig. 3F). One specimen (CORBIDI 14902) has the gular region and ventral surface of neck black with a cream blotch on the middle. One of the five female paratypes (CORBIDI 14672) has a dusty brown dorsum with dark brown chevrons along the vertebral line and the loreal and subocular regions bright cream like adult males. One juvenile female (CORBIDI 9913) differs from the adult females by having a light cream dorsolateral stripe (gray in adult females). Distribution and natural history observations. Stenocercus diploauris is known from four localities in the SDF of the Mantaro River Valley at elevations between 1,678 to 2,920 m asl, Huancavelica Department, Peru (Fig. 5). Stenocercus diploauris inhabits a seasonal dry forest, according to Linares-Palomino (2004) classification, and Central Andean Yungas according to Olson et al. (2001). Limonal (1,678 m asl), corresponds to the locality with the lowest elevation record of S. diploauris, its general landscape is that of a typical dry forest but with scattered crop lands with plantations of corn Zea mays, avocado pear Persea sp., citrus fruit trees and several other species of fruit trees. The agriculture is more intensive in the surrounding areas of Pichiu village (2,090 m asl); however, the remnants of native vegetation are dominated by shrubs, Opuntia and other cacti species. Chupto, at 2,328 m asl, is a very steep area dominated by shrubs and grasses. The locality of Jatuspata is located within an evergreen forest at elevations between 2,609 m and 2,920 m asl. The holotype of Stenocercus diploauris was collected by chance, it was found dead but not decomposed in a Sherman trap. Most paratypes were collected inactive under rocks, among cacti and shrubs, during cloudy days. Only CORBIDI 14672 was collected while active and moving among shrubs. Stenocercus diploauris and S. nigrobarbatus are known to co-occur at Chupto and Pichiu. Other species of Squamate reptiles collected at the same localities of S. diploauris include Ameiva reticulata, Mastigodryas boddaerti, Micrurus sp., and Wilsonosaura josyi. Etymology. The specific epithet “ diploauris ” is a noun (in apposition) in the nominative singular and derives from the Greek word diploos (= double) and the Latin word auris (= ear). It refers to the depression on both sides of the neck of the new species, appearing like a second tympanic opening.

Stenocercus nigrobarbatus sp. nov. Figures 6���10, Table 1. Holotype. CORBIDI 13725, adult male from Fundici��n (12��17���36.41���S, 74��39���39.67��� W) at 2,158 m asl, Tayacaja Province, Huancavelica Region, Peru, collected on January 31, 2014 by L. Y. Echevarr��a. Paratypes. Peru: Huancavelica Department: Tayacaja Province: CORBIDI 13637 a juvenile female from Jatuspata (12��15���14.617������ S, 74��41��� 16.93������ W) at 2,972 m asl, collected on September 3, 2013 by C. Landauro; CORBIDI 13638 a juvenile female from Pichiu (12��19���11.815������ S, 74��39���3.588������ W) at 2,036 m asl, collected on September 13, 2013 by C. Landauro; CORBIDI 13640 juvenile female from Pichiu, collected on September 12, 2013 by C. Landauro; CORBIDI 13641 adult female from Barropata (12��17���21.943������ S, 74��41���19.035������ W) at 1,725 m asl, collected on October 2, 2013 by C. Landauro; CORBIDI 13639 and CORBIDI 13642 female juveniles from Fundici��n (12��17���32.347������ S, 74��39���45.433������ W), at 2,169 m asl, collected on September 28, 2013 by C. Landauro; CORBIDI 13718 adult male from Chupto (12��19���52.96��� S, 74��39���13��� W) at 2,058 m asl, collected on January 23, 2014 by L. Y. Echevarr��a; CORBIDI 13719 adult male from Chupto (12��18���22.19��� S, 74��39���18.42��� W) at 2,314 m asl, on January 24, 2014 by L. Y. Echevarr��a; CORBIDI 13722 adult female from Chupto (12��19���52.96��� S, 74��39���13��� W) at 2,058 m asl, collected on January 21, 2014 by L. Y. Echevarr��a; CORBIDI 13728 adult female from Barropata (12��17���34.95��� S, 74��41���34.08��� W) at 1,704 m asl, collected on February 2, 2014 by L. Y. Echevarr��a; CORBIDI 13729 adult female from Barropata (12��17���22.11��� S, 74��41���19.07��� W) at 1,783 m asl, collected on February 3, 2014 by L. Y. Echevarr��a; CORBIDI 14669 adult female from Chupto (12��18���48.18��� S, 74��39���1.83��� W) at 2,345 m asl, collected on July, 1 2014 by L. Y. Echevarr��a; CORBIDI 14674 adult female from type locality, collected on July 6, 2014 by L. Y. Echevarr��a; CORBIDI 14676 juvenile female and CORBIDI 14677 adult male from Barropata (12��17���37.66��� S, 74��41���39.18��� W) at 1,693 m asl, collected on July 7, 2014 by L. Y. Echevarr��a; CORBIDI 15765���66 (adult male and female, respectively) and CORBIDI 15767 (juvenile male) from Pichiu (12��15���31.57��� S, 75��6���3.6��� W) at 2,206 m asl, collected on January 14, 2015 by L. Lujan; CORBIDI 16033 adult male from Pichiu (12��19���50��� S, 74��39���13.07��� W) at 2,090 m asl, collected on July 9, 2015 by J. Malqui; CORBIDI 20546���47 two adult males from San Luis de Estanque (12��19���8.62��� S, 74��44���24.19��� W) at 2,636 m asl., collected on Jun 17, 2019 by L.A. Garc��a-Ayachi. Diagnosis. Adult males of Stenocercus nigrobarbatus can be easily distinguished from all species of Stenocercus by having a continuous black patch covering the infralabials, throat, chest, ventral surfaces of forelimbs, belly (as a black midventral line), ventral surfaces of hind limbs, and pelvic region (Figs. 6B, 8B, 9B and 9C). However, among the 69 currently described species of Stenocercus, S. nigrobarbatus resembles S. frittsi Torres-Carvajal, 2005, and S. variabilis Boulenger, 1901, by the following combinations of characters: (1) granular scales on the posterior surface of thighs, (2) imbricate and keeled lateral body scales, (3) a distinct row of enlarged vertebral scales, (4) unnotched gular scales, (5) three caudal whorls per autotomic segment, (6) brown dorsal ground color, and (7) distinct oblique neck fold, and antegular fold not continuous medially. Stenocercus nigrobarbatus differs from the geographically close S. frittsi (character state of latter in parenthesis) by having a postfemoral mite pocket composed by one or more vertical folds or ridges, Type 1 of Torres-Carvajal (2007a) (postfemoral pocket absent in S. frittsi), 2���4 postrostrals (5���7), and two pairs of postxiphisternal inscriptional ribs, long, not in contact midventrally, pattern 1A of Torres-Carvajal (2004) (three pairs of postxiphisternal inscriptional ribs, two pairs long and the last pair short, not in contact midventrally; pattern 2B of Torres-Carvajal 2004). Stenocercus nigrobarbatus can be distinguished from S. variabilis (character state in parenthesis) by having 2���4 postrostrals (6), postfemoral pocket composed by one or more vertical folds or ridges, Type 1 of Torres-Carvajal (2007a) (slit-like opening, Type 2 of Torres-Carvajal 2007a), and inguinal groove absent (present). In addition, lateral body scales in S. nigrobarbatus are slightly smaller than dorsals, while S. frittsi and S. variabilis have lateral body scales approximately half the size of dorsals. Characterization. (1) Maximum SVL in males 67.73 mm (n = 4); (2) maximum SVL in females 62.29 mm (n = 5); (3) vertebrals 47���73; (4) paravertebrals 69���87; (5) scales around midbody 57���74; (6) supraoculars 5���7; (7) internasals 3���5; (8) postrostrals 2���4; (9) loreals 3���7; (10) gulars 19���30; (11) lamellae on Finger IV 18���23; (12) lamellae on Toe IV 27���34; (13) posthumeral pocket as one or more vertical folds or ridges, Type 1 of Torres-Carvajal (2007a); (14) postfemoral pocket composed by one or more vertical folds or ridges, Type 1 of Torres-Carvajal (2007a); (15) parietal eye not visible; (16) occipital scales small, smooth, juxtaposed; (17) no projecting angulate temporals; (18) supraoculars of varying sizes; (19) scales in frontonasal region slightly imbricate anteriorly; (20) short preauricular fringe present; (21) antehumeral, antegular, gular, oblique, suprauricular and postauricular neck folds present; (22) nuchal mite pocket absent; (23) lateral nuchals less than half size of dorsal nuchals; (24) posterior gulars in adults smooth, imbricate, not mucronate; (25) lateral scales slightly smaller than dorsal body scales; (26) vertebrals larger than adjacent paravertebrals; (27) dorsolateral crest absent; (28) ventrals in adults smooth, imbricate, not mucronate; (29) scales on posterior surfaces of thighs granular; (30) inguinal granular pocket absent; (31) inguinal groove absent; (32) preanals not projecting; (33) tail not compressed laterally in adult males; (34) tail length 62���70% of total length; (35) three caudal whorls per autotomic segment; (36) caudals not spinose; (37) dark stripe extending anterodorsally from subocular region to supraciliaries present; (38) gular region of adult females dark or densely pigmented in all specimens; (39) gular region of adult males dark or densely pigmented in all specimens; (40) black blotch on ventral surface of neck in adult males absent; (41) bold black midventral line present in males; (42) black patch on ventral surface of thighs present in males; (43) dorsal ground color brown, in females and males; and (44) postxiphisternal inscriptional ribs not articulating midventrally, both pairs long, Pattern 1A of Torres-Carvajal (2004). Description of holotype. Male (Fig. 6-8); SVL= 66.02 mm; TL = 130.0 mm; maximum head width = 12.24; head length = 15.87; head height = 8.63; scales on parietal and occipital regions small, smooth, juxtaposed; parietal eye not visible; supraoculars in six rows, smooth, slightly imbricate, of varying sizes; canthals two; anterior most canthal separated from nasal by one minute scale; scales in frontonasal region slightly imbricate; internasals four; postrostrals three, subequal in size, wider than long; supralabials seven; infralabials seven; loreals six; lorilabials in one row; preocular not divided, in contact with posterior canthal; lateral temporals imbricate, weakly keeled; gulars in 29 rows between tympanic openings; all gulars smooth, imbricate; second infralabial in contact with first two sublabials; mental in contact with first pair of infralabials and first pair of postmentals; lateral and dorsal scales of body and dorsal scales of neck keeled, imbricate; lateral scales of neck granular; scales around midbody 63; vertebrals large, in 68 rows, not forming a prominent serrate vertebral crest; paravertebrals 86; ventrals smooth, imbricate; preauricular fringe short, composed of two enlarged scales, both same size; antehumeral, antegular, gular, oblique, suprauricular, postauricular neck folds and ventrolateral body fold present; limb scales keeled, imbricate; ventral scales of hind limbs and upper arms smooth; lamellae on Finger IV 23; lamellae on Toe IV 31; tail not compressed laterally; caudals keeled, imbricate; basal subcaudals smooth, imbricate; vertebral crest extending more than half length of tail; tail length 1.9 times SVL; posthumeral pocket shallow (Type 1 Torres-Carvajal 2007a) with vertically oriented fold approximately 2.57 mm long; postfemoral pocket composed by one vertical fold (type 1 of Torres-Carvajal 2007a); postfemoral region composed of imbricate, smooth scales, becoming smaller toward insertion of hind limbs (Fig. 10A). Color in life of the holotype. Dorsal surfaces of head and neck cinnamon brown with yellow spots, denser on neck; lorilabials and supralabials yellow; infralabials black; ventrolateral regions of head and neck black; dorsal surfaces of body, limbs and tail dusty brown with pale yellow spots; flanks light blue, densely covered with cream spots; limbs and tail with thin dark brown marks on dorsal surfaces (Fig. 8A). Ventral surface of head, throat, chest black; ventral surfaces of forelimbs black, although pigmentation is faint at elbow joint; a bold black midventral line, continuous with chest pigmentation, flanked by pink; ventral surfaces of hind limbs, and pelvic region black, except by the cloaca; posterior surfaces of thighs pink; pre-cloacal region dark yellow and post post-cloacal region pink; ventral surface of tail pink, with a yellow blotch and blackish speckles at the base (Fig. 8B). All black pigmentation on ventral surfaces constitutes a continuous patch, except black pigmentation on tail. Coloration of holotype in ethanol 70%. Dorsally, all yellow spots turned to pale yellow. Dorsal surfaces of head and neck turned brown. Ventrally, pink and yellow pigmentation turned to a less vivid (pale) shade, almost vanished on cloaca. Variation. Scale counts and measurements for Stenocercus nigrobarbatus are presented in Table 1. Loreals 3���7; supralabials 6���8; infralabials 5���8; postrostrals 2���4; second infralabial not in contact third sublabial in all specimens; first pair of postmentals in contact in all specimens. Dorsal coloration in adult males of Stenocercus nigrobarbatus is the same in all adult male paratypes (n = 5) (Fig. 9A). The black coloration on ventral surfaces as a continuous patch is exclusive to adult males (Fig. 9 B��� C). The juvenile specimen CORBIDI 13718 has a faint black midventral line on the belly (Fig. 9 D���E). Another juvenile specimen (CORBIDI 13719) has the gular region and ventral surface of neck covered by a dark gray patch and the midventral region of the belly has scattered dark gray dots (Fig. 9 F���G). These immature males (CORBIDI 13718 and 13719) have the pelvic region and the base of tail yellow and the ventral surface of forelimbs cream (Fig. 9 D���G). In both specimens the pink coloration on the flanks of the belly is also less intense than in the holotype. In the adult male paratype (CORBIDI 20547) the base of the tail has a black blotch and the pre-cloacal region is yellow with black speckles (Fig. 9C). Sexual dimorphism is evident in coloration and size (maximum SVL in males 67.0 mm versus 62.0 mm in females). Dorsal coloration in adult females can be gray or brown with irregular dark brown blotches over the vertebral line and a light brown dorsolateral stripe from the temporal region to the base of tail (Fig. 9H); dorsal surface of limbs has dark brown or gray transversal bars; sides of head brown or gray with a thin postocular stripe and some scattered dark brown dots; sides of neck yellowish brown with light yellow flecks; jaw and ventrolateral region of neck black or dark gray; body flanks covered by a dark brown reticulation. Ventrally all adult females have the gular region and neck dark gray, belly and limbs are light cream, and the tail is light orange (Fig. 9I). Juvenile females, between 43.0���51.0 mm of SVL, have the same dorsal and ventral pattern than adult females, only two specimens (CORBIDI 13637 and 13642) have the gular region gray with dark gray speckles, and chest and belly light cream with gray speckles. The smallest juvenile female (CORBIDI 14676) has the gular region covered by gray flecks. The juvenile male CORBIDI 13640 (SVL = 43.0 mm) has the dorsum with dark brown dorsal marks as adult and juvenile females, however, the gular region and chest are gray and the belly is grayish cream. The juvenile male CORBIDI 15767 (SVL = 53.0 mm) has the same dorsal coloration as adult males but ventrally the gular region and chest are gray, and has a faint gray midventral line on belly flanked by a weak pink hue; pelvic region, ventral surface of hindlimbs and tail are light cream. Distribution and natural history observations. Stenocercus nigrobarbatus is known from four localities in SDF of the Mantaro River Valley at elevations between 1,693 and 2,345 m asl, Huancavelica Department, Peru (Fig. 5). Stenocercus nigrobarbatus inhabits a seasonal dry forest, according to classification of Linares-Palomino (2004), and Central Andean Yungas according to Olson et al. (2001). In most localities the habitat has been modified by crops of Zea mays, Opuntia, and Persea sp., roads and the construction of a hydroelectric plant. However, this species is very conspicuous in most localities. Specimens of Stenocercus nigrobarbatus were found inactive under rocks during cloudy days and active, basking or running, during warm days, among rocks, cacti or shrubs. Other species of Squamate reptiles collected at the same localities of S. nigrobarbatus include Ameiva reticulata, Epictia sp., Mastigodryas boddaerti, Micrurus sp., and Oxyrhopus cf. erdisii, and Stenocercus diploauris. Etymology. The specific epithet ��� nigrobarbatus ��� is a noun derived from the Latin words ��� nigro ��� (= black) and ��� barbatus ��� (= bearded). It refers to the black patch covering the gular region and ventral surface of neck, including the jaw angle, of adult males which resembles a black beard., Published as part of Venegas, Pablo J., Echevarr��a, Lourdes Y., Garc��a-Ayachi, Luis A. & Landauro, Caroll Z., 2020, Two new sympatric species of Stenocercus (Squamata: Iguania) from the inter- Andean valley of the Mantaro River, Peru, pp. 555-575 in Zootaxa 4858 (4) on pages 564-571, DOI: 10.11646/zootaxa.4858.4.5, http://zenodo.org/record/4412698, {"references":["Torres-Carvajal, O. (2005) New Species of Stenocercus (Squamata: Iguania) from the Andes of Central Peru with a Redescription of Stenocercus variabilis. Journal of Herpetology, 39 (3), 471 - 477. https: // doi. org / 10.1670 / 26 - 05 a. 1","Torres-Carvajal, O. (2007 a) A taxonomic revision of South American Stenocercus (Squamata: Iguania) lizards. Herpetological Monographs, 21, 76 - 178. https: // doi. org / 10.1655 / 06 - 001.1","Torres-Carvajal, O. (2004) The abdominal skeleton of tropidurid lizards (Squamata: Tropiduridae). Herpetologica, 60, 75 - 83. https: // doi. org / 10.1655 / 03 - 15","Linares-Palomino, R. (2004) Los Bosques Tropicales Estacionalmente Secos: II. Fitogeografia y Composicion Floristica. Arnaldoa, 11 (1), 103 - 138.","Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N. D., Powell, G. V. N., Underwood, E. C., D'amico, J. A., Itoua, I., Strand, H. E., Morrison, J. C., Loucks, C. J., Allnutt, T. F., Ricketts, T. H., Kura, Y., Lamoreux, J. F., Wettengel, W. W., Hedao, P. & Kassem, K. R. (2001) Terrestrial Ecoregions of the World: A New Map of Life on Earth. BioScience, 51 (11), 933 - 938. https: // doi. org / 10.1641 / 0006 - 3568 (2001) 051 [0933: teotwa] 2.0. co; 2"]}

Phrynopus mariellaleo sp. nov. FIGURES 2–4. Holotype. CORBIDI 11668, AN ADULT fEMALE, fROM SEDRUSCO, VISTA ALEGRE DISTRICT, RODRÍGUEZ DE MENDOZA PROVINCE, DEPARTAMENTO AMAZONAS, PERU (6°6’42.9’’S, 77°26’24’’W, 2575 M ASL.; FIG. 5) ObTAINED ON 3 AUGUST 2012 bY PAbLO J. VENEGAS AND VILMA DURáN. Paratypes. SIX fEMALES (CORBIDI 11669, 11657, 11692, 11707, 11796, 11799), fIVE MALES (CORBIDI 11658, 11797–98, 11800–01, AND fOUR jUVENILES (CORBIDI 11670–71, 11802, 11804), ALL COLLECTED AT THE SAME LOCATION AS THE HOLOTYPE bETWEEN 1–10 AUGUST 2012 bY PAbLO J. VENEGAS AND VILMA DURáN. Diagnosis. (1) SKIN ON DORSUM SHAGREEN WITH SCATTERED LOW TUbERCLES, “V” OR “Y” SHAPED IRREGULAR (RIDGED) fOLDS IN SCAPULAR REGION, AND WITH OR WITHOUT A WEAK MIDDORSAL RIDGE; SKIN ON fLANKS SMOOTH WITH LOW fLAT WARTS; SKIN ON VENTER SMOOTH; DISCOIDAL fOLD WEAK, THORACIC fOLD PRESENT; DORSOLATERAL fOLDS PRESENT, COMPLETE; (2) TYMPANIC MEMbRANE AND TYMPANIC ANNULUS PRESENT AND PROMINENT; SUPRATYMPANIC fOLD PROMINENT, THICK AND LONG, EXTENDING fROM THE POSTERIOR MARGIN Of THE EYELID TO bEHIND THE ARM INSERTION; (3) SNOUT ROUNDED IN DORSAL AND LATERAL VIEWS; (4) UPPER EYELID bEARING SMALL, LOW, ROUND TUbERCLES; UPPER EYELID WIDTH SLIGHTLY NARROWER THAN IOD; CRANIAL CRESTS AbSENT; (5) DENTIGEROUS PROCESSES Of VOMERS PROMINENT, ObLIqUE, NARROWLY SEPARATED, SITUATED POSTEROMEDIAL TO CHOANAE; (6) VOCAL SLITS AND NUPTIAL PADS AbSENT; (7) FINGERS I AND II EqUAL IN LENGTH; DISCS ON fINGERS NARROW, ROUNDED, bEARING WEAK CIRCUMfERENTIAL GROOVES; (8) fINGERS LACKING LATERAL fRINGES; (9) ULNAR TUbERCLES AbSENT; (10) HEEL AND OUTER EDGE TARSUS WITHOUT TUbERCLES; INNER TARSAL fOLD PRESENT, SHORT, ObLIqUE; (11) INNER METATARSAL TUbERCLE PROMINENT, ELLIPTICAL, THREE TIMES LARGER THAN THE LOWER, OVAL OUTER METATARSAL TUbERCLE; PLANTAR SURfACE SMOOTH; (12) TOES LACKING LATERAL fRINGES; WEbbING AbSENT; TOE III AND TOE V AbOUT EqUAL IN SIZE; TOE DISCS NARROW, ROUND, bEARING WEAK MARGINAL GROOVES, EqUAL IN SIZE AS THOSE ON fINGERS; (13) DORSAL SURfACES LIGHT TO DARK bROWN WITH DARK bROWN TO bOLD bLACK bLOTCHES, WITH A DARK INTERORbITAL bAR, A PROMINENT DARK MIDDORSAL SPOT, AND DARK DIAGONAL bARS ON LIMbS; SIDES Of HEAD PALE TO DARK bROWN OR bLACK; CANTHUS ROSTRALIS AND TYMPANIC fOLD bOLD bLACK, OUTLINED AbOVE bY A CONTINUOUS THIN AND LIGHT STRIPE THAT RUNS fROM THE SNOUT TO THE END Of THE TYMPANIC fOLD bEHIND THE ARM; fLANKS Of SAME COLOR AS DORSUM OR SLIGHTLY LIGHTER THAN DORSUM, WITH SCATTERED DARK bROWN bLOTCHES OR DOTS AND WITH OR WITHOUT WHITE fLECKS; GROIN bROWN TO REDDISH bROWN, TRANSLUCENT, WITH DIffUSE bROWN SPOTS; VENTRAL SURfACE PALE PINK, PINKISH bROWN OR bROWN WITH DARK bROWN MOTTLING, ESPECIALLY ON THE THROAT, OR DARK bROWN WITH WHITE RETICULATIONS ON THE bELLY; (14) SVL 23.4–30.0 MM (x̅ = 27.7; n = 7) IN ADULTS MALES, 25.4–39.7 MM IN ADULT fEMALES (x̅ = 34.4; n = 8). Comparison with other species. Phrynopus mariellaleo DIffERS fROM ALL OTHER SPECIES Of Phrynopus EXCEPT P. auriculatus DUELLMAN AND HEDGES, AND P. peruanus PETERS, bY HAVING A DISTINCT TYMPANIC MEMbRANE AND TYMPANIC ANNULUS. Phrynopus auriculatus AND P. peruanus CAN bE DISTINGUISHED fROM THE NEW SPECIES bY HAVING FINGER I LONGER THAN FINGER II (EqUAL IN SIZE IN THE NEW SPECIES). ADDITIONALLY, P. auriculatus IS bY fAR SMALLER THAN P. mariellaleo sp. nov. [MAXIMUM SVL 13.4 MM IN MALES AND 14.5 MM IN fEMALES Of P. auriculatus VS. SVL Of 23.4– 30.0 MM (x̅ = 27.7; n = 7) IN MALES AND 25.4–39.7 MM (x̅ = 34.4; n = 8) IN fEMALES Of P. mariellaleo sp. nov.]. FURTHERMORE, THE bELLY Of P. mariellaleo sp. nov. IS SMOOTH, WHILE ALL OTHER SPECIES Of Phrynopus HAVE EITHER AREOLATE VENTER (P. anancites RODRÍGUEZ AND CATENAZZI, P. barthlenae LEHR AND AGUILAR, P. bufoides LEHR, LUNDbERG, AND AGUILAR, P. capitalis RODRÍGUEZ AND CATENAZZI, P. chaparroi MAMANI AND MALqUI, P. daemon CHáVEZ, SANTA- CRUZ, RODRÍGUEZ, AND LEHR, P. dagmarae LEHR, AGUILAR, AND KöHLER, P. dumicola RODRÍGUEZ AND CATENAZZI, P. heimorum LEHR, P. horstpauli LEHR, KöHLER, AND PONCE, P. kotosh LEHR, P. lapidoides LEHR AND RODRÍGUEZ, P. miroslawae CHAPARRO, PADIAL, AND DE LA RIVA, P. montium SHREVE, P. paucari LEHR, LUNDbERG, AND AGUILAR, P. peruanus, P. pesantesi LEHR, LUNDbERG, AND AGUILAR, P. tautzorum LEHR AND AGUILAR, P. thompsoni DUELLMAN, P. unchog LEHR AND RODRÍGUEZ, AND P. vestigiatus LEHR AND OROZ) OR fINELY OR WEAKLY AREOLATE VENTER (P. badius LEHR, MORAVEC AND CUSI, P. interstinctus LEHR AND OROZ, P. lechriorhynchus TRUEb AND LEHR, P. oblivius LEHR, P. personatus RODRÍGUEZ AND CATENAZZI, AND P. tribulosus DUELLMAN AND HEDGES). Phrynopus mariellaleo sp. nov. IS MORPHOLOGICALLY VERY SIMILAR TO SPECIES Of Hypodactylus. HOWEVER, IT IS EASILY DISTINGUISHED fROM ALL SPECIES Of Hypodactylus bY HAVING A PROMINENT, bROAD, AND LONG SUPRATYMPANIC fOLD EXTENDING fROM bEHIND THE EYELID TO THE POSTERIOR END Of THE ARM INSERTION, AND A bOLD bLACK fACEMASK. THERE ARE CURRENTLY fOUR SPECIES Of Hypodactylus RECORDED IN PERU (H. araiodactylus DUELLMAN AND PRAMUK, H. fallaciosus DUELLMAN, H. lucida CANNATELLA, AND H. nigrovittatus ANDERSSON) (DUELLMAN & LEHR 2009). THE GEOGRAPHICALLY CLOSER SPECIES TO P. mariellaleo sp. nov. ARE H. fallaciosus AND H. araiodactylus, bOTH fROM AMAZONAS DEPARTMENT IN THE NORTHERN PART Of THE CORDILLERA CENTRAL IN NORTHERN PERU (DUELLMAN & LEHR 2009). Hypodactylus araiodactylus IS ONLY KNOWN bY A SINGLE SPECIMEN COLLECTED IN 1972 fOR WHICH THERE ARE NO PICTURES OR NOTES AbOUT ITS COLORATION IN LIfE (DUELLMAN AND PRAMUK 1999). LIKE P. mariellaleo sp. nov., H. araiodactylus HAS DORSOLATERAL fOLDS AND FINGER I EqUALS IN SIZE FINGER II. HOWEVER, P. mariellaleo DIffERS fROM H. araiodactylus bY LACKING LATERAL fRINGES ON fINGERS AND TOES (PRESENT IN H. araiodactylus) AND bY HAVING A “V” OR “Y” SHAPED fOLD IN THE SCAPULAR REGION. Phrynopus mariellaleo sp. nov. ALSO CAN bE DISTINGUISHED fROM OTHER PERUVIAN SPECIES Of Hypodactylus, LIKE H. fallaciosus, H. lucida AND H. nigrovittatus, bY HAVING COMPLETE DORSOLATERAL fOLDS, WHICH ARE INCOMPLETE IN H. lucida AND AbSENT IN THE REST Of SPECIES. Hypodactylus lucida HAS A SMALL TYMPANIC MEMbRANE AND A SHORT SUPRATYMPANIC fOLD, AND LACKS MIDDORSAL fOLDS. Hypodactylus nigrovittatus HAS A SMALL, bARELY PERCEPTIbLE TYMPANIC MEMbRANE AND AN INCONSPICUOUS SUPRATYMPANIC fOLD, AND EXTREMELY SHORT HEAD WITH SUbACUMINATE SNOUT IN DORSAL VIEW (bLUNTLY ROUNDED IN P. mariellaleo sp. nov.). Description of the holotype. ADULT fEMALE WITH RObUST bODY; HEAD SLIGHTLY NARROWER THAN bODY, WIDER THAN LONG; HW 41 % Of SVL; HL 36 % Of SVL; SNOUT SHORT, LACKING TERMINAL TUbERCLE, bLUNTLY ROUNDED IN DORSAL AND LATERAL VIEWS; E–N 73 % Of EYE DIAMETER; NOSTRILS ROUNDED, DIRECTED DORSOLATERALLY; CANTHUS ROSTRALIS SLIGHTLY CURVED IN DORSAL VIEW, CURVED IN PROfILE; LOREAL REGION CONCAVE; LIPS ROUNDED; UPPER EYELID WITH fEW fLATTENED LITTLE TUbERCLES; EW 70 % Of IOD; TYMPANIC ANNULUS PRESENT; TYMPANIC MEMbRANE DISTINCT WITH DORSAL MARGIN ObSCURED bY SUPRATYMPANIC fOLD; SUPRATYMPANIC fOLD PROMINENT, EXTENDING fROM EYELIDS TO bEHIND ARM INSERTION; TY 35 % Of EYE DIAMETER, ED AbOUT 1.5 TIMES TYMPANUM DIAMETER; SEVERAL MINUTE POSTRICTAL TUbERCLES PRESENT. CHOANAE SMALL, OVOID, PARTLY CONCEALED bY PALATAL SHELf Of MAXILLA; DENTIGEROUS PROCESSES Of VOMERS PROMINENT, ObLIqUE, NARROWLY SEPARATED, WITH INDISTINCT NUMbER Of TEETH, SITUATED POSTEROMEDIAL TO CHOANAE; TONGUE AS LONG AS WIDE, NOTCHED bEHIND, HALf Of ITS LENGTH fREE POSTERIORLY. SKIN ON HEAD SMOOTH WITH SOME SCATTERED MINUTE TUbERCLES fROM THE INTERORbITAL REGION TO TIP Of THE SNOUT; DORSUM SMOOTH WITH A WEAKLY DEfINED MIDDORSAL fOLD; “V” OR “Y” SHAPED SCAPULAR fOLD PRESENT; DORSOLATERAL fOLD PRESENT, PROMINENT; SKIN ON fLANKS WITH SEVERAL fLATTENED TUbERCLES; SKIN ON THROAT, CHEST AND VENTER SMOOTH; DISCOIDAL fOLD AbSENT; CLOACAL SHEATH SHORT; CLOACAL REGION TUbERCULATE. ULNAR TUbERCLES COALESCED INTO A LOW fOLD; PALMAR TUbERCLES fLESHY, ELEVATED, OUTER PALMAR TUbERCLE bIfID, APPROXIMATELY TWICE SIZE Of OVOID, THENAR TUbERCLE; DISTAL SUbARTICULAR TUbERCLES WEAKLY DEfINED, EXCEPT PROXIMALLY, ROUND IN VENTRAL AND LATERAL VIEWS; SUPERNUMERARY TUbERCLE AT bASES Of FINGERS II AND III INDISTINCT; fINGERS LACKING LATERAL fRINGES; FINGER I AND FINGER II EqUAL IN SIZE; DISCS ON fINGERS NARROW, WITH ROUNDED TIPS; PADS ON fINGERS bEARING WEAK CIRCUMfERENTIAL GROOVES. HIND LIMbS SLENDER, TL 55 % Of SVL; FL 53 % Of SVL; UPPER SURfACES Of HIND LIMbS AREOLATE; POSTERIOR AND VENTRAL SURfACES Of THIGHS SMOOTH; HEEL LACKING TUbERCLES; OUTER SURfACE Of TARSUS LACKING TUbERCLES; SHORT INNER TARSAL fOLD PRESENT; INNER METATARSAL TUbERCLE PROMINENT, ELLIPTICAL, THREE TIMES bIGGER THAN SIZE Of LOW, INDISTINCT OUTER METATARSAL TUbERCLE; PLANTAR SURfACE SMOOTH WITH SOME INDISTINCT SUPERNUMERARY TUbERCLES; SUbARTICULAR TUbERCLES WELL DEfINED, ROUND IN VENTRAL AND LATERAL VIEWS; TOES LACKING LATERAL fRINGES; WEbbING AbSENT; DISCS ON TOES NARROW WITH ROUNDED TIPS, LIKE THOSE ON fINGERS; TOES LACKING CIRCUMfERENTIAL GROOVES; RELATIVE LENGTHS Of TOES: 1 Measurements of the holotype (in mm). SVL, 39.7; TL, 22.0; FL, 21.3; HL, 14.4; HW, 16.6; ED, 4.5; TY, 1.6; IOD, 5.1; EW 3.6; IND, 4.1; E–N, 3.3. Color of holotype in life. DORSALLY, HEAD CREAM WITH A DARK bROWN INTERORbITAL STRIPE (FIG. 3); SIDES Of HEAD DARK bROWN, LIPS WHITISH CREAM, TYMPANIC MEMbRANE DARK bROWN WITH YELLOWISH bROAD RING AROUND, SUPRATYMPANIC fOLD AND CANTHUS ROSTRALIS bOLD bLACK, OUTLINED bY A THIN CREAM STRIPE AbOVE; DORSUM CREAM WITH DIffUSE GRAYISHbROWN bLOTCHES, “V” SHAPED SCAPULAR fOLD AND DORSOLATERAL fOLD WITH DARK bROWN (ALMOST bLACK) DOTS, AND A DARK bROWN MIDDORSAL SPOT; fLANKS AND GROIN PALER THAN DORSUM, WITH SCATTERED bROWN WARTS; DORSAL SURfACE Of LIMbS GRAYISH bROWN WITH A DARK bROWN bLOTCH ON THE fOREARM AND DARK bROWN TRANSVERSAL bARS WITH CREAM MARGINS ON THE HINDLIMbS; ANTERIOR AND POSTERIOR SURfACE Of THIGHS bROWNISH CREAM; DORSAL SURfACE Of fINGERS CREAM WITH bROWN bLOTCHES; DORSAL SURfACE Of TOES LIGHT CREAM. VENTRALLY, THROAT PALE ORANGE WITH bROWN MOTTLING ANTERIORLY AND bROWN bLOTCHES ALONG THE EDGE Of CHIN; CHEST PALE LAVENDER; bELLY WITH WHITE AND bROWN MOTTLING; VENTRAL SURfACE Of THIGHS PALE LAVENDER WITH fAINT WHITE fLECKS; PALMS, SOLES AND METATARSUS DARK bROWN, AND TOES CREAM SUffUSED WITH bROWN. IRIS COPPER-COLORED IN THE UPPER HALf AND DARK bROWN IN THE LOWER HALf. Color of holotype in preservative. THE DORSAL PATTERNS ARE IDENTICAL TO THOSE IN LIfE (FIG. 2), bUT THE bACKGROUND LIGHT COLORATION bECAME DARKER, AND DARK MARKS bECAME PALER. VENTRALLY, ALL LIGHT AREAS bECAME YELLOW, AND ALL MARKS bECAME bROWN OR PALE bROWN. Variation. MALES ARE SMALLER THAN fEMALES (TAbLE 2). THE SKIN TEXTURE IN THE NEW SPECIES IS SLIGHTLY VARIAbLE. THE “V” SHAPED SCAPULAR fOLD CAN bE WELL DEfINED IN SOME SPECIMENS (E.G. CORBIDI 11658) OR POORLY DEfINED (E.G. CORBIDI 11657); ONLY IN SOME SPECIMENS (CORBIDI 11669–70, AND 11707) THE “V” CONNECTS WITH A SHORT MIDDORSAL fOLD fORMING A “Y” SHAPED fOLD. SPECIMEN CORBIDI 11658 HAS A MIDDORSAL fOLD THAT EXTENDS fROM THE END Of THE SCAPULAR REGION TO THE VENT. A bLACK MIDDORSAL TUbERCLE IS PRESENT IN SOME SPECIMENS (E.G., CORBIDI 11658 AND 11692) AND ONLY A bLACK MIDDORSAL bLOTCH IN OTHERS (E.G. CORBIDI 11657 AND 11669), AND THERE IS ONE SPECIMEN WITHOUT MIDDORSAL TUbERCLE OR bLOTCH (CORBIDI 11801). SOME SPECIMENS, AS CORBIDI 11658, HAVE A LONGITUDINAL RIDGED fOLD ON THE MIDDLE Of THE HEAD. IN LIfE, DORSAL SURfACES CAN bE CREAM, LIKE IN THE HOLOTYPE, OR PRESENT DIffERENT SHADES Of bROWN LIKE CINNAMON bROWN (CORBIDI 11669), DARK bROWN (CORBIDI 11657), OR GRAYISH bROWN (CORBIDI 11658). JUVENILES ARE GRAY DORSALLY (CORBIDI 11671). DORSAL MARKS ARE HIGHLY VARIAbLE; THEY CAN bE fAINT, LIKE IN THE HOLOTYPE AND MOST SPECIMENS, OR AbUNDANT AND DISTINCT (E.G., CORBIDI 11657). DORSAL MARKS INCLUDE DARK bROWN bLOTCHES, SPOTS, fLECKS, AN INTERORbITAL bAR, A CHEVRON MARK ON THE POSTERIOR HALf Of DORSUM, A MIDDORSAL SPOT, AND DIAGONAL MARKS ON LIMbS. THE VENTRAL HALf Of fLANKS IS SUffUSED WITH CREAM AND bROWN, bUT SOME SPECIMENS HAVE SCATTERED bROWN bLOTCHES (CORBIDI 11657) AND OTHER ARE SPRINKLED WITH WHITE fLECKS (CORBIDI 11692). THE GROIN IS PALER THAN THE DORSUM IN ALL SPECIMENS, AND ONE SPECIMEN (CORBIDI 11658) HAS A RED GROIN. THE DARK bROWN fACIAL MASK IS PRESENT IN ADULTS bUT AbSENT IN jUVENILES (CORBIDI 11670 AND 11671). IN MOST SPECIMENS, THE DORSAL fOLDS ARE DISTINCTLY PALER THAN THE bACKGROUND COLORATION (E.G. CORBIDI 11707), bUT IN SOME SPECIMENS fOLDS ARE OUTLINED WITH DARK bROWN TO bLACK MARKS (CORBIDI 11657–8). THE LIPS ARE SLIGHTLY LIGHTER (E.G. CORBIDI 11657 AND 11692), OR DISTINCTLY LIGHTER (CORBIDI 11658, 11669, 11796), AND COLORED IN WHITE OR CREAM. VENTRALLY, THE THROAT IS USUALLY PINKISH bROWN bUT IT IS DARK bROWN IN SOME SPECIMENS (CORBIDI 11657). THE CHEST, bELLY AND VENTRAL SURfACE Of THIGHS ARE DARK PINK WITH A bROWN AND WHITE MOTTLING OR WHITISH CREAM WITH A bROWN MOTTLING (E.G. CORBIDI 11671). PALMS AND SOLES ARE bROWN OR PINKISH bROWN LIKE fINGERS AND TOES, bUT SOME SPECIMENS HAVE fINGERS OR TOES DISTINCTIVELY CREAM (CORBIDI 11657). THE IRIS IS bICOLORED IN ALL SPECIMENS bUT THE UPPER HALf CAN bE DARK COPPER OR GOLDEN COPPER WHILE THE LOWER HALf IS ALWAYS bROWN. Distribution and natural history. THE NEW SPECIES IS KNOWN ONLY fROM THE TYPE LOCALITY IN THE HEADWATERS Of THE MAYO RIVER, AMAZONAS DEPARTMENT, AT 2575 M ASL (FIG. 5). THIS AREA LIES ON THE AMAZONIAN SLOPES Of THE EXTREME NORTHEASTERN PORTION Of THE CENTRAL ANDES IN NORTHERN PERU (FIG. 5). THE ECOSYSTEM IS COMPOSED Of HUMID MONTANE fORESTS WITH SOME SCATTERED PATCHES Of Chusquea SPP. THAT GROW ON ROCKY fLOORS COVERED bY A THICK LAYER Of LEAf LITTER MIXED WITH MOSS AND ROOTS (FIG. 6). THIS HAbITAT IS CLASSIfIED AS YUNGAS (500–2300 M) ACCORDING TO BRACK (1986) AND PEÑA-HERRERA DEL ÁGUILA (1989), OR PERUVIAN YUNGAS ACCORDING TO OLSON et al. (2001). Phrynopus mariellaleo sp. nov. WAS fOUND ON THE fOREST fLOOR WITHIN THE LEAf LITTER. THE fIRST COLLECTED SPECIMEN WAS fOUND bY DAY WHILE REMOVING THE LEAf LITTER AND ROOTS TO SET THE CAMPING SITE. SPECIMENS jUMPED WHEN THE MOSS WAS REMOVED. THREE SPECIMENS WERE CAPTURED IN PITfALL TRAPS AND ANOTHER WAS fOUND INSIDE A SHERMAN TRAP fOR RODENTS. THE REST Of SPECIMENS WERE SPOTTED IN THE MIDDLE Of A bROAD TRAIL WITH SCARCE LEAf LITTER WHILE SEARCHING AT NIGHT. OTHER ANURANS COLLECTED IN THIS AREA WERE Pristimantis corrugatus, P. schultei, Pristimantis SP., AND Gastrotheca SP. SOME SPECIMENS SHOW SCRATCHES ON THE bACK RESEMbLING THOSE ObSERVED IN MALES Of GLADIATOR fROGS (Boana pulchellus GROUP) AND THAT IN THOSE fROGS ARE PRODUCED DURING MALE-MALE COMbATS USING THE PREPOLLICAL SPINES. Etymology. THE SPECIfIC NAME “MARIELLALEO” IS A PATRONYM (USED AS A SUbSTANTIVE IN APPOSITION) fOR MARIELLA LEO, IN RECOGNITION Of HER TIRELESS EffORTS TO PRESERVE bIOLOGICAL DIVERSITY IN PERU. SINCE 1982 SHE HAS bEEN WORKING fOR THE ASOCIACIÓN PERUANA PARA LA CONSERVACIÓN DE LA NATURALEZA (APECO), ONE Of THE MOST IMPORTANT NON-PROfIT ORGANIZATIONS DEDICATED TO bIOLOGICAL CONSERVATION IN PERU. WITH APECO, MARIELLA CONTINUES TO WORK fOR THE PROTECTION Of MONTANE ECOSYSTEMS IN AMAZONAS DEPARTMENT, INCLUDING THE AREA WHERE THE NEW SPECIES WAS DISCOVERED.

Phrynopus mariellaleo sp. nov. FIGURES 2���4. Holotype. CORBIDI 11668, AN ADULT fEMALE, fROM SEDRUSCO, VISTA ALEGRE DISTRICT, RODR��GUEZ DE MENDOZA PROVINCE, DEPARTAMENTO AMAZONAS, PERU (6��6���42.9������S, 77��26���24������W, 2575 M ASL.; FIG. 5) ObTAINED ON 3 AUGUST 2012 bY PAbLO J. VENEGAS AND VILMA DUR��N. Paratypes. SIX fEMALES (CORBIDI 11669, 11657, 11692, 11707, 11796, 11799), fIVE MALES (CORBIDI 11658, 11797���98, 11800���01, AND fOUR jUVENILES (CORBIDI 11670���71, 11802, 11804), ALL COLLECTED AT THE SAME LOCATION AS THE HOLOTYPE bETWEEN 1���10 AUGUST 2012 bY PAbLO J. VENEGAS AND VILMA DUR��N. Diagnosis. (1) SKIN ON DORSUM SHAGREEN WITH SCATTERED LOW TUbERCLES, ���V��� OR ���Y��� SHAPED IRREGULAR (RIDGED) fOLDS IN SCAPULAR REGION, AND WITH OR WITHOUT A WEAK MIDDORSAL RIDGE; SKIN ON fLANKS SMOOTH WITH LOW fLAT WARTS; SKIN ON VENTER SMOOTH; DISCOIDAL fOLD WEAK, THORACIC fOLD PRESENT; DORSOLATERAL fOLDS PRESENT, COMPLETE; (2) TYMPANIC MEMbRANE AND TYMPANIC ANNULUS PRESENT AND PROMINENT; SUPRATYMPANIC fOLD PROMINENT, THICK AND LONG, EXTENDING fROM THE POSTERIOR MARGIN Of THE EYELID TO bEHIND THE ARM INSERTION; (3) SNOUT ROUNDED IN DORSAL AND LATERAL VIEWS; (4) UPPER EYELID bEARING SMALL, LOW, ROUND TUbERCLES; UPPER EYELID WIDTH SLIGHTLY NARROWER THAN IOD; CRANIAL CRESTS AbSENT; (5) DENTIGEROUS PROCESSES Of VOMERS PROMINENT, ObLIqUE, NARROWLY SEPARATED, SITUATED POSTEROMEDIAL TO CHOANAE; (6) VOCAL SLITS AND NUPTIAL PADS AbSENT; (7) FINGERS I AND II EqUAL IN LENGTH; DISCS ON fINGERS NARROW, ROUNDED, bEARING WEAK CIRCUMfERENTIAL GROOVES; (8) fINGERS LACKING LATERAL fRINGES; (9) ULNAR TUbERCLES AbSENT; (10) HEEL AND OUTER EDGE TARSUS WITHOUT TUbERCLES; INNER TARSAL fOLD PRESENT, SHORT, ObLIqUE; (11) INNER METATARSAL TUbERCLE PROMINENT, ELLIPTICAL, THREE TIMES LARGER THAN THE LOWER, OVAL OUTER METATARSAL TUbERCLE; PLANTAR SURfACE SMOOTH; (12) TOES LACKING LATERAL fRINGES; WEbbING AbSENT; TOE III AND TOE V AbOUT EqUAL IN SIZE; TOE DISCS NARROW, ROUND, bEARING WEAK MARGINAL GROOVES, EqUAL IN SIZE AS THOSE ON fINGERS; (13) DORSAL SURfACES LIGHT TO DARK bROWN WITH DARK bROWN TO bOLD bLACK bLOTCHES, WITH A DARK INTERORbITAL bAR, A PROMINENT DARK MIDDORSAL SPOT, AND DARK DIAGONAL bARS ON LIMbS; SIDES Of HEAD PALE TO DARK bROWN OR bLACK; CANTHUS ROSTRALIS AND TYMPANIC fOLD bOLD bLACK, OUTLINED AbOVE bY A CONTINUOUS THIN AND LIGHT STRIPE THAT RUNS fROM THE SNOUT TO THE END Of THE TYMPANIC fOLD bEHIND THE ARM; fLANKS Of SAME COLOR AS DORSUM OR SLIGHTLY LIGHTER THAN DORSUM, WITH SCATTERED DARK bROWN bLOTCHES OR DOTS AND WITH OR WITHOUT WHITE fLECKS; GROIN bROWN TO REDDISH bROWN, TRANSLUCENT, WITH DIffUSE bROWN SPOTS; VENTRAL SURfACE PALE PINK, PINKISH bROWN OR bROWN WITH DARK bROWN MOTTLING, ESPECIALLY ON THE THROAT, OR DARK bROWN WITH WHITE RETICULATIONS ON THE bELLY; (14) SVL 23.4���30.0 MM (x�� = 27.7; n = 7) IN ADULTS MALES, 25.4���39.7 MM IN ADULT fEMALES (x�� = 34.4; n = 8). Comparison with other species. Phrynopus mariellaleo DIffERS fROM ALL OTHER SPECIES Of Phrynopus EXCEPT P. auriculatus DUELLMAN AND HEDGES, AND P. peruanus PETERS, bY HAVING A DISTINCT TYMPANIC MEMbRANE AND TYMPANIC ANNULUS. Phrynopus auriculatus AND P. peruanus CAN bE DISTINGUISHED fROM THE NEW SPECIES bY HAVING FINGER I LONGER THAN FINGER II (EqUAL IN SIZE IN THE NEW SPECIES). ADDITIONALLY, P. auriculatus IS bY fAR SMALLER THAN P. mariellaleo sp. nov. [MAXIMUM SVL 13.4 MM IN MALES AND 14.5 MM IN fEMALES Of P. auriculatus VS. SVL Of 23.4��� 30.0 MM (x�� = 27.7; n = 7) IN MALES AND 25.4���39.7 MM (x�� = 34.4; n = 8) IN fEMALES Of P. mariellaleo sp. nov.]. FURTHERMORE, THE bELLY Of P. mariellaleo sp. nov. IS SMOOTH, WHILE ALL OTHER SPECIES Of Phrynopus HAVE EITHER AREOLATE VENTER (P. anancites RODR��GUEZ AND CATENAZZI, P. barthlenae LEHR AND AGUILAR, P. bufoides LEHR, LUNDbERG, AND AGUILAR, P. capitalis RODR��GUEZ AND CATENAZZI, P. chaparroi MAMANI AND MALqUI, P. daemon CH��VEZ, SANTA- CRUZ, RODR��GUEZ, AND LEHR, P. dagmarae LEHR, AGUILAR, AND K��HLER, P. dumicola RODR��GUEZ AND CATENAZZI, P. heimorum LEHR, P. horstpauli LEHR, K��HLER, AND PONCE, P. kotosh LEHR, P. lapidoides LEHR AND RODR��GUEZ, P. miroslawae CHAPARRO, PADIAL, AND DE LA RIVA, P. montium SHREVE, P. paucari LEHR, LUNDbERG, AND AGUILAR, P. peruanus, P. pesantesi LEHR, LUNDbERG, AND AGUILAR, P. tautzorum LEHR AND AGUILAR, P. thompsoni DUELLMAN, P. unchog LEHR AND RODR��GUEZ, AND P. vestigiatus LEHR AND OROZ) OR fINELY OR WEAKLY AREOLATE VENTER (P. badius LEHR, MORAVEC AND CUSI, P. interstinctus LEHR AND OROZ, P. lechriorhynchus TRUEb AND LEHR, P. oblivius LEHR, P. personatus RODR��GUEZ AND CATENAZZI, AND P. tribulosus DUELLMAN AND HEDGES). Phrynopus mariellaleo sp. nov. IS MORPHOLOGICALLY VERY SIMILAR TO SPECIES Of Hypodactylus. HOWEVER, IT IS EASILY DISTINGUISHED fROM ALL SPECIES Of Hypodactylus bY HAVING A PROMINENT, bROAD, AND LONG SUPRATYMPANIC fOLD EXTENDING fROM bEHIND THE EYELID TO THE POSTERIOR END Of THE ARM INSERTION, AND A bOLD bLACK fACEMASK. THERE ARE CURRENTLY fOUR SPECIES Of Hypodactylus RECORDED IN PERU (H. araiodactylus DUELLMAN AND PRAMUK, H. fallaciosus DUELLMAN, H. lucida CANNATELLA, AND H. nigrovittatus ANDERSSON) (DUELLMAN & LEHR 2009). THE GEOGRAPHICALLY CLOSER SPECIES TO P. mariellaleo sp. nov. ARE H. fallaciosus AND H. araiodactylus, bOTH fROM AMAZONAS DEPARTMENT IN THE NORTHERN PART Of THE CORDILLERA CENTRAL IN NORTHERN PERU (DUELLMAN & LEHR 2009). Hypodactylus araiodactylus IS ONLY KNOWN bY A SINGLE SPECIMEN COLLECTED IN 1972 fOR WHICH THERE ARE NO PICTURES OR NOTES AbOUT ITS COLORATION IN LIfE (DUELLMAN AND PRAMUK 1999). LIKE P. mariellaleo sp. nov., H. araiodactylus HAS DORSOLATERAL fOLDS AND FINGER I EqUALS IN SIZE FINGER II. HOWEVER, P. mariellaleo DIffERS fROM H. araiodactylus bY LACKING LATERAL fRINGES ON fINGERS AND TOES (PRESENT IN H. araiodactylus) AND bY HAVING A ���V��� OR ���Y��� SHAPED fOLD IN THE SCAPULAR REGION. Phrynopus mariellaleo sp. nov. ALSO CAN bE DISTINGUISHED fROM OTHER PERUVIAN SPECIES Of Hypodactylus, LIKE H. fallaciosus, H. lucida AND H. nigrovittatus, bY HAVING COMPLETE DORSOLATERAL fOLDS, WHICH ARE INCOMPLETE IN H. lucida AND AbSENT IN THE REST Of SPECIES. Hypodactylus lucida HAS A SMALL TYMPANIC MEMbRANE AND A SHORT SUPRATYMPANIC fOLD, AND LACKS MIDDORSAL fOLDS. Hypodactylus nigrovittatus HAS A SMALL, bARELY PERCEPTIbLE TYMPANIC MEMbRANE AND AN INCONSPICUOUS SUPRATYMPANIC fOLD, AND EXTREMELY SHORT HEAD WITH SUbACUMINATE SNOUT IN DORSAL VIEW (bLUNTLY ROUNDED IN P. mariellaleo sp. nov.). Description of the holotype. ADULT fEMALE WITH RObUST bODY; HEAD SLIGHTLY NARROWER THAN bODY, WIDER THAN LONG; HW 41 % Of SVL; HL 36 % Of SVL; SNOUT SHORT, LACKING TERMINAL TUbERCLE, bLUNTLY ROUNDED IN DORSAL AND LATERAL VIEWS; E���N 73 % Of EYE DIAMETER; NOSTRILS ROUNDED, DIRECTED DORSOLATERALLY; CANTHUS ROSTRALIS SLIGHTLY CURVED IN DORSAL VIEW, CURVED IN PROfILE; LOREAL REGION CONCAVE; LIPS ROUNDED; UPPER EYELID WITH fEW fLATTENED LITTLE TUbERCLES; EW 70 % Of IOD; TYMPANIC ANNULUS PRESENT; TYMPANIC MEMbRANE DISTINCT WITH DORSAL MARGIN ObSCURED bY SUPRATYMPANIC fOLD; SUPRATYMPANIC fOLD PROMINENT, EXTENDING fROM EYELIDS TO bEHIND ARM INSERTION; TY 35 % Of EYE DIAMETER, ED AbOUT 1.5 TIMES TYMPANUM DIAMETER; SEVERAL MINUTE POSTRICTAL TUbERCLES PRESENT. CHOANAE SMALL, OVOID, PARTLY CONCEALED bY PALATAL SHELf Of MAXILLA; DENTIGEROUS PROCESSES Of VOMERS PROMINENT, ObLIqUE, NARROWLY SEPARATED, WITH INDISTINCT NUMbER Of TEETH, SITUATED POSTEROMEDIAL TO CHOANAE; TONGUE AS LONG AS WIDE, NOTCHED bEHIND, HALf Of ITS LENGTH fREE POSTERIORLY. SKIN ON HEAD SMOOTH WITH SOME SCATTERED MINUTE TUbERCLES fROM THE INTERORbITAL REGION TO TIP Of THE SNOUT; DORSUM SMOOTH WITH A WEAKLY DEfINED MIDDORSAL fOLD; ���V��� OR ���Y��� SHAPED SCAPULAR fOLD PRESENT; DORSOLATERAL fOLD PRESENT, PROMINENT; SKIN ON fLANKS WITH SEVERAL fLATTENED TUbERCLES; SKIN ON THROAT, CHEST AND VENTER SMOOTH; DISCOIDAL fOLD AbSENT; CLOACAL SHEATH SHORT; CLOACAL REGION TUbERCULATE. ULNAR TUbERCLES COALESCED INTO A LOW fOLD; PALMAR TUbERCLES fLESHY, ELEVATED, OUTER PALMAR TUbERCLE bIfID, APPROXIMATELY TWICE SIZE Of OVOID, THENAR TUbERCLE; DISTAL SUbARTICULAR TUbERCLES WEAKLY DEfINED, EXCEPT PROXIMALLY, ROUND IN VENTRAL AND LATERAL VIEWS; SUPERNUMERARY TUbERCLE AT bASES Of FINGERS II AND III INDISTINCT; fINGERS LACKING LATERAL fRINGES; FINGER I AND FINGER II EqUAL IN SIZE; DISCS ON fINGERS NARROW, WITH ROUNDED TIPS; PADS ON fINGERS bEARING WEAK CIRCUMfERENTIAL GROOVES. HIND LIMbS SLENDER, TL 55 % Of SVL; FL 53 % Of SVL; UPPER SURfACES Of HIND LIMbS AREOLATE; POSTERIOR AND VENTRAL SURfACES Of THIGHS SMOOTH; HEEL LACKING TUbERCLES; OUTER SURfACE Of TARSUS LACKING TUbERCLES; SHORT INNER TARSAL fOLD PRESENT; INNER METATARSAL TUbERCLE PROMINENT, ELLIPTICAL, THREE TIMES bIGGER THAN SIZE Of LOW, INDISTINCT OUTER METATARSAL TUbERCLE; PLANTAR SURfACE SMOOTH WITH SOME INDISTINCT SUPERNUMERARY TUbERCLES; SUbARTICULAR TUbERCLES WELL DEfINED, ROUND IN VENTRAL AND LATERAL VIEWS; TOES LACKING LATERAL fRINGES; WEbbING AbSENT; DISCS ON TOES NARROW WITH ROUNDED TIPS, LIKE THOSE ON fINGERS; TOES LACKING CIRCUMfERENTIAL GROOVES; RELATIVE LENGTHS Of TOES: 1Measurements of the holotype (in mm). SVL, 39.7; TL, 22.0; FL, 21.3; HL, 14.4; HW, 16.6; ED, 4.5; TY, 1.6; IOD, 5.1; EW 3.6; IND, 4.1; E���N, 3.3. Color of holotype in life. DORSALLY, HEAD CREAM WITH A DARK bROWN INTERORbITAL STRIPE (FIG. 3); SIDES Of HEAD DARK bROWN, LIPS WHITISH CREAM, TYMPANIC MEMbRANE DARK bROWN WITH YELLOWISH bROAD RING AROUND, SUPRATYMPANIC fOLD AND CANTHUS ROSTRALIS bOLD bLACK, OUTLINED bY A THIN CREAM STRIPE AbOVE; DORSUM CREAM WITH DIffUSE GRAYISHbROWN bLOTCHES, ���V��� SHAPED SCAPULAR fOLD AND DORSOLATERAL fOLD WITH DARK bROWN (ALMOST bLACK) DOTS, AND A DARK bROWN MIDDORSAL SPOT; fLANKS AND GROIN PALER THAN DORSUM, WITH SCATTERED bROWN WARTS; DORSAL SURfACE Of LIMbS GRAYISH bROWN WITH A DARK bROWN bLOTCH ON THE fOREARM AND DARK bROWN TRANSVERSAL bARS WITH CREAM MARGINS ON THE HINDLIMbS; ANTERIOR AND POSTERIOR SURfACE Of THIGHS bROWNISH CREAM; DORSAL SURfACE Of fINGERS CREAM WITH bROWN bLOTCHES; DORSAL SURfACE Of TOES LIGHT CREAM. VENTRALLY, THROAT PALE ORANGE WITH bROWN MOTTLING ANTERIORLY AND bROWN bLOTCHES ALONG THE EDGE Of CHIN; CHEST PALE LAVENDER; bELLY WITH WHITE AND bROWN MOTTLING; VENTRAL SURfACE Of THIGHS PALE LAVENDER WITH fAINT WHITE fLECKS; PALMS, SOLES AND METATARSUS DARK bROWN, AND TOES CREAM SUffUSED WITH bROWN. IRIS COPPER-COLORED IN THE UPPER HALf AND DARK bROWN IN THE LOWER HALf. Color of holotype in preservative. THE DORSAL PATTERNS ARE IDENTICAL TO THOSE IN LIfE (FIG. 2), bUT THE bACKGROUND LIGHT COLORATION bECAME DARKER, AND DARK MARKS bECAME PALER. VENTRALLY, ALL LIGHT AREAS bECAME YELLOW, AND ALL MARKS bECAME bROWN OR PALE bROWN. Variation. MALES ARE SMALLER THAN fEMALES (TAbLE 2). THE SKIN TEXTURE IN THE NEW SPECIES IS SLIGHTLY VARIAbLE. THE ���V��� SHAPED SCAPULAR fOLD CAN bE WELL DEfINED IN SOME SPECIMENS (E.G. CORBIDI 11658) OR POORLY DEfINED (E.G. CORBIDI 11657); ONLY IN SOME SPECIMENS (CORBIDI 11669���70, AND 11707) THE ���V��� CONNECTS WITH A SHORT MIDDORSAL fOLD fORMING A ���Y��� SHAPED fOLD. SPECIMEN CORBIDI 11658 HAS A MIDDORSAL fOLD THAT EXTENDS fROM THE END Of THE SCAPULAR REGION TO THE VENT. A bLACK MIDDORSAL TUbERCLE IS PRESENT IN SOME SPECIMENS (E.G., CORBIDI 11658 AND 11692) AND ONLY A bLACK MIDDORSAL bLOTCH IN OTHERS (E.G. CORBIDI 11657 AND 11669), AND THERE IS ONE SPECIMEN WITHOUT MIDDORSAL TUbERCLE OR bLOTCH (CORBIDI 11801). SOME SPECIMENS, AS CORBIDI 11658, HAVE A LONGITUDINAL RIDGED fOLD ON THE MIDDLE Of THE HEAD. IN LIfE, DORSAL SURfACES CAN bE CREAM, LIKE IN THE HOLOTYPE, OR PRESENT DIffERENT SHADES Of bROWN LIKE CINNAMON bROWN (CORBIDI 11669), DARK bROWN (CORBIDI 11657), OR GRAYISH bROWN (CORBIDI 11658). JUVENILES ARE GRAY DORSALLY (CORBIDI 11671). DORSAL MARKS ARE HIGHLY VARIAbLE; THEY CAN bE fAINT, LIKE IN THE HOLOTYPE AND MOST SPECIMENS, OR AbUNDANT AND DISTINCT (E.G., CORBIDI 11657). DORSAL MARKS INCLUDE DARK bROWN bLOTCHES, SPOTS, fLECKS, AN INTERORbITAL bAR, A CHEVRON MARK ON THE POSTERIOR HALf Of DORSUM, A MIDDORSAL SPOT, AND DIAGONAL MARKS ON LIMbS. THE VENTRAL HALf Of fLANKS IS SUffUSED WITH CREAM AND bROWN, bUT SOME SPECIMENS HAVE SCATTERED bROWN bLOTCHES (CORBIDI 11657) AND OTHER ARE SPRINKLED WITH WHITE fLECKS (CORBIDI 11692). THE GROIN IS PALER THAN THE DORSUM IN ALL SPECIMENS, AND ONE SPECIMEN (CORBIDI 11658) HAS A RED GROIN. THE DARK bROWN fACIAL MASK IS PRESENT IN ADULTS bUT AbSENT IN jUVENILES (CORBIDI 11670 AND 11671). IN MOST SPECIMENS, THE DORSAL fOLDS ARE DISTINCTLY PALER THAN THE bACKGROUND COLORATION (E.G. CORBIDI 11707), bUT IN SOME SPECIMENS fOLDS ARE OUTLINED WITH DARK bROWN TO bLACK MARKS (CORBIDI 11657���8). THE LIPS ARE SLIGHTLY LIGHTER (E.G. CORBIDI 11657 AND 11692), OR DISTINCTLY LIGHTER (CORBIDI 11658, 11669, 11796), AND COLORED IN WHITE OR CREAM. VENTRALLY, THE THROAT IS USUALLY PINKISH bROWN bUT IT IS DARK bROWN IN SOME SPECIMENS (CORBIDI 11657). THE CHEST, bELLY AND VENTRAL SURfACE Of THIGHS ARE DARK PINK WITH A bROWN AND WHITE MOTTLING OR WHITISH CREAM WITH A bROWN MOTTLING (E.G. CORBIDI 11671). PALMS AND SOLES ARE bROWN OR PINKISH bROWN LIKE fINGERS AND TOES, bUT SOME SPECIMENS HAVE fINGERS OR TOES DISTINCTIVELY CREAM (CORBIDI 11657). THE IRIS IS bICOLORED IN ALL SPECIMENS bUT THE UPPER HALf CAN bE DARK COPPER OR GOLDEN COPPER WHILE THE LOWER HALf IS ALWAYS bROWN. Distribution and natural history. THE NEW SPECIES IS KNOWN ONLY fROM THE TYPE LOCALITY IN THE HEADWATERS Of THE MAYO RIVER, AMAZONAS DEPARTMENT, AT 2575 M ASL (FIG. 5). THIS AREA LIES ON THE AMAZONIAN SLOPES Of THE EXTREME NORTHEASTERN PORTION Of THE CENTRAL ANDES IN NORTHERN PERU (FIG. 5). THE ECOSYSTEM IS COMPOSED Of HUMID MONTANE fORESTS WITH SOME SCATTERED PATCHES Of Chusquea SPP. THAT GROW ON ROCKY fLOORS COVERED bY A THICK LAYER Of LEAf LITTER MIXED WITH MOSS AND ROOTS (FIG. 6). THIS HAbITAT IS CLASSIfIED AS YUNGAS (500���2300 M) ACCORDING TO BRACK (1986) AND PE��A-HERRERA DEL ��GUILA (1989), OR PERUVIAN YUNGAS ACCORDING TO OLSON et al. (2001). Phrynopus mariellaleo sp. nov. WAS fOUND ON THE fOREST fLOOR WITHIN THE LEAf LITTER. THE fIRST COLLECTED SPECIMEN WAS fOUND bY DAY WHILE REMOVING THE LEAf LITTER AND ROOTS TO SET THE CAMPING SITE. SPECIMENS jUMPED WHEN THE MOSS WAS REMOVED. THREE SPECIMENS WERE CAPTURED IN PITfALL TRAPS AND ANOTHER WAS fOUND INSIDE A SHERMAN TRAP fOR RODENTS. THE REST Of SPECIMENS WERE SPOTTED IN THE MIDDLE Of A bROAD TRAIL WITH SCARCE LEAf LITTER WHILE SEARCHING AT NIGHT. OTHER ANURANS COLLECTED IN THIS AREA WERE Pristimantis corrugatus, P. schultei, Pristimantis SP., AND Gastrotheca SP. SOME SPECIMENS SHOW SCRATCHES ON THE bACK RESEMbLING THOSE ObSERVED IN MALES Of GLADIATOR fROGS (Boana pulchellus GROUP) AND THAT IN THOSE fROGS ARE PRODUCED DURING MALE-MALE COMbATS USING THE PREPOLLICAL SPINES. Etymology. THE SPECIfIC NAME ���MARIELLALEO��� IS A PATRONYM (USED AS A SUbSTANTIVE IN APPOSITION) fOR MARIELLA LEO, IN RECOGNITION Of HER TIRELESS EffORTS TO PRESERVE bIOLOGICAL DIVERSITY IN PERU. SINCE 1982 SHE HAS bEEN WORKING fOR THE ASOCIACI��N PERUANA PARA LA CONSERVACI��N DE LA NATURALEZA (APECO), ONE Of THE MOST IMPORTANT NON-PROfIT ORGANIZATIONS DEDICATED TO bIOLOGICAL CONSERVATION IN PERU. WITH APECO, MARIELLA CONTINUES TO WORK fOR THE PROTECTION Of MONTANE ECOSYSTEMS IN AMAZONAS DEPARTMENT, INCLUDING THE AREA WHERE THE NEW SPECIES WAS DISCOVERED., Published as part of Venegas, Pablo J., Barboza, Andy C., Riva, Ignacio De La & Padial, Jos�� M., 2018, A new species of Phrynopus from the northeastern Andes of Peru, its phylogenetic position, and notes on the relationships of Holoadeninae (Anura: Craugastoridae), pp. 501-524 in Zootaxa 4446 (4) on pages 504-510, DOI: 10.11646/zootaxa.4446.4.5, http://zenodo.org/record/1444442, {"references":["Duellman, W. E. & Lehr, E. (2009) Terrestrial-breeding frogs (Strabomantidae) in Peru. Natur und Tier-Verlag, Naturwissenschaft, Munster, 382 pp.","Duellman, W. E. & Pramuk, J. B. (1999) Frogs of the genus Eleutherodactylus (Anura: Leptodactylidae) in the Andes of northern Peru. Scientific Papers of the Natural History Museum, University of Kansas, 13, 1 - 78.","Brack, A. (1986) Las Ecorregiones del Peru. Boletin de Lima, 44, 57 - 70.","Penaherrera del Aguila, C. (1989) Atlas del Peru. Instituto Geografico Nacional, Lima, 400 pp."]}

Euspondylus excelsum sp. nov. (Figs. 1���4) Unnamed Clade 1���Torres-Carvajal et al. 2016: 70 (Fig. 2) Holotype. (Figs. 1���2) CORBIDI 16457, adult male, from Quebrada Tambo (-9.70483�� Lat, -75.81383�� Long, WGS 84, 1147 m above sea level): Hu��nuco Province, Hu��nuco Department, Peru; collected by Lesly Luj��n on 15 October 2015. Paratopotype. CORBIDI 16450 subadult female; collected by Lesly Luj��n on 12 October 2015. Paratypes. (Figs. 3���4) Two males, five females and three juveniles, all from Hu��nuco Department, Peru. CORBIDI 14666, adult male, from Zaria, Pachitea Province (-9.68905�� Lat, -75.84300�� Long, WGS 84, 1023 m above sea level); collected by Vilma Dur��n on 16 July 2014. CORBIDI 15573, adult female, from Campamento La garganta de la Bella, Tingo Maria National Park, Leoncio Prado Province (-9.33841�� Lat, -76.00206�� Long, WGS 84, 1095 m above sea level); collected by Diego V��squez and Germ��n Ch��vez on 21 November 2014. CORBIDI 16396, female, from Zaria, Huanuco Province (-9.71467�� Lat; -75.81592�� Long, WGS 84, 1157 m above sea level); collected by Diego V��squez on 7 September 2015. CORBIDI 16459, juvenile female from Santa Elena, Pachitea Province (-9.73027�� Lat, 75.80503�� Long, WGS 84, 1144 m above sea level); collected by Lesly Luj��n on 18 October 2015. CORBIDI 16463, adult female from Shincay, Huanuco Province (-9.72563�� Lat, -75.81497�� Long, WGS 84, 1149 m.a.s.l); collected by Lesly Luj��n on 27 October 2015. CORBIDI 16464, adult female from Zaria, Huanuco Province (-9.71830�� Lat; -75.81575�� LongWGS 84, 1159 m above sea level); collected by Lesly Luj��n on 28 October 2015. CORBIDI 16465 adult male, 16466 adult female, 16467- 68 juveniles, from Zaria, Huanuco province (-9.71897�� Lat, 75.8144�� Long, WGS 84 1137 m above sea level); collected by Lesly Luj��n on 5 November 2015. Generic placement. We assign tentatively the new species to Euspondylus on the basis of morphological comparisons. The presence of a tongue homogeneously covered with imbricate-like papillae, of well-developed eyelids, a divided lower palpebral disc, elongated nose, striated and subimbricate dorsal scales, lateral scales reduced when are closer to ventrals, limbs clawed and well-developed, and a compact body (all this characters in accordance with the type species of the genus, E. maculatus) leads us to allocate the new species to the genus Euspondylus (see Table 1 for comparisons with types of E. maculatus). Additionally, a recent phylogeny (Torres- Carvajal et al. 2016) that includes paratype CORBIDI 15573 (referred as Cercosaurini CORBIDI 15573 into Unnamed clade 1) shows that this taxon is closely related to Pholidobolus and Macropholidus, and it is nested within the paraphyletic clade of Anadia. Diagnosis. The new species can be identified by the following combination of characters: 1) body robust, moderate size, maximum SVL 83.7 mm in males, 81.4 mm in females; 2) head flat, elongated, about 1.6���1.9 times longer than wide; 3) ear openings distinct, slightly recessed; 6) nasals separated by frontonasal; frontonasal undivided; 7) prefrontals, frontal, frontoparietals, parietals, postparietals and interparietal present; 8) parietals longer than wide; 9) supraoculars four, anteriormost fused with anteriormost superciliar; 10) superciliar series complete, five, occasionally four; 11) uncomplete suture from narina to posterodorsal suture of nasal present; 12) loreal in large contact with second supralabial; 13) supralabials six; infralabials five; 14) genials in three pairs, first and second pair always in contact, third pair usually in contact; 15) collar fold present; 16) dorsals in 32���34 rows in both sexes, rectangular, about three times longer than wide, striated, subimbricate; 17) ventrals in 21���23 rows in both sexes, quadrangular, smooth, juxtaposed; 18) scales around mid-body 40���42; 19) lateral scales at mid-body reduced in 6���9 lines; 20) limbs pentadactyl, all digits clawed, forelimb reaching anteriorly to sixth supralabial; 21) subdigital lamellae under finger IV 15���18, under toe IV 20���24; 22) femoral pores in males 10���12, in females 4���7, four scales between femoral pores; 23) preanal plate scales seven; 24) tail up to 2.5 times longer than body; 25) caudals subimbricate, striated dorsally, smooth ventrally; 26) lower palpebral disc transparent, divided in four sections; 27) coloration in life golden-brown with scattered black spots or blotches on head and dorsum, with a black stripe running from the nasal plate to supratympanic temporals, only interrupted by the eye; dorsum with dorsolateral dark-bordered stripes, red-orange in males, brown in females; males with 7���14 pairs of lateral ocelli, from anterior part of the insertion of forelimbs to posterior part of the insertion of hindlimbs; limbs golden-brown; dorsal scales of tail golden-brown with pale flecks arranged dorsolaterally; venter, ventral surfaces of limbs and tail creamy white with black spots; iris copper-red. Comparisons. Euspondylus excelsum sp. nov. resembles in scutellation and measurements to E. guentheri and E. maculatus, from which it can be easily differentiated by the strong fusion of anteriormost supraocular with anteriormost superciliar (fusion absent in E. guentheri and E. maculatus) (See Fig. 5). Furthermore, E. excelsum can be distinguished from both species by the following characters (conditions for E. guentheri and E. maculatus in parenthesis): body robust (vs. slender in E. guentheri), the presence of small rounded irregular spots (vs. large, transversal black spots on dorsum in E. guentheri), by having striated dorsal scales (vs. smooth in E. guentheri); third pair of chin shields usually in contact (vs. separated in E. guentheri and E. maculatus); head 1.6���1.9 longer than wide (vs. 1.5 in E. maculatus); eye���nose distance is 0.3���0.4 times of head length (vs. 0.5 in E. guentheri); longitudinal dorsal count 32���34 (vs. 38 in E. maculatus) ventral scales rows 21���23 (vs. 25 in E. maculatus); 10���12 femoral pores in adult males (vs. 17 in E. guentheri); and 7���8 preanal scales (vs. 5 in E. guentheri and E. maculatus). Furthermore, E. excelsum sp. nov. can be differentiated from other congeners (those with the same condition on the lower palpebral disc) by having a longer body with a maximum SVL of 83.7 mm (vs. 53.5 mm in E. acutirostris, 59.6 mm in E. auyanensis, 46.0 mm in E. monsfumus); head 1.6���1.9 longer than wide (vs. 1.5 in E. sp. 1, 1.7 in E. acutirostris and E. guentheri, 1.8 in E. monsfumus); first supraocular fused with first superciliar (vs. not fused in E. acutirostris, E. monsfumus); most dorsal temporal 3���3.5 times longer than small lateral temporals (vs. 4���4.5 times longer than small lateral temporals in E. sp. 2); a lower number of postparietals having 2���3 (vs. 3 in E. auyanensis, 3���4 in E. sp. 1); 2���3 pairs of genials in contact (vs. 2 pairs in contact in E. acutirostris and E. sp. 2); a lower number of ventral scales rows having 21���23 (vs. 26���33 in E. acutirostris, 24 in E. auyanensis, 28 in E. monsfumus); a higher number of scales around mid-body 40���42 (vs. 34���40 in E. acutirostris, 32 in E. auyanensis, 36 in E. monsfumus); a lower number of femoral pores having 3���7 on each side (vs. 8���11 in E. acutirostris, 15���18 in E. auyanensis, 9���10 in E. monsfumus); a higher number of preanal scales 7���8 (vs. 5 in E. sp. 1). Description of the holotype (Figs 1���2). Adult male (CORBIDI 16457); SVL = 69.4 mm, tail (regenerated) length = 55.6 mm; axilla to groin distance = 32.6 mm; head length = 18.1; head width = 11.2 mm; shank length = 11.0 mm. Head scales smooth, glossy; rostral scale wider (2.7 mm) than long (1.5 mm), slightly higher than adjacent supralabials, in contact with frontonasal, nasals, and first supralabials posteriorly; frontonasal pentagonal, slightly wider (2.8 mm) than long (2.6 mm), posterior suture angular with point directed posteriorly, in contact with nasal and laterally, prefrontals posteriorly, nasoloreal suture uncomplete; prefrontals present, in contact with each other medially, in contact with fused anteriormost superciliary-anteriormost supraocular, frontal posteriorly; frontal pentagonal, longer (3.6 mm) than wide (2.7 mm), anterior suture angular with point directed anteriorly, lateral sutures straight, posterior suture angular with point slightly directed posteriorly, in contact with second supraocular laterally, frontoparietals posteriorly; frontoparietals pentagonal, in contact with third and fourth supraocular, parietals and interparietal posteriorly; supraoculars four, none in contact with ciliaries; superciliary series complete, anteriormost superciliary fused with anteriormost supraocular; interparietal pentagonal, longer (3.6 mm) than wide (2.8 mm), lateral sutures concave, in contact with parietals laterally, postparietals posteriorly; parietals polygonal, in contact with fourth supraocular anterolaterally, temporal scales laterally, dorsalmost postocular, postparietals posteriorly; postparietals four, polygonal; palpebral disc oval, unpigmented, divided in four sections; frenocular triangular, in contact with loreal anteriorly; postoculars three; temporals polygonal; supratympanic temporals two; supralabials six; infralabials five; mental wider (2.8 mm) than long (1.7 mm), in contact with first infralabials, postmental posteriorly; postmental single, pentagonal, posterior suture angular, point slightly directed posteriorly, in contact with first and second infralabials; genials in three pairs, anterior pair subquadrangular, in contact with second and third infralabials, medial pair of genials subcuadrangular, in contact with third and fourth infralabials laterally, posterior pair of genials trapezoid, in contact with fourth and fifth infralabials laterally; scale rows between genials and collar fold (along midventral line) 15; medialmost scales of three penultimate gular scale rows cuadrangular; posteriormost gular row enfolded posteriorly, concealing two granular scale rows; lateral neck scales rounded, smooth. Dorsal scales rectangular, longer than wide, juxtaposed, striated, 33 in a longitudinal count, homogeneously arranged; transverse dorsal count (enlarged rows at mid-body) at fifth transverse ventral scale row 10, at 10th transverse ventral scale row 12, at 15th transverse ventral scale row 14; lateral scale rows at fifth transverse ventral scale row 13, at 10th transverse ventral scale row 9, at 15th transverse ventral scale row 10; lateral scales on body near insertion of forelimb small to granular; ventrals cuadrangular and juxtaposed; complete longitudinal ventral count 21; longitudinal ventral scale rows at mid-body 10; 41 scales around mid-body; anterior preanal plate scales two; posterior preanal plate scales seven, all the scales at the same size; scales on tail rectangular and juxtaposed, striated; mid-ventral subcaudals squarish. Limbs pentadactyl; digits clawed; forelimb reaching anteriorly to sixth supralabial; dorsal brachial scales polygonal, of twice the size of anterodorsal midbrachial scale, imbricate, smooth; midbrachial anterodorsal scale small, at least twice shorter than adjacent scales, smooth; anteroventral, ventral, and posteroventral scales roundish, imbricate, smooth; antebrachial scales polygonal, at least as large as midbrachial anterodorsal scale; medial antebrachial scales larger, polygonal, smooth; dorsal manus scales polygonal, subimbricate; palmar scales granular, small, oval; dorsal scales on fingers smooth, quadrangular, covering dorsal half of digit, overhanging supradigital scales, four on I, seven on II, nine on III, 11/10 on IV, six on V; subdigital scales seven on I, 14/13 on II, seventeen on III, 16/17 on IV, eleven on V; anterodorsal thigh scales polygonal, at least three times as large as adjacent scales, becoming smaller ventrally, smooth; posterodorsal thigh scales small, rounded, arranged irregularly; anterior and anteromedial shank scales polygonal, subimbricate, smooth, anteriormost scales many times larger than lateral, posterolateral, and posteromedial shank scales; lateral, posterolateral, and posteromedial shank scales polygonal or roundish, subimbricate, smooth; dorsal pes scales polygonal, subimbricate, smooth; scales on dorsal surface of digits single, quadrangular, smooth, overhanging supradigital scales, four on I, seven on II, thirteen on III, 16/15 on IV, eleven on V; subdigital scales single, seven on I, 14/13 on II, nineteen on III, twentyfour on IV, 16/17 on V; femoral pores series not continued, 5/7; four scales between medialmost femoral pores. Hemipenial morphology. (Fig. 6) The right hemipenis was everted during preservation and prepared posteriorly. The organ extends along approximately 11 mm. The hemipenis is completely expanded and the lobes of the organ are partially everted. Hemipenial body in light bulb shape, with the basis distinctly narrower than the rest of the organ, and ending in two small lobes with apical folds in the apex. The s ulcus spermaticus is central in position, originating at the base of the organ, and proceeding in a straight line towards the lobes. The sulcus is broader in the region of the lobular crotch, where it is divided by a small bulky fold; branches running on the medial region of the lobes, and ending in their tip among folds. The sulcate face of the hemipenial body presents two fleshy nude areas, stained with blue, parallel to the sulcus spermaticus that run through all the hemipenial body extension. Laterals and asulcate face of the hemipenis are ornamented with series of roughly equidistant flounces with calcareous spinules, all of them stained in red with Alizarin. Twenty-four rows of flounces extend along the body of the organ. There are three proximal rows restricted to a central position on the basal asulcate face of the hemipenis, all of them are roughly chevron-shaped. The five proximalmost flounces in the laterals are diagonally positioned, with the second to fifth flounces separated from a complementary flounce positioned in the asulcate face and orientated in an inverse diagonal. The nude area between the proximalmost fifth flounce is narrower than the nude area between the other four proximal flounces and its complementary ones. The subsequent flounces towards the lobes cross the laterals of the organ from the sulcate to the asulcate face, in chevron-shape with vertexes in the central region of each lateral pointed towards the basis of the organ. These chevron-shaped rows reduce in size progressively towards the hemipenial apex. The lateral flounces are separated in two groups by a nude area in the central asulcate face that increases in size in the apical region acquiring a Y-shape. The regions between the asulcate face and the laterals in this species are marked by a conspicuous unevenness forming a bulge shared by other related species such as Macropholidus annectens, M. huancabambae, M. ruthveni, Pholidobolus affinis, P. macbrydei, P. montium, P. prefrontalis (Nunes, 2011), P. hillisi (Torres-Carvajal et al. 2014), and P. ulisesi (Venegas et al. 2016). Color of holotype in life. (Fig. 2) Head golden-brown with scattered black spots, down to inferior level of supralabials, infralabials creamy white with black spots; a dark stripe running from the nasal plate to the anterior angle of the eye and from the posterior angle of the eye to the supratympanic temporals, iris copper-red. Dorsum golden-brown with scattered black spots, a dorsolateral dark-bordered red-orange stripe from nuchals to the base of the tail, dark border turning paler and fading entirely towards the base of tail, seven pairs of lateral ocelli. Dorsal surface of tail golden brown with pale dorsolateral spots. Throat, chest, belly and ventral surfaces of limbs creamy white, with scattered black spots or blotches. Precloacal and cloacal scales creamy white with scattered black spots; ventral surface of tail creamy white with scattered black blotches. Variation and sexual dimorphism. (Fig. 3, 4) Variation in measurements and scutellation is summarized in Table 1. The largest male is CORBIDI 16465 (81.7 mm), the largest female is CORBIDI 16466 (81.4 mm). Females CORBIDI 15573 and 16466 have two genials in contact, not three as the rest of type series. Six specimens have the supraocular-superciliar scale fusion divided by a diagonal, weakly defined, complete or uncomplete suture (CORBIDI 16396, 16459, 16463, 16465���66, 16468); also a juvenile (CORBIDI 16468) has six superciliars. Dorsolateral red-orange dark bordered stripe is extended to 10th verticile of the tail in male CORBIDI 16465. Juveniles CORBIDI 16467���68 have yellow coloration on belly and ventral surfaces of hindlimbs and tail. The sex can be distinguished by a higher number of femoral pores in males (10���12) than females (4���6), only juvenile male CORBIDI 16459 has 6 femoral pores, furthermore males can be differentiated by the presence of 7���14 pairs of lateral ocelli (absent in females), and by the presence of black scattered spots or blotches on ventral surface of tail in males, instead black spots or blotches arranged ventrolaterally on tails of females. Additionally, male CORBIDI 16465 has an atypical high number of irregular black spots on belly, remarkably higher than the rest of males of the type series. Etymology. The specific epithet ��� excelsum ��� is from Latin and means ���tall���, in reference to the habitat of the new species in the canopy of the tropical rainforest. Distribution and natural History. Euspondylus excelsum sp. nov. is known from localities on both sides of the Huallaga river, including within Tingo Maria National Park (Fig. 7, 8), at distances as far as 48.5 km between the northernmost and southernmost collecting sites. Specimens were collected in the canopy of primary and secondary forests. Specimens CORBIDI 14666 and 15573 were observed during diurnal Visual Encounter Surveys climbing trunks at 1.5 m above leaf litter. The holotype and specimens CORBIDI 16396, 16450, 16459, 16463-65, were collected in the canopy of Couma spp. at night; specimens CORBIDI 16466���68 were collected under leaves of bromeliads gathered on crags. Male CORBIDI 16465 started swimming underwater during an escape attempt prior to capture. After dissection for laboratory analyses, two eggs were observed into the ventral cavity of three females of type series collected at November 2014 and October-November 2015, eggs lengths were (measurements of egg in right oviduct is presented first): 18.0 mm and 18.0 mm in CORBIDI 15573 (23.2% of the SVL), 16.6 mm and 16.5 mm in CORBIDI 16463 (20.2% and 20.4% o, Published as part of Ch��vez, Germ��n, Catenazzi, Alessandro & Venegas, Pablo J., 2017, A new species of arboreal microteiid lizard of the genus Euspondylus (Gymnophtalmidae: Cercosaurinae) from the Andean slopes of central Peru with comments on Peruvian Euspondylus, pp. 301-316 in Zootaxa 4350 (2) on pages 303-314, DOI: 10.11646/zootaxa.4350.2.6, http://zenodo.org/record/1053168, {"references":["Nunes, P. M. S. (2011) Morfologia hemipeniana dos lagartos microteideos e suas implicacoes nas relacoes filogeneticas da familia Gymnophthalmidae (Teioidea: Squamata). PhD Thesis, Departamento de Zoologia, Universidade de Sao Paulo, Sao Paulo, 88 pp.","Torres-Carvajal, O., Venegas, P. J., Lobos, S. E., Mafla-Endara, P. & Sales Nunes, P. M. (2014) A new species of Pholidobolus (Squamata: Gymnophthalmidae) from the Andes of southern Ecuador. Amphibian & reptile conservation, 8 (1), 76 - 88.","Goicoechea, N., Padial, J. M., Chaparro, J. C., Castroviejo-Fisher, S. & De la Riva, I. (2012) Molecular phylogenetics, species diversity, and biogeography of the Andean lizards of the genus Proctoporus (Squamata: Gymnophtalmidae). Molecular Phylogenetics and Evolution, 65 (3), 953 - 964. https: // doi. org / 10.1016 / j. ympev. 2012.08.017","Mijares-Urrutia, C. A., Searis, J. C. & Arends, A. (2001) Taxonomia de algunos Microteidos (Squamata) de Venezuela, I: Variacion y distribucion geografica de Euspondylus acutirostris con la descripcion de un nuevo Euspondylus. Revista de Biologia Tropical, 48 (2 / 3), 671 - 680.","Uzzell, T. (1973) A revision of lizards of the genus Prionodactylus with a new genus for P. leucosticus and notes on the genus Euspondylus (Sauria, Teiidae). Postilla Peabody Museum Yale University, 159, 1 - 67.","Kohler, G. (2003) Two new species of Euspondylus (Squamata: Gymnophtalmidae) from Peru. Salamadra, 39 (1), 5 - 20.","Tschudi, J. J. von (1845) Reptilium conspectum quae in republica Peruana reperiuntur er pleraque observata vel collecta sunt in itenere. Archuv fur Naturegeschichte, 11 (1), 150 - 170. https: // doi. org / 10.5962 / bhl. part. 7963","Kohler, G. & Lehr, E. (2004) Comments on Euspondylus and Proctoporus (Squamata: Gymnophtalmidae) from Peru, with the description of three new species and a key to the Peruvian species. Herpetologica, 60 (4), 501 - 518. https: // doi. org / 10.1655 / 03 - 93","Myers, C. W., Rivas-Fuenmayor, G. & Jadin, R. C. (2009) New species of lizards from Auyantepui and La Escalera in the Venezuelan Guyana, with notes on the \" microteiid \" hemipenes (Squamata, Gymnophtalmidae). American Museum Novitates, 3660, 1 - 32. https: // doi. org / 10.1206 / 657.1","O´Shaugnessy, A. W. E. (1881) An account of the collection of lizards made by Mr. Buckley in Ecuador, and now in the British Museum, with descriptions of the new species. Proceedings of the Zoological Society of London, 1881, 227 - 245.","Boulenger, G. A. (1885) Catalogue of the lizards of the British Museum of Natural History. Vol. 2. 2 nd Edition. Printed by order of the Trustees, London, xiii + 497 pp.","Oftedal, O. T. (1974) A revision of the genus Anadia (Sauria, Teiidae). Arquivos de Zoologia, 25 (4), 203 - 265. https: // doi. org / 10.11606 / issn. 2176 - 7793. v 25 i 4 p 203 - 265","Rivas, G. A., Sales-Nunes, P. M., Dixon, J. R., Schargel, W. E., Caidedo, J. R., Barro, T. R., Camargo, E. G. & Barrio-Amoros, C. L. (2012) Taxonomy, hemipenial morphology, and natural history of two poorly known species of Anadia (Gymnophtalmidae) from northern South America. Journal of Herpetology, 46 (1), 33 - 40. https: // doi. org / 10.1670 / 10 - 139","Venegas, P. J., Duran, V. & Garcia-Burneo, K. (2013) A new species of arboreal iguanid lizard, genus Stenocercus (Squamata: Iguania), from central Peru. Zootaxa, 3609 (3), 291 - 301. https: // doi. org / 10.11646 / zootaxa. 3609.3.3","Chavez, G. & Catenazzi, A. (2016) A new species of frog of the genus Pristimantis from Tingo Maria National Park, Huanuco Department, central Peru (Anura, Craugastoridae). Zookeys, 610, 113 - 130. https: // doi. org / 10.3897 / zookeys. 610.8507"]}

Chávez, Germán, Catenazzi, Alessandro, Venegas, Pablo J. (2017): A new species of arboreal microteiid lizard of the genus Euspondylus (Gymnophtalmidae: Cercosaurinae) from the Andean slopes of central Peru with comments on Peruvian Euspondylus. Zootaxa 4350 (2): 301-316, DOI: https://doi.org/10.11646/zootaxa.4350.2.6

Euspondylus excelsum sp. nov. (Figs. 1–4) Unnamed Clade 1—Torres-Carvajal et al. 2016: 70 (Fig. 2) Holotype. (Figs. 1–2) CORBIDI 16457, adult male, from Quebrada Tambo (-9.70483° Lat, -75.81383° Long, WGS 84, 1147 m above sea level): Huánuco Province, Huánuco Department, Peru; collected by Lesly Luján on 15 October 2015. Paratopotype. CORBIDI 16450 subadult female; collected by Lesly Luján on 12 October 2015. Paratypes. (Figs. 3–4) Two males, five females and three juveniles, all from Huánuco Department, Peru. CORBIDI 14666, adult male, from Zaria, Pachitea Province (-9.68905° Lat, -75.84300° Long, WGS 84, 1023 m above sea level); collected by Vilma Durán on 16 July 2014. CORBIDI 15573, adult female, from Campamento La garganta de la Bella, Tingo Maria National Park, Leoncio Prado Province (-9.33841° Lat, -76.00206° Long, WGS 84, 1095 m above sea level); collected by Diego Vásquez and Germán Chávez on 21 November 2014. CORBIDI 16396, female, from Zaria, Huanuco Province (-9.71467° Lat; -75.81592° Long, WGS 84, 1157 m above sea level); collected by Diego Vásquez on 7 September 2015. CORBIDI 16459, juvenile female from Santa Elena, Pachitea Province (-9.73027° Lat, 75.80503° Long, WGS 84, 1144 m above sea level); collected by Lesly Luján on 18 October 2015. CORBIDI 16463, adult female from Shincay, Huanuco Province (-9.72563° Lat, -75.81497° Long, WGS 84, 1149 m.a.s.l); collected by Lesly Luján on 27 October 2015. CORBIDI 16464, adult female from Zaria, Huanuco Province (-9.71830° Lat; -75.81575° LongWGS 84, 1159 m above sea level); collected by Lesly Luján on 28 October 2015. CORBIDI 16465 adult male, 16466 adult female, 16467- 68 juveniles, from Zaria, Huanuco province (-9.71897° Lat, 75.8144° Long, WGS 84 1137 m above sea level); collected by Lesly Luján on 5 November 2015. Generic placement. We assign tentatively the new species to Euspondylus on the basis of morphological comparisons. The presence of a tongue homogeneously covered with imbricate-like papillae, of well-developed eyelids, a divided lower palpebral disc, elongated nose, striated and subimbricate dorsal scales, lateral scales reduced when are closer to ventrals, limbs clawed and well-developed, and a compact body (all this characters in accordance with the type species of the genus, E. maculatus) leads us to allocate the new species to the genus Euspondylus (see Table 1 for comparisons with types of E. maculatus). Additionally, a recent phylogeny (Torres- Carvajal et al. 2016) that includes paratype CORBIDI 15573 (referred as Cercosaurini CORBIDI 15573 into Unnamed clade 1) shows that this taxon is closely related to Pholidobolus and Macropholidus, and it is nested within the paraphyletic clade of Anadia. Diagnosis. The new species can be identified by the following combination of characters: 1) body robust, moderate size, maximum SVL 83.7 mm in males, 81.4 mm in females; 2) head flat, elongated, about 1.6–1.9 times longer than wide; 3) ear openings distinct, slightly recessed; 6) nasals separated by frontonasal; frontonasal undivided; 7) prefrontals, frontal, frontoparietals, parietals, postparietals and interparietal present; 8) parietals longer than wide; 9) supraoculars four, anteriormost fused with anteriormost superciliar; 10) superciliar series complete, five, occasionally four; 11) uncomplete suture from narina to posterodorsal suture of nasal present; 12) loreal in large contact with second supralabial; 13) supralabials six; infralabials five; 14) genials in three pairs, first and second pair always in contact, third pair usually in contact; 15) collar fold present; 16) dorsals in 32–34 rows in both sexes, rectangular, about three times longer than wide, striated, subimbricate; 17) ventrals in 21–23 rows in both sexes, quadrangular, smooth, juxtaposed; 18) scales around mid-body 40–42; 19) lateral scales at mid-body reduced in 6–9 lines; 20) limbs pentadactyl, all digits clawed, forelimb reaching anteriorly to sixth supralabial; 21) subdigital lamellae under finger IV 15–18, under toe IV 20–24; 22) femoral pores in males 10–12, in females 4–7, four scales between femoral pores; 23) preanal plate scales seven; 24) tail up to 2.5 times longer than body; 25) caudals subimbricate, striated dorsally, smooth ventrally; 26) lower palpebral disc transparent, divided in four sections; 27) coloration in life golden-brown with scattered black spots or blotches on head and dorsum, with a black stripe running from the nasal plate to supratympanic temporals, only interrupted by the eye; dorsum with dorsolateral dark-bordered stripes, red-orange in males, brown in females; males with 7–14 pairs of lateral ocelli, from anterior part of the insertion of forelimbs to posterior part of the insertion of hindlimbs; limbs golden-brown; dorsal scales of tail golden-brown with pale flecks arranged dorsolaterally; venter, ventral surfaces of limbs and tail creamy white with black spots; iris copper-red. Comparisons. Euspondylus excelsum sp. nov. resembles in scutellation and measurements to E. guentheri and E. maculatus, from which it can be easily differentiated by the strong fusion of anteriormost supraocular with anteriormost superciliar (fusion absent in E. guentheri and E. maculatus) (See Fig. 5). Furthermore, E. excelsum can be distinguished from both species by the following characters (conditions for E. guentheri and E. maculatus in parenthesis): body robust (vs. slender in E. guentheri), the presence of small rounded irregular spots (vs. large, transversal black spots on dorsum in E. guentheri), by having striated dorsal scales (vs. smooth in E. guentheri); third pair of chin shields usually in contact (vs. separated in E. guentheri and E. maculatus); head 1.6–1.9 longer than wide (vs. 1.5 in E. maculatus); eye–nose distance is 0.3–0.4 times of head length (vs. 0.5 in E. guentheri); longitudinal dorsal count 32–34 (vs. 38 in E. maculatus) ventral scales rows 21–23 (vs. 25 in E. maculatus); 10–12 femoral pores in adult males (vs. 17 in E. guentheri); and 7–8 preanal scales (vs. 5 in E. guentheri and E. maculatus). Furthermore, E. excelsum sp. nov. can be differentiated from other congeners (those with the same condition on the lower palpebral disc) by having a longer body with a maximum SVL of 83.7 mm (vs. 53.5 mm in E. acutirostris, 59.6 mm in E. auyanensis, 46.0 mm in E. monsfumus); head 1.6–1.9 longer than wide (vs. 1.5 in E. sp. 1, 1.7 in E. acutirostris and E. guentheri, 1.8 in E. monsfumus); first supraocular fused with first superciliar (vs. not fused in E. acutirostris, E. monsfumus); most dorsal temporal 3–3.5 times longer than small lateral temporals (vs. 4–4.5 times longer than small lateral temporals in E. sp. 2); a lower number of postparietals having 2–3 (vs. 3 in E. auyanensis, 3–4 in E. sp. 1); 2–3 pairs of genials in contact (vs. 2 pairs in contact in E. acutirostris and E. sp. 2); a lower number of ventral scales rows having 21–23 (vs. 26–33 in E. acutirostris, 24 in E. auyanensis, 28 in E. monsfumus); a higher number of scales around mid-body 40–42 (vs. 34–40 in E. acutirostris, 32 in E. auyanensis, 36 in E. monsfumus); a lower number of femoral pores having 3–7 on each side (vs. 8–11 in E. acutirostris, 15–18 in E. auyanensis, 9–10 in E. monsfumus); a higher number of preanal scales 7–8 (vs. 5 in E. sp. 1). Description of the holotype (Figs 1–2). Adult male (CORBIDI 16457); SVL = 69.4 mm, tail (regenerated) length = 55.6 mm; axilla to groin distance = 32.6 mm; head length = 18.1; head width = 11.2 mm; shank length = 11.0 mm. Head scales smooth, glossy; rostral scale wider (2.7 mm) than long (1.5 mm), slightly higher than adjacent supralabials, in contact with frontonasal, nasals, and first supralabials posteriorly; frontonasal pentagonal, slightly wider (2.8 mm) than long (2.6 mm), posterior suture angular with point directed posteriorly, in contact with nasal and laterally, prefrontals posteriorly, nasoloreal suture uncomplete; prefrontals present, in contact with each other medially, in contact with fused anteriormost superciliary-anteriormost supraocular, frontal posteriorly; frontal pentagonal, longer (3.6 mm) than wide (2.7 mm), anterior suture angular with point directed anteriorly, lateral sutures straight, posterior suture angular with point slightly directed posteriorly, in contact with second supraocular laterally, frontoparietals posteriorly; frontoparietals pentagonal, in contact with third and fourth supraocular, parietals and interparietal posteriorly; supraoculars four, none in contact with ciliaries; superciliary series complete, anteriormost superciliary fused with anteriormost supraocular; interparietal pentagonal, longer (3.6 mm) than wide (2.8 mm), lateral sutures concave, in contact with parietals laterally, postparietals posteriorly; parietals polygonal, in contact with fourth supraocular anterolaterally, temporal scales laterally, dorsalmost postocular, postparietals posteriorly; postparietals four, polygonal; palpebral disc oval, unpigmented, divided in four sections; frenocular triangular, in contact with loreal anteriorly; postoculars three; temporals polygonal; supratympanic temporals two; supralabials six; infralabials five; mental wider (2.8 mm) than long (1.7 mm), in contact with first infralabials, postmental posteriorly; postmental single, pentagonal, posterior suture angular, point slightly directed posteriorly, in contact with first and second infralabials; genials in three pairs, anterior pair subquadrangular, in contact with second and third infralabials, medial pair of genials subcuadrangular, in contact with third and fourth infralabials laterally, posterior pair of genials trapezoid, in contact with fourth and fifth infralabials laterally; scale rows between genials and collar fold (along midventral line) 15; medialmost scales of three penultimate gular scale rows cuadrangular; posteriormost gular row enfolded posteriorly, concealing two granular scale rows; lateral neck scales rounded, smooth. Dorsal scales rectangular, longer than wide, juxtaposed, striated, 33 in a longitudinal count, homogeneously arranged; transverse dorsal count (enlarged rows at mid-body) at fifth transverse ventral scale row 10, at 10th transverse ventral scale row 12, at 15th transverse ventral scale row 14; lateral scale rows at fifth transverse ventral scale row 13, at 10th transverse ventral scale row 9, at 15th transverse ventral scale row 10; lateral scales on body near insertion of forelimb small to granular; ventrals cuadrangular and juxtaposed; complete longitudinal ventral count 21; longitudinal ventral scale rows at mid-body 10; 41 scales around mid-body; anterior preanal plate scales two; posterior preanal plate scales seven, all the scales at the same size; scales on tail rectangular and juxtaposed, striated; mid-ventral subcaudals squarish. Limbs pentadactyl; digits clawed; forelimb reaching anteriorly to sixth supralabial; dorsal brachial scales polygonal, of twice the size of anterodorsal midbrachial scale, imbricate, smooth; midbrachial anterodorsal scale small, at least twice shorter than adjacent scales, smooth; anteroventral, ventral, and posteroventral scales roundish, imbricate, smooth; antebrachial scales polygonal, at least as large as midbrachial anterodorsal scale; medial antebrachial scales larger, polygonal, smooth; dorsal manus scales polygonal, subimbricate; palmar scales granular, small, oval; dorsal scales on fingers smooth, quadrangular, covering dorsal half of digit, overhanging supradigital scales, four on I, seven on II, nine on III, 11/10 on IV, six on V; subdigital scales seven on I, 14/13 on II, seventeen on III, 16/17 on IV, eleven on V; anterodorsal thigh scales polygonal, at least three times as large as adjacent scales, becoming smaller ventrally, smooth; posterodorsal thigh scales small, rounded, arranged irregularly; anterior and anteromedial shank scales polygonal, subimbricate, smooth, anteriormost scales many times larger than lateral, posterolateral, and posteromedial shank scales; lateral, posterolateral, and posteromedial shank scales polygonal or roundish, subimbricate, smooth; dorsal pes scales polygonal, subimbricate, smooth; scales on dorsal surface of digits single, quadrangular, smooth, overhanging supradigital scales, four on I, seven on II, thirteen on III, 16/15 on IV, eleven on V; subdigital scales single, seven on I, 14/13 on II, nineteen on III, twentyfour on IV, 16/17 on V; femoral pores series not continued, 5/7; four scales between medialmost femoral pores. Hemipenial morphology. (Fig. 6) The right hemipenis was everted during preservation and prepared posteriorly. The organ extends along approximately 11 mm. The hemipenis is completely expanded and the lobes of the organ are partially everted. Hemipenial body in light bulb shape, with the basis distinctly narrower than the rest of the organ, and ending in two small lobes with apical folds in the apex. The s ulcus spermaticus is central in position, originating at the base of the organ, and proceeding in a straight line towards the lobes. The sulcus is broader in the region of the lobular crotch, where it is divided by a small bulky fold; branches running on the medial region of the lobes, and ending in their tip among folds. The sulcate face of the hemipenial body presents two fleshy nude areas, stained with blue, parallel to the sulcus spermaticus that run through all the hemipenial body extension. Laterals and asulcate face of the hemipenis are ornamented with series of roughly equidistant flounces with calcareous spinules, all of them stained in red with Alizarin. Twenty-four rows of flounces extend along the body of the organ. There are three proximal rows restricted to a central position on the basal asulcate face of the hemipenis, all of them are roughly chevron-shaped. The five proximalmost flounces in the laterals are diagonally positioned, with the second to fifth flounces separated from a complementary flounce positioned in the asulcate face and orientated in an inverse diagonal. The nude area between the proximalmost fifth flounce is narrower than the nude area between the other four proximal flounces and its complementary ones. The subsequent flounces towards the lobes cross the laterals of the organ from the sulcate to the asulcate face, in chevron-shape with vertexes in the central region of each lateral pointed towards the basis of the organ. These chevron-shaped rows reduce in size progressively towards the hemipenial apex. The lateral flounces are separated in two groups by a nude area in the central asulcate face that increases in size in the apical region acquiring a Y-shape. The regions between the asulcate face and the laterals in this species are marked by a conspicuous unevenness forming a bulge shared by other related species such as Macropholidus annectens, M. huancabambae, M. ruthveni, Pholidobolus affinis, P. macbrydei, P. montium, P. prefrontalis (Nunes, 2011), P. hillisi (Torres-Carvajal et al. 2014), and P. ulisesi (Venegas et al. 2016). Color of holotype in life. (Fig. 2) Head golden-brown with scattered black spots, down to inferior level of supralabials, infralabials creamy white with black spots; a dark stripe running from the nasal plate to the anterior angle of the eye and from the posterior angle of the eye to the supratympanic temporals, iris copper-red. Dorsum golden-brown with scattered black spots, a dorsolateral dark-bordered red-orange stripe from nuchals to the base of the tail, dark border turning paler and fading entirely towards the base of tail, seven pairs of lateral ocelli. Dorsal surface of tail golden brown with pale dorsolateral spots. Throat, chest, belly and ventral surfaces of limbs creamy white, with scattered black spots or blotches. Precloacal and cloacal scales creamy white with scattered black spots; ventral surface of tail creamy white with scattered black blotches. Variation and sexual dimorphism. (Fig. 3, 4) Variation in measurements and scutellation is summarized in Table 1. The largest male is CORBIDI 16465 (81.7 mm), the largest female is CORBIDI 16466 (81.4 mm). Females CORBIDI 15573 and 16466 have two genials in contact, not three as the rest of type series. Six specimens have the supraocular-superciliar scale fusion divided by a diagonal, weakly defined, complete or uncomplete suture (CORBIDI 16396, 16459, 16463, 16465–66, 16468); also a juvenile (CORBIDI 16468) has six superciliars. Dorsolateral red-orange dark bordered stripe is extended to 10th verticile of the tail in male CORBIDI 16465. Juveniles CORBIDI 16467–68 have yellow coloration on belly and ventral surfaces of hindlimbs and tail. The sex can be distinguished by a higher number of femoral pores in males (10–12) than females (4–6), only juvenile male CORBIDI 16459 has 6 femoral pores, furthermore males can be differentiated by the presence of 7–14 pairs of lateral ocelli (absent in females), and by the presence of black scattered spots or blotches on ventral surface of tail in males, instead black spots or blotches arranged ventrolaterally on tails of females. Additionally, male CORBIDI 16465 has an atypical high number of irregular black spots on belly, remarkably higher than the rest of males of the type series. Etymology. The specific epithet “ excelsum ” is from Latin and means “tall”, in reference to the habitat of the new species in the canopy of the tropical rainforest. Distribution and natural History. Euspondylus excelsum sp. nov. is known from localities on both sides of the Huallaga river, including within Tingo Maria National Park (Fig. 7, 8), at distances as far as 48.5 km between the northernmost and southernmost collecting sites. Specimens were collected in the canopy of primary and secondary forests. Specimens CORBIDI 14666 and 15573 were observed during diurnal Visual Encounter Surveys climbing trunks at 1.5 m above leaf litter. The holotype and specimens CORBIDI 16396, 16450, 16459, 16463-65, were collected in the canopy of Couma spp. at night; specimens CORBIDI 16466–68 were collected under leaves of bromeliads gathered on crags. Male CORBIDI 16465 started swimming underwater during an escape attempt prior to capture. After dissection for laboratory analyses, two eggs were observed into the ventral cavity of three females of type series collected at November 2014 and October-November 2015, eggs lengths were (measurements of egg in right oviduct is presented first): 18.0 mm and 18.0 mm in CORBIDI 15573 (23.2% of the SVL), 16.6 mm and 16.5 mm in CORBIDI 16463 (20.2% and 20.4% o

Stenocercus omari sp. nov. Figs. 1–5, Table 1. Holotype. CORBIDI 0 0 577, an adult male from Abra Chanchillo (06˚47'07.00'' S, 77˚56'32.50'' W), 2786 m, Chachapoyas province, Amazonas department, Peru, collected by P. J. Venegas on 17 June 2007. Paratypes. Peru: Amazonas department: Chachapoyas province: CORBIDI 00569–70, 00578–80, adult females, and CORBIDI 00571–76, adult males, collected with the holotype; Luya province: CORBIDI 11961, adult male, and CORBIDI 11962, adult female, from Pircopampa (06°19'32.17'' S, 78°13'1.99'' W), 2378 m, collected by K. García-Burneo on 23 September 2012; La Libertad department: Bolivar province: CORBIDI 05793–94, sub adult female and adult male, respectively, and ZFMK 90835 adult female from Bolivar (07°10'00.00'' S, 77°42'46.50'' W), 2772–3035 m, collected by A. García-Bravo and C. Koch on 21 April 2009. Diagnosis. Stenocercus omari is distinguished from other species in the genus, except for S. amydrorhytus, S. chrysopygus, S. cupreus, S. johaberfellneri, S. latebrosus, S. melanopygus, S. modestus, S. ornatissimus, S. orientalis, and S. stigmosus by having granular scales on the posterior surfaces of thighs, a conspicuous antehumeral fold and by lacking a vertebral crest. Stenocercus omari is easily distinguished from the aforementioned species, except for S. orientalis, by the presence of prominently keeled dorsal head scales. The new species differs from S. orientalis in lacking a prominent oblique neck fold with a mite pocket under it, a distinctive character in S. orientalis, and also present in S. latebrosus and S. ornatissimus. Another important difference between the new species and S. orientalis is the presence of a longitudinal neck fold in the former, absent in S. orientalis; such fold is also present in S. melanopygus and S. stigmosus; however, unlike the new species described herein, both species have smooth or slightly keeled dorsal head scales. Additionally, the new species has a distinct deep postfemoral mite pocket with a posteroventrally oriented slit-like opening, whereas S. orientalis has one or more vertical folds instead (Fig. 4). The new species can be distinguished from S. orientalis in the field by color pattern. Adult males of S. omari have distinct dark triangular marks longitudinally arranged on each side of the vertebral line (Fig. 1 A, 3A, 5A) and S. orientalis usually has a tabby dorsal pattern (Fig. 5 D) that in some individuals is indistinct. Although males of both species share a yellow ventral coloration, males of the new species have a blue throat with dark flecks, whereas males of S. orientalis have a yellow throat with or without dark flecks and a bluish center in some specimens (Fig. 5 E). Adult females of S. omari and S. orientalis are similar in coloration pattern; however, females of S. omari, have dorsolateral and ventrolateral light stripes on each side of the neck and females of S. orientalis have only one dorsolateral light stripe on each side of the neck (Fig. 5 F). Characterization. (1) Maximum SVL in males 68 mm (N =8); (2) maximum SVL in females 69 mm (N =7); (3) vertebrals 36–47; (4) paravertebrals 38–52; (5) scales around midbody 42–54; (6) supraoculars 4–7; (7) internasals 4; (8) postrostrals 2–4; (9) loreals 2–5; (10) gulars 13–25; (11) lamellae on Finger IV 18–21; (12) lamellae on Toe IV 18–29; (13) posthumeral pocket present as one vertical fold; (14) postfemoral mite pocket present as a distinct deep pocket with a posteroventrally oriented slit-like opening [Type 2 of Torres-Carvajal (2007a)]; (15) parietal eye not visible; (16) occipital scales small, keeled, subimbricate; (17) projecting angulate temporal absent; (18) supraoculars subequal in size; (19) scales in frontonasal region subimbricate, keeled; (20) short preauricular fringe present; (21) antehumeral and longitudinal neck folds present; (22) lateral nuchals half the size of dorsal nuchals; (23) posterior gulars in adults smooth, imbricate; (24) lateral scales slightly smaller than dorsal body scales; (25) vertebral crest absent; (26) dorsolateral crests absent; (27) ventrals in adults broad, lanceolate, smooth, imbricate; (28) scales on posterior surfaces of thighs granular; (29) prefemoral fold absent; (30) inguinal groove absent; (31) preanals not projecting; (32) tail not compressed laterally in adult males; (33) tail length 53–70 % of total length; (34) caudal whorls per autotomic segment three; (35) caudals not spinose; (36) dark stripe extending anterodorsally from subocular region to supraciliaries absent; (37) gular region of adult females pigmented with or without dark spots; (38) gular region of adult males pigmented with dark spots; (39) black blotch on ventral surface of neck in adult males absent; (40) thin black or dark brown midventral line absent; (41) black patch on ventral surface of thighs absent; (42) dorsal ground color brown with darker triangular marks longitudinally arranged on each side of vertebral line in females and males; (43) two xiphisternal and two postxiphisternal pairs of inscriptional ribs, the two postxiphisternal pairs of ribs short and not in contact midventrally (Pattern 1C of Torres-Carvajal, 2004a). Description of the holotype. Male (Figs. 1–3); SVL 66 mm; TL 157 mm; maximum head width 10.3 mm; head length 15.3 mm; head height 9.2 mm; occipitals, parietals, interparietals, and postparietals small, keeled, imbricate; parietal eye not visible; supraoculars in six rows, keeled, imbricate, subequal in size; canthals two; anterior most canthal in contact with the nasal; scales on frontonasal region slightly imbricate and keeled; internasals four; postrostrals four, two most lateral wider than long on each side, medial postrostrals as long as wide; supralabials five; infralabials six; loreals three; lorilabials in one row; preocular one, in contact with posterior canthal; lateral temporals keeled, imbricate; gulars in 22 rows between tympanic openings; all gulars smooth, imbricate, apical pit absent; second infralabial not in contact with second and third sublabials; mental in contact with first pair of infralabials; lateral and dorsal scales of body and neck keeled, imbricate, mucronate; lateral scales slightly smaller than dorsal body scales; scales around midbody 44; vertebrals same size as dorsals, 39 scales on vertebral row, prominent serrate vertebral crest absent; paravertebrals 45; ventrals broad, lanceolate, smooth, imbricate; preauricular fringe short, composed of three enlarged scales, all same size; antegular, gular, postauricular, oblique and supraauricular neck folds absent; longitudinal and antehumeral neck folds present; limb scales keeled, imbricate; ventral scales of hind limbs smooth and upper arms keeled; lamellae on Finger IV 18; lamellae on Toe IV 18; tail not compressed laterally; caudals keeled, imbricate, mucronate; basal subcaudals smooth, imbricate; tail length 2.4 times SVL; posthumeral pocket present as one vertical fold; postfemoral mite pocket present as a distinct deep pocket with a posteroventrally oriented slit-like opening [Type 2 of Torres- Carvajal (2007a)]; postfemoral region composed of imbricate, smooth scales that become smaller towards insertion of hind limb. Color in life. Head and neck dark brown (almost black) with white flecks. A black vertical blotch on the antehumeral fold and a black rounded blotch on the side of the neck. Dorsum brown with dark brown triangular marks longitudinally arranged on each side of the vertebral line, and flanks yellowish cream with dark brown bands. Forelimbs dark brown with yellow flecks and hind limbs brownish yellow with dark brown transverse bands. Tail brown with dark brown marks along it. Ventrally, throat and chest sky blue with dark blue marks on the throat; ventral surface of thighs, cloacal region, and base of tail dark yellow. Color in ethanol 70%. Head brown, white flecks turned cream. Dorsal background color of body and base of tail brownish cream with dark brown triangular marks (but lighter than in life). Sides of neck brownish cream with brown, instead of black, blotches. Flanks brownish cream flecked by brown and light cream scales, and ventrolateral region sky blue. Limbs brown with dark marks and distal portion of tail brown with light transverse bands. Ventrally the yellow surfaces turned cream. Stenocercus omari sp. nov. Variation. Some measurements and scutellation characters of Stenocercus omari are presented in Table 1. Second infralabial not in contact with third sublabial in 93.8% of specimens, and first pair of postmentals in contact medially in all specimens. All male paratypes are similar to the holotype, varying only in the tone of ventral coloration; although the adult male specimen from Bolivar (CORBIDI 05794) lacks the black blotch on the sides of neck. Adult females have a cinnamon or dark brown dorsum with distinct or faint darker triangular marks longitudinally arranged on each side of the vertebral line; sides of the head and flanks brown (darker than the dorsum); supralabials white or cream, continuous with a white or cream dorsolateral stripe along the neck, becoming pale brown along the dorsum; a white or cream stripe along the edge of the jaw continues as a white ventrolateral stripe along the neck, becoming dirty cream on the flanks; flanks with or without vertical bands; dorsal surface of tail with or without transverse bands. Ventral surface dirty cream or white with or without dark flecks on the throat. Sexual dimorphism is evident in adult individuals. Females can be easily recognized by lacking white flecks on the dorsal surface of head and neck, and by having dark brown flanks (yellowish in males), and a dirty cream or white ventral coloration (throat sky blue, chest and belly yellow in males). Distribution and natural history observations. Stenocercus omari is known from three localities along the western slope of the northern portion of the Cordillera Central at the Marañón river basin in the departments of Amazonas and La Libertad, at elevations of 2300–3035 m. According to Brack (1986) the distribution of S. omari lies inside the ecoregion of the Serrania esteparia and according to Olson et al. (2001) this species is distributed in the Peruvian Yungas ecoregion. The habitat at the type locality is a steep area located on the right side of the Marañón river with the presence of corn plantations, cattle pasture and some scattered patches of montane forest along the slope. Stenocercus omari was very common along the sides of the Leimebamba-Balsas road, a steep band covered by herbaceous vegetation with big boulders and scattered bushes. All collected individuals were found active in daylight (between 11:00 and 15:00) basking on rocks, fallen tree trunks and at the base of bushes. A gravid female (CORBIDI 00570) with SVL 65 mm collected on June contained eight yolked follicles, four in each ovary. Another gravid female (CORBIDI 11962) with SVL 69 mm collected on September contained seven well developed eggs four in the left oviduct and three in the right one, eggs ranged from 13.96–15.15 mm in length, 7.56–8.87 mm in width, and 422.33–613.98 mm 3 in volume. Etymology. The specific epithet omari is a noun in the genitive case and is a patronym for our appreciated friend, the Ecuadorian herpetologist Omar Torres-Carvajal, who has published important contributions to the taxonomy and systematics of Neotropical lizards, especially of the genus Stenocercus.

Phyllodactylus pachamama sp. nov. Phyllodactylus reissii— Koch, Venegas & Böhme, Salamandra 42(2/3): 148. — 2006 Phyllodactylus reissii— Venegas, Townsend, Koch & Böhme, Journal of Herpetology 42(2): 393. — 2008 Phyllodactylus reissii— Koch & Beraun, Check List 7(3): 272. — 2011 Phyllodactylus reissii— Aurich, Koch & Böhme, North-Western Journal of Zoology 7(2): 310. — 2011 Phyllodactylus reissii (partim) ― Torres- Carvajal, Barnes, Pozo- Andrade, Tapia & Nicholls, Journal of Biogeography 41: 1887. — 2014 Phyllodactylus reissii (partim) — Aurich, Koch & Böhme, Phyllomedusa 14(1): 55. — 2015 Holotype. An adult male (ZFMK 90886) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6°50’45.0’’S, 77°59’47.9’’W, 1271 m a.s.l.), collected on 26 April 2009 by A. Garcia Bravo and C. Koch. Paratypes (24). An adult male (ZFMK 85004) and an adult female (ZFMK 85005) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6°48’26’’S, 77°59’29’’W, 1000 m a.s.l.), collected on 10 July 2005 by P. Venegas and C. Koch; three adult females (CORBIDI 5700, 5701; ZFMK 90885) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6°51’ S, 78°01’ W, 896—900 m a.s.l.), collected on 15 April 2009 by A. Garcia Bravo and C. Koch; an adult male (CORBIDI 7710) and a subadult male (ZFMK 91727) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6°51’ S, 78°02’ W, 870—919 m a.s.l.), collected on 30 March 2010 by J. Aurich and L. Y. Echevarria Espinosa; an adult female (ZFMK 91725) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6°50’56.7’’ S, 78°01’32.4’’ W, 895 m a.s.l.), collected on 15 April 2010 by J. Aurich and L. Y. Echevarria Espinosa; four adult males (CORBIDI 7711, 7712; ZFMK 91724, 91726) and a subadult female (ZFMK 91728) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6°51’ S, 78°01’ W, 920 m a.s.l.), collected on 27—29 May 2010 by J. Aurich and C. Laandauro Sanabria; two adult males (CORBIDI 5703; ZFMK 90890), two adult females (CORBIDI 5704, 5705), and a juvenile (ZFMK 90889) from Zapatalgo, Province of Utcubamba, Department of Amazonas, Peru (6°05’ S, 78°29’ W, 830—1030 m a.s.l.) collected on 6—8 December 2009 by A. Garcia Bravo and C. Koch; an adult male (CORBIDI 5706) and a subadult female (ZFMK 90892) from Zapatalgo, Province of Utcubamba, Department of Amazonas, Peru (6°04’ S, 78°30’ W, 907 m a.s.l.) collected on 10 December 2009 by A. Garcia Bravo and C. Koch; a juvenile (ZFMK 90891) from Zapatalgo, Province of Utcubamba, Department of Amazonas, Peru (6°06’05.2’’ S, 78°29’54.5’’ W, 521 m a.s.l.) collected on 9 December 2009 by A. Garcia Bravo and C. Koch; two adult males (ZFMK 90887, 90888) and an adult female (CORBIDI 5702) from San Vicente /Pusac, Province of Bolívar, Department of La Libertad, Peru (6°59’ S, 77°55’ W, 1425—1432 m a.s.l.) collected on 22 April 2009 by A. Garcia Bravo and C. Koch. Diagnosis and comparison. Phyllodactylus pachamama sp. nov. is a comparatively large species with a maximum SVL of 77 mm. Among species of Phyllodactylus found in mainland South America only P. delsolari, P. dixoni, P. reissii and P. v en t r al i s exceed 70 mm SVL. Thus, this species can be distinguished from all other species by its larger size. In lacking an enlarged postanal plate it further differs from P. thompsoni and by the absence of a preanal plate it can be distinguished from P. angustidigitus, P. gerrhopygus, and P. heterurus. By having 10—16 well-defined rows of enlarged, trihedral keeled tubercles, the new species differs from P. angustidigitus, P. gerrhopygus, P. heterurus (dorsal tubercles absent in all three species), P. delsolari, P. inaequalis (fewer than 10 poorly defined rows of small, smooth, round tubercles in both species), P. microphyllus (dorsal tubercular rows indistinct, composed of small flat, oval tubercles), and P. thompsoni (10). By having 43—56 paravertebral tubercles between rear of head and posterior edge of thigh it differs from P. interandinus (54—83), P. k of ord i (31—36), P. pumilus (37—43), P. sentosus (26—31), and P. thompsoni (36—40). Phyllodactylus pachamama can be distinguished from P. heterurus, P. kofordi, P. pumilus, and P. sentosus by lacking tubercles on the base of tail. By the presence of tubercles on the tibia this species can further be differentiated from P. angustidigitus, P. clinatus, P. delsolari, P. gerrhopygus, P. inaequalis, P. interandinus, P. lepidopygus, and P. microphyllus. By the absence of tubercles on the thigh it can be distinguished from P. d i x o ni, P. k o f o rd i, P. pu m i l u s, P. s e n t o s u s, and P. ventralis. By the absence of tubercles on the forearm it can be differentiated from P. dixoni, P. kofordi, P. sentosus, and P. ventralis. From P. angustidigitus, P. microphyllus, and P. sentosus the new species further differs by having large terminal lamellae. It can be distinguished by the number of lamellae under the fourth toe (14—17, mean 15.1) from P. dixoni (11—13, mean 11.3), P. inaequalis (10—12, mean 10.7), P. johnwrighti (7—14, mean 10.5), P. k o f o rd i (11—13, mean 11.5), P. leoni (9—13, mean 10.9), P. lepidopygus (10—14, mean 11.5), and P. pumilus (11—13, mean 12.0). 40 % of specimens have 3 postmentals compared to only 1.9 % in P. reissii. In contrast to P. johnwrighti and P. pumilus (dark stripe from nostril to arm insertion always present), the postnasal and postorbital brown stripes in P. pachamama is usually absent or, if present, poorly defined. Description of holotype (Fig. 5). Head distinct from neck, covered with small granules that are largest on snout and smallest on interparietal region; rostral 3.3 mm wide, 2 mm high, somewhat rectangular, with a median groove along its posterior half; two internasals, in broad medial contact, each bordered posteriorly by two granules and a postnasal; nostril surrounded by rostral, first supralabial, two postnasals and internasal; 11 loreals between postnasal and anterior edge of orbit; scales in loreal region about three times larger than scales in interorbital region; 24 scales across snout at level of third supralabial, 17 at level of second supralabial; 12 supralabials, six to a point below the center of eye; eyes large, pupil vertically elliptical, with corrugated edges; orbital diameter 4.6 mm; eyelid with three visible rows of granules on left side and five on right side, granules of median row pointed in posterior part; 25 midorbital granules; interorbital distance 1.8 mm; 29 granules between posterior edge of eye and anterior edge of ear, eye to ear distance 7.1 mm; ear opening elliptical, posterodorsally oriented, 2.4 mm high and 0.7 mm wide, its anterior and posterior edges denticulated; rear of head intermixed with slightly larger tubercles; mental bell-shaped, larger than infralabials, 4 mm long, and 4 mm wide at its widest point; two postmentals, in contact with first pair of infralabials laterally and five scales posteriorly; nine infralabials, six to a point below center of eye, first two pairs largest, adjacent pairs of infralabials descreasing in size posteriard; scales on chin largest and juxtaposed in anterior part, decreasing in size and becoming more imbricate towards gular region. Body depressed; dorsal surface of neck with small granules that are slightly smaller than those on the head; dorsum covered with small granules and 10 to 12 longitudinal rows of enlarged, trihedral keeled tubercles at midbody; six rows of tubercles from occiput to base of tail; paravertebral row with 44 tubercles between occiput and cloaca, 32 between axilla and groin; paravertebral rows separated from each other by about six granules; tubercles in paravertebral row separated from each other by 1—2 granules; 2—4 granules between other longitudinal tubercular rows; 110 scales around the midbody; ventral surface with smooth, imbricate scales, 74 from posterior gular region to cloaca, 26 transversely across midbody; dorsal surface of forearm with flattened, imbricate scales and without tubercles; thigh with flat, imbricate scales, largest in size anterodorsally and smallest posterodorsally, without tubercles; dorsal surface of tibia with a few scattered, hardly visible, cone-shaped or pulvinate tubercles, which are much smaller than the large tubercles on the back and only slightly larger than other scales on tibia; subdigital lamellae on left fingers (I—V) 8—10—12—13—11; subdigital lamellae on left toes (I—V) 8—11—15—15—13; terminal lamellae paired, greatly enlarged, truncated, the tip of the claw hardly visible between the two terminal pads, not projecting the terminal pads; tail complete, 1.18 times the SVL, covered with smooth and imbricate scales, on dorsal surface of tail about 3—4 times larger than dorsal body granules; medial subcaudals broadly enlarged, about two times as wide as long in anterior part of tail, decreasing in width posteriard. Measurements in mm. — Snout-vent length 71; tail length 84; axilla-groin length 29; right forelimb length 24.0; right hindlimb length 31.9; head length 21.6; maximum head width 15.9; midorbital head width 12.3; head height 8.9. Coloration of holotype in ethanol (Fig. 5). Dorsal ground color of head, body and limbs pale brown, with irregular small brown flecks and blotches on the dorsal surface of the body and limbs; indistinct brown line extending laterally on each side from nostril through eye to above insertion of the forearm; lateral surface of the body cream with brown mottling on each scale; ventral surface of head and body cream, with brown mottling on each scale of the chest region and towards the venterolateral parts of head and body, scales on median part of venter and gular region almost without mottling; dorsal surface of tail with alternating cream, brown and dark brown blotches; subcaudal ground color cream, with dense grayish brown mottling; lamellae under fingers and toes grayish. Variation (see also Table 3). SVL of male paratypes ranges from 48–75 mm (68.8 ± 4.3, n = 9), and SVL of female paratypes ranges from 44–77 mm (63.7 ± 11.6, n = 10). Tail length/total length is 0.43–0.56 (0.51 ± 0.03, n = 18). Male specimens possess 3—4 postanal tubercles on each side, which are slightly elevated. In females, postanal tubercles are lacking or only hardly visible and usually less pronounced as in males. The number of postmentals is 2—4 (2.5 ± 0.64). In one specimen (ZFMK 90887), postmentals contact first and second infralabials on each side, in all other specimens only first infralabial contacts postmentals, scales bordering postmentals 5—8 (6.4 ± 1). Two paratypes (ZFMK 90887, CORBIDI 5705) have 3 internasals, all others have 2 (2.1 ± 0.29). Lamellae on fourth finger 10—14 (12.5 ± 0.86); lamellae on fourth toe 14—17 (15.1 ± 0.81); supralabials 8—13 (10.2 ± 1.4), to a point below the center of eye 6—9 (7.7 ± 0.83); infralabials 7—11 (8.7 ± 1.03), to a point below the center of eye 5—7 (6.2 ± 0.7); loreals between postnasal and anterior edge of orbit 9—13 (11.6 ± 1.3); granules between posterior edge of eye and anterior edge of ear 24—28 (26.4 ± 1.1); scales across snout at level of third supralabial 22—26 (24.1 ± 1.4), at level of second supralabial 15—19 (17.1 ± 1.6); midorbital granules 23—25 (23.9 ± 0.86); longitudinal rows of dorsal tubercles 10—16 (13.2 ± 1.4); paravertebral tubercles between rear of head and cloaca 43—56 (49.6 ± 3.3), between axilla and groin 25—32 (29.1 ± 2.2); scales around midbody 92—116 (103 ± 7.4); ventral scales from gular region to cloaca 67—87 (73.5 ± 5.7), across midbody 26—36 (29.4 ± 3). In life (Fig. 6), dorsal ground color of head, body and limbs of paratypes varies between yellowish-white, grayish-white, cream and pale brown; irregular median or dark brown flecks and blotches on the dorsal surface of the body and limbs may either be almost lacking or quite prominent and forming transverse dorsal bands or a more or less reticulate pattern, that may be discontinuous in some specimens resulting in a pale cream or light brown middorsal line; postnasal and postorbital brown stripe lacking in most specimens; ventral surface of head and body cream or yellowish, with or without brown mottling; dorsal surface of unregenerated tails with alternating cream or light brown, median brown and dark or blackish brown blotches; ventral surface of tail varying from uniformly cream or yellowish to a grayish brown reticulation with strong brown mottling; pupil silver-gray or slightly bronzecolored. In ethanol, dorsal ground color of head, body and limbs of paratypes varies between pale brown and cream; ventral surface of head and body cream; ventral surface of tail varying from uniformly cream to grayish with strong grayish-brown mottling. Etymology. The specific epithet pachamama is derived from the Quechuan language and can be translated as Mother Earth. It represents the name of a goddess revered by indigenous people from South America, and in particular from Andean regions. According to many believers, problems will arise when people take too much from nature, because they are stealing it from Pachamama. This name was chosen to emphasize the need for protected areas along the Marañón river and blames the numerous ongoing mining and dam construction activities for exploiting this unique habitat. Distribution and natural history. Phyllodactylus pachamama sp. nov. is endemic to the upper valley of the Marañón River and some of its tributaries in the Northern Peruvian Andes from Zapatalgo, Province of Utcubamba, Department of Amazonas to San Vicente, Province of Bolívar, Department of La Libertad. It inhabits the seasonally dry forest, occurring in xeric habitats dominated by less dense shrubs and cacti (e.g. Armatocereus, Browningia, Espostoa) at elevations between 521—1432 m a.s.l. Individuals of this nocturnal gecko were active between 6.25 pm and 11.35 pm and mostly found on rock walls or house walls in heights up to about 3 m above the ground. Some individuals were also found running over sandy ground next to the Marañón River. Only one male individual (CORBIDI 5703) was found during the day hidden under some logs. During their active hours, air temperatures ranged from 23.0 to 29.7° C, substrate temperatures ranged from 20.7 to 30.0° C and air humidity ranged from 41 to 71%. Aurich et al. (2011) examined the stomach contents of 8 individuals of P. pachamama from Balsas (referred to as P. reissii), which contained exclusively arthropod material, dominated by coleopterans, which made up 40% of all prey items. Three gravid females (CORBIDI 5700; ZFMK 90885, 91725) were collected in mid April (2009 and 2010), each contained two eggs. On 10 July 2005, a gravid female (ZFMK 85005) was collected which contained only one egg. Two juveniles with a SVL of 31 mm (ZFMK 90889) and 32 mm (ZFMK 90891) were found in December 2009. Most gravid females of the species examined by Aurich et al. (2015) also contained two eggs. In Balsas and San Vicente this species occurs sympatrically with the congeners P. delsolari and P. thompsoni, and with Phyllopezus maranjonensis, whereas in Zapatalgo it was found sympatrically with the sphaerodactylid Pseudogonatodes barbouri.

Phyllodactylus reissii Phyllodactylus reissii Peters, Monatsberichte der K��niglichen Preussische Akademie der Wissenschaften zu Berlin: 626.―1862 Phyllodactylus reissii��� Boulenger, Catalogue of the lizards in the British Museum (Nat. Hist.): 80. ��� 1885 Phyllodactylus baessleri Werner, Abhandl. Ber. Zool. Anthrop. Mus. Dresden, 9 (2): 2. ��� 1901 Phyllodactylus guayaquilensis Werner, Mitt. Naturh. Mus. Hamburg 27 (2): 4. ��� 1910 Phyllodactylus abrupteseriatus Werner, Mitt. Naturh. Mus. Hamburg 30, 2. Beiheft: 4. ��� 1913 Phyllodactylus abrupteseriatus��� Burt & Burt, Bulletin American Museum of Natural History, 61: 249. ��� 1931 Phyllodactylus magister (partim) Burt & Burt, Bulletin American Museum of Natural History, 61: 250. ��� 1931 Phyllodactylus reissii (partim) ��� Dixon & Huey, Los Angeles County Museum Contributions in Science 192: 50. ��� 1970 Phyllodactylus reissii (partim) ��� Peters & Donoso-Barros, United States Nat. Mus. Bull. 297: 242. ��� 1970 Phyllodactylus reissii (partim) ��� Peters & Donoso-Barros, Smithsonian Institution Press, Washington D.C. & London: 242. ��� 1986 Phyllodactylus reissii (partim) ― Torres- Carvajal, Barnes, Pozo- Andrade, Tapia & Nicholls, Journal of Biogeography 41: 1887. ��� 2014 Phyllodactylus reissii (partim) ��� Aurich, Koch & B��hme, Phyllomedusa 14(1): 55. ��� 2015 Lectotype. ZMB 4567 (designated by Dixon & Huey 1970), adult female with a SVL of 64 mm, collected from Guayaquil, Ecuador by C. Reiss. Paralectotypes (5). All paralectotypes (ZMB 3734 (4), ZMB 4567 (1)) were collected by C. Reiss together with the Lectotype; date of collection unkown. New collected material (18). See Appendix 2. Additional material examined (21). See Appendix 2. Diagnosis and comparison. Phyllodactylus reissii is a comparatively large species, probably reaching a SVL of 75 mm (according to data of Dixon & Huey 1970). Thus, by its larger size this species can be distinguished from all species of Phyllodactylus found in mainland South America except P. delsolari, P. dixoni, P. pachamama and P. ventralis. It can further be distinguished from P. thompsoni by the lack of an enlarged postanal plate and from P. angustidigitus, P. heterurus, and P. gerrhopygus by the absence of a preanal plate. By having ���10 well-defined rows of enlarged, trihedral keeled tubercles on the dorsum, P. reissii differs from P. angustidigitus, P. gerrhopygus, P. heterurus (dorsal tubercles absent in all three species), P. delsolari, P. inaequalis (fewer than 10 poorly defined rows of small, smooth, round tubercles in both species), and P. microphyllus (dorsal tubercular rows indistinct, composed of small flat, oval tubercles. Phyllodactylus sentosus (26���31) generally has a lower number of paravertebral tubercles between rear of head and cloaca. By the presence of tubercles on the tibia, P. reissii can further be differentiated from P. angustidigitus, P. clinatus, P. delsolari, P. gerrhopygus, P. inaequalis, P. interandinus, P. lepidopygus, and P. microphyllus, and by the absence of tubercles on the forearm it can be differentiated from P. dixoni, P. kofordi, P. sentosus, and P. ventralis. The absence of tubercles on the tail further differentiates it from P. he t er ur u s, P. k of o rd i, P. p u m i l u s, and P. sentosus. In contrast to P. angustidigitus, P. microphyllus and P. sentosus, P. reissii possesses large terminal lamellae. It further differs from P. m a gi s t e r by the absence of small tubercles on the thigh (present in about half of the in P. magister). On average the number of scales around midbody (69���102, mean 84.1) is lower when compared to P. magister (78���114, mean 90.3) and P. pachamama (92���116, mean 103.0). Compared to P. magister the anterior edge of the ear opening is strongly denticulated with pointed scales (smooth or only slightly denticulated in most specimens of P. magister). Redescription. A detailed redescription of the species is given in Dixon & Huey (1970) on the basis of their designated lectotype. Variation. Dixon & Huey (1970) provide detailed information on the variation of this species, but unfortunately their variation section is also based on specimens belonging to P. magister and some specimens from populations that may represent so far undescribed species. Table 3 shows the morphological variation of some characters we examined in 39 specimens of Phyllodactylus reissii (see Appendix 2 for locality information). Distribution and natural history. Phyllodactylus reissii is widely distributed in coastal and western Andean regions in northern Peru and southern Ecuador from sea level to at least 1305 m a.s.l.. The species was recently introduced on the Gal��pagos islands (Hoogmoed 1989) and seems to have already formed established populations (Torres-Carvajal & Tapia 2011). One population was found in the western part of the Mara����n valley in northern Peru. Gravid females of Phyllodactylus reissii were detected end of March (CORBIDI 5693) and in July (ZFMK 88744, 88745), each containing two eggs. Three juveniles with a SVL of 28 mm (CORBIDI 5689), 31 mm (ZFMK 90877), and 34 mm (CORBIDI 5690) were found in March 2009. Two subadults (CORBIDI 5687, 5688) with a SVL of 41 mm and 36 mm, respectively, were collected in March 2009. In Pucar�� we found P. reissii in sympatry with the congener P. johnwrighti., Published as part of Koch, Claudia, Flecks, Morris, Venegas, Pablo J., Bialke, Patrick, Valverde, Sebastian & R��dder, Dennis, 2016, Applying n-dimensional hypervolumes for species delimitation: unexpected molecular, morphological, and ecological diversity in the Leaf-Toed Gecko Phyllodactylus reissii Peters, 1862 (Squamata: Phyllodactylidae) from northern Peru, pp. 41-80 in Zootaxa 4161 (1) on pages 56-57, DOI: 10.11646/zootaxa.4161.1.2, http://zenodo.org/record/256497, {"references":["Dixon, J. R. & Huey, R. B. (1970) Systematics of the lizards of the gekkonid genus Phyllodactylus of mainland South America. Los Angeles County Museum Contributions in Science, 192, 1 - 78.","Hoogmoed, M. S. (1989) Introduced geckos in Puerto Ayora, Santa Cruz, with remarks on other areas. Noticias de Galapagos, 47, 12 - 16.","Torres-Carvajal, O. & Tapia, W. (2011) First record of the common house gecko Hemidactylus frenatus Schlegel, 1836 and distribution extension of Phyllodactylus reissii Peters, 1862 in the Galapagos. Check List, 7 (4), 470 - 472."]}

Phyllodactylus pachamama sp. nov. Phyllodactylus reissii��� Koch, Venegas & B��hme, Salamandra 42(2/3): 148. ��� 2006 Phyllodactylus reissii��� Venegas, Townsend, Koch & B��hme, Journal of Herpetology 42(2): 393. ��� 2008 Phyllodactylus reissii��� Koch & Beraun, Check List 7(3): 272. ��� 2011 Phyllodactylus reissii��� Aurich, Koch & B��hme, North-Western Journal of Zoology 7(2): 310. ��� 2011 Phyllodactylus reissii (partim) ― Torres- Carvajal, Barnes, Pozo- Andrade, Tapia & Nicholls, Journal of Biogeography 41: 1887. ��� 2014 Phyllodactylus reissii (partim) ��� Aurich, Koch & B��hme, Phyllomedusa 14(1): 55. ��� 2015 Holotype. An adult male (ZFMK 90886) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6��50���45.0������S, 77��59���47.9������W, 1271 m a.s.l.), collected on 26 April 2009 by A. Garcia Bravo and C. Koch. Paratypes (24). An adult male (ZFMK 85004) and an adult female (ZFMK 85005) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6��48���26������S, 77��59���29������W, 1000 m a.s.l.), collected on 10 July 2005 by P. Venegas and C. Koch; three adult females (CORBIDI 5700, 5701; ZFMK 90885) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6��51��� S, 78��01��� W, 896���900 m a.s.l.), collected on 15 April 2009 by A. Garcia Bravo and C. Koch; an adult male (CORBIDI 7710) and a subadult male (ZFMK 91727) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6��51��� S, 78��02��� W, 870���919 m a.s.l.), collected on 30 March 2010 by J. Aurich and L. Y. Echevarria Espinosa; an adult female (ZFMK 91725) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6��50���56.7������ S, 78��01���32.4������ W, 895 m a.s.l.), collected on 15 April 2010 by J. Aurich and L. Y. Echevarria Espinosa; four adult males (CORBIDI 7711, 7712; ZFMK 91724, 91726) and a subadult female (ZFMK 91728) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6��51��� S, 78��01��� W, 920 m a.s.l.), collected on 27���29 May 2010 by J. Aurich and C. Laandauro Sanabria; two adult males (CORBIDI 5703; ZFMK 90890), two adult females (CORBIDI 5704, 5705), and a juvenile (ZFMK 90889) from Zapatalgo, Province of Utcubamba, Department of Amazonas, Peru (6��05��� S, 78��29��� W, 830���1030 m a.s.l.) collected on 6���8 December 2009 by A. Garcia Bravo and C. Koch; an adult male (CORBIDI 5706) and a subadult female (ZFMK 90892) from Zapatalgo, Province of Utcubamba, Department of Amazonas, Peru (6��04��� S, 78��30��� W, 907 m a.s.l.) collected on 10 December 2009 by A. Garcia Bravo and C. Koch; a juvenile (ZFMK 90891) from Zapatalgo, Province of Utcubamba, Department of Amazonas, Peru (6��06���05.2������ S, 78��29���54.5������ W, 521 m a.s.l.) collected on 9 December 2009 by A. Garcia Bravo and C. Koch; two adult males (ZFMK 90887, 90888) and an adult female (CORBIDI 5702) from San Vicente /Pusac, Province of Bol��var, Department of La Libertad, Peru (6��59��� S, 77��55��� W, 1425���1432 m a.s.l.) collected on 22 April 2009 by A. Garcia Bravo and C. Koch. Diagnosis and comparison. Phyllodactylus pachamama sp. nov. is a comparatively large species with a maximum SVL of 77 mm. Among species of Phyllodactylus found in mainland South America only P. delsolari, P. dixoni, P. reissii and P. v en t r al i s exceed 70 mm SVL. Thus, this species can be distinguished from all other species by its larger size. In lacking an enlarged postanal plate it further differs from P. thompsoni and by the absence of a preanal plate it can be distinguished from P. angustidigitus, P. gerrhopygus, and P. heterurus. By having 10���16 well-defined rows of enlarged, trihedral keeled tubercles, the new species differs from P. angustidigitus, P. gerrhopygus, P. heterurus (dorsal tubercles absent in all three species), P. delsolari, P. inaequalis (fewer than 10 poorly defined rows of small, smooth, round tubercles in both species), P. microphyllus (dorsal tubercular rows indistinct, composed of small flat, oval tubercles), and P. thompsoni (10). By having 43���56 paravertebral tubercles between rear of head and posterior edge of thigh it differs from P. interandinus (54���83), P. k of ord i (31���36), P. pumilus (37���43), P. sentosus (26���31), and P. thompsoni (36���40). Phyllodactylus pachamama can be distinguished from P. heterurus, P. kofordi, P. pumilus, and P. sentosus by lacking tubercles on the base of tail. By the presence of tubercles on the tibia this species can further be differentiated from P. angustidigitus, P. clinatus, P. delsolari, P. gerrhopygus, P. inaequalis, P. interandinus, P. lepidopygus, and P. microphyllus. By the absence of tubercles on the thigh it can be distinguished from P. d i x o ni, P. k o f o rd i, P. pu m i l u s, P. s e n t o s u s, and P. ventralis. By the absence of tubercles on the forearm it can be differentiated from P. dixoni, P. kofordi, P. sentosus, and P. ventralis. From P. angustidigitus, P. microphyllus, and P. sentosus the new species further differs by having large terminal lamellae. It can be distinguished by the number of lamellae under the fourth toe (14���17, mean 15.1) from P. dixoni (11���13, mean 11.3), P. inaequalis (10���12, mean 10.7), P. johnwrighti (7���14, mean 10.5), P. k o f o rd i (11���13, mean 11.5), P. leoni (9���13, mean 10.9), P. lepidopygus (10���14, mean 11.5), and P. pumilus (11���13, mean 12.0). 40 % of specimens have 3 postmentals compared to only 1.9 % in P. reissii. In contrast to P. johnwrighti and P. pumilus (dark stripe from nostril to arm insertion always present), the postnasal and postorbital brown stripes in P. pachamama is usually absent or, if present, poorly defined. Description of holotype (Fig. 5). Head distinct from neck, covered with small granules that are largest on snout and smallest on interparietal region; rostral 3.3 mm wide, 2 mm high, somewhat rectangular, with a median groove along its posterior half; two internasals, in broad medial contact, each bordered posteriorly by two granules and a postnasal; nostril surrounded by rostral, first supralabial, two postnasals and internasal; 11 loreals between postnasal and anterior edge of orbit; scales in loreal region about three times larger than scales in interorbital region; 24 scales across snout at level of third supralabial, 17 at level of second supralabial; 12 supralabials, six to a point below the center of eye; eyes large, pupil vertically elliptical, with corrugated edges; orbital diameter 4.6 mm; eyelid with three visible rows of granules on left side and five on right side, granules of median row pointed in posterior part; 25 midorbital granules; interorbital distance 1.8 mm; 29 granules between posterior edge of eye and anterior edge of ear, eye to ear distance 7.1 mm; ear opening elliptical, posterodorsally oriented, 2.4 mm high and 0.7 mm wide, its anterior and posterior edges denticulated; rear of head intermixed with slightly larger tubercles; mental bell-shaped, larger than infralabials, 4 mm long, and 4 mm wide at its widest point; two postmentals, in contact with first pair of infralabials laterally and five scales posteriorly; nine infralabials, six to a point below center of eye, first two pairs largest, adjacent pairs of infralabials descreasing in size posteriard; scales on chin largest and juxtaposed in anterior part, decreasing in size and becoming more imbricate towards gular region. Body depressed; dorsal surface of neck with small granules that are slightly smaller than those on the head; dorsum covered with small granules and 10 to 12 longitudinal rows of enlarged, trihedral keeled tubercles at midbody; six rows of tubercles from occiput to base of tail; paravertebral row with 44 tubercles between occiput and cloaca, 32 between axilla and groin; paravertebral rows separated from each other by about six granules; tubercles in paravertebral row separated from each other by 1���2 granules; 2���4 granules between other longitudinal tubercular rows; 110 scales around the midbody; ventral surface with smooth, imbricate scales, 74 from posterior gular region to cloaca, 26 transversely across midbody; dorsal surface of forearm with flattened, imbricate scales and without tubercles; thigh with flat, imbricate scales, largest in size anterodorsally and smallest posterodorsally, without tubercles; dorsal surface of tibia with a few scattered, hardly visible, cone-shaped or pulvinate tubercles, which are much smaller than the large tubercles on the back and only slightly larger than other scales on tibia; subdigital lamellae on left fingers (I���V) 8���10���12���13���11; subdigital lamellae on left toes (I���V) 8���11���15���15���13; terminal lamellae paired, greatly enlarged, truncated, the tip of the claw hardly visible between the two terminal pads, not projecting the terminal pads; tail complete, 1.18 times the SVL, covered with smooth and imbricate scales, on dorsal surface of tail about 3���4 times larger than dorsal body granules; medial subcaudals broadly enlarged, about two times as wide as long in anterior part of tail, decreasing in width posteriard. Measurements in mm. ��� Snout-vent length 71; tail length 84; axilla-groin length 29; right forelimb length 24.0; right hindlimb length 31.9; head length 21.6; maximum head width 15.9; midorbital head width 12.3; head height 8.9. Coloration of holotype in ethanol (Fig. 5). Dorsal ground color of head, body and limbs pale brown, with irregular small brown flecks and blotches on the dorsal surface of the body and limbs; indistinct brown line extending laterally on each side from nostril through eye to above insertion of the forearm; lateral surface of the body cream with brown mottling on each scale; ventral surface of head and body cream, with brown mottling on each scale of the chest region and towards the venterolateral parts of head and body, scales on median part of venter and gular region almost without mottling; dorsal surface of tail with alternating cream, brown and dark brown blotches; subcaudal ground color cream, with dense grayish brown mottling; lamellae under fingers and toes grayish. Variation (see also Table 3). SVL of male paratypes ranges from 48���75 mm (68.8 �� 4.3, n = 9), and SVL of female paratypes ranges from 44���77 mm (63.7 �� 11.6, n = 10). Tail length/total length is 0.43���0.56 (0.51 �� 0.03, n = 18). Male specimens possess 3���4 postanal tubercles on each side, which are slightly elevated. In females, postanal tubercles are lacking or only hardly visible and usually less pronounced as in males. The number of postmentals is 2���4 (2.5 �� 0.64). In one specimen (ZFMK 90887), postmentals contact first and second infralabials on each side, in all other specimens only first infralabial contacts postmentals, scales bordering postmentals 5���8 (6.4 �� 1). Two paratypes (ZFMK 90887, CORBIDI 5705) have 3 internasals, all others have 2 (2.1 �� 0.29). Lamellae on fourth finger 10���14 (12.5 �� 0.86); lamellae on fourth toe 14���17 (15.1 �� 0.81); supralabials 8���13 (10.2 �� 1.4), to a point below the center of eye 6���9 (7.7 �� 0.83); infralabials 7���11 (8.7 �� 1.03), to a point below the center of eye 5���7 (6.2 �� 0.7); loreals between postnasal and anterior edge of orbit 9���13 (11.6 �� 1.3); granules between posterior edge of eye and anterior edge of ear 24���28 (26.4 �� 1.1); scales across snout at level of third supralabial 22���26 (24.1 �� 1.4), at level of second supralabial 15���19 (17.1 �� 1.6); midorbital granules 23���25 (23.9 �� 0.86); longitudinal rows of dorsal tubercles 10���16 (13.2 �� 1.4); paravertebral tubercles between rear of head and cloaca 43���56 (49.6 �� 3.3), between axilla and groin 25���32 (29.1 �� 2.2); scales around midbody 92���116 (103 �� 7.4); ventral scales from gular region to cloaca 67���87 (73.5 �� 5.7), across midbody 26���36 (29.4 �� 3). In life (Fig. 6), dorsal ground color of head, body and limbs of paratypes varies between yellowish-white, grayish-white, cream and pale brown; irregular median or dark brown flecks and blotches on the dorsal surface of the body and limbs may either be almost lacking or quite prominent and forming transverse dorsal bands or a more or less reticulate pattern, that may be discontinuous in some specimens resulting in a pale cream or light brown middorsal line; postnasal and postorbital brown stripe lacking in most specimens; ventral surface of head and body cream or yellowish, with or without brown mottling; dorsal surface of unregenerated tails with alternating cream or light brown, median brown and dark or blackish brown blotches; ventral surface of tail varying from uniformly cream or yellowish to a grayish brown reticulation with strong brown mottling; pupil silver-gray or slightly bronzecolored. In ethanol, dorsal ground color of head, body and limbs of paratypes varies between pale brown and cream; ventral surface of head and body cream; ventral surface of tail varying from uniformly cream to grayish with strong grayish-brown mottling. Etymology. The specific epithet pachamama is derived from the Quechuan language and can be translated as Mother Earth. It represents the name of a goddess revered by indigenous people from South America, and in particular from Andean regions. According to many believers, problems will arise when people take too much from nature, because they are stealing it from Pachamama. This name was chosen to emphasize the need for protected areas along the Mara����n river and blames the numerous ongoing mining and dam construction activities for exploiting this unique habitat. Distribution and natural history. Phyllodactylus pachamama sp. nov. is endemic to the upper valley of the Mara����n River and some of its tributaries in the Northern Peruvian Andes from Zapatalgo, Province of Utcubamba, Department of Amazonas to San Vicente, Province of Bol��var, Department of La Libertad. It inhabits the seasonally dry forest, occurring in xeric habitats dominated by less dense shrubs and cacti (e.g. Armatocereus, Browningia, Espostoa) at elevations between 521���1432 m a.s.l. Individuals of this nocturnal gecko were active between 6.25 pm and 11.35 pm and mostly found on rock walls or house walls in heights up to about 3 m above the ground. Some individuals were also found running over sandy ground next to the Mara����n River. Only one male individual (CORBIDI 5703) was found during the day hidden under some logs. During their active hours, air temperatures ranged from 23.0 to 29.7�� C, substrate temperatures ranged from 20.7 to 30.0�� C and air humidity ranged from 41 to 71%. Aurich et al. (2011) examined the stomach contents of 8 individuals of P. pachamama from Balsas (referred to as P. reissii), which contained exclusively arthropod material, dominated by coleopterans, which made up 40% of all prey items. Three gravid females (CORBIDI 5700; ZFMK 90885, 91725) were collected in mid April (2009 and 2010), each contained two eggs. On 10 July 2005, a gravid female (ZFMK 85005) was collected which contained only one egg. Two juveniles with a SVL of 31 mm (ZFMK 90889) and 32 mm (ZFMK 90891) were found in December 2009. Most gravid females of the species examined by Aurich et al. (2015) also contained two eggs. In Balsas and San Vicente this species occurs sympatrically with the congeners P. delsolari and P. thompsoni, and with Phyllopezus maranjonensis, whereas in Zapatalgo it was found sympatrically with the sphaerodactylid Pseudogonatodes barbouri., Published as part of Koch, Claudia, Flecks, Morris, Venegas, Pablo J., Bialke, Patrick, Valverde, Sebastian & R��dder, Dennis, 2016, Applying n-dimensional hypervolumes for species delimitation: unexpected molecular, morphological, and ecological diversity in the Leaf-Toed Gecko Phyllodactylus reissii Peters, 1862 (Squamata: Phyllodactylidae) from northern Peru, pp. 41-80 in Zootaxa 4161 (1) on pages 49-53, DOI: 10.11646/zootaxa.4161.1.2, http://zenodo.org/record/256497, {"references":["Aurich, J., Koch, C. & Bohme, W. (2011) Ecology of a unique gecko assemblage (Phyllodactylidae: Squamata) from northern Peru. Northwestern Journal of Zoology, 7 (2), 310 - 317.","Aurich, J., Koch, C. & Bohme, W. (2015) Reproduction of a gecko assemblage (Squamata: Phyllodactylidae) in the Maranon Region (Peru) and comments on the largest gekkonid in the New World. Phyllomedusa, 14 (1), 53 - 62. http: // dx. doi. org / 10.11606 / issn. 2316 - 9079. v 14 i 1 p 53 - 62"]}

Phyllodactylus magister Phyllodactylus magister Noble, Occasional Papers of the Boston Society of Natural History 5:110. ―1924 Phyllodactylus magister (partim) ��� Burt & Burt, Bulletin American Museum of Natural History, 61: 250. ��� 1931 Phyllodactylus reissii (partim) ��� Dixon & Huey, Los Angeles County Museum Contributions in Science 192: 50. ―1970 Phyllodactylus reissii (partim) ��� Peters & Donoso-Barros, United States Nat. Mus. Bull. 297: 242. ��� 1970 Phyllodactylus reissii (partim) ��� Peters & Donoso-Barros, Smithsonian Institution Press, Washington D.C. & London: 242. ��� 1986 Phyllodactylus reissii (partim) ― Torres- Carvajal, Barnes, Pozo- Andrade, Tapia & Nicholls, Journal of Biogeography 41: 1887. ��� 2014 Phyllodactylus reissii (partim) ��� Aurich, Koch & B��hme, Phyllomedusa 14(1): 55. ��� 2015 Holotype. MCZ 17974, adult male, collected in deserted huts in the Chinchipe valley near Perico, Province of Ja��n, Department of Cajamarca by G.K. Noble during the Harvard Peruvian Expedition in September 1916. (Pictures of the preserved holotype are available at http://mczbase.mcz.harvard.edu/MediaSet.cfm?media_id=10875) Paratypes (153). All paratypes collected by G.K. Noble during the Harvard Peruvian Expedition in 1916 form the Province of Ja��n, Department of Cajamarca: MCZ 18126���18129, 18141, 18142, 126238���126375, 169013 from Bellavista; MCZ 18143���18150 from Perico. New collected material (47). See Appendix 2. Diagnosis and comparison. According to our data Phyllodactylus magister reaches a maximum SVL of 69 mm. Compared to other species of mainland South America it is thus slightly smaller than P. pachamama, P. delsolari, P. dixoni, P. reissii and P. ve n t r a l i s (all exceeding 70 mm SVL) and larger than P. angustidigitus (SVL ��� 57 mm), P. clinatus (SVL ��� 46 mm), P. gerrhopygus (SVL ��� 56 mm), P. heterurus (SVL of the single known specimen is 38 mm), P. inaequalis (SVL ��� 42 mm), P. interandinus (SVL ��� 49 mm), P. johnwrighti (SVL ��� 44 mm), P. kofordi (SVL ��� 46 mm), P. leoni (SVL ��� 55 mm), P. lepidopygus (SVL ��� 55 mm), P. microphyllus (SVL ��� 58 mm), P. pumilus (SVL ��� 51 mm), P. sentosus (SVL ��� 56 mm), and P. thompsoni (SVL ��� 42 mm). The species can be distinguished from P. thompsoni by the lack of an enlarged postanal plate. It further differs from P. angustidigitus, P. gerrhopygus, and P. heterurus by the absence of a preanal plate. By having 12���16 well-defined rows of enlarged, trihedral keeled tubercles, P. m a gi s t e r differs from P. angustidigitus, P. gerrhopygus, P. heterurus (dorsal tubercles absent in all three species), P. delsolari, P. inaequalis (fewer than 10 poorly defined rows of small, smooth, round tubercles in both species), P. microphyllus (dorsal tubercular rows indistinct, composed of small flat, oval tubercles), and P. thompsoni (10). Having 40���60 paravertebral tubercles between rear of head and posterior edge of thigh differentiates the species from P. ko f o rd i (31���36), P. interandinus (54���83) and P. sentosus (26���31). Phyllodactylus magister can further be differentiated from P. angustidigitus, P. clinatus, P. gerrhopygus, P. inaequalis, P. lepidopygus, and P. microphyllus by the presence of tubercles on the tibia. By the absence of tubercles on the forearm it can be differentiated from P. dixoni, P. kofordi, P. sentosus, and P. ventralis and by the absence of tubercles on the tail it further differs from P. heterurus, P. kofordi, P. pumilus and P. sentosus. In contrast to P. angustidigitus, P. microphyllus, and P. sentosus the species possesses large terminal lamellae. By having 12��� 18 (mean 14.7) lamellae under the fourth toe it can be distinguished from P. inaequalis (10���12, mean 10.7), and P. leoni (9���13, mean 10.9). It further differs from P. reissii and P. pachamama by the presence of small tubercles on the thigh in about half of the specimens (always absent in P. reissii and P. pachamama), and besides it differs from P. pachamama by having always 2 postmentals (2���4, mean 2.5 in P. pachamama), and by an on average smaller number of scales around midbody (78���114, mean 90.3 in P. m ag i s t e r vs. 92���116, mean 103.0 in P. pachamama). Compared to P. reissii the anterior edge of the ear opening is smooth or only slightly denticulated in most specimens (strongly denticulated with pointed scales in P. reissii). Redescription. The following redescription is based on the adult female ZFMK 90883 from the type locality Perico. Head distinct from neck, covered with small granules that are largest on snout and smallest on interparietal region; rostral 2.8 mm wide, 1.5 mm high, rectangular, with a median groove along its posterior one third; two internasals, in broad medial contact, each bordered posteriorly by two granules and a postnasal; nostril surrounded by rostral, first supralabial, two postnasals and internasal; 11 loreals between postnasal and anterior edge of orbit; granules in loreal region about three times larger than scales in interorbital region; 21 scales across snout at level of third supralabial, 17 at level of second supralabial; 11 supralabials, six to a point below the center of eye; eyes large, pupil vertically elliptical, with corrugated edges; orbital diameter 3.3 mm; eyelid with three visible rows of granules on both sides, granules of median row strongly pointed in posterior part; 23 midorbital granules; interorbital distance 2.0 mm; 22 granules between posterior edge of eye and anterior edge of ear, eye to ear distance 5.0 mm; ear opening elliptical, posterodorsally oriented, 1.9 mm high and 0.7 mm wide, its anterior edge almost smooth and the posterior edge only very slightly denticulated; rear of head intermixed with small, round, pulvinate tubercles; mental bell-shaped, larger than infralabials, about as long as wide; two postmentals, in contact with first pair of infralabials laterally and seven scales posteriorly; nine infralabials, six to a point below center of eye, first two pairs largest, adjacent pairs of infralabials continuously descreasing in size posteriard; scales on chin largest and juxtaposed in anterior part, decreasing in size and becoming more imbricate towards gular region. Body depressed; dorsal surface of neck with small granules that are slightly smaller than those on the head; dorsum covered with small granules and 14 longitudinal rows of enlarged, trihedral and slightly keeled tubercles at midbody; eight rows of tubercles reach to occiput and six rows reach to base of tail; paravertebral row with 45 tubercles between occiput and cloaca, 29 between axilla and groin; paravertebral rows separated from each other by 4���7 granules; tubercles in paravertebral row usually separated from each other by 1���2 granules; 1���4, mostly 2 granules between other longitudinal tubercular rows; 90 scales around the midbody; ventral surface with smooth, imbricate scales, constantly increasing in size towards vent, 65 from posterior gular region to cloaca, 28 tr��nsversely across midbody; dorsal surface of forearm with flattened, imbricate scales and without tubercles; thigh with flat, imbricate scales, largest in size anterodorsally and smallest posterodorsally, with at least small tubercles; dorsal surface of tibia scattered with small cone-shaped or pulvinate tubercles, which are much smaller than the large tubercles on the back and about 3���4 times larger than adjacent granules on tibia; subdigital lamellae on left fingers (I���V) 8���10���11���12���9; subdigital lamellae on left toes (I���V) 9���11���13���14���13; terminal and penultimate lamellae paired, terminal lamellae greatly enlarged, truncated, the tip of the claw hardly visible, slightly projecting terminal pads; tail complete and unregenerate, 1.2 times the SVL, covered with smooth and imbricate scales, heterogeneous in size; most scales in median row of subcaudals slightly to moderately enlarged. Measurements in mm. ��� Snout-vent length 53; tail length 63.7; axilla-groin length 26; right forelimb length 21.3; right hindlimb length 24; head length 15.5; maximum head width 10.8; midorbital head width 8.8; head height 6.3. Coloration of ZFMK 90883 in ethanol. Dorsal ground color of head, body, limbs and tail pale grayish brown or cream, most scales mottled with dark brown, forming some irregular small brown flecks and blotches on the dorsal surface of the body and limbs; no facial stripe visible; ventral surface of head and body cream, with a trace of a brown mottling on about half of the scales; subcaudal ground color cream, with grayish brown mottling that becomes denser towards tip; lamellae under fingers and toes pale grayish. Variation (see also Table 3; based on the paratypes MCZ 18142 and MCZ 18145 and 47 newly collected specimens). ��� SVL of adult males ranges from 51���69 mm (58.6 �� 4.2, n = 16), and SVL of adult females ranges from 50���66 mm (54.4 �� 4.2, n = 9). Tail length/total length is 0.45���0.56 (0.52 �� 0.02, n = 36). Male specimens possess 3���5 (3.6 �� 0.69) postanal tubercles on each side. The number of postmentals is 2 in all specimens. In the majority of specimens postmentals contacting first infralabial on each side, scales bordering postmentals 5���7 (6.0 �� 0.62). Internasals 0���3 (2.1 �� 0.56). Lamallae on fourth finger 10���14 (12.4 �� 1.00); lamellae on fourth toe 12���18 (14.7 �� 1.11); supralabials 7���13 (10.0 �� 1.44), to a point below the center of eye 6���8 (7.2 �� 0.73); infralabials 6���12 (8.2 �� 1.2), to a point below the center of eye 6���8 (6.2 �� 0.53); loreals between postnasal and anterior edge of orbit 9���15 (11.8 �� 1.45); granules between posterior edge of eye and anterior edge of ear 21���30 (25.1 �� 2.55); anterior edge of the ear opening is smooth or only slightly denticulated in most specimens; scales across snout at level of third supralabial 20���27 (22.6 �� 1.62), at level of second supralabial 14���18 (15.8 �� 1.14); midorbital granules 19���26 (22.7 �� 1.76); longitudinal rows of dorsal tubercles 12���16 (14.2 �� 1.27); paravertebral tubercles between rear of head and cloaca 40���60 (49.2 �� 3.74), between axilla and groin 24���35 (28.4 �� 2.94); tubercles on tibia are present in all specimens, but may be very small and very few in some specimens; very small and few tubercles on thigh present in about half of the specimens; scales around midbody 78���114 (90.3 �� 7.53); ventral scales from gular region to cloaca 63���80 (70.0 �� 4.21), across midbody 23���32 (26.8 �� 1.77). In life, dorsal ground color of head, body and limbs varies between grayish-white, yellowish-cream, cream and pale brown; irregular whitish, brown or dark brown flecks and blotches on dorsal surface of the body and limbs may either be hardly visible or quite prominent and forming transverse dorsal bands or a more or less reticulated pattern (in the majority of specimens), that may be discontinuous in some specimens resulting in a pale cream or light brown middorsal line; dark brown line extending laterally on each side from nostril through eye to above insertion of the forearm conspicuous in most specimens, faint or absent in others; ventral surface of head and body cream to bright yellow, with or without brown mottling; dorsal surface of unregenerated tails with alternating cream or light brown, median brown and dark or blackish brown blotches; ventral surface of tail varying from uniformly cream or bright yellow to a grayish brown with strong brown mottling; pupil golden brown or coppery; tails of most juveniles annulated with yellow and brown. In ethanol, dorsal ground color of head, body and limbs varies between pale gray, brown and cream; ventral surface of head and body cream or grayish; ventral surface of tail varying from uniformly cream to grayish brown with a grayish-brown mottling. Distribution and natural history. Phyllodactylus magister is endemic to the seasonally dry forest vegetation of the upper Mara����n River basin and some of its tributaries in the Northern Peruvian Andes in the Departments Cajamarca and Amazonas from Perico to Puerto Malleta, at elevations between 419���1247 m a.s.l.. The habitat of Phyllodactylus magister is composed of drought-resistant trees (e.g. Acacia, Anadenanthera, Ceiba, Cordia, Prosopis), dense shrubs (e.g. Croton, Mimosa) and ground vegetation layer (e.g. Opuntia, Poaceae). During daytime surveys we found several specimens of this nocturnal species hidden under stones or inside deserted huts. Active individuals were recorded between 6.45 pm and 0 1.00 am, primarily on rock walls, sandy walls besides paths, roads or dry creek beds, as well as house walls from ground level up to about 2.5 m above the ground. Air temperatures during the active hours of this species ranged from 19.2 to 29.9�� C, substrate temperatures ranged from 17.4 to 31.9�� C and air humidity ranged from 52 to 75%. Gravid females of P. m ag i s t e r were collected end of May (ZFMK 88739), and in mid December (CORBIDI 5708, ZFMK 90898), containing two eggs, except for ZFMK 90898 which had only one egg. On the 8th of May 2008 at 11.15 pm a gravid female was discovered (not collected) directly after it had deposited an egg in the soft soil of the sandy walls of a dry creekbed. One juvenile with a SVL of 31 mm (ZFMK 90881) was found in March 2009 and another juvenile with a SVL of 33 mm (CORBIDI 1812) was found in May 2008. Subadults with SVL between 37���45 mm were collected in March 2009 (ZFMK 90880, 90882), April 2009 (CORBIDI 5696, 5697), May 2008 (CORBIDI 1829, 1833, 1836), July 2008 (CORBIDI 1819, ZFMK 88741), and in December 2009 (ZFMK 90895, 90896, 90901, 90902). We found Phyllodactylus magister in sympatry with the congener P. interandinus in Ja��n, Bellavista and Puerto Malleta, and according to Dixon & Huey (1970) the two species also occur sympatrically in Bagua Chica, Bagua Grande, and Perico. In Ja��n and Santa Rosa de la Yunga, we observed P. magister in the same habitat as the sphaerodactylid species Gonatodes atricucullaris, and Noble (1921) also adds Bellavista as a contact zone of both species. According to him, a second sphaerodactylid species, Pseudogonatodes barbouri, also occurs in Bellavista, but we did not find this species in sympatry with P. m ag i s t e r in Perico., Published as part of Koch, Claudia, Flecks, Morris, Venegas, Pablo J., Bialke, Patrick, Valverde, Sebastian & R��dder, Dennis, 2016, Applying n-dimensional hypervolumes for species delimitation: unexpected molecular, morphological, and ecological diversity in the Leaf-Toed Gecko Phyllodactylus reissii Peters, 1862 (Squamata: Phyllodactylidae) from northern Peru, pp. 41-80 in Zootaxa 4161 (1) on pages 53-55, DOI: 10.11646/zootaxa.4161.1.2, http://zenodo.org/record/256497, {"references":["Dixon, J. R. & Huey, R. B. (1970) Systematics of the lizards of the gekkonid genus Phyllodactylus of mainland South America. Los Angeles County Museum Contributions in Science, 192, 1 - 78.","Noble, G. K. (1921) Some new lizards from northwestern Peru. Annals of the New York Academy of Sciences, 29, 133 - 139. http: // dx. doi. org / 10.1111 / j. 1749 - 6632.1920. tb 55353. x"]}

Stenocercus omari sp. nov. Figs. 1���5, Table 1. Holotype. CORBIDI 0 0 577, an adult male from Abra Chanchillo (06˚47'07.00'' S, 77˚56'32.50'' W), 2786 m, Chachapoyas province, Amazonas department, Peru, collected by P. J. Venegas on 17 June 2007. Paratypes. Peru: Amazonas department: Chachapoyas province: CORBIDI 00569���70, 00578���80, adult females, and CORBIDI 00571���76, adult males, collected with the holotype; Luya province: CORBIDI 11961, adult male, and CORBIDI 11962, adult female, from Pircopampa (06��19'32.17'' S, 78��13'1.99'' W), 2378 m, collected by K. Garc��a-Burneo on 23 September 2012; La Libertad department: Bolivar province: CORBIDI 05793���94, sub adult female and adult male, respectively, and ZFMK 90835 adult female from Bolivar (07��10'00.00'' S, 77��42'46.50'' W), 2772���3035 m, collected by A. Garc��a-Bravo and C. Koch on 21 April 2009. Diagnosis. Stenocercus omari is distinguished from other species in the genus, except for S. amydrorhytus, S. chrysopygus, S. cupreus, S. johaberfellneri, S. latebrosus, S. melanopygus, S. modestus, S. ornatissimus, S. orientalis, and S. stigmosus by having granular scales on the posterior surfaces of thighs, a conspicuous antehumeral fold and by lacking a vertebral crest. Stenocercus omari is easily distinguished from the aforementioned species, except for S. orientalis, by the presence of prominently keeled dorsal head scales. The new species differs from S. orientalis in lacking a prominent oblique neck fold with a mite pocket under it, a distinctive character in S. orientalis, and also present in S. latebrosus and S. ornatissimus. Another important difference between the new species and S. orientalis is the presence of a longitudinal neck fold in the former, absent in S. orientalis; such fold is also present in S. melanopygus and S. stigmosus; however, unlike the new species described herein, both species have smooth or slightly keeled dorsal head scales. Additionally, the new species has a distinct deep postfemoral mite pocket with a posteroventrally oriented slit-like opening, whereas S. orientalis has one or more vertical folds instead (Fig. 4). The new species can be distinguished from S. orientalis in the field by color pattern. Adult males of S. omari have distinct dark triangular marks longitudinally arranged on each side of the vertebral line (Fig. 1 A, 3A, 5A) and S. orientalis usually has a tabby dorsal pattern (Fig. 5 D) that in some individuals is indistinct. Although males of both species share a yellow ventral coloration, males of the new species have a blue throat with dark flecks, whereas males of S. orientalis have a yellow throat with or without dark flecks and a bluish center in some specimens (Fig. 5 E). Adult females of S. omari and S. orientalis are similar in coloration pattern; however, females of S. omari, have dorsolateral and ventrolateral light stripes on each side of the neck and females of S. orientalis have only one dorsolateral light stripe on each side of the neck (Fig. 5 F). Characterization. (1) Maximum SVL in males 68 mm (N =8); (2) maximum SVL in females 69 mm (N =7); (3) vertebrals 36���47; (4) paravertebrals 38���52; (5) scales around midbody 42���54; (6) supraoculars 4���7; (7) internasals 4; (8) postrostrals 2���4; (9) loreals 2���5; (10) gulars 13���25; (11) lamellae on Finger IV 18���21; (12) lamellae on Toe IV 18���29; (13) posthumeral pocket present as one vertical fold; (14) postfemoral mite pocket present as a distinct deep pocket with a posteroventrally oriented slit-like opening [Type 2 of Torres-Carvajal (2007a)]; (15) parietal eye not visible; (16) occipital scales small, keeled, subimbricate; (17) projecting angulate temporal absent; (18) supraoculars subequal in size; (19) scales in frontonasal region subimbricate, keeled; (20) short preauricular fringe present; (21) antehumeral and longitudinal neck folds present; (22) lateral nuchals half the size of dorsal nuchals; (23) posterior gulars in adults smooth, imbricate; (24) lateral scales slightly smaller than dorsal body scales; (25) vertebral crest absent; (26) dorsolateral crests absent; (27) ventrals in adults broad, lanceolate, smooth, imbricate; (28) scales on posterior surfaces of thighs granular; (29) prefemoral fold absent; (30) inguinal groove absent; (31) preanals not projecting; (32) tail not compressed laterally in adult males; (33) tail length 53���70 % of total length; (34) caudal whorls per autotomic segment three; (35) caudals not spinose; (36) dark stripe extending anterodorsally from subocular region to supraciliaries absent; (37) gular region of adult females pigmented with or without dark spots; (38) gular region of adult males pigmented with dark spots; (39) black blotch on ventral surface of neck in adult males absent; (40) thin black or dark brown midventral line absent; (41) black patch on ventral surface of thighs absent; (42) dorsal ground color brown with darker triangular marks longitudinally arranged on each side of vertebral line in females and males; (43) two xiphisternal and two postxiphisternal pairs of inscriptional ribs, the two postxiphisternal pairs of ribs short and not in contact midventrally (Pattern 1C of Torres-Carvajal, 2004a). Description of the holotype. Male (Figs. 1���3); SVL 66 mm; TL 157 mm; maximum head width 10.3 mm; head length 15.3 mm; head height 9.2 mm; occipitals, parietals, interparietals, and postparietals small, keeled, imbricate; parietal eye not visible; supraoculars in six rows, keeled, imbricate, subequal in size; canthals two; anterior most canthal in contact with the nasal; scales on frontonasal region slightly imbricate and keeled; internasals four; postrostrals four, two most lateral wider than long on each side, medial postrostrals as long as wide; supralabials five; infralabials six; loreals three; lorilabials in one row; preocular one, in contact with posterior canthal; lateral temporals keeled, imbricate; gulars in 22 rows between tympanic openings; all gulars smooth, imbricate, apical pit absent; second infralabial not in contact with second and third sublabials; mental in contact with first pair of infralabials; lateral and dorsal scales of body and neck keeled, imbricate, mucronate; lateral scales slightly smaller than dorsal body scales; scales around midbody 44; vertebrals same size as dorsals, 39 scales on vertebral row, prominent serrate vertebral crest absent; paravertebrals 45; ventrals broad, lanceolate, smooth, imbricate; preauricular fringe short, composed of three enlarged scales, all same size; antegular, gular, postauricular, oblique and supraauricular neck folds absent; longitudinal and antehumeral neck folds present; limb scales keeled, imbricate; ventral scales of hind limbs smooth and upper arms keeled; lamellae on Finger IV 18; lamellae on Toe IV 18; tail not compressed laterally; caudals keeled, imbricate, mucronate; basal subcaudals smooth, imbricate; tail length 2.4 times SVL; posthumeral pocket present as one vertical fold; postfemoral mite pocket present as a distinct deep pocket with a posteroventrally oriented slit-like opening [Type 2 of Torres- Carvajal (2007a)]; postfemoral region composed of imbricate, smooth scales that become smaller towards insertion of hind limb. Color in life. Head and neck dark brown (almost black) with white flecks. A black vertical blotch on the antehumeral fold and a black rounded blotch on the side of the neck. Dorsum brown with dark brown triangular marks longitudinally arranged on each side of the vertebral line, and flanks yellowish cream with dark brown bands. Forelimbs dark brown with yellow flecks and hind limbs brownish yellow with dark brown transverse bands. Tail brown with dark brown marks along it. Ventrally, throat and chest sky blue with dark blue marks on the throat; ventral surface of thighs, cloacal region, and base of tail dark yellow. Color in ethanol 70%. Head brown, white flecks turned cream. Dorsal background color of body and base of tail brownish cream with dark brown triangular marks (but lighter than in life). Sides of neck brownish cream with brown, instead of black, blotches. Flanks brownish cream flecked by brown and light cream scales, and ventrolateral region sky blue. Limbs brown with dark marks and distal portion of tail brown with light transverse bands. Ventrally the yellow surfaces turned cream. Stenocercus omari sp. nov. Variation. Some measurements and scutellation characters of Stenocercus omari are presented in Table 1. Second infralabial not in contact with third sublabial in 93.8% of specimens, and first pair of postmentals in contact medially in all specimens. All male paratypes are similar to the holotype, varying only in the tone of ventral coloration; although the adult male specimen from Bolivar (CORBIDI 05794) lacks the black blotch on the sides of neck. Adult females have a cinnamon or dark brown dorsum with distinct or faint darker triangular marks longitudinally arranged on each side of the vertebral line; sides of the head and flanks brown (darker than the dorsum); supralabials white or cream, continuous with a white or cream dorsolateral stripe along the neck, becoming pale brown along the dorsum; a white or cream stripe along the edge of the jaw continues as a white ventrolateral stripe along the neck, becoming dirty cream on the flanks; flanks with or without vertical bands; dorsal surface of tail with or without transverse bands. Ventral surface dirty cream or white with or without dark flecks on the throat. Sexual dimorphism is evident in adult individuals. Females can be easily recognized by lacking white flecks on the dorsal surface of head and neck, and by having dark brown flanks (yellowish in males), and a dirty cream or white ventral coloration (throat sky blue, chest and belly yellow in males). Distribution and natural history observations. Stenocercus omari is known from three localities along the western slope of the northern portion of the Cordillera Central at the Mara����n river basin in the departments of Amazonas and La Libertad, at elevations of 2300���3035 m. According to Brack (1986) the distribution of S. omari lies inside the ecoregion of the Serrania esteparia and according to Olson et al. (2001) this species is distributed in the Peruvian Yungas ecoregion. The habitat at the type locality is a steep area located on the right side of the Mara����n river with the presence of corn plantations, cattle pasture and some scattered patches of montane forest along the slope. Stenocercus omari was very common along the sides of the Leimebamba-Balsas road, a steep band covered by herbaceous vegetation with big boulders and scattered bushes. All collected individuals were found active in daylight (between 11:00 and 15:00) basking on rocks, fallen tree trunks and at the base of bushes. A gravid female (CORBIDI 00570) with SVL 65 mm collected on June contained eight yolked follicles, four in each ovary. Another gravid female (CORBIDI 11962) with SVL 69 mm collected on September contained seven well developed eggs four in the left oviduct and three in the right one, eggs ranged from 13.96���15.15 mm in length, 7.56���8.87 mm in width, and 422.33���613.98 mm 3 in volume. Etymology. The specific epithet omari is a noun in the genitive case and is a patronym for our appreciated friend, the Ecuadorian herpetologist Omar Torres-Carvajal, who has published important contributions to the taxonomy and systematics of Neotropical lizards, especially of the genus Stenocercus., Published as part of Venegas, Pablo J., Echevarr��a, Lourdes Y., Garc��a-Burneo, Karla & Koch, Claudia, 2016, A new species of iguanid lizard, genus Stenocercus (Squamata, Iguania), from the Central Andes in Peru in Zootaxa 4205 (1), DOI: 10.11646/zootaxa.4205.1.4, http://zenodo.org/record/192540, {"references":["Torres-Carvajal, O. (2007 a) A taxonomic revision of South American Stenocercus (Squamata: Iguania) lizards. Herpetological Monographs, 21, 76 - 178. http: // dx. doi. org / 10.1655 / 06 - 001.1","Torres-Carvajal, O. (2004 a) The abdominal skeleton of tropidurid lizards (Squamata: Tropiduridae). Herpetologica, 60, 75 - 83. http: // dx. doi. org / 10.1655 / 03 - 15","Brack, A. (1986) Las Ecoregiones del Peru. Boletin de Lima, 44, 57 - 70.","Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N. D., Powel, l G. V. N., Underwood, E. C., D'amico, J. A., Itoua, I., Strand, H. E., Morrison, J. C., Loucks, C. J., Allnutt, T. F., Ricketts, T. H., Kura, Y., Lamoreux, J. F., Wettengel, W. W., Hedao, P. & Kassem, K. R. (2001) Terrestrial Ecoregions of the World: A New Map of Life on Earth. BioScience, 51, (11), 933 - 938. http: // dx. doi. org / 10.1641 / 0006 - 3568 (2001) 051 [0933: TEOTWA] 2.0. C"]}

Epictia antoniogarciai sp. nov. (Figures 2 G–I, 6) Holotype: CORBIDI 7678, from the vicinities of Santa Rosa de la Yunga Village, Jaén Province, Cajamarca Region, Peru (S 05° 25 ’ 53.3 ’’, W 078° 33 ’47.0’’, 1268 m.a.s.l.), collected by M. Enciso, S. Duran and C. Koch on 4 May 2009. Paratypes (N = 4): three specimens from the vicinities Zapatalgo Village, Utcubamba Province, Amazonas Region, Peru: ZFMK 96676 (S 06°04’ 47.7 ’’, W 078° 29 ’ 18.7 ’’, 934 m. a.s.l.), CORBIDI 5670, ZFMK 90934 (S 06°04’44.0’’, W 078° 29 ’ 16.7 ’’, 968 m. a.s.l.), collected by A. Garcia Bravo and C. Koch between 07–09 December 2009; and one specimen from the type locality: CORBIDI 2069, collected in August 2008 by N. Monsalve. Diagnosis. (1) 14 midbody scale rows; (2) 10 midtail scale rows; (3) 2 supralabials, first large and in broad contact with supraocular; (4) 14–18 subcaudals; (5) 195–208 mid-dorsal scale rows; (6) dorsal scales black with yellow margins; (7) rostral bright yellow or yellowish-white dorsally, and grayish-brown or blackish ventrally; (8) terminal spine and surrounding scales yellow; (9) ventral surface of head, body and tail almost completely grayishbrown or light-gray scattered with dark grayish-brown scales. Comparisons [conditions for other Epictia in brackets]: By having 195–208 mid-dorsal scales this species has a higher number than E. collaris [155–166] and E. vanwallachi [188], and a lower number than E. albipuncta [213–285], E. alfredschmidti [267–279], E. borapeliotes [256–282], E. columbi [240–265], E. melanura [395– 396], E. rufidorsa [255–270], E. septemlineata [257], E. striatula [216–265], E. subcrotilla [318–333], E. teaguei [232–259], E. tesselata [258–283] and E. tricolor [285–310]. By having 14–18 subcaudal scales it further differs from E. columbi [22–25], E. melanura [18–20], E. munoai [10–14], E. nasalis [21] and E. tricolor [18–23]. By having a different color pattern with dorsal scales black with thin yellowish-white or bright yellow margins, and rostral and terminal portion of tail yellowish-white or bright yellow, it further differs from E. septemlineata [dorsum with seven black longitudinal stripes], E. rubrolineata [dorsal coloration red with 5 longitudinal stripes], E. rufidorsa [striped pattern with a red dorsal longitudinal stripe], E. teaguei [tricolor dorsal pattern of red, black and yellow stripes], E. tenella [dorsum dark brown with lateral edges of the scales lighter, forming a pattern of serrated longitudinal light lines] and E. vanwallachi [dorsal scales brown with thin white or yellowish margins, rostral grayish-brown]. Description of holotype: An adult specimen with SVL of 129 mm; TAL of 14.5 mm; MB of 3.8 mm; MT of 3.0 mm; TL/TAL of 9.9; TL/MB of 37.8; HW of 2.8 mm; HH of 2.2 mm; HL of 3.4 mm; DSN of 0.9 mm; DNE of 0.8 mm; ED of 0.6 mm. Head subcylindrical, slightly depressed dorsoventrally, indistinguishable from neck; body cylindrical; not tapered cranially or caudally. Snout rounded in lateral and ventral view. Rostral visible in dorsal view, about 1.5 times longer than wide, almost squared ventrally with dorsal termination (apex) sharply rounded, almost reaching the imaginary transverse line between the anterior borders of the eyes, contacting upper and lower nasal laterally and frontal dorsally. Nasal completely divided horizontally by a suture slightly oblique, reaching rostral and first supralabial; ellipsoid nostril located almost in the center of the suture between upper and lower nasal, having the major axis oriented along the suture; supranasal about 1.9 times higher than wide and about 1.3 times higher than infranasal, contacting infranasal ventrally, first supralabial and supraocular posteriorly, and frontal dorsally; infranasal 1.7 times higher than wide, about 1.5 times wider than anterior supralabial, contacting first supralabial posteriorly; two supralabial scales, first positioned anteriorly and second posteriorly to ocular scale (1 + 1); upper lip border formed by rostral, infranasal, anterior supralabial, ocular and posterior supralabial; first supralabial about 2.8 times higher than wide, exceeding nostril, slightly exceeding central level of eye, in contact with supraocular scale dorsally and ocular posteriorly; ocular scale pentagonal with dorsal apex acuminate, 1.5 times higher than wide, contacting supraocular anterodorsally, parietal posterodorsally and second supralabial posteriorly; eye located at level of maximum width of ocular and with lower eye margin at nostril level, positioned anteriorly and almost contacting scale sutures; eyes partly visible in dorsal view; second supralabial subtrapezoidal, about as high as wide, slightly exceeding central level of eye, as high as anterior supralabial, 2.5 times wider than anterior supralabial at widest point; posterior margin of second supralabial in broad contact with temporal and in contact with first scale of lateral body row; dorsal margin of second supralabial in contact with parietal; temporal scale of same size as dorsal scales of lateral rows; supraocular scale almost spindle-shaped, oriented oblique, about twice as long as wide, contacting parietal posteriorly, and frontal and postfrontal dorsally; supraocular, parietal and occipital scales visible in lateral view; mid-dorsal head plates (frontal, postfrontal, interparietal and interoccipital) slightly imbricate, hardly decreasing in size posteriorly, pentagonal or round in dorsal view, higher and narrower than posterior mid-dorsal scales; frontal contacting postfrontal posteriorly; postfrontal contacting supraoculars anteriorly, parietals and interparietal posteriorly; interparietal contacting parietals and occipitals laterally, and interoccipital posteriorly; interoccipital contacting occipitals laterally, and nuchal and three dorsal body scales posteriorly; parietal about 1.8 times higher than wide, as high as occipital, about 1.5 times wider than occipital, both scales almost rectangular in shape; occipital about 2.5 times higher than wide; lower margin of parietal contacting upper border of temporal, posterior margin in broad contact with occipital; lower margin of occipital contacting temporal and first lateral body scale, posterior margin in broad contact with first paravertebral and first dorsolateral scale (latter both scales fused on left side); mental small with a median depression, followed by six infralabials per side, slightly subequal in size; first pair of infralabials rectangular, slightly larger than other infralabials; chin, gular, and dorsal and lateral head scales porous. Dorsal scales imbricate, smooth, homogeneous, rhomboid or elliptical in shape, about 1.4 times wider than long; 202 MDS; 14 – 14 – 14 D; 184 V; 10 TS; Anal plate large, 1.7 times wider than long, subtriangular in shape with distal apex rounded, bordered anteriorly and posteriorly by five rows of scales; 16 SC, becoming slightly narrower distally, except for ultimate four scales which are each fused with adjacent scales; each of last three scales on the dorsal surface of the tail fused with adjacent dorsolateral scales; terminal spine conical and strongly pointed, with stout base about as wide as long. Color of holotype in life: Anterior portion of head with a large bright yellow blotch, covering completely the dorsal surface of frontal and rostral scales, and partially the supranasal scales; rostral dark grayish-brown ventrally; remaining head and body dorsal scales black, with striking bright yellow margins; terminal spine and surrounding scales (two rows dorsally and three rows ventrally) bright yellow; ventral surface of head, body and tail light-gray, scattered with numerous darker grayish-brown scales. Color of holotype in preservative: Coloration of blotch on anterior dorsal scales of the head and terminal portion of tail changed to cream; rostral changed to dark gray ventrally; dorsal scales from head and body changed from dark brown to blackish with cream margins; ventral surface of head, body and tail changed to cream except for some scattered grayish-brown scales. Variation. Morphometric and pholidosis characters of the four paratypes are given in Table 1. The paratypes further vary from the holotype in the following characters: the yellow color pattern is distinctly less striking in three of the paratypes (ZFMK 96676, CORBIDI 5670, ZFMK 90934) than in the holotype: the blotch on the anterodorsal surface of the head is cream or yellowish-white; the dorsals are dark brown to black, most scales with thin, indistinct, yellowish margins; terminal spine pale yellow; ventral surface of head, body and tail almost uniformly grayish-brown, somewhat lighter than dorsal scales, or sometimes scattered with darker scales. Etymology. This species is dedicated to Antonio Garcia Bravo, Peruvian biologist, in recognition of both, his support in the investigation of the Peruvian herpetofauna and his continued and unattenuated efforts in the conservation of the dry forests along the Marañón River. Distribution and natural history. This species is known from the interandean part of the Equatorial Dry Forest from Santa Rosa de la Yunga in the North to about 80 km southwards (Fig. 8). The holotype (CORBIDI 7678) was found on 4 May 2009 at 3: 30 pm on a track (Fig. 7 A). Air temperature was 23.6 °C, ground temperature was 25.6 °C and air humidity was 73 %. ZFMK 96676 was found on 7 December 2009 at 10: 40 am lying dead on dry and hot soil (45.5 °C). Despite the high ground temperature the specimen was in a good condition indicating that it was only dead for a short time. Two days later at 2: 10 pm two further specimens (CORBIDI 5670, ZFMK 90934) were found under stones at the roadside (Fig. B). While lifting the stone CORBIDI 5670 was accidentally killed and its head was destroyed. Air temperature was between 30.5 °C– 33.8 °C, temperature under the stones was 33.6 °C and air humidity was between 57 %– 64 %.

Epictia vanwallachi sp. nov. (Figures 2 D–F, 4) Holotype: CORBIDI 14682, from Vijus Village, Pataz Province, La Libertad Region, Peru (S 07° 43 ’ 11.6 ’’, W 077° 39 ’ 51.1 ’’, 1290 m.a.s.l.), collected by E. Hoyas Granda, A. Beraún and C. Koch on 10 January 2010. Diagnosis. (1) 14 midbody scale rows; (2) 10 midtail scale rows; (3) 2 supralabials, first large and in broad contact with supraocular; (4) 16 subcaudals; (5) 188 mid-dorsal scale rows; (6) dorsal scales of head, body and tail brown with thin white or yellowish margins; (7) rostral uniformly grayish-brown; (8) terminal spine dorsally and ventrally and last three subcaudals yellow; (9) ventral scales of head, body and tail grayish-brown with creamishwhite margins. Comparisons [conditions for other Epictia in brackets]: By having 188 mid-dorsal scales, the new species has a higher number than E. collaris [155–166] and a lower number than E. albipuncta [213–285], E. alfredschmidti [267–279], E. borapeliotes [256–282], E. columbi [240–265], E. diaplocia [205–233], E. subcrotilla [318–333], E. melanura [395–396], E. rufidorsa [255–270], E. septemlineata [257], E. striatula [216–265], E. teaguei [232–259], E. tenella [198–299], E. tesselata [258–283] and E. tricolor [285–310]. By having 16 subcaudal scales it further differs from E. columbi [22–25], E. melanura [18–20], E. munoai [10–14], E. nasalis [21] and E. tricolor [18–23]. By lacking a yellow dorsal blotch on the anterior region of the head this species differs from Epictia septemlineata, E. alfredschmidti, E. borapeliotes, E. clinorostris, E. collaris, E. diaplocia, E. goudotii, E. magnamaculata, E. nasalis, E. peruviana, E. rubrolineata, E. signata, E. striatula, E. subcrotilla, E. teaguei, E. tenella, E. tesselata, E. tricolor, E, undecimstriata and E. vellardi. It further differs from E. septemlineata [dorsum with seven black longitudinal stripes] and E. rubrolineata [dorsum red with 5 longitudinal black stripes] by having a color pattern with grayish-brown dorsal scales with thin white or yellowish margins. Description of holotype: An adult specimen with SVL of 99 mm; TAL of 8.1 mm; MB of 2.9 mm; MT of 2.2 mm; TL/TAL of 13.2; TL/MB of 36.9; HW of 2.6 mm; HH of 1.7 mm; HL of 2.6 mm; DSN of 0.8 mm; DNE of 0.6 mm; ED of 0.4 mm. Head subcylindrical, slightly depressed dorsoventrally, hardly distinguishable from neck; body cylindrical; slightly tapered cranially and caudally. Snout slightly sloped in lateral view, rounded in ventral view. Rostral visible in dorsal view, about 1.4 times longer than wide, almost squared ventrally, triangular dorsally with dorsal termination (apex) acute, almost reaching the imaginary transverse line between the anterior borders of eyes, contacting upper and lower nasal laterally and frontal dorsally. Nasal completely divided horizontally by a suture slightly oblique, reaching rostral and first supralabial; ellipsoid nostril located in the center of the suture and having the major axis oriented along the suture; supranasal about 1.5 times higher than wide and about 1.3 times higher than infranasal, contacting infranasal ventrally, first supralabial and supraocular posteriorly, and frontal dorsally; infranasal slightly higher than wide, about 1.7 times wider than anterior supralabial, contacting first supralabial posteriorly; two supralabial scales, first positioned anteriorly and second posteriorly to ocular scale (1 + 1); upper lip border formed by rostral, infranasal, anterior supralabial, ocular and posterior supralabial; first supralabial about three times higher than wide, exceeding nostril, almost reaching central level of eye, in contact with supraocular scale dorsally and ocular posteriorly; ocular scale pentagonal with dorsal apex acuminate, 1.4 times higher than wide, contacting supraocular anterodorsally, parietal posterodorsally and second supralabial posteriorly; eye located slightly above level of maximum width of ocular and with lower eye margin at nostril level, positioned anteriorly without contacting scale sutures; eyes entirely visible in dorsal view; second supralabial triangular, about as wide as high, reaching central level of eye, as high as anterior supralabial, 3.3 times wider than anterior supralabial at widest point; posterior margin of second supralabial in broad contact with temporal and in contact with first ventrolateral scale; dorsal margin of second supralabial in contact with parietal; temporal scale of same size as dorsal scales of lateral rows; supraocular scale almost spindle-shaped, oriented oblique, 2.5 times longer than wide, contacting parietal posteriorly, and frontal and postfrontal dorsally; supraocular, parietal and occipital scales visible in lateral view; mid-dorsal head plates (frontal, postfrontal, interparietal and interoccipital) only marginally imbricate, not decreasing in size posteriorly, hexagonal in dorsal view, except for pentagonal frontal, slightly higher but of same width than posterior mid-dorsal scales; frontal contacting postfrontal posteriorly; postfrontal contacting supraoculars anteriorly, parietals and interparietal posteriorly; interparietal contacting parietals and occipitals laterally, and interoccipital posteriorly; interoccipital contacting occipitals laterally, and nuchal and first pair of paravertebral dorsal scales posteriorly; parietal about 2.4 times higher than wide, marginally larger than occipital, both almost ovoid, oriented slightly oblique; lower margin of parietal contacting upper border temporal, posterior margin in broad contact with occipital; lower margin of occipital contacting temporal and first lateral body scale, posterior margin in broad contact with first paravertebral and first dorsolateral body scales; six infralabials per side, subequal in size; mental scale bell-shaped, excluded from infralabial border by the first two pairs of infralabial scales; first pair of infralabials triangular, distinctly larger than other infralabials; chin, gular, and dorsal and lateral head scales porous. Dorsal scales slightly imbricate, smooth, homogeneous, rhomboid in shape, about 1.6 times wider than long; 188 MDS; 14 – 14 – 14 D; 171 V; 10 TS; Anal plate large, twice as wide as long, almost triangular in shape, bordered anteriorly and posteriorly by five rows of scales; 16 SC, becoming smaller distally; each of last four scales on the dorsal surface of the tail fused with adjacent dorsolateral scales; terminal spine conical and strongly pointed, with stout base about as wide as long. Color of holotype in life: Rostral uniformly grayish-brown; dorsal scales from head, body, and tail brown; ventral scales from head, body and tail grayish-brown; both dorsal and ventral scales of head body and tail with thin creamish-white margins; terminal spine and last three subcaudals yellow, strongly mottled with brown dorsally and ventrally. Color of holotype in preservative: Rostral gray; dorsal scales from head and body brown with cream margins, yellow regions of terminal portion of tail changed to whitish cream; ventral surface of head, body and tail changed to sandy brown with cream margins. Etymology. This species name is a patronym for Van Wallach, American herpetologist, in recognition of his outstanding contributions towards systematics of the snake family Leptotyphlopidae. Distribution and natural history. Epictia vanwallachi is only known from the type locality in the interandean part of the Equatorial Dry Forest (Figs. 5 and 8). The specimen was collected under a stone on 10 January 2010 at 9: 55 pm, with air temperature of 25.8 °C and air humidity of 60 %. Remarks. This species does not have striped pattern and multiple colors as mentioned by Adalsteinsson et al. (2009) as diagnostic characters for the genus Epictia, but it shares all other diagnostic characters with its congeners.

Landauro, Caroll Z., García-Bravo, Antonio, Venegas, Pablo J. (2015): An endemic new species of Ameiva (Squamata: Teiidae) from an isolated dry forest in southern Peru. Zootaxa 3946 (3): 387-400, DOI: http://dx.doi.org/10.11646/zootaxa.3946.3.6

Koch, Claudia, Venegas, Pablo J., Böhme, Wolfgang (2015): Three new endemic species of Epictia Gray, 1845 (Serpentes: Leptotyphlopidae) from the dry forest of northwestern Peru. Zootaxa 3964 (2): 228-244, DOI: http://dx.doi.org/10.11646/zootaxa.3964.2.4

Epictia septemlineata sp. nov. (Figures 1, 2 A���C) Holotype: CORBIDI 14683, from Limon Village, Celend��n Province, Cajamarca Region, Peru (S 06�� 52 ��� 34.2 ������, W 078��05��� 10.5 ������, 2053 m.a.s.l.), collected by A. Garcia Bravo and C. Koch on 28 April 2009. Diagnosis. (1) 14 midbody scale rows; (2) 10 midtail scale rows; (3) 2 supralabials, first large and in broad contact with supraocular; (4) 16 subcaudals; (5) 257 mid-dorsal scale rows; (6) Dorsum with seven black longitudinal stripes, outermost interspaces bright yellow along the body, medial interspaces yellow near the head and tail, and midbody interspaces reddish-brown; (7) rostral yellowish-white dorsally and cream ventrally; (8) terminal spine black; (9) ventral surface of head and body cream except for a soft dark longitudinal dotted line running along the center of each ventral scale row, anal plate cream with two lateral irregular dark blotches, and ventral surface of the tail cream, with three longitudinal rows of dark spots that merge distally and form a large irregular triangle. Comparisons [conditions for other Epictia in brackets]: By having 257 mid-dorsal scales this species has a higher number than E. peruviana [185���199], E. collaris [155���166], E. diaplocia [205���233] and E. munoai [184��� 230], and a lower number than E. alfredschmidti [267���279], E. subcrotilla [318���333], E. melanura [395���396], and E. tricolor [285���310]. By having 16 subcaudal scales it further differs from E. columbi [22���25], E. melanura [18��� 20], E. munoai [10���14], E. nasalis [21] and E. tricolor [18���23]. By having a tricolor pattern of dorsal longitudinal stripes (reddish-brown, black and yellow) it differs from all members of the tesselata group except for E. alfredschmidti, E. teaguei and E. tricolor. By lacking a yellow terminal spine this species differs from E. alfredschmidti, E.australis, E. borapeliotes, E. clinorostris, E. collaris, E. diaplocia, E. goudotii, E. magnamaculata, E. nasalis, E. peruviana, E. rubrolineata, E. signata, E. striatula, E. subcrotilla, E. teaguei, E. tenella, E. tesselata, E. tricolor, E. undecimstriata and E. vellardi. Description of holotype: An adult specimen with SVL of 172 mm; TAL of 9 mm; MB of 3.7 mm; MT of 3 mm; TL/TAL of 20.1; TL/MB of 48.9; HW of 3.1 mm; HH of 2.1 mm; HL of 4.1 mm; DSN of 1.2 mm; DNE of 0.8 mm; ED of 0.5 mm. Head subcylindrical, slightly depressed dorsoventrally, indistinguishable from neck; body cylindrical; slightly tapered cranially and caudally. Snout rounded in lateral view, slightly angled in ventral view. Rostral visible in dorsal view, about twice as long as wide, rectangular ventrally, triangular dorsally with dorsal termination (apex) acute, reaching the imaginary transverse line between the anterior borders of the eyes, contacting upper and lower nasal laterally and frontal dorsally. Nasal completely divided horizontally by oblique suture, reaching rostral and first supralabial; ovoid nostril located in the center of the suture and having the major axis oriented along the suture; supranasal about twice as high as wide and about twice higher than infranasal, contacting infranasal ventrally, first supralabial and supraocular posteriorly, and frontal dorsally; infranasal slightly higher than wide, about 1.5 times wider than anterior supralabial, contacting first supralabial posteriorly; two supralabial scales, first positioned anteriorly and second posteriorly to ocular scale (1 + 1); upper lip border formed by rostral, infranasal, anterior supralabial, ocular and posterior supralabial; first supralabial three times higher than wide, exceeding nostril, slightly exceeding central level of eye, in contact with supraocular scale dorsally and ocular posteriorly; ocular scale pentagonal with dorsal apex acuminate, 1.5 times higher than wide, contacting supraocular anterodorsally, parietal posterodorsally and second supralabial posteriorly; eye located at level of maximum width of ocular and almost at nostril level, positioned anteriorly without contacting scale sutures; eyes placed laterally, but partly visible in dorsal view; second supralabial subtrapezoidal, about as wide as high, reaching central level of eye and almost as high as anterior supralabial, 2.5 times wider than anterior supralabial at widest point; posterior margin of second supralabial in broad contact with temporal and in small contact with first ventrolateral scale; dorsal margin of second supralabial in contact with parietal; temporal scale of same size as dorsal scales of lateral rows; supraocular scale almost spindle-shaped, oriented oblique, three times longer than wide, contacting parietal posteriorly, and frontal and postfrontal dorsally; supraocular, parietal and occipital scales visible in lateral view; mid-dorsal head plates (frontal, postfrontal, interparietal and interoccipital) imbricate, slightly decreasing in size posteriorly, subcircular in dorsal view, except for ellipsoid interoccipital, less wider than posterior mid-dorsal scales; frontal contacting postfrontal posteriorly; postfrontal contacting supraoculars anteriorly, parietals and interparietal posteriorly; interparietal contacting parietals and occipitals laterally, and interoccipital posteriorly; interoccipital contacting occipitals laterally, and nuchal and first pair of paravertebral dorsal scales posteriorly; parietal almost 2.5 times higher than wide, marginally larger than occipital, both almost rectangular, oriented slightly oblique; lower margin of parietal contacting upper border of temporal, posterior margin in broad contact with occipital; lower margin of occipital contacting temporal and first lateral body scale, posterior margin in broad contact with first paravertebral and first dorsolateral body scales; six infralabials per side, slightly subequal in size; mental scale with the same width at lip border as rostral; first pair of infralabials with exactly the same width at lip border as lower nasal; dorsal and lateral head scales porous. Dorsal scales imbricate, smooth, homogeneous, cycloid in shape, almost twice as wide as long; 257 MDS; 14 ��� 14 ��� 14 D; 241 V; 10 TS; Anal plate large, almost twice as wide as long, triangular in shape with distal apex rounded, bordered anteriorly and posteriorly by three rows of scales; 16 SC, becoming slightly smaller distally; each of last four scales on the dorsal surface of the tail fused with adjacent dorsolateral scales; terminal spine conical, with stout base slightly wider than long. Color of holotype in life: Rostral scale and infranasals yellowish-white dorsally and cream ventrally; dorsal head scales mostly blackish, except for thin indistinct white margins in the mid-dorsal head plates (frontal, postfrontal, interparietal and interoccipital); whitish lateral stripe covering entire infranasal, and lower parts of supralabials and ocular, dorsal part of supralabials and ocular blackish; dorsum with seven black longitudinal stripes, which run along the middle of each dorsal scale row; vertebral stripe slightly thicker than paravetebral stripes; outermost interspaces bright yellow, medial interspaces yellow near the head and tail, fading into reddishbrown at midbody; terminal spine black; ventral surface of head, body and tail cream except for a soft dark longitudinal dotted line running along the middle of each ventral scale row, outermost thickest, two dark indistinct blotches on the anal plate, three longitudinal rows of dark indistinct blotches on the subcaudals, and a triangleshaped blotch formed by the fusion of the three longitudinal rows in the distal portion of the ventral surface of the tail, with the apex oriented caudally. Color of holotype in preservative: The pattern of longitudinal black stripes and other dark markings on the body remains unchanged, likewise the dark coloration of the head scales; the yellowish-white dot on rostral and infranasals changed to whitish cream; the yellow interspaces changed to cream, and the reddish-brown interspaces changed to sandy brown. Etymology. The specific name is derived from the Latin septem = seven and lineata = striped and refers to the diagnostic pattern of seven longitudinal black stripes on the dorsum. Distribution and natural history:. So far only known from the type locality in the interandean part of the Equatorial Dry Forest (Figs. 3 and 8). The single specimen was collected on the end of April 2009 at 5: 30 pm under a stone on soft soil of a recently tilled grainfield (Fig. 3). Temperature on the soil under the stone was 25.9 ��C, air temperature was 23 ��C and air humidity was 57 %., Published as part of Koch, Claudia, Venegas, Pablo J. & B��hme, Wolfgang, 2015, Three new endemic species of Epictia Gray, 1845 (Serpentes: Leptotyphlopidae) from the dry forest of northwestern Peru, pp. 228-244 in Zootaxa 3964 (2) on pages 230-232, DOI: 10.11646/zootaxa.3964.2.4, http://zenodo.org/record/244441

Epictia vanwallachi sp. nov. (Figures 2 D���F, 4) Holotype: CORBIDI 14682, from Vijus Village, Pataz Province, La Libertad Region, Peru (S 07�� 43 ��� 11.6 ������, W 077�� 39 ��� 51.1 ������, 1290 m.a.s.l.), collected by E. Hoyas Granda, A. Bera��n and C. Koch on 10 January 2010. Diagnosis. (1) 14 midbody scale rows; (2) 10 midtail scale rows; (3) 2 supralabials, first large and in broad contact with supraocular; (4) 16 subcaudals; (5) 188 mid-dorsal scale rows; (6) dorsal scales of head, body and tail brown with thin white or yellowish margins; (7) rostral uniformly grayish-brown; (8) terminal spine dorsally and ventrally and last three subcaudals yellow; (9) ventral scales of head, body and tail grayish-brown with creamishwhite margins. Comparisons [conditions for other Epictia in brackets]: By having 188 mid-dorsal scales, the new species has a higher number than E. collaris [155���166] and a lower number than E. albipuncta [213���285], E. alfredschmidti [267���279], E. borapeliotes [256���282], E. columbi [240���265], E. diaplocia [205���233], E. subcrotilla [318���333], E. melanura [395���396], E. rufidorsa [255���270], E. septemlineata [257], E. striatula [216���265], E. teaguei [232���259], E. tenella [198���299], E. tesselata [258���283] and E. tricolor [285���310]. By having 16 subcaudal scales it further differs from E. columbi [22���25], E. melanura [18���20], E. munoai [10���14], E. nasalis [21] and E. tricolor [18���23]. By lacking a yellow dorsal blotch on the anterior region of the head this species differs from Epictia septemlineata, E. alfredschmidti, E. borapeliotes, E. clinorostris, E. collaris, E. diaplocia, E. goudotii, E. magnamaculata, E. nasalis, E. peruviana, E. rubrolineata, E. signata, E. striatula, E. subcrotilla, E. teaguei, E. tenella, E. tesselata, E. tricolor, E, undecimstriata and E. vellardi. It further differs from E. septemlineata [dorsum with seven black longitudinal stripes] and E. rubrolineata [dorsum red with 5 longitudinal black stripes] by having a color pattern with grayish-brown dorsal scales with thin white or yellowish margins. Description of holotype: An adult specimen with SVL of 99 mm; TAL of 8.1 mm; MB of 2.9 mm; MT of 2.2 mm; TL/TAL of 13.2; TL/MB of 36.9; HW of 2.6 mm; HH of 1.7 mm; HL of 2.6 mm; DSN of 0.8 mm; DNE of 0.6 mm; ED of 0.4 mm. Head subcylindrical, slightly depressed dorsoventrally, hardly distinguishable from neck; body cylindrical; slightly tapered cranially and caudally. Snout slightly sloped in lateral view, rounded in ventral view. Rostral visible in dorsal view, about 1.4 times longer than wide, almost squared ventrally, triangular dorsally with dorsal termination (apex) acute, almost reaching the imaginary transverse line between the anterior borders of eyes, contacting upper and lower nasal laterally and frontal dorsally. Nasal completely divided horizontally by a suture slightly oblique, reaching rostral and first supralabial; ellipsoid nostril located in the center of the suture and having the major axis oriented along the suture; supranasal about 1.5 times higher than wide and about 1.3 times higher than infranasal, contacting infranasal ventrally, first supralabial and supraocular posteriorly, and frontal dorsally; infranasal slightly higher than wide, about 1.7 times wider than anterior supralabial, contacting first supralabial posteriorly; two supralabial scales, first positioned anteriorly and second posteriorly to ocular scale (1 + 1); upper lip border formed by rostral, infranasal, anterior supralabial, ocular and posterior supralabial; first supralabial about three times higher than wide, exceeding nostril, almost reaching central level of eye, in contact with supraocular scale dorsally and ocular posteriorly; ocular scale pentagonal with dorsal apex acuminate, 1.4 times higher than wide, contacting supraocular anterodorsally, parietal posterodorsally and second supralabial posteriorly; eye located slightly above level of maximum width of ocular and with lower eye margin at nostril level, positioned anteriorly without contacting scale sutures; eyes entirely visible in dorsal view; second supralabial triangular, about as wide as high, reaching central level of eye, as high as anterior supralabial, 3.3 times wider than anterior supralabial at widest point; posterior margin of second supralabial in broad contact with temporal and in contact with first ventrolateral scale; dorsal margin of second supralabial in contact with parietal; temporal scale of same size as dorsal scales of lateral rows; supraocular scale almost spindle-shaped, oriented oblique, 2.5 times longer than wide, contacting parietal posteriorly, and frontal and postfrontal dorsally; supraocular, parietal and occipital scales visible in lateral view; mid-dorsal head plates (frontal, postfrontal, interparietal and interoccipital) only marginally imbricate, not decreasing in size posteriorly, hexagonal in dorsal view, except for pentagonal frontal, slightly higher but of same width than posterior mid-dorsal scales; frontal contacting postfrontal posteriorly; postfrontal contacting supraoculars anteriorly, parietals and interparietal posteriorly; interparietal contacting parietals and occipitals laterally, and interoccipital posteriorly; interoccipital contacting occipitals laterally, and nuchal and first pair of paravertebral dorsal scales posteriorly; parietal about 2.4 times higher than wide, marginally larger than occipital, both almost ovoid, oriented slightly oblique; lower margin of parietal contacting upper border temporal, posterior margin in broad contact with occipital; lower margin of occipital contacting temporal and first lateral body scale, posterior margin in broad contact with first paravertebral and first dorsolateral body scales; six infralabials per side, subequal in size; mental scale bell-shaped, excluded from infralabial border by the first two pairs of infralabial scales; first pair of infralabials triangular, distinctly larger than other infralabials; chin, gular, and dorsal and lateral head scales porous. Dorsal scales slightly imbricate, smooth, homogeneous, rhomboid in shape, about 1.6 times wider than long; 188 MDS; 14 ��� 14 ��� 14 D; 171 V; 10 TS; Anal plate large, twice as wide as long, almost triangular in shape, bordered anteriorly and posteriorly by five rows of scales; 16 SC, becoming smaller distally; each of last four scales on the dorsal surface of the tail fused with adjacent dorsolateral scales; terminal spine conical and strongly pointed, with stout base about as wide as long. Color of holotype in life: Rostral uniformly grayish-brown; dorsal scales from head, body, and tail brown; ventral scales from head, body and tail grayish-brown; both dorsal and ventral scales of head body and tail with thin creamish-white margins; terminal spine and last three subcaudals yellow, strongly mottled with brown dorsally and ventrally. Color of holotype in preservative: Rostral gray; dorsal scales from head and body brown with cream margins, yellow regions of terminal portion of tail changed to whitish cream; ventral surface of head, body and tail changed to sandy brown with cream margins. Etymology. This species name is a patronym for Van Wallach, American herpetologist, in recognition of his outstanding contributions towards systematics of the snake family Leptotyphlopidae. Distribution and natural history. Epictia vanwallachi is only known from the type locality in the interandean part of the Equatorial Dry Forest (Figs. 5 and 8). The specimen was collected under a stone on 10 January 2010 at 9: 55 pm, with air temperature of 25.8 ��C and air humidity of 60 %. Remarks. This species does not have striped pattern and multiple colors as mentioned by Adalsteinsson et al. (2009) as diagnostic characters for the genus Epictia, but it shares all other diagnostic characters with its congeners., Published as part of Koch, Claudia, Venegas, Pablo J. & B��hme, Wolfgang, 2015, Three new endemic species of Epictia Gray, 1845 (Serpentes: Leptotyphlopidae) from the dry forest of northwestern Peru, pp. 228-244 in Zootaxa 3964 (2) on pages 233-235, DOI: 10.11646/zootaxa.3964.2.4, http://zenodo.org/record/244441, {"references":["Adalsteinsson, S. A., Branch, W. R., Trape, S., Vitt, L. J. & Hedges, S. B. (2009) Molecular phylogeny, classification, and biogeography of snakes of the Family Leptotyphlopidae (Reptilia, Squamata). Zootaxa, 2244, 1 - 50."]}

Ameiva reticulata sp. nov. Figs. 1 –4, 5 (A–D) Holotype. CORBIDI 0 8816 (Figs. 1–2), an adult male collected on 31 July 2010 by A. García in the Valle Seco del Mantaro (12 ° 5 ' 26.916 ''S, 74 ° 41 ' 55.968 ''W, WGS 84) at 1411 m.a.s.l., District of Surcubamba, Province of Tayacaja, Region of Huancavelica, Peru. Paratypes. A total of 35 specimens, all from the Province of Tayacaja, Region of Huancavelica, Peru: one adult male (CORBIDI 1095) from Barropata (12 ° 17 ' 34.812 ''S, 74 ° 40 ' 58.826 ''W, WGS 84), at 1528 m.a.s.l., District of Surcubamba, collected on 28 April 2008 by C.Z. Landauro; one adult male (CORBIDI 9919) from Intivilca (12 ° 19 ' 51.4 ''S, 74 ° 37 ' 53.1 ''W, WGS 84) at 2238 m.a.s.l., District of Colcabamba, collected on 0 4 April 2011 by D. Amaya; one adult male (CORBIDI 9917) and three adult females (CORBIDI 9918, 9920, 9921) from Jatuspata (12 ° 15 ' 1.2 ''S, 74 ° 41 ' 33.6 ''W, WGS 84) at 2609 m.a.s.l., District of Surcubamba, collected on 0 7 April 2011 by D. Amaya; four adult males (CORBIDI 10084, 10086, 10090, 10092), four juvenile males (CORBIDI 10081, 10082, 10087, 10094), seven adult females (CORBIDI 10075, 10076, 10078, 10085, 10088, 10089, 10093) and five juvenile females (CORBIDI 10077, 10079, 10080, 10083, 10091), all from Valle Seco del Mantaro (12 ° 5 ' 43.073 ''S, 74 ° 42 ' 45.002 '' W, WGS 84) at 1180 m.a.s.l., District of Surcubamba, collected on 0 1 and 0 2 November 2011 by A. Urbay and C.Z. Landauro; one adult female (CORBIDI 13623) from Campamento Limonal (12 ° 14 ' 5.184 '' S, 74 ° 41 ' 52.100 '' W, WGS 84) at 1438 m.a.s.l., District of Surcubamba, collected on 31 August 2013 by A. Escóbar; eight specimens from Pichiu District of Colcabamba: one juvenile (CORBIDI 13622) at 1974 m.a.s.l. (12 ° 20 ' 9.890 '' S, 74 ° 39 ' 26.755 '' W, WGS 84), collected on 10 September 2013 by C.Z. Landauro; two juveniles (CORBIDI 13620, 13621) at 2003 m.a.s.l. (12 ° 19 ' 53.815 '' S, 74 ° 39 ' 6.943 '' W, WGS 84), collected on 11 September 2013 by C.Z. Landauro; one adult male (CORBIDI 13625) and one adult female (CORBIDI 13628) at 2169 m.a.s.l. (12 ° 19 ' 15.902 '' S, 74 ° 38 ' 51.498 '' W, WGS 84), collected on 14 September 2013 by A. Escóbar; one adult female (CORBIDI 13629) at 2169 m.a.s.l. (12 ° 19 ' 32.913 '' S, 74 ° 39 ' 5.017 '' W, WGS 84), collected on 15 September 2013 by A. Escóbar; one adult male (CORBIDI 13626) and one adult female (CORBIDI 13624) at 1868 m.a.s.l (12 ° 21 ' 48.634 '' S, 74 ° 37 ' 27.249 '' W, WGS 84), collected on 18 September 2013 by A. Escóbar. Diagnosis. A medium-sized Ameiva distinguished from all congeners by the following combination of characters: (1) maximum SVL in males 149 mm; (2) dorsal head scales smooth; (3) frontal single; (4) frontoparietal and parietal plates in contact with interparietals; (5) 17–24 (both sides) scales, usually in single row between supraoculars and supraciliaries; (6) 8–17 occipitals, usually subequal to first dorsal row; (7) 19–25 anterior gulars; (8) middle anterior gulars polygonal and usually distinctly enlarged; (9) patch of distinctly enlarged posterior gulars usually present; (10) 8–14 posterior gulars between antegular and gular folds; (11) enlarged mesoptychial scales subequal or larger than largest gulars; (12) postbrachials moderately to distinctly enlarged; (13) 181–237 scales between interparietals and base of tail; (14) 102–137 dorsal scales across midbody; (15) ventrals in 28–32 transverse rows, and in eight longitudinal rows; (16) in life, adult individuals with head, arms, dorsum and flanks pale brown or grayish brown, legs and tail bright green or turquoise, flanks with light green or turquoise ocelli; (17) throat in adults white; (18) dorsum finely reticulated without a pale vertebral stripe along dorsum; (19) in life, dorsum and arms of juveniles brown with an olive green tint, legs and tail turquoise, and flanks with pale yellow ocelli; (20) associated with interandean dry forest. Comparison with other species. Ameiva reticulata shares the characters described by Harvey et al. (2012) for the A. ameiva complex (e.g. A. ameiva, A. atrigularis, A. pantherina, and A. praesignis): frontal entire, its posterior suture usually contacting second supraocular or aligned with suture between second and third supraoculars; frontal ridge absent; interparietal entire; parietal series (including interparietal) composed of five relatively long plates; narial suture passes through nostril; first supralabial usually curved; first subocular usually separated from supralabials by anterior expansion of the second subocular; intertympanic crease present; pectoral sulcus absent; ventral scales in 10–12 longitudinal rows; plate-like antebrachials continuous with or narrowly separated from brachial scales; combined femoral and abdominal pores number 28–45; fifth toe reduced; complete row of scales separate supradigital and subdigital lamellae along postaxial edge of each toe; tip of snout of adult males never reddish; adults without dorsolateral and vertebral stripes, or vertebral stripe only apparent on posterior dorsum; flanks with pale ocelli. Within the Ameiva ameiva complex, A. reticulata most resembles A. ameiva, a species widely distributed throughout Amazonia (Ugueto & Harvey 2011), and differs from it as follows (characteristics of A. ameiva in parentheses): maximum SVL in males 149 mm (163 mm) and in females 129 mm (146 mm); frontal usually pentagonal (hexagonal or roughly pentagonal in some specimens); fourth supraocular separated from parietals by one or two rows of circumorbital scales (up to four rows of circumorbital scales); 17–24 scales laterally between supraoculars and supraciliaries laterally (19–31 scales); ventrals in eight longitudinal rows (ten longitudinal rows); 181–237, mean= 199.9 ± 36.52, dorsal scales along the middorsal line (209–287, mean = 252.6 ± 16.65); 102–137, mean = 123.4 ± 8.24 dorsal scales at midbody in a transverse row (135–175, mean = 154.3 ± 8.8); 19–25, mean = 22.27 ± 1.56 anterior gulars (18–32, mean = 25.6 ± 2.64); preanal shield with four rows of enlarged scales (5–7); in juveniles, dorsal surface brown or olive green with black reticulation and white pale ocelli on flanks (first third or half of dorsum green and rest brown, flanks dark brown without ocelli; Fig. 5 E); in adults, dorsal surface of body brown or greenish-brown with a soft black reticulation, flanks completely brown or brown fading into greenish or turquoise towards hind limbs and with ill-defined light ocelli, tail turquoise (first third or half of dorsum brown with dark brown flecks, dots or blotches and remaining part of dorsum and tail green; Fig. 5 F). From the other species of the Ameiva ameiva complex such as A. atrigularis, A. pantherina (both from Venezuela) and A. praesignis ( from Costa Rica, Panama and Colombia) (Ugueto & Harvey 2011), A. reticulata can be separated by having: a lower scale count with 181–237, mean = 252.6 dorsals along the middorsal line and 102– 137, mean = 123.4 dorsals across the midbody in a transverse row (263–361, mean = 309 and 134–179, mean = 151.3 in A. atrigularis; 291–343, mean = 311.3 and 137–163, mean = 147.4 in A. pantherina; and 237–348, mean = 289.3 and 111–157, mean = 133 in A. praesignis). Ameiva atrigularis and A. praesignis are longer than A. reticulata, with a maximum SVL in males of 186 mm and 243 mm, respectively (149 mm in A. reticulata). Further, throat coloration in adults is dark gray or black in A. atrigularis, dark gray in A. pantherina, cream or blue in A. praesignis (white in A. reticulata). Other species of the genus Ameiva that occur in Peru such as A. aggerescusans, A. concolor and A. nodam, are all restricted to the upper Marañón basin in northern Peru (Koch et al. 2013), and share a transversely divided frontal plate, while it is entire in A. reticulata. Description. In males maximum SVL is 149.1 mm (CORBIDI 9919) and maximum total length is 481.1 mm (CORBIDI 9919); in females maximum SVL is 129.1 mm (CORBIDI 10089) and maximum total length is 357.1 mm (CORBIDI 10089). Head pyramidal, 0.25–0.29 (0.27 ± 0.01, n = 16) times SVL in males; 0.23–0.30 (0.26 ± 0.016, n = 20) times SVL in females. Snout elongate, bluntly pointed; canthus rostralis distinct. Neck narrower than head and body. Body cylindrical. Limbs well developed; tibia 0.11–0.18 (0.15 ± 0.02, n = 16) times SVL in males; 0.12–0.16 (0.14 ± 0.01, n = 20) times SVL in females; foot 0.30–0.42 (0.37 ± 0.03, n = 16) times SVL in males; 0.28–0.41 (0.36 ± 0.03, n = 20) times SVL in females. Tail round in cross section, tapers toward tip, 1.93– 2.50 (2.26 ± 0.19, n = 11) times SVL in males; 1.76–2.40 (2.14 ± 0.24, n = 13) times SVL in females. Rostral pentagonal, slightly wider than high, visible from above, bordered posteriorly by nasals, which form a short medial suture. Each nasal divided by an oblique suture. Nostril in lower part of suture and anterior to it, directed lateroposteriorly, slightly taller than long. Frontonasal octagonal, in contact with nasals, loreal and prefrontals. Prefrontals paired, roughly pentagonal, with a very long medial suture about twice as long as that between nasals, laterally in contact with loreal, first supraocular and first supraciliar. Frontal pentagonal (rectangular in CORBIDI 10082, CORBIDI 10092), longer than wide and wider anteriorly, its sutures with prefrontals form a wide angle, slightly straight, its sutures with frontoparietals almost forming a straight line; frontal laterally in contact with first and second supraocular. Pair of trapezoidal frontoparietals (divided into two posteriorly in CORBIDI 9918, 1007677, 10080, 10084, 10087, 10094 and 10095 and with a small scale accessory between frontal and frontoparietals in CORBIDI 0 9919, 10090, 10082, 10087, 10089 and 10094), wider than long and with a long medial suture; frontoparietals laterally in contact with second and third supraocular, and small circumorbital scales bordering fourth up to middle portion of third supraocular or beginning of second supraocular. Interparietal irregularly hexagonal, higher than wide, often narrower (but sometimes slightly wider) than adjacent parietals; interparietal longitudinally divided; sutures with parietals slightly oblique and curved; interparietal bordered at each side by two large, irregular parietals divided by an oblique suture (parietals divided into two in CORBIDI 10078); outermost parietal oval in shape and larger than inner parietal. Parietal series composed of five scales including interparietal. Occipitals 8–17 (13.61 ± 2.25, n = 36), irregular and heterogeneous in size; occipitals usually subequal in size to first dorsal row, less often moderately to distinctly larger. Supraoculars four at each side. Circumorbital semicircles formed by 4–8 scales at each side, 9–15 (11.25 ± 1.46, n = 36) combining both sides, reaching middle, posterior and anterior portion of third and second supraocular (extending completely around supraoculars except first supraocular); only one scale row between third supraocular and prefrontals; fourth supraocular separated from parietals by one or two rows of circumorbital scales. Laterally, all supraoculars except first separated from supraciliaries by a single row of small rectangular scales; 17–24 (20.72 ± 1.59, n = 36) combining both sides. Supraciliaries five or seven (on one side), first highest, second longest. Loreal very large, trapezoidal and single, in contact with nasal, frontonasal, first supraciliary, first and second subocular, and third and fourth supralabial. Suboculars four; first subocular in preocular position, irregularly trapezoidal, longer than wide and slightly wider than second subocular; in contact with loreal, first supraciliary, small scales in ocular region, and second subocular; margin with loreal slightly curved or straight. Third subocular longest, all, except first, in contact with supralabials. Continuous keel from first subocular through entire length of third subocular. Postoculars small, approximately in one row of four or five irregularly trapezoidal scales. Palpebral disc opaque with enlarged scales on lower eyelid. Supralabials seven, sixth below center of eye. Row of enlarged supratemporals decreasing in size posteriad. Temporal region with polygonal scales, slightly smaller centrally than peripherally. External auditory meatus large, vertically oval, bordered by granular scales; anterior margin semicircular, posterior one straight. Tympanum recessed. All dorsal and lateral head scales juxtaposed and smooth. Symphysal anteriorly ellipsoid, its posterior sutures forming wide angles with infralabials and postsymphysal. Postsymphysal single and pentagonal; in contact with first and second infralabial, followed by five enlarged chinshields. First pair in ample medial contact, only the first in contact with infralabials. Remaining chinshields separated from infralabials by one row of sublabials. Medial chin scales moderately small, elongate, convex, smooth, juxtaposed, subhexagonal, in slightly oblique rows, all subequal in size. Infralabials six, fifth below center of eye followed to commissure of mouth by smaller scales. Gular region divided into two areas: anterior region with polygonal and flat scales in slightly oblique rows that usually remain subequal or rarely grade to larger scales medially, delimited posteriorly by a line uniting the lower margins of the ear openings; middle anterior gular scales usually small, rarely moderately enlarged, 19–25 (22.27 ± 1.56, n = 36) scales along midline of chin from anteriormost to posteriormost anterior gular. Posterior gular region covered by smaller polygonal scales in transverse rows; posteriormost median gulars usually forming medial patch of moderately enlarged scales, 8–14 (10.72 ± 1.64, n = 36) scales midventrally from anteriormost posterior gular to antegular fold. Mesoptychial scales moderately enlarged (larger or, less often, subequal to largest anterior gulars), in about one or two rows, hexagonal, flat, smooth, and juxtaposed. Scales on nape and sides of neck similar to dorsals. Dorsals and scales on flanks granular (slightly larger on dorsum than laterally), round, smooth, subimbricate, in approximately transverse rows; 181–237 (199.9 ± 36.52, n = 36) scales between occipitals and base of tail. Scales around midbody (excluding ventrals) 102–137 (123.47 ± 8.24, n = 36). Ventrals large, smooth, rectangular (wider than long), imbricate, in eight longitudinal (at midbody) and 28–32 (30.08 ± 0.93, n = 36) transverse rows; transition between ventrals and scales on flanks sharp. Preanal shield with four rows of enlarged scales; pattern of terminal scales of the preanal plate somewhat variable; often large median scale followed posteriorly by two subequal scales. Preanal plate surrounded anteriorly and laterally by smaller scales; posteriorly by much smaller scales. Femoral pores in continuous row along each thigh, 16–20 pores on each leg, 32–40 (35.88 ± 2.13, n = 36) pores combining both legs; each pore surrounded by four scales. Scales on base of tail trapezoidal, smaller than ventrals, longer than wide, mostly keeled (smooth on base of tail on ventral surface). Caudal scales imbricate, in transverse rows, continuous around tail. Distal caudals longer and narrower, distinctly keeled, in transverse and approximately longitudinal rows. Arms with rows of very large, smooth, slightly imbricate, trapezoidal antebrachial scales on anterodorsal aspect of forearms and similar but smaller brachial scales on upper arms that extend almost to shoulder. Brachial row distinctly enlarged with smaller accessory row bordering upper margin and grading into granules posteriorly. Antebrachials and brachials in contact or separated by smaller scales at elbow. Dorsoposterior, posterior, and ventral aspect of arms with scales similar to dorsals, but slightly larger, except for a group (approximately one or two rows) of postbrachial scales on posterior aspect of upper arms, which are slightly more irregular. Legs with large, smooth, imbricate scales on anterior and ventral aspects of thighs, and ventral aspects of shanks. One row of large, trapezoidal scales anteriorly on thigh, gradually becoming smaller and irregular toward pores. On ventral aspect of shanks, four rows of large scales, anterior two trapezoidal, posterior one rhomboidal, decreasing in size from anterior toward posterior row. Elsewhere on legs scales similar to dorsals. Subdigital lamellae transversely enlarged and single, moderately to distinctly tuberculate towards base. Tubercles most prominent under base of third toe, lamellae of outer toe continuing to heel. On palms, lamellae of inner finger continuing to wrist as large scales, increasing in size toward it; single large scale on postaxial side, following line of fifth finger, at short distance from wrist. Lamellae under fourth finger 14–17 (15.83 ± 0.69, n = 36), under fourth toe 27–36 (31.41 ± 1.79, n = 36). Data of holotype. SVL 136.5 mm; head length 36.9 mm; tibia length 19.7 mm; foot length 45.5 mm; hand length 13.35 mm; tail length 285 mm; supralabials 7 / 7; infralabials 6 / 6; supraoculars 4 / 4; circumorbitals 5 / 5 extending to mid third supraocular; 10 / 11 scales between supraorbital and supraciliary scales; parietals 5; occipitals 16; occipitals larger than first dorsal row; 22 anterior gulars; mid anterior gulars small; 14 posterior gulars, midposterior gulars moderately enlarged; mesoptychial scales smaller than largest anterior gulars; dorsals between occipitals and base of tail 185; 120 scales around midbody (excluding ventrals); 8 longitudinal ventral rows; 30 transverse ventral rows; femoral pores 18 / 18; four scales forming preanal plate; 16 lamellae under fourth finger on right hand; 31 lamellae under fourth toe on right foot. Color in preservative. Young specimens with dorsal surface of head and dorsal part of body uniformly brownish with medium-sized black spots at each side of vertebral region (most conspicuous towards sacrum). Sides of head uniformly brownish. Some specimens with almost half of vertebral region skyblue or turquoise. Dorsolateral surface black with regular turquoise spots, extending to base of tail. Upper limbs brownish with some small black scales. Hind limbs black with turquoise spots or turquoise with black spots. Throat pale, chest, ventral aspect of arms and most of abdomen semi pale (with a bluish tint). Ventral aspect of legs, posteriormost portion of abdomen, anal and subcaudal regions whitish with some dark scales. Adult males have a turquoise brown dorsum with black reticulation or flecks (Fig. 3; more visible in some females: Fig. 4); these markings remain conspicuous along dorsum from nape to base of tail. Flanks with small, regular, slightly scattered turquoise ocelli with distinct black margins more or less arranged in vertical rows; ocelli never extend onto dorsum. Top and sides

Epictia antoniogarciai sp. nov. (Figures 2 G���I, 6) Holotype: CORBIDI 7678, from the vicinities of Santa Rosa de la Yunga Village, Ja��n Province, Cajamarca Region, Peru (S 05�� 25 ��� 53.3 ������, W 078�� 33 ���47.0������, 1268 m.a.s.l.), collected by M. Enciso, S. Duran and C. Koch on 4 May 2009. Paratypes (N = 4): three specimens from the vicinities Zapatalgo Village, Utcubamba Province, Amazonas Region, Peru: ZFMK 96676 (S 06��04��� 47.7 ������, W 078�� 29 ��� 18.7 ������, 934 m. a.s.l.), CORBIDI 5670, ZFMK 90934 (S 06��04���44.0������, W 078�� 29 ��� 16.7 ������, 968 m. a.s.l.), collected by A. Garcia Bravo and C. Koch between 07���09 December 2009; and one specimen from the type locality: CORBIDI 2069, collected in August 2008 by N. Monsalve. Diagnosis. (1) 14 midbody scale rows; (2) 10 midtail scale rows; (3) 2 supralabials, first large and in broad contact with supraocular; (4) 14���18 subcaudals; (5) 195���208 mid-dorsal scale rows; (6) dorsal scales black with yellow margins; (7) rostral bright yellow or yellowish-white dorsally, and grayish-brown or blackish ventrally; (8) terminal spine and surrounding scales yellow; (9) ventral surface of head, body and tail almost completely grayishbrown or light-gray scattered with dark grayish-brown scales. Comparisons [conditions for other Epictia in brackets]: By having 195���208 mid-dorsal scales this species has a higher number than E. collaris [155���166] and E. vanwallachi [188], and a lower number than E. albipuncta [213���285], E. alfredschmidti [267���279], E. borapeliotes [256���282], E. columbi [240���265], E. melanura [395��� 396], E. rufidorsa [255���270], E. septemlineata [257], E. striatula [216���265], E. subcrotilla [318���333], E. teaguei [232���259], E. tesselata [258���283] and E. tricolor [285���310]. By having 14���18 subcaudal scales it further differs from E. columbi [22���25], E. melanura [18���20], E. munoai [10���14], E. nasalis [21] and E. tricolor [18���23]. By having a different color pattern with dorsal scales black with thin yellowish-white or bright yellow margins, and rostral and terminal portion of tail yellowish-white or bright yellow, it further differs from E. septemlineata [dorsum with seven black longitudinal stripes], E. rubrolineata [dorsal coloration red with 5 longitudinal stripes], E. rufidorsa [striped pattern with a red dorsal longitudinal stripe], E. teaguei [tricolor dorsal pattern of red, black and yellow stripes], E. tenella [dorsum dark brown with lateral edges of the scales lighter, forming a pattern of serrated longitudinal light lines] and E. vanwallachi [dorsal scales brown with thin white or yellowish margins, rostral grayish-brown]. Description of holotype: An adult specimen with SVL of 129 mm; TAL of 14.5 mm; MB of 3.8 mm; MT of 3.0 mm; TL/TAL of 9.9; TL/MB of 37.8; HW of 2.8 mm; HH of 2.2 mm; HL of 3.4 mm; DSN of 0.9 mm; DNE of 0.8 mm; ED of 0.6 mm. Head subcylindrical, slightly depressed dorsoventrally, indistinguishable from neck; body cylindrical; not tapered cranially or caudally. Snout rounded in lateral and ventral view. Rostral visible in dorsal view, about 1.5 times longer than wide, almost squared ventrally with dorsal termination (apex) sharply rounded, almost reaching the imaginary transverse line between the anterior borders of the eyes, contacting upper and lower nasal laterally and frontal dorsally. Nasal completely divided horizontally by a suture slightly oblique, reaching rostral and first supralabial; ellipsoid nostril located almost in the center of the suture between upper and lower nasal, having the major axis oriented along the suture; supranasal about 1.9 times higher than wide and about 1.3 times higher than infranasal, contacting infranasal ventrally, first supralabial and supraocular posteriorly, and frontal dorsally; infranasal 1.7 times higher than wide, about 1.5 times wider than anterior supralabial, contacting first supralabial posteriorly; two supralabial scales, first positioned anteriorly and second posteriorly to ocular scale (1 + 1); upper lip border formed by rostral, infranasal, anterior supralabial, ocular and posterior supralabial; first supralabial about 2.8 times higher than wide, exceeding nostril, slightly exceeding central level of eye, in contact with supraocular scale dorsally and ocular posteriorly; ocular scale pentagonal with dorsal apex acuminate, 1.5 times higher than wide, contacting supraocular anterodorsally, parietal posterodorsally and second supralabial posteriorly; eye located at level of maximum width of ocular and with lower eye margin at nostril level, positioned anteriorly and almost contacting scale sutures; eyes partly visible in dorsal view; second supralabial subtrapezoidal, about as high as wide, slightly exceeding central level of eye, as high as anterior supralabial, 2.5 times wider than anterior supralabial at widest point; posterior margin of second supralabial in broad contact with temporal and in contact with first scale of lateral body row; dorsal margin of second supralabial in contact with parietal; temporal scale of same size as dorsal scales of lateral rows; supraocular scale almost spindle-shaped, oriented oblique, about twice as long as wide, contacting parietal posteriorly, and frontal and postfrontal dorsally; supraocular, parietal and occipital scales visible in lateral view; mid-dorsal head plates (frontal, postfrontal, interparietal and interoccipital) slightly imbricate, hardly decreasing in size posteriorly, pentagonal or round in dorsal view, higher and narrower than posterior mid-dorsal scales; frontal contacting postfrontal posteriorly; postfrontal contacting supraoculars anteriorly, parietals and interparietal posteriorly; interparietal contacting parietals and occipitals laterally, and interoccipital posteriorly; interoccipital contacting occipitals laterally, and nuchal and three dorsal body scales posteriorly; parietal about 1.8 times higher than wide, as high as occipital, about 1.5 times wider than occipital, both scales almost rectangular in shape; occipital about 2.5 times higher than wide; lower margin of parietal contacting upper border of temporal, posterior margin in broad contact with occipital; lower margin of occipital contacting temporal and first lateral body scale, posterior margin in broad contact with first paravertebral and first dorsolateral scale (latter both scales fused on left side); mental small with a median depression, followed by six infralabials per side, slightly subequal in size; first pair of infralabials rectangular, slightly larger than other infralabials; chin, gular, and dorsal and lateral head scales porous. Dorsal scales imbricate, smooth, homogeneous, rhomboid or elliptical in shape, about 1.4 times wider than long; 202 MDS; 14 ��� 14 ��� 14 D; 184 V; 10 TS; Anal plate large, 1.7 times wider than long, subtriangular in shape with distal apex rounded, bordered anteriorly and posteriorly by five rows of scales; 16 SC, becoming slightly narrower distally, except for ultimate four scales which are each fused with adjacent scales; each of last three scales on the dorsal surface of the tail fused with adjacent dorsolateral scales; terminal spine conical and strongly pointed, with stout base about as wide as long. Color of holotype in life: Anterior portion of head with a large bright yellow blotch, covering completely the dorsal surface of frontal and rostral scales, and partially the supranasal scales; rostral dark grayish-brown ventrally; remaining head and body dorsal scales black, with striking bright yellow margins; terminal spine and surrounding scales (two rows dorsally and three rows ventrally) bright yellow; ventral surface of head, body and tail light-gray, scattered with numerous darker grayish-brown scales. Color of holotype in preservative: Coloration of blotch on anterior dorsal scales of the head and terminal portion of tail changed to cream; rostral changed to dark gray ventrally; dorsal scales from head and body changed from dark brown to blackish with cream margins; ventral surface of head, body and tail changed to cream except for some scattered grayish-brown scales. Variation. Morphometric and pholidosis characters of the four paratypes are given in Table 1. The paratypes further vary from the holotype in the following characters: the yellow color pattern is distinctly less striking in three of the paratypes (ZFMK 96676, CORBIDI 5670, ZFMK 90934) than in the holotype: the blotch on the anterodorsal surface of the head is cream or yellowish-white; the dorsals are dark brown to black, most scales with thin, indistinct, yellowish margins; terminal spine pale yellow; ventral surface of head, body and tail almost uniformly grayish-brown, somewhat lighter than dorsal scales, or sometimes scattered with darker scales. Etymology. This species is dedicated to Antonio Garcia Bravo, Peruvian biologist, in recognition of both, his support in the investigation of the Peruvian herpetofauna and his continued and unattenuated efforts in the conservation of the dry forests along the Mara����n River. Distribution and natural history. This species is known from the interandean part of the Equatorial Dry Forest from Santa Rosa de la Yunga in the North to about 80 km southwards (Fig. 8). The holotype (CORBIDI 7678) was found on 4 May 2009 at 3: 30 pm on a track (Fig. 7 A). Air temperature was 23.6 ��C, ground temperature was 25.6 ��C and air humidity was 73 %. ZFMK 96676 was found on 7 December 2009 at 10: 40 am lying dead on dry and hot soil (45.5 ��C). Despite the high ground temperature the specimen was in a good condition indicating that it was only dead for a short time. Two days later at 2: 10 pm two further specimens (CORBIDI 5670, ZFMK 90934) were found under stones at the roadside (Fig. B). While lifting the stone CORBIDI 5670 was accidentally killed and its head was destroyed. Air temperature was between 30.5 ��C��� 33.8 ��C, temperature under the stones was 33.6 ��C and air humidity was between 57 %��� 64 %., Published as part of Koch, Claudia, Venegas, Pablo J. & B��hme, Wolfgang, 2015, Three new endemic species of Epictia Gray, 1845 (Serpentes: Leptotyphlopidae) from the dry forest of northwestern Peru, pp. 228-244 in Zootaxa 3964 (2) on pages 236-240, DOI: 10.11646/zootaxa.3964.2.4, http://zenodo.org/record/244441

Epictia septemlineata sp. nov. (Figures 1, 2 A–C) Holotype: CORBIDI 14683, from Limon Village, Celendín Province, Cajamarca Region, Peru (S 06° 52 ’ 34.2 ’’, W 078°05’ 10.5 ’’, 2053 m.a.s.l.), collected by A. Garcia Bravo and C. Koch on 28 April 2009. Diagnosis. (1) 14 midbody scale rows; (2) 10 midtail scale rows; (3) 2 supralabials, first large and in broad contact with supraocular; (4) 16 subcaudals; (5) 257 mid-dorsal scale rows; (6) Dorsum with seven black longitudinal stripes, outermost interspaces bright yellow along the body, medial interspaces yellow near the head and tail, and midbody interspaces reddish-brown; (7) rostral yellowish-white dorsally and cream ventrally; (8) terminal spine black; (9) ventral surface of head and body cream except for a soft dark longitudinal dotted line running along the center of each ventral scale row, anal plate cream with two lateral irregular dark blotches, and ventral surface of the tail cream, with three longitudinal rows of dark spots that merge distally and form a large irregular triangle. Comparisons [conditions for other Epictia in brackets]: By having 257 mid-dorsal scales this species has a higher number than E. peruviana [185–199], E. collaris [155–166], E. diaplocia [205–233] and E. munoai [184– 230], and a lower number than E. alfredschmidti [267–279], E. subcrotilla [318–333], E. melanura [395–396], and E. tricolor [285–310]. By having 16 subcaudal scales it further differs from E. columbi [22–25], E. melanura [18– 20], E. munoai [10–14], E. nasalis [21] and E. tricolor [18–23]. By having a tricolor pattern of dorsal longitudinal stripes (reddish-brown, black and yellow) it differs from all members of the tesselata group except for E. alfredschmidti, E. teaguei and E. tricolor. By lacking a yellow terminal spine this species differs from E. alfredschmidti, E.australis, E. borapeliotes, E. clinorostris, E. collaris, E. diaplocia, E. goudotii, E. magnamaculata, E. nasalis, E. peruviana, E. rubrolineata, E. signata, E. striatula, E. subcrotilla, E. teaguei, E. tenella, E. tesselata, E. tricolor, E. undecimstriata and E. vellardi. Description of holotype: An adult specimen with SVL of 172 mm; TAL of 9 mm; MB of 3.7 mm; MT of 3 mm; TL/TAL of 20.1; TL/MB of 48.9; HW of 3.1 mm; HH of 2.1 mm; HL of 4.1 mm; DSN of 1.2 mm; DNE of 0.8 mm; ED of 0.5 mm. Head subcylindrical, slightly depressed dorsoventrally, indistinguishable from neck; body cylindrical; slightly tapered cranially and caudally. Snout rounded in lateral view, slightly angled in ventral view. Rostral visible in dorsal view, about twice as long as wide, rectangular ventrally, triangular dorsally with dorsal termination (apex) acute, reaching the imaginary transverse line between the anterior borders of the eyes, contacting upper and lower nasal laterally and frontal dorsally. Nasal completely divided horizontally by oblique suture, reaching rostral and first supralabial; ovoid nostril located in the center of the suture and having the major axis oriented along the suture; supranasal about twice as high as wide and about twice higher than infranasal, contacting infranasal ventrally, first supralabial and supraocular posteriorly, and frontal dorsally; infranasal slightly higher than wide, about 1.5 times wider than anterior supralabial, contacting first supralabial posteriorly; two supralabial scales, first positioned anteriorly and second posteriorly to ocular scale (1 + 1); upper lip border formed by rostral, infranasal, anterior supralabial, ocular and posterior supralabial; first supralabial three times higher than wide, exceeding nostril, slightly exceeding central level of eye, in contact with supraocular scale dorsally and ocular posteriorly; ocular scale pentagonal with dorsal apex acuminate, 1.5 times higher than wide, contacting supraocular anterodorsally, parietal posterodorsally and second supralabial posteriorly; eye located at level of maximum width of ocular and almost at nostril level, positioned anteriorly without contacting scale sutures; eyes placed laterally, but partly visible in dorsal view; second supralabial subtrapezoidal, about as wide as high, reaching central level of eye and almost as high as anterior supralabial, 2.5 times wider than anterior supralabial at widest point; posterior margin of second supralabial in broad contact with temporal and in small contact with first ventrolateral scale; dorsal margin of second supralabial in contact with parietal; temporal scale of same size as dorsal scales of lateral rows; supraocular scale almost spindle-shaped, oriented oblique, three times longer than wide, contacting parietal posteriorly, and frontal and postfrontal dorsally; supraocular, parietal and occipital scales visible in lateral view; mid-dorsal head plates (frontal, postfrontal, interparietal and interoccipital) imbricate, slightly decreasing in size posteriorly, subcircular in dorsal view, except for ellipsoid interoccipital, less wider than posterior mid-dorsal scales; frontal contacting postfrontal posteriorly; postfrontal contacting supraoculars anteriorly, parietals and interparietal posteriorly; interparietal contacting parietals and occipitals laterally, and interoccipital posteriorly; interoccipital contacting occipitals laterally, and nuchal and first pair of paravertebral dorsal scales posteriorly; parietal almost 2.5 times higher than wide, marginally larger than occipital, both almost rectangular, oriented slightly oblique; lower margin of parietal contacting upper border of temporal, posterior margin in broad contact with occipital; lower margin of occipital contacting temporal and first lateral body scale, posterior margin in broad contact with first paravertebral and first dorsolateral body scales; six infralabials per side, slightly subequal in size; mental scale with the same width at lip border as rostral; first pair of infralabials with exactly the same width at lip border as lower nasal; dorsal and lateral head scales porous. Dorsal scales imbricate, smooth, homogeneous, cycloid in shape, almost twice as wide as long; 257 MDS; 14 – 14 – 14 D; 241 V; 10 TS; Anal plate large, almost twice as wide as long, triangular in shape with distal apex rounded, bordered anteriorly and posteriorly by three rows of scales; 16 SC, becoming slightly smaller distally; each of last four scales on the dorsal surface of the tail fused with adjacent dorsolateral scales; terminal spine conical, with stout base slightly wider than long. Color of holotype in life: Rostral scale and infranasals yellowish-white dorsally and cream ventrally; dorsal head scales mostly blackish, except for thin indistinct white margins in the mid-dorsal head plates (frontal, postfrontal, interparietal and interoccipital); whitish lateral stripe covering entire infranasal, and lower parts of supralabials and ocular, dorsal part of supralabials and ocular blackish; dorsum with seven black longitudinal stripes, which run along the middle of each dorsal scale row; vertebral stripe slightly thicker than paravetebral stripes; outermost interspaces bright yellow, medial interspaces yellow near the head and tail, fading into reddishbrown at midbody; terminal spine black; ventral surface of head, body and tail cream except for a soft dark longitudinal dotted line running along the middle of each ventral scale row, outermost thickest, two dark indistinct blotches on the anal plate, three longitudinal rows of dark indistinct blotches on the subcaudals, and a triangleshaped blotch formed by the fusion of the three longitudinal rows in the distal portion of the ventral surface of the tail, with the apex oriented caudally. Color of holotype in preservative: The pattern of longitudinal black stripes and other dark markings on the body remains unchanged, likewise the dark coloration of the head scales; the yellowish-white dot on rostral and infranasals changed to whitish cream; the yellow interspaces changed to cream, and the reddish-brown interspaces changed to sandy brown. Etymology. The specific name is derived from the Latin septem = seven and lineata = striped and refers to the diagnostic pattern of seven longitudinal black stripes on the dorsum. Distribution and natural history:. So far only known from the type locality in the interandean part of the Equatorial Dry Forest (Figs. 3 and 8). The single specimen was collected on the end of April 2009 at 5: 30 pm under a stone on soft soil of a recently tilled grainfield (Fig. 3). Temperature on the soil under the stone was 25.9 °C, air temperature was 23 °C and air humidity was 57 %.

Ameiva reticulata sp. nov. Figs. 1 ���4, 5 (A���D) Holotype. CORBIDI 0 8816 (Figs. 1���2), an adult male collected on 31 July 2010 by A. Garc��a in the Valle Seco del Mantaro (12 �� 5 ' 26.916 ''S, 74 �� 41 ' 55.968 ''W, WGS 84) at 1411 m.a.s.l., District of Surcubamba, Province of Tayacaja, Region of Huancavelica, Peru. Paratypes. A total of 35 specimens, all from the Province of Tayacaja, Region of Huancavelica, Peru: one adult male (CORBIDI 1095) from Barropata (12 �� 17 ' 34.812 ''S, 74 �� 40 ' 58.826 ''W, WGS 84), at 1528 m.a.s.l., District of Surcubamba, collected on 28 April 2008 by C.Z. Landauro; one adult male (CORBIDI 9919) from Intivilca (12 �� 19 ' 51.4 ''S, 74 �� 37 ' 53.1 ''W, WGS 84) at 2238 m.a.s.l., District of Colcabamba, collected on 0 4 April 2011 by D. Amaya; one adult male (CORBIDI 9917) and three adult females (CORBIDI 9918, 9920, 9921) from Jatuspata (12 �� 15 ' 1.2 ''S, 74 �� 41 ' 33.6 ''W, WGS 84) at 2609 m.a.s.l., District of Surcubamba, collected on 0 7 April 2011 by D. Amaya; four adult males (CORBIDI 10084, 10086, 10090, 10092), four juvenile males (CORBIDI 10081, 10082, 10087, 10094), seven adult females (CORBIDI 10075, 10076, 10078, 10085, 10088, 10089, 10093) and five juvenile females (CORBIDI 10077, 10079, 10080, 10083, 10091), all from Valle Seco del Mantaro (12 �� 5 ' 43.073 ''S, 74 �� 42 ' 45.002 '' W, WGS 84) at 1180 m.a.s.l., District of Surcubamba, collected on 0 1 and 0 2 November 2011 by A. Urbay and C.Z. Landauro; one adult female (CORBIDI 13623) from Campamento Limonal (12 �� 14 ' 5.184 '' S, 74 �� 41 ' 52.100 '' W, WGS 84) at 1438 m.a.s.l., District of Surcubamba, collected on 31 August 2013 by A. Esc��bar; eight specimens from Pichiu District of Colcabamba: one juvenile (CORBIDI 13622) at 1974 m.a.s.l. (12 �� 20 ' 9.890 '' S, 74 �� 39 ' 26.755 '' W, WGS 84), collected on 10 September 2013 by C.Z. Landauro; two juveniles (CORBIDI 13620, 13621) at 2003 m.a.s.l. (12 �� 19 ' 53.815 '' S, 74 �� 39 ' 6.943 '' W, WGS 84), collected on 11 September 2013 by C.Z. Landauro; one adult male (CORBIDI 13625) and one adult female (CORBIDI 13628) at 2169 m.a.s.l. (12 �� 19 ' 15.902 '' S, 74 �� 38 ' 51.498 '' W, WGS 84), collected on 14 September 2013 by A. Esc��bar; one adult female (CORBIDI 13629) at 2169 m.a.s.l. (12 �� 19 ' 32.913 '' S, 74 �� 39 ' 5.017 '' W, WGS 84), collected on 15 September 2013 by A. Esc��bar; one adult male (CORBIDI 13626) and one adult female (CORBIDI 13624) at 1868 m.a.s.l (12 �� 21 ' 48.634 '' S, 74 �� 37 ' 27.249 '' W, WGS 84), collected on 18 September 2013 by A. Esc��bar. Diagnosis. A medium-sized Ameiva distinguished from all congeners by the following combination of characters: (1) maximum SVL in males 149 mm; (2) dorsal head scales smooth; (3) frontal single; (4) frontoparietal and parietal plates in contact with interparietals; (5) 17���24 (both sides) scales, usually in single row between supraoculars and supraciliaries; (6) 8���17 occipitals, usually subequal to first dorsal row; (7) 19���25 anterior gulars; (8) middle anterior gulars polygonal and usually distinctly enlarged; (9) patch of distinctly enlarged posterior gulars usually present; (10) 8���14 posterior gulars between antegular and gular folds; (11) enlarged mesoptychial scales subequal or larger than largest gulars; (12) postbrachials moderately to distinctly enlarged; (13) 181���237 scales between interparietals and base of tail; (14) 102���137 dorsal scales across midbody; (15) ventrals in 28���32 transverse rows, and in eight longitudinal rows; (16) in life, adult individuals with head, arms, dorsum and flanks pale brown or grayish brown, legs and tail bright green or turquoise, flanks with light green or turquoise ocelli; (17) throat in adults white; (18) dorsum finely reticulated without a pale vertebral stripe along dorsum; (19) in life, dorsum and arms of juveniles brown with an olive green tint, legs and tail turquoise, and flanks with pale yellow ocelli; (20) associated with interandean dry forest. Comparison with other species. Ameiva reticulata shares the characters described by Harvey et al. (2012) for the A. ameiva complex (e.g. A. ameiva, A. atrigularis, A. pantherina, and A. praesignis): frontal entire, its posterior suture usually contacting second supraocular or aligned with suture between second and third supraoculars; frontal ridge absent; interparietal entire; parietal series (including interparietal) composed of five relatively long plates; narial suture passes through nostril; first supralabial usually curved; first subocular usually separated from supralabials by anterior expansion of the second subocular; intertympanic crease present; pectoral sulcus absent; ventral scales in 10���12 longitudinal rows; plate-like antebrachials continuous with or narrowly separated from brachial scales; combined femoral and abdominal pores number 28���45; fifth toe reduced; complete row of scales separate supradigital and subdigital lamellae along postaxial edge of each toe; tip of snout of adult males never reddish; adults without dorsolateral and vertebral stripes, or vertebral stripe only apparent on posterior dorsum; flanks with pale ocelli. Within the Ameiva ameiva complex, A. reticulata most resembles A. ameiva, a species widely distributed throughout Amazonia (Ugueto & Harvey 2011), and differs from it as follows (characteristics of A. ameiva in parentheses): maximum SVL in males 149 mm (163 mm) and in females 129 mm (146 mm); frontal usually pentagonal (hexagonal or roughly pentagonal in some specimens); fourth supraocular separated from parietals by one or two rows of circumorbital scales (up to four rows of circumorbital scales); 17���24 scales laterally between supraoculars and supraciliaries laterally (19���31 scales); ventrals in eight longitudinal rows (ten longitudinal rows); 181���237, mean= 199.9 �� 36.52, dorsal scales along the middorsal line (209���287, mean = 252.6 �� 16.65); 102���137, mean = 123.4 �� 8.24 dorsal scales at midbody in a transverse row (135���175, mean = 154.3 �� 8.8); 19���25, mean = 22.27 �� 1.56 anterior gulars (18���32, mean = 25.6 �� 2.64); preanal shield with four rows of enlarged scales (5���7); in juveniles, dorsal surface brown or olive green with black reticulation and white pale ocelli on flanks (first third or half of dorsum green and rest brown, flanks dark brown without ocelli; Fig. 5 E); in adults, dorsal surface of body brown or greenish-brown with a soft black reticulation, flanks completely brown or brown fading into greenish or turquoise towards hind limbs and with ill-defined light ocelli, tail turquoise (first third or half of dorsum brown with dark brown flecks, dots or blotches and remaining part of dorsum and tail green; Fig. 5 F). From the other species of the Ameiva ameiva complex such as A. atrigularis, A. pantherina (both from Venezuela) and A. praesignis ( from Costa Rica, Panama and Colombia) (Ugueto & Harvey 2011), A. reticulata can be separated by having: a lower scale count with 181���237, mean = 252.6 dorsals along the middorsal line and 102��� 137, mean = 123.4 dorsals across the midbody in a transverse row (263���361, mean = 309 and 134���179, mean = 151.3 in A. atrigularis; 291���343, mean = 311.3 and 137���163, mean = 147.4 in A. pantherina; and 237���348, mean = 289.3 and 111���157, mean = 133 in A. praesignis). Ameiva atrigularis and A. praesignis are longer than A. reticulata, with a maximum SVL in males of 186 mm and 243 mm, respectively (149 mm in A. reticulata). Further, throat coloration in adults is dark gray or black in A. atrigularis, dark gray in A. pantherina, cream or blue in A. praesignis (white in A. reticulata). Other species of the genus Ameiva that occur in Peru such as A. aggerescusans, A. concolor and A. nodam, are all restricted to the upper Mara����n basin in northern Peru (Koch et al. 2013), and share a transversely divided frontal plate, while it is entire in A. reticulata. Description. In males maximum SVL is 149.1 mm (CORBIDI 9919) and maximum total length is 481.1 mm (CORBIDI 9919); in females maximum SVL is 129.1 mm (CORBIDI 10089) and maximum total length is 357.1 mm (CORBIDI 10089). Head pyramidal, 0.25���0.29 (0.27 �� 0.01, n = 16) times SVL in males; 0.23���0.30 (0.26 �� 0.016, n = 20) times SVL in females. Snout elongate, bluntly pointed; canthus rostralis distinct. Neck narrower than head and body. Body cylindrical. Limbs well developed; tibia 0.11���0.18 (0.15 �� 0.02, n = 16) times SVL in males; 0.12���0.16 (0.14 �� 0.01, n = 20) times SVL in females; foot 0.30���0.42 (0.37 �� 0.03, n = 16) times SVL in males; 0.28���0.41 (0.36 �� 0.03, n = 20) times SVL in females. Tail round in cross section, tapers toward tip, 1.93��� 2.50 (2.26 �� 0.19, n = 11) times SVL in males; 1.76���2.40 (2.14 �� 0.24, n = 13) times SVL in females. Rostral pentagonal, slightly wider than high, visible from above, bordered posteriorly by nasals, which form a short medial suture. Each nasal divided by an oblique suture. Nostril in lower part of suture and anterior to it, directed lateroposteriorly, slightly taller than long. Frontonasal octagonal, in contact with nasals, loreal and prefrontals. Prefrontals paired, roughly pentagonal, with a very long medial suture about twice as long as that between nasals, laterally in contact with loreal, first supraocular and first supraciliar. Frontal pentagonal (rectangular in CORBIDI 10082, CORBIDI 10092), longer than wide and wider anteriorly, its sutures with prefrontals form a wide angle, slightly straight, its sutures with frontoparietals almost forming a straight line; frontal laterally in contact with first and second supraocular. Pair of trapezoidal frontoparietals (divided into two posteriorly in CORBIDI 9918, 1007677, 10080, 10084, 10087, 10094 and 10095 and with a small scale accessory between frontal and frontoparietals in CORBIDI 0 9919, 10090, 10082, 10087, 10089 and 10094), wider than long and with a long medial suture; frontoparietals laterally in contact with second and third supraocular, and small circumorbital scales bordering fourth up to middle portion of third supraocular or beginning of second supraocular. Interparietal irregularly hexagonal, higher than wide, often narrower (but sometimes slightly wider) than adjacent parietals; interparietal longitudinally divided; sutures with parietals slightly oblique and curved; interparietal bordered at each side by two large, irregular parietals divided by an oblique suture (parietals divided into two in CORBIDI 10078); outermost parietal oval in shape and larger than inner parietal. Parietal series composed of five scales including interparietal. Occipitals 8���17 (13.61 �� 2.25, n = 36), irregular and heterogeneous in size; occipitals usually subequal in size to first dorsal row, less often moderately to distinctly larger. Supraoculars four at each side. Circumorbital semicircles formed by 4���8 scales at each side, 9���15 (11.25 �� 1.46, n = 36) combining both sides, reaching middle, posterior and anterior portion of third and second supraocular (extending completely around supraoculars except first supraocular); only one scale row between third supraocular and prefrontals; fourth supraocular separated from parietals by one or two rows of circumorbital scales. Laterally, all supraoculars except first separated from supraciliaries by a single row of small rectangular scales; 17���24 (20.72 �� 1.59, n = 36) combining both sides. Supraciliaries five or seven (on one side), first highest, second longest. Loreal very large, trapezoidal and single, in contact with nasal, frontonasal, first supraciliary, first and second subocular, and third and fourth supralabial. Suboculars four; first subocular in preocular position, irregularly trapezoidal, longer than wide and slightly wider than second subocular; in contact with loreal, first supraciliary, small scales in ocular region, and second subocular; margin with loreal slightly curved or straight. Third subocular longest, all, except first, in contact with supralabials. Continuous keel from first subocular through entire length of third subocular. Postoculars small, approximately in one row of four or five irregularly trapezoidal scales. Palpebral disc opaque with enlarged scales on lower eyelid. Supralabials seven, sixth below center of eye. Row of enlarged supratemporals decreasing in size posteriad. Temporal region with polygonal scales, slightly smaller centrally than peripherally. External auditory meatus large, vertically oval, bordered by granular scales; anterior margin semicircular, posterior one straight. Tympanum recessed. All dorsal and lateral head scales juxtaposed and smooth. Symphysal anteriorly ellipsoid, its posterior sutures forming wide angles with infralabials and postsymphysal. Postsymphysal single and pentagonal; in contact with first and second infralabial, followed by five enlarged chinshields. First pair in ample medial contact, only the first in contact with infralabials. Remaining chinshields separated from infralabials by one row of sublabials. Medial chin scales moderately small, elongate, convex, smooth, juxtaposed, subhexagonal, in slightly oblique rows, all subequal in size. Infralabials six, fifth below center of eye followed to commissure of mouth by smaller scales. Gular region divided into two areas: anterior region with polygonal and flat scales in slightly oblique rows that usually remain subequal or rarely grade to larger scales medially, delimited posteriorly by a line uniting the lower margins of the ear openings; middle anterior gular scales usually small, rarely moderately enlarged, 19���25 (22.27 �� 1.56, n = 36) scales along midline of chin from anteriormost to posteriormost anterior gular. Posterior gular region covered by smaller polygonal scales in transverse rows; posteriormost median gulars usually forming medial patch of moderately enlarged scales, 8���14 (10.72 �� 1.64, n = 36) scales midventrally from anteriormost posterior gular to antegular fold. Mesoptychial scales moderately enlarged (larger or, less often, subequal to largest anterior gulars), in about one or two rows, hexagonal, flat, smooth, and juxtaposed. Scales on nape and sides of neck similar to dorsals. Dorsals and scales on flanks granular (slightly larger on dorsum than laterally), round, smooth, subimbricate, in approximately transverse rows; 181���237 (199.9 �� 36.52, n = 36) scales between occipitals and base of tail. Scales around midbody (excluding ventrals) 102���137 (123.47 �� 8.24, n = 36). Ventrals large, smooth, rectangular (wider than long), imbricate, in eight longitudinal (at midbody) and 28���32 (30.08 �� 0.93, n = 36) transverse rows; transition between ventrals and scales on flanks sharp. Preanal shield with four rows of enlarged scales; pattern of terminal scales of the preanal plate somewhat variable; often large median scale followed posteriorly by two subequal scales. Preanal plate surrounded anteriorly and laterally by smaller scales; posteriorly by much smaller scales. Femoral pores in continuous row along each thigh, 16���20 pores on each leg, 32���40 (35.88 �� 2.13, n = 36) pores combining both legs; each pore surrounded by four scales. Scales on base of tail trapezoidal, smaller than ventrals, longer than wide, mostly keeled (smooth on base of tail on ventral surface). Caudal scales imbricate, in transverse rows, continuous around tail. Distal caudals longer and narrower, distinctly keeled, in transverse and approximately longitudinal rows. Arms with rows of very large, smooth, slightly imbricate, trapezoidal antebrachial scales on anterodorsal aspect of forearms and similar but smaller brachial scales on upper arms that extend almost to shoulder. Brachial row distinctly enlarged with smaller accessory row bordering upper margin and grading into granules posteriorly. Antebrachials and brachials in contact or separated by smaller scales at elbow. Dorsoposterior, posterior, and ventral aspect of arms with scales similar to dorsals, but slightly larger, except for a group (approximately one or two rows) of postbrachial scales on posterior aspect of upper arms, which are slightly more irregular. Legs with large, smooth, imbricate scales on anterior and ventral aspects of thighs, and ventral aspects of shanks. One row of large, trapezoidal scales anteriorly on thigh, gradually becoming smaller and irregular toward pores. On ventral aspect of shanks, four rows of large scales, anterior two trapezoidal, posterior one rhomboidal, decreasing in size from anterior toward posterior row. Elsewhere on legs scales similar to dorsals. Subdigital lamellae transversely enlarged and single, moderately to distinctly tuberculate towards base. Tubercles most prominent under base of third toe, lamellae of outer toe continuing to heel. On palms, lamellae of inner finger continuing to wrist as large scales, increasing in size toward it; single large scale on postaxial side, following line of fifth finger, at short distance from wrist. Lamellae under fourth finger 14���17 (15.83 �� 0.69, n = 36), under fourth toe 27���36 (31.41 �� 1.79, n = 36). Data of holotype. SVL 136.5 mm; head length 36.9 mm; tibia length 19.7 mm; foot length 45.5 mm; hand length 13.35 mm; tail length 285 mm; supralabials 7 / 7; infralabials 6 / 6; supraoculars 4 / 4; circumorbitals 5 / 5 extending to mid third supraocular; 10 / 11 scales between supraorbital and supraciliary scales; parietals 5; occipitals 16; occipitals larger than first dorsal row; 22 anterior gulars; mid anterior gulars small; 14 posterior gulars, midposterior gulars moderately enlarged; mesoptychial scales smaller than largest anterior gulars; dorsals between occipitals and base of tail 185; 120 scales around midbody (excluding ventrals); 8 longitudinal ventral rows; 30 transverse ventral rows; femoral pores 18 / 18; four scales forming preanal plate; 16 lamellae under fourth finger on right hand; 31 lamellae under fourth toe on right foot. Color in preservative. Young specimens with dorsal surface of head and dorsal part of body uniformly brownish with medium-sized black spots at each side of vertebral region (most conspicuous towards sacrum). Sides of head uniformly brownish. Some specimens with almost half of vertebral region skyblue or turquoise. Dorsolateral surface black with regular turquoise spots, extending to base of tail. Upper limbs brownish with some small black scales. Hind limbs black with turquoise spots or turquoise with black spots. Throat pale, chest, ventral aspect of arms and most of abdomen semi pale (with a bluish tint). Ventral aspect of legs, posteriormost portion of abdomen, anal and subcaudal regions whitish with some dark scales. Adult males have a turquoise brown dorsum with black reticulation or flecks (Fig. 3; more visible in some females: Fig. 4); these markings remain conspicuous along dorsum from nape to base of tail. Flanks with small, regular, slightly scattered turquoise ocelli with distinct black margins more or less arranged in vertical rows; ocelli never extend onto dorsum. Top and sides, Published as part of Landauro, Caroll Z., Garc��a-Bravo, Antonio & Venegas, Pablo J., 2015, An endemic new species of Ameiva (Squamata: Teiidae) from an isolated dry forest in southern Peru, pp. 387-400 in Zootaxa 3946 (3) on pages 388-397, DOI: 10.11646/zootaxa.3946.3.6, http://zenodo.org/record/238213, {"references":["Harvey, M. B., Ugueto, G. N. & Gutberlet, R. L. Jr. (2012) Review of Teiid Morphology with a Revised Taxonomy and Phylogeny of the Teiidae (Lepidosauria: Squamata). Zootaxa, 3459, 1 - 156.","Ugueto, G. N. & Harvey, M. B. (2011) Revision of Ameiva ameiva Linnaeus (Squamata: Teiidae) in Venezuela: Recognition of Four Species and Status of Introduced Populations in Southern Florida, USA. Herpetological Monographs, 25 (1), 113 - 170. http: // dx. doi. org / 10.1655 / herpmonographs-d- 10 - 00007.1","Koch, C., Venegas, P. J., Rodder, D., Flecks, M. & Bohme, W. (2013) Two new endemic species of Ameiva (Squamata: Teiidae) from the dry forest of northwestern Peru and additional information on Ameiva concolor Ruthven, 1924. Zootaxa, 3745 (2), 263 - 295. http: // dx. doi. org / 10.11646 / zootaxa. 3745.2.6"]}

Duellman, William E., Barley, Anthony J., Venegas, Pablo J. (2014): Cryptic species diversity in marsupial frogs (Anura: Hemiphractidae: Gastrotheca) in the Andes of northern Peru. Zootaxa 3768 (2): 159-177, DOI: http://dx.doi.org/10.11646/zootaxa.3768.2.4

Venegas, Pablo J., Echevarria, Lourdes Y., Alvarez, Silvana C. (2014): A new species of spiny-tailed iguanid lizard (Iguania: Stenocercus) from northwestern Peru. Zootaxa 3753 (1): 47-58, DOI: http://dx.doi.org/10.11646/zootaxa.3753.1.4

Pristimantis luscombei (Duellman and Lehr, 1995). (Figs. 1���2) Eleutherodactylus luscombei ��� Duellman and Mendelson, 1995, Univ. Kansas Sci. Bull., 55: 354. Holotype: MHNURP 120 (W.E. Duellman 59957), by original designation. Type locality: "study zone at 1.5 km N Teniente Lopez, (02�� 35 ��� 39.6 ��� S, 76 �� 06��� 55.0��� W, 312 m), Provincia Loreto, Departamento Loreto, Peru ". Eleutherodactylus (Eleutherodactylus) luscombei ��� Lynch and Duellman, 1997, Univ. Kansas Mus. Nat. Hist. Spec. Publ., 23: 227. Pristimantis luscombei ��� Heinicke, Duellman, and Hedges, 2007, Proc. Natl. Acad. Sci. USA, Suppl., 104: Table 2. Pristimantis luscombei ��� Hedges, Duellman, and Heinicke, 2008, Zootaxa, 1737: 128. Pristimantis luscombei ��� Duellman and Lehr, 2009, Nature und Tier Verlag: 190. Pristimantis luscombei ��� Ortega-Andrade and Valencia, 2010, Herpetology Notes, 3: 251���256. Pristimantis achuar ���New synonymy, Elmer and Cannatella, 2008, Zootaxa, 1784: 13. Holotype: QCAZ 25463, by original designation. Type locality: "Kapawi Jungle Lodge, Pastaza province, Ecuador (S 02�� 32.32 ���, W 76 �� 51.50 ���, altitude 239 m)". Remarks. A list of diagnostic characters, character variation, distribution and comparison with similar species in the Amazon basin for Pristimantis luscombei are provided by Elmer and Cannatella (2008) as P. achuar. Overall, P. luscombei is most similar to P. kichwarum, but is clearly distinguished form the latter by lacking a dark canthal stripe and by a blunter snout. Moreover, P. luscombei is slightly smaller in size (Elmer & Cannatella 2008). This species is known from several localities along the upper Amazon Basin in Morona Santiago and Pastaza provinces in Ecuador (17 localities), and Amazonas and Loreto Departments in Peru (28 localities). Except for the record of this species in Distrito R��o Santiago, Quebrada Kampankis (CORBIDI 11393, S 4.04308, W 77.54119, 1212 m a.s.l.), all localities correspond to lowland evergreen forest at elevations between 100 and 800 m a.s.l (Fig. 4). This area covers 437,500 km 2 along the northern part of the upper Amazon Basin. Lynch (1980 b) reported this species as P. ockendeni morph B (Elmer & Cannatella 2008), from a series collected by B. Malkin at Cusime, R��o Cusime, Morona Santiago province, southeastern Ecuador. We reviewed specimens studied by Lynch (1980 b) and additional material collected in a nearby locality from Ashuara village on r��o Macuma, Morona Santiago province. Among the 64 specimens studied, 74 % correspond to P. luscombei, whereas the rest are P. ki c hw ar u m (Appendix III). Furthermore, L��pez-Rojas et al. (2013) reported P. luscombei (as P. achuar) for western Brazil., Published as part of Ortega-Andrade, H. Mauricio & Venegas, Pablo J., 2014, A new synonym for Pristimantis luscombei (Duellman and Mendelson 1995) and the description of a new species of Pristimantis from the upper Amazon basin (Amphibia: Craugastoridae), pp. 31-57 in Zootaxa 3895 (1) on pages 34-35, DOI: 10.11646/zootaxa.3895.1.2, http://zenodo.org/record/287609, {"references":["Duellman, W. E. & Mendelson, J. (1995) Amphibians and reptiles from northern Departamento Loreto, Peru: Taxonomy and biogeography. The University of Kansas Science Bulletin, 55, 329 - 376.","Lynch, J. & Duellman, W. E. (1997) Frogs of Genus Eleutherodactylus (Leptodactylidae) in Western Ecuador: Systematic, ecology and biogeography. The University of Kansas Museum of Natural History. Special Publication N ° 23, Lawrence, Kansas, 236 pp.","Heinicke, M. P., Duellman, W. E. & Hedges, B. (2007) Major Caribbean and Central American frog faunas originated by ancient oceanic dispersal. PNAS, 104, 10092 - 10097. http: // dx. doi. org / 10.1073 / pnas. 0611051104","Hedges, S. B., Duellman, W. E. & Heinicke, M. P. (2008) New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 2008, 1 - 182.","Duellman, W. E. & Lehr, E. (2009) Terrestrial breeding frogs (Strabomantidae) in Peru. Nature und Tier Verlag, Munster, Germany, 382 pp.","Ortega-Andrade, H. M. & Valencia, J. (2010) First country records of Pristimantis luscombei (Duellman and Mendelson) and Syncope tridactyla (Duellman and Mendelson) in eastern lowlands of Ecuador (Amphibia: Anura: Strabomantidae, Microhylidae). Herpetology Notes, 3, 251 - 256.","Elmer, K. & Cannatella, D. (2008) Three new species of leaflitter frogs from the upper Amazon forests: cryptic diversity within Pristimantis \" ockendeni \" (Anura: Strabomantidae) in Ecuador. Zootaxa, 1784, 11 - 38.","Lynch, J. (1980 b) A taxonomic and distributional synopsis of the Amazonian frogs of the genus Eleutherodactylus. American Museum Novitates, 2696, 1 - 24.","Lopez-Rojas, J. J., Ramalho, W. P., da Silveira Suscuarana, M. & de Souza, M. B. (2013) Three new records of Pristimantis (Amphibia: Anura: Craugastoridae) for Brazil and a comment of the advertisement call of Pristimantis orcus. Check List, 9, 1548 - 1551."]}

Gastrotheca aratia new species Holotype: KU 212067, adult female, from 8 km (by road) NW of Cutervo, 6 ˚ 14 '00"S, 78 ˚ 51 ' 24 "W, 2560 m, Provincia Cutervo, Departamento Cajamarca, Peru, collected by J. J. Wiens on 27 February 1989. Paratypes: KU 212055 and KU 212057, adult males, from vicinity of Cutervo, 6 ˚ 22 '01"S, 78 ˚ 51 '00"W, 2620 m, Provincia Cutervo, Departamento Cajamarca, Peru, collected by F. M. Cuadros and J. J Wiens on 26 February 1989; KU 212060,and KU 212062 ��� 66, adult males, collected by J. J. Wiens on 28 February 1989. Referred specimens: KU 212056 and KU 212058 ��� 59, juveniles; MUSM 6188 ��� 72, adult males, and MUSM 6194 ��� 95, juveniles all from vicinity of Cutervo, 2620 m, Departamento Cajamarca, Peru, collected by M. E. Morrison and J. J. Wiens on 26���28 February 1989 l MCZ 5328 ��� 30, adult males, from Querocotillo, 6 ˚ 16 ' 26 "S, 79 ˚02' 16 "W, 2875 m, Provincia Cutervo, Departamento Cajamarca, Peru, collected by G. K. Noble in 1916. Diagnosis. A moderately large species���SVL 42.8���55.7 mm (x = 48.1 �� 4.6, n = 8 males), 56.8 (n = 1 female)���with (1) tibia length 40���51 % SVL, about same length as foot; (2) interorbital distance slightly less than width of upper eyelid; (3) skin on dorsum granular, smooth to slightly pustular, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus smooth; (7) Finger I shorter than II with discs slightly wider than digits; (8) fingers unwebbed; (9) webbing extending maximally to penultimate subarticular tubercle on Toe IV and nearly to penultimate subarticular tubercle on Toe V; (10) dorsum green or brown with darker paravertebral marks and inverted V-shaped mark posterior to sacrum; (11) head markings consisting of dark canthal stripe and pale labial stripe; (12) pale dorsolateral stripe usually present; (13) flanks and anterior and posterior surfaces of thighs lacking dark markings; (14) venter cream with dark flecks or spots; (15) brood pouch single, dorsal. Gastrotheca aratia most closely resembles four other species in northern Peru. In contrast to the much larger G. monticola, G. aratia lacks a pale supracloacal stripe (67 % of the adults examined) and black spots on the flanks and anterior and posterior surfaces of the thighs (88 % of the adults examined) (Table 1; Fig. 3). The slightly smaller G. aguaruna��� SVL 41.6���46.8 mm (x = 45.1 �� 2.17, n = 6 males), 49.4���50.8 (x = 50.1, n = 3 females)��� differs by having dark marks in the groin, green vocal sac, tarsal fold extending full length of tarsus, and webbing extending only to the antepenultimate subarticular tubercle on Toe IV. Gastrotheca dysprosita differs from G. aratia by having a green dorsum with a narrow, yellow middorsal stripe and green flanks with small yellow spots; furthermore G. dysprosita lacks a dark canthal stripe and pale labial stripe and has a granular tympanic annulus. Gastrotheca lateonota differs from G. a r at i a by having a truncate snout in profile, smooth skin on the dorsum, and a nearly uniform grayish brown venter. Six other species of Gastrotheca in the Andes in northern Peru include G. peruana, G. phalarosa, and G. phelloderma, all of which have strongly pustular skin on the dorsum. In G. ossilaginis, the skin on the skull is co-ossified with the underlying dermal bones. Additionally, G. abdita, differs by having an acuminate snout in dorsal view, and G. galeata that differs from all the rest by having a spatulate labium; the latter two also produce eggs that undergo direct development. Description of holotype. Adult female; body robust; SVL 56.8 mm; head slightly wider than long; snout rounded in dorsal view and bluntly rounded in profile, extending slightly beyond margin of lower lip; canthus rostralis acutely rounded in section; loreal region barely concave; lips rounded; top of head flat; interorbital distance 85 % of width of upper eyelid; internarial area flat; nostrils not protuberant at terminus of canthus rostralis posterior to level of anterior margin of lower jaw; diameter of eye slightly greater than its distance from nostril; tympanum round, separated from eye by distance about equal to length of tympanum; tympanic annulus distinct, smooth; supratympanic fold weak, extending from posterior corner of orbit to point posterior to tympanum. Arm robust; row of ulnar tubercles absent; hand moderately large; fingers short, unwebbed; discs moderately large, rounded, width of disc on Finger III equal to half length of tympanum; relative lengths of fingers I II 2 ��� 2 III 2��� 3 IV 3 ��� 1 V; subarticular tubercles small, rounded; supernumerary tubercles absent. Skin on dorsum and flanks graular; skin on throat, belly, and ventral surfaces of thighs granular, on other surfaces of smooth; cloacal tubercles and folds absent; pouch opening puckered; pouch filled with eggs. Dentigerous processes of the vomers inclined posteromedially, narrowly separated medially, between round choanae, bearing six teeth each. Coloration in preservative: The dorsum of the head, body, and limbs are dark gray. The flanks are dark brown. A dark brown canthal stripe is present; a white labial stripe borders the entire margin of the upper lip and extends posteriorly to the base of the forelimb. The posterior surfaces of the thighs are gray with faintly darker irregular markings. all ventral surfaces are dull tan with scattered small gray spots on the chest and belly. Color in life: The dorsum is green with faint darker green paravertebral marks originating on the upper eyelids and becoming indistinct on the anterior part of the body (Fig. 6 A). A narrow brown canthal stripe and a distinct white labial stripe are present. The postorbital region and the flanks are mottled green and tan. The iris is a silvery bronze with fine black flecks and reticulations. Measurements (in mm): SVL 56.8, tibia length 25.4, foot length 25.1, head length 18.8, head width 19.5 interorbital distance 5.3, eyelid width 6.3, internarial distance 3.4, eye���nostril 5.0, eye diameter 6.9, tympanum diameter 4.4, orbit���jaw 2.6, nostril���jaw 4.1, thumb length 10.7, third finger length 18.0, width of disc on third finger 2.2. Variation. There is little variation in size and proportions of the eight male paratypes (Table 2). The head width is slightly greater than the head length, except in KU 212062 in which the ratio is 1.12. Likewise, in all specimens the interorbital distance is 0.83���0.98 % (x = 0.91 %) and the width of the upper eyelid, and the diameter of the tympanum is 0.55���0.76 % (x = 0.65 %) of the diameter of the eye. Males have 4���5 (x = 4.8) teeth on each vomerine process; vocal slits extend posterolaterally from the base of the tongue. Calling males have unpigmented nuptial excrescences. In preservative, the dorsum is brown to pale gray with darker gray or brown markings consisting of broad paravertebral marks extending from the eyelids to the sacrum and a narrower V-shaped postsacral mark with the apex anteriorly. The dorsal surfaces of the limbs vary from no definite markings to transverse bars nearly as wide as the interspaces and as many as two on the forearm, four on the thigh, three on the shank, and two on the tarsus. A white labial stripe and dark brown canthal stripe and broad postorbital bar are present; in some specimens the postorbital bar extends to the middle of the flank. Three individuals have a narrow cream dorsolateral stripe extending from the margin of the upper eyelid to the groin. Two specimens have a discontinuous supracloacal white stripe. The venter is creamy tan with or without small dark brown flecks on the chest; the vocal sac in calling males is brown. Color in life was obtained from a single color photograph of an adult male (KU 212055) that shows a tan dorsum with brown markings (Fig. 6 B). A narrow pale cream dorsolateral stripe is barely evident, but the white labial stripe is evident. The iris is pale bronze with fine black reticulations. Distribution and ecology: Gastrotheca aratia is known from elevations of 2560���2875 m in the northern part of the Cordillera Occidental of the Andes in northwestern Peru (Fig. 5). The type locality, Cutervo is in the valley of the R��o Chatano, a tributary of the R��o Chamaya, which flows eastward into the R��o Mara����n. Much of the terrain in the immediate vicinity of Cutervo is cultivated. The holotype was turned up in a plowed field by day. Three males were calling from low bushes during a rain at night, and two young individuals were found in a cistern at night. All other specimens were extracted from terrestrial spiny bromeliads by day. Other anurans found in the vicinity of Cutervo include Hyloxalus elachyhistus, H. pulcherrimus, and Telmatobius latirostris. Three faded specimens (MCZ 5328 ��� 30) are from Querocotillo, a village northwest of Cutervo. Life history: A brooding female contains a large number of small eggs in the brood pouch. This indicates that eggs hatch as tadpoles that complete their development in ponds. Etymology. The specific name is derived from the Latin noun aratio, meaning plowed field. All specimens were found in, or at the edge of, plowed fields. Remarks. Gastrotheca aratia seems to be surviving in cultivated areas, but possible use of herbicides or chemical fertilizers would endanger the species. Thus, G. aratia should be categorized as Vulnerable., Published as part of Duellman, William E., Barley, Anthony J. & Venegas, Pablo J., 2014, Cryptic species diversity in marsupial frogs (Anura: Hemiphractidae: Gastrotheca) in the Andes of northern Peru, pp. 159-177 in Zootaxa 3768 (2) on pages 170-172, DOI: 10.11646/zootaxa.3768.2.4, http://zenodo.org/record/227977

Redefinition of Gastrotheca monticola As shown in the phylogenetic analysis, topotypic Gastrotheca monticola are in the same clade as those from Pomacochas, from where a large series is available for the following redefinition; however, there is considerable variation in dorsal coloration (Fig. 2). Also, the analysis shows that specimens from Ecuador are distinct; for these the name Gastrotheca lojana Parker is available. This species is being redefined by Carvajal-Endara et al. (in prep.) and, except for the sequence data, is not included herein. Diagnosis. A moderately large species���SVL 49.0��� 55.7 mm (x = 51.7 �� 2.62, n = 11 males), 48.0��� 66.3 mm (x = 58.4 �� 5.11, n = 16 females)���with (1) tibia length 47���55 % SVL, about same length as foot; (2) interorbital distance much greater than width of upper eyelid; (3) skin on dorsum shagreen to finely granular, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus smooth; (7) Finger I slightly shorter than II with discs slightly wider than digits; (8) fingers unwebbed; (9) webbing extending maximally to penultimate subarticular tubercle on Toes IV and V; (10) dorsum green or brown with darker middorsal mark extending to sacrum or paravertebral marks usually connected in occipital region; (11) head markings consisting of dark canthal stripe and pale labial stripe; (12) pale dorsolateral stripe usually absent; (13) flanks and anterior and posterior surfaces of thighs with dark markings; (14) venter cream with dark spots; (15) brood pouch single, dorsal. Gastrotheca monticola most closely resembles four other species in northern Peru. Gastrotheca aratia lacks black spots on the flanks and anterior and posterior surfaces of the thighs (Table 1); the nuptial excrescences are unpigmented. Additionally, the following phenotypic characters (although partially overlapping) can help distinguishing these species. Specimens of G. aratia usually lack a pale supracloacal stripe (absent in 67 % of specimens studied) and are smaller (SVLs of males = 42.8���55.7 mm). Gastrotheca aguaruna differs by being much smaller (SVLs of males = 41.6���46.8 mm) than G. monticola and also by lacking a pale supracloacal stripe and black spots on the flanks and anterior and posterior surfaces of the thighs, and by having a green vocal sac. Gastrotheca dysprosita is like G. monticola in having a bluntly rounded snout in profile but differs from G. monticola by having a green dorsum with a narrow, diffuse, yellow middorsal stripe and green flanks with small yellow spots; furthermore, G. dysprosita has coarsely granular skin on the dorsum and a granular tympanic annulus and lacks a dark canthal stripe and pale labial stripe (Fig. 2). Gastrotheca lateonota is like G. monticola in size and most proportions, but G. lateonota differs from G. monticola by having a truncate snout in profile, smooth skin on the dorsum, and a nearly uniform grayish brown venter. Six other species of Gastrotheca inhabit the Andes in northern Peru. Of these, G. peruana, G. phalarosa, and G. phelloderma have pustular skin on the dorsum. In G. ossilaginis the skin on the head is co-ossified with the underlying bones of the skull. Gastrotheca abdita differs by having an acuminate snout in dorsal view, and G. galeata differs by having a spatulate labium; the latter two species also produce eggs that undergo direct development. Gastrotheca monticola also is similar to G. litonedis Duellman and Hillis and G. lojana Parker in southern Ecuador; both of these species differ from G. monticola by having Fingers I and II equal in length, whereas Finger I is shorter than Finger II in G. monticola. Furthermore, G. litonedis has smooth skin on the dorsum and a uniformly pale grayish brown venter, in contrast to shagreen to granular skin on the dorsum and a cream venter with dark spots in G. monticola. Distribution and ecology. Gastrotheca monticola, as recognized here, is known from elevations of 1900��� 1960 m in the vicinity of the type locality in the valley of the R��o Huancabamba, which flows southward into the R��o Chamaya, a tributary of the R��o Mara����n. The species occurs at Ayabaca (2700 m) northwest of the type locality. Its range extends southeastward to several localities at elevations of 2360���3235 m in the northern part of the Cordillera Central in Departamento Amazonas. Individuals have been found in arboreal bromeliads by day; the species thrives in anthropogenic conditions, as was evident of individuals found at night on trees within the village of Pomacochas., Published as part of Duellman, William E., Barley, Anthony J. & Venegas, Pablo J., 2014, Cryptic species diversity in marsupial frogs (Anura: Hemiphractidae: Gastrotheca) in the Andes of northern Peru, pp. 159-177 in Zootaxa 3768 (2) on pages 162-164, DOI: 10.11646/zootaxa.3768.2.4, http://zenodo.org/record/227977

Ortega-Andrade, H. Mauricio, Venegas, Pablo J. (2014): A new synonym for Pristimantis luscombei (Duellman and Mendelson 1995) and the description of a new species of Pristimantis from the upper Amazon basin (Amphibia: Craugastoridae). Zootaxa 3895 (1): 31-57, DOI: http://dx.doi.org/10.11646/zootaxa.3895.1.2

Gastrotheca aguaruna new species Holotype: KU 212022, an adult female, from Molinapampa, 6 ˚ 10 ' 50 "S, 77 ˚ 37 '01"W, 2400 m, Provincia Chachapoyas, Departamento Amazonas, Peru, one of a series collected by S. S. Barantes, F. M. Cuadros, W. E. Duellman, M. E. Morrison, and J. J. Wiens on 26 January 1989. Paratypes: KU 212023 ���212025, adult males, 212026���212027, adult females, 212028 and 212030 ��� 31, adult males collected with the holotype, and KU 212021, adult male, from 5 km N Levanto, 6 ˚ 17 ' 59 "S, 77 ˚ 53 ' 55 "W, 2850 m, Provincia Chachapoyas, Departamento Amazonas, Peru, collected by W. E. Duellman on 25 January 1989. Referred specimens: CORBIDI 380, adult female, from ACP Huiquilla, 6 ˚ 22 ' 45 "S, 77 ˚ 58 ' 42 "W, 2935 m, Provincia Luya, Departamento Amazonas, Peru, collected by G. Ch��vez 1 February 2008. CORBIDI 592, adult female, from Camino Leimebamba���Los Chilchos, 6 ˚ 39 ' 51.2 "S, 77 ˚ 41 ' 23.9 "W, 3240 m, Provincia Mariscal Caceres, Departamento San Mart��n, Peru, collected by P. J. Venegas on 23 January 2008; CORBIDI 10933, adult male, from Tragadero���Las Pi��as, 6 ˚ 39 ' 45.5 "S, 77 ˚ 44 ' 28.8 "W, 3235 m, Provincia San Mart��n, Departamento San Mart��n, Peru, collected by P. J. Venegas and L. Echevarr��a on 10 May 2012; CORBIDI 11747, 11752, 11763, 11767, 11770, adult males, 11773, juvenile, 11774, adult male, 11788 ��� 89, adult females, 11792 ��� 93, adult males, 11794, adult female from Hornillo, 6 ˚08' 30 "S, 77 ˚ 29 '04.9"W, 3308 m, Distrito de Vista Alegre, Provincia Rodr��guez de Mendoza, Departamento Amazonas, Peru, collected by P. J. Venegas and V. Duran on 23 August��� 2 September 2012; KU 138238 ��� 40, adult males, and 128241, juvenile, from Chachapoyas, 6 �� 12 ' 63 "S, 77 �� 51 ' 22 " W, 2340 m, Provincia Chachapoyas, Departamento Amazonas, Peru, collected by T. H. and P. R. Fritts on 1 May 1970; KU 181741, juvenile, from 20.5 km (by road) WSW of Leimebamba, 6 �� 55 ' 40 "S, 77 �� 54 ' 48 " W, 3220 m, Provincia Chachapoyas, Departamento Amazonas, Peru, collected by D. C. Cannatella on 6 March 1979; KU 212029, adult female (skeleton), MUSM 6116 ��� 21, adult males and MUSM 6292, juvenile, all from the type locality, collected with the holotype; KU 215627, juvenile, from Chachapoyas, 6 �� 12 ' 63 "S, 77 �� 51 ' 22 " W, 2360 m, Provincia Chachapoyas, Departamento Amazonas, Peru, collected by F. M. Cuadros and J. J. Wiens on 25 January 1989. Diagnosis. A moderately large species (SVL 41.6���46.8 mm in males, 49.4���50.8 mm in females) with (1) tibia length 45���54 % SVL, about same length as foot; (2) interorbital distance about equal to width of upper eyelid; (3) skin on dorsum weakly granular to smooth, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus smooth; (7) Finger I> II, with discs slightly wider than digits; (8) fingers unwebbed; (9) webbing extending maximally to antepenultimate subarticular tubercle on Toe IV and nearly to penultimate subarticular tubercle on Toe V; (10) dorsum green with or without faintly darker paravertebral marks; (11) head markings consisting of a pale labial stripe and dark canthal stripe in some; (12) pale dorsolateral stripe present or not; (13) flanks and anterior surfaces of thighs lacking dark markings, but black spots or reticulations present in groin; (14) venter cream with dark flecks or spots; (15) brood pouch single, dorsal. Gastrotheca aguaruna most closely resembles four other species in northern Peru. In contrast to the much larger G. monticola��� SVL 49.0��� 55.7 mm (x = 51.7 �� 2.62, n = 11 males), 48.0��� 66.3 mm (x = 58.4 �� 5.11, n = 16 females)��� G. aguaruna lacks a pale supracloacal stripe and black spots on the flanks and anterior surfaces of the thighs (Table 1; Fig. 3). The slightly larger G. aratia��� SVL 42.8���55.7 mm (x = 48.1 �� 4.6, n = 8 males), 56.8 (n = 1 female)���differs by lacking black marks in the groin, having unpigmented nuptial excrescences, a brown vocal sac, and by having slightly more webbing on the foot; the web extends to the penultimate subarticular tubercle of Toe IV. In addition, G. aratia differs from G. aguaruna by having a tarsal fold extending the full length of the tarsus. Gastrotheca dysprosita Duellman differs from G. aguaruna by having a green dorsum with a narrow, diffuse, yellow middorsal stripe, green flanks with small yellow spots, and a granular tympanic annulus. Gastrotheca lateonota Duellman and differs from G. aguaruna by having a truncate snout in profile, smooth skin on the dorsum, and a nearly uniform grayish brown venter. Six other species of Gastrotheca in the Andes in northern Peru include G. peruana (Boulenger), G. phalarosa Duellman and Venegas, and G. phelloderma Lehr and Catenazzi, all of which have pustular skin on the dorsum. In G. ossilaginis Duellman and Venegas the skin on the skull is co-ossified with the underlying dermal bones. Gastrotheca abdita Duellman, differs by having an acuminate snout in dorsal view. Lastly, G. galeata Trueb and Duellman differs from all the others by having a spatulate labium; the latter two also produce eggs that undergo direct development. The Ecuadorian G. pseustes Duellman and Hillis differs from G. aguaruna by having an anteroventrally inclined snout in profile and a pale supracloacal stripe. Description of holotype. Adult female; body robust; SVL 50.8 mm; head as wide as long; snout rounded in dorsal view and in profile, extending slightly beyond margin of lower lip; canthus rostralis acutely rounded in section; loreal region concave; lips rounded; top of head flat; interorbital distance equal to width of upper eyelid; internarial area flat; nostrils not protuberant near terminus of canthus rostralis, posterior to level of anterior margin of lower jaw; diameter of eye much greater than its distance from nostril; tympanum round, separated from eye by distance about one and a half times length of tympanum; tympanic annulus distinct (barely evident on left side), smooth; supratympanic fold moderately heavy, extending from posterior corner of orbit to point posterior to tympanum, obscuring upper edge of tympanum. Arm robust; row of ulnar tubercles absent; hand moderately large; fingers short, unwebbed; discs small, rounded, slightly wider than digits; width of disc on Finger III about equal to half length of tympanum; relative lengths of fingers I> II II 1��� 2 III 1.5��� 2 IV 2 ��� 1 V; subarticular tubercles small, rounded; supernumerary tubercles present on proximal segments of digits. Skin on dorsum and flanks weakly granular to smooth; skin on throat, belly, and ventral surfaces of thighs granular, on other surfaces of smooth; cloacal tubercles and folds absent; pouch opening puckered; pouch filled with eggs. Dentigerous processes of the vomers inclined posteromedially, narrowly separated medially, between round choanae, bearing five teeth each. Coloration in preservative: The dorsum of the head, body, and limbs are gray. A pair of slightly darker paravertebral marks are barely discernible. The flanks are dull brown; the groin is gray with irregular black markings. A fragmented, faint pale gray dorsolateral stripe is bordered below by a fragmented, narrow black stripe; these stripes extend from the thoracic region and the groin. A dark brown canthal stripe is present; a white labial stripe borders the entire margin of the upper lip and extends posteriorly to the base of the forelimb. The posterior surfaces of the thighs are gray with faintly darker small spots or irregular markings. All ventral surfaces are dull tan with small gray spots on the throat and chest. Coloration in life: No color notes were recorded for the holotype. The coloration is most similar to the individual in Figure 4 C. Measurements (in mm): SVL 50.8, tibia length 23.3, foot length 23.3, head length 14.8, head width 16.5 interorbital distance 5.0, eyelid width 5.0, internarial distance 3.1, eye���nostril 4.9, eye diameter 5.3, tympanum diameter 4.0, orbit���jaw 2.2, nostril���jaw 4.0, thumb length 10.3, third finger length 17.3, width of disc on third finger 2.3. G. aguaruna (6 ♂, 3 ♀) G. aratia (8 ♂, 1 ♀) Character Sex Mean Range SD Mean Range SD Snout-vent ♂ 45.1 41.6���46.8 2.17 * 48.1 42.8���55.7 4.6 length ♀ 50.1 49.4���50.8 ��� 56.8 ��� ��� Tibia length ♂ 21.8 20.0��� 22.8 1.17 * 22.4 19.7���27.7 2.72 ♀ 25.0 23.3���26.8 ��� 25.4 ��� ��� Foot length ♂ 22.0 20.1 ���24.0 1.39 22.4 20.1���26.3 2.33 ♀ 24.4 23.3���25.8 ��� 25.1 ��� ��� Head length ♂ 16.5 13.9���25.4 4.41 16.3 15.0���18.0 1.09 ♀ 15.8 14.8���16.6 ��� 18.8 ��� ��� Head width ♂ 17.4 14.8���26.4 4.43 16.8 14.6 ���19.0 1.54 ♀ 17.2 16.5 ���18.0 ��� 19.5 ��� ��� Thumb length ♂ 8.4 7.9���9.5 0.56 * 9.0 8.0��� 10.4 1.03 ♀ 9.9 9.3���10.3 ��� 10.7 ��� ��� Third finger ♂ 14.9 13.2���15.8 1.10 * 15.5 13.5���18.5 1.92 length ♀ 17.2 17.2���17.3 ��� 18.0 ��� ��� Interorbital ♂ 4.6 4.3���4.8 0.26 4.6 3.8���5.5 0.34 distance ♀ 5.1 5.0��� 5.3 ��� 5.3 ��� ��� Internarial ♂ 3.3 2.9���3.7 0.33 3.5 2.9���3.9 0.34 distance ♀ 2.9 2.7���3.1 ��� 3.4 ��� ��� Eye-nostril ♂ 4.0 3.8���4.2 0.13 4.4 3.5���5.8 0.74 ♀ 4.7 4.1 ���5.0 ��� 5.0 ��� ��� Eye diameter ♂ 4.7 4.3���5.5 0.43 *** 5.5 5.0��� 5.9 0.31 ♀ 5.4 5.3���5.5 ��� 6.9 ��� ��� Tympanum ♂ 2.9 2.5���3.3 0.33 ** 3.6 3.1���4.5 0.49 diameter ♀ 3.3 2.8 ���4.0 ��� 4.4 ��� ��� Eyelid width ♂ 4.4 4.1 ���5.0 0.34 5.0 4.4���5.9 0.60 ♀ 5.0 4.9 ���5.0 ��� 6.3 ��� ��� Orbit-jaw ♂ 2.1 1.9���2.3 0.15 ** 2.3 2.2���2.4 0.10 ♀ 2.4 2.2���2.6 ��� 2.6 ��� ��� Nostril-jaw ♂ 3.1 2.7���3.7 0.35 * 3.0 2.8���3.5 0.23 ♀ 3.9 3.8 ���4.0 ��� 4.1 ��� ��� Disc width ♂ 1.7 1.5���1.9 0.14 *** 2.1 1.8���2.5 0.23 ♀ 2.3 2.2���2.3 ��� 2.2 ��� ��� * Differences between means of sexes in G. aguaruna significant (ANOVA, PG aguaruna significant and differences between means for males of the two species significant (ANOVA, PVariation. The maximum SVL in six males is 46.7 mm and 50.8 mm in three females (Table 2). The basic dorsal color is green with or without creamy yellow labial and dorsolateral stripes. The dorsum is green with or without faintly distinct darker green or brown paravertebral marks (Fig. 4). Small black flecks are present on the dorsum in some individuals. There is a faint tinge of pale blue on the posterior surfaces of the thighs and in the groin, where small black marks are present. The venter is cream with or without black spots (Fig. 4); the vocal sac in calling males is pale green. The iris is a copper color. The distinctness of a dorsal pattern of a pair of paravertebral marks is shown in figure 3, in which no pattern is evident in KU 212021 (Fig. 4 A); a faint outline is present in KU 212022 (Fig. 4 B). The paravertebral marks are faint green, slightly darker than the rest of the dorsum in KU 212024 (Fig. 4 C); and distinct brown paravertebral marks are evident in KU 212025 (Fig. 4 D). Breeding males have small brown nuptial excrescences on the medial surface of the thumb; the vocal slits extend posterolaterally from the base of the tongue. In juveniles from the type locality, the dorsum is uniform green, green with darker green or brown paravertebral marks, or tan with dark brown paravertebral marks The venter is dirty white; the iris is copper. A juvenile from Chachapoyas had a tan dorsum with pale brown blotches edged with dark brown. A subadult found at the edge of a grassy pond 20 km WSW of Leimebamba had a tan dorsum with bright green markings; the venter was cream, and the labial stripe was creamy white. Distribution and ecology: Gastrotheca aguaruna is known from several localities at elevations of 2360���3308 m in the northern part of the Cordillera Central in Peru (Fig. 5). The type locality is in a valley where the frogs were found on bushes along, and on emergent vegetation in, a sluggish slowly moving marshy stream in a pasture by day. At Chachapoyas three individuals were found on low bushes, and under a clod of dirt by day. At the locality north of Levanto one adult was found under sod in a potato field in cloud forest. Hyloxalus leucophaeus and Scinax oreites were syntopic with G. aguaruna at the type locality; the latter and Pristimantis schultei were found in bromeliads at the locality where G. aguaruna was found north of Levanto. The referred specimen CORBIDI 380 from the ��rea de Conservaci��n Privada Huiquilla was in a small pond by day in humid puna above the tree line. CORBIDI 509 and 10933 from Camino Leimebamba-Los Chilchos were basking on leaves of terrestrial bromeliads by day in the humid puna. All specimens from Hornillo were in humid puna above the tree line. Gravid females (CORBIDI 11788 ��� 89 and 11794) were basking on the ground by day. At night, males were calling on bushes close to streams. Hornillo and Camino Leimebamba-Los Chilchos are in grassy regions dominated by bunch grass (Stipa) with scattered bushes, terrestrial bromeliads, and some scattered patches of elfin forest. In ACP Huiquilla and Camino Leimebamba-Los Chilchos, G. aguaruna was sympatric with Pristimantis melanogaster. At Hornillo, G. aguaruna was sympatric with G. phalarosa, G. phelloderma, and an unidentified species of Pristimantis of the P. orestes group. Life history: The large number of small eggs in the brood pouches of two brooding females are indicative of eggs that hatch as tadpoles that complete their development in ponds (Duellman et al. 2001). Metamorphosing young with tail stubs collected at the type locality substantiate this mode of life history. Etymology. The specific name is a noun in apposition. Aguaruna is the name of the indigenous people of the Andes of northern Peru including the Departamento Amazonas; they never were successfully conquered by the Inca. Remarks. This species was first discovered at Chachapoyas in May 1970 (KU 138238 ��� 41). Subsequently, a few green Gastrotheca were found throughout the geographic range of G. monticola in northern Peru. These specimens were considered to be a color variant of G. monticola until genetic data revealed that they were a separate species. This species apparently is abundant in agricultural areas, as well as in high Andean regions not suitable to agriculture. Thus, we suggest that Gastrotheca aguaruna should be categorized as Least Concern., Published as part of Duellman, William E., Barley, Anthony J. & Venegas, Pablo J., 2014, Cryptic species diversity in marsupial frogs (Anura: Hemiphractidae: Gastrotheca) in the Andes of northern Peru, pp. 159-177 in Zootaxa 3768 (2) on pages 164-170, DOI: 10.11646/zootaxa.3768.2.4, http://zenodo.org/record/227977, {"references":["Duellman, W. E., Lehr, E. & Aguilar, C. (2001) A new species of marsupial frog (Anura: Hylidae: Gastrotheca) from the Cordillera Azul in Peru. Scientific Papers Natural History Museum University of Kansas, 22, 1 - 10."]}

Stenocercus arndti sp. nov. (Figs. 1���4) Proposed standard English name: La Granja whorltail lizard Proposed standard Spanish name: Cap��n de La Granja Holotype. CORBIDI 0 1680 (Figs. 1���3), an adult male from Quebrada Checos (6 �� 21��12.63 ���� S, 79 ��06��41.15 ���� W), at 1997 m elevation, La Granja District, Chota Province, Cajamarca Region, Peru, collected by P.J. Venegas on 22 August 2008. Paratypes. CORBIDI 0 1681 and 0 1685, an adult male and adult female, respectively, collected with the holotype; CORBIDI 0 1682 and 0 1688, adult males; CORBIDI 0 1692, adult female; CORBIDI 0 1686, 0 1687, and 0 1690, subadult males; CORBIDI 0 1683 and 0 1689, subadult females; CORBIDI 0 1691, a juvenile male; CORBIDI 0 1684, a hatchling, all from Quebrada La Iraca, (6 �� 22��09.9���� S, 79 ��08��04.61���� W), at 2213 m elevation, La Granja District, Chota Province, Cajamarca Region, Peru, collected by P.J. Venegas on 9 August 2008. Referred specimens (photo vouchers). Two adult females from Ka��aris (6 ��03��26.18 ���� S, 79 �� 16��00.35���� W), at 2318 m elevation, Ferre��afe Province, Lambayeque Region, Peru, captured and released by P. J. Venegas on 25 May 2007. Diagnosis. Stenocercus arndti differs from all other species of Stenocercus, except for S. bolivarensis Castro & Ayala 1982, S. carrion Parker 1934, S. chlorostictus Cadle 1991, S. crassicaudatus Tschudi 1845, S. empetrus Fritts 1972, S. eunetopsis Calde 1991, S. simonsii Boulenger 1885, and S. torquatus Boulenger 1885, in having granular scales on the posterior surface of the thighs, two caudal whorls per autotomic segment, mucronate caudal scales, and a distinct longitudinal row of enlarged vertebral scales. However, the new species is easily distinguished from these species in having a bold black transverse band at midbody that extends ventrolaterally in adult males. Furthermore, S. carrioni, S. chlorostichus and S. euneptopsis differ from S. arndti (in parentheses) in having the dorsal scales of the neck keeled and imbricate (slightly keeled and subimbricate) and the dorsal scales of the trunk keeled (feebly keeled). Additionally, S. carrioni and S. euneptopsis have fewer scales at midbody (66���96, x = 82.43 �� 8.13 in S. carrioni and 60���80, x = 70.62 �� 5.38 in S. euneptopsis, versus 85���100, x = 91.54 �� 6.32 in the new species), and S. chlorostichus has a strong sexual dichromatism, with the dorsal background color green in males and brown in females (males and females gray or brown). Stenocercus empetrus differs from S. arndti in having a venter of yellowish-orange with black reticulations (white, pale gray, or pale orange without reticulations), caudal scales without strongly projected mucrons (strongly projected mucrons present), and attaining a larger size, with a maximum SVL= 103 mm in males and 90 mm in females (maximum SVL= 90 mm in males and 77 mm in females). Stenocercus simonsii differs from S. arndti in having the dorsal scales of the neck almost coarsely granular, while in the new species these scales are slightly keeled and subimbricate. Stenocercus crassicaudatus and S. torquatus differ from the new species in having the dorsal scales of the neck granular (slightly keeled and subimbricate) and more scales around the midbody (97���121, x = 108.87 �� 5.99 in S. crassicaudatus and 102���137, x = 116.96 �� 8.21 in S. torquatus, versus 85���100, x = 91.54 �� 6.32 in S. arndti). Moreover, S. torquatus differs from S. arndti in having a black antehumeral collar complete middorsally in adult males (incomplete), as well as two transverse bands anterior to the antehumeral collar (absent), and the ability to change color between emerald green and dark brown or gray (unable to change color). Stenocercus bolivarensis has strongly keeled and imbricate lateral body scales (Torres-Carvajal 2007 b), which are smooth in the new species, and fewer scales around the midbody (67���82, x = 73.08 �� 4.63 in S. bolivarensis, versus 85���100, x = 91.54 �� 6.32 in S. arndti). Characterization. (1) Maximum total length in males 90 mm (n = 7); (2) maximum total length in females 77 mm (n = 3); (3) vertebrals 74���94; (4) paravertebrals 87���108; (5) scales around midbody 85���100; (6) supraoculars 5���7; (7) internasals 3���4; (8) postrostrals 4; (9) loreals 2; (10) gulars 49���60; (11) lamellae on Finger IV 22���28; (12) lamellae on Toe IV 23���30; (13) posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007 b)]; (14) postfemoral mite pocket present as a distinct pocket slightly depth with a posteroventrally oriented slit-like opening [Type 2 of Torres-Carvajal (2007 b)]; (15) parietal eye absent; (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 absent; (21) antegular, antehumeral, gular, longitudinal, oblique, postauricular, supraauricular, and transverse antegular neck folds present; (22) lateral nuchals slightly 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 dorsals, forming inconspicuous longitudinal row between fore- and hind-limbs; (26) dorsolateral crests absent; (27) ventrals smooth, imbricate; (28) scales on posterior surfaces of thighs granular; (29) prefemoral fold present; (30) inguinal groove present; (31) preanals not projected; (32) tail not compressed laterally in adult males; (33) tail length 53���64 % of total length; (34) two caudal whorls per autotomic segment; (35) caudals spinose; (36) dark stripe that extends anterodorsally from subocular region to supraciliaries absent; (37) color pattern of gular region in adult females with clear marks, similar to ventral color pattern; (38) color pattern of gular region in adult males with clear marks; (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; (43) postxiphisternal inscriptional ribs not in contact midventrally (Pattern 2 B and 4 A of Torres-Carvajal 2004). Description of the holotype. Male (Figs. 1���3); SVL 79 mm; TL 101 mm; maximum head width 15.3 mm; head length 18.9 mm; head height 11.5 mm; scales on parietal and occipital regions small, slightly wrinkled, juxtaposed (Fig. 2); parietal eye not visible; supraoculars in five rows, smooth, with the lateral most three rows less than half the size of the medial adjacent rows; distinct circumorbitals present; canthals two; anterior-most canthal in contact with nasal; internasals four; postrostrals four, approximately as wide as long; supralabials seven; infralabials six; loreals three; lorilabials in one row; preocular divided into four scales, the two dorsals in contact with posterior canthal; lateral temporals granular; gulars in 49 rows between tympanic openings; all gulars cycloid, smooth, imbricate, apical pit absent; second infralabial in contact with the second and third sublabials; first pair of postmentals in contact medially; mental separated from the infralabials by the first pair of postmentals; dorsal and lateral scales of neck granular; lateral scales of body conspicuously smaller than dorsals, but not granular, slightly imbricate; scales around midbody 88; vertebrals enlarged, keeled, imbricate, in 74 rows, forming distinct vertebral row; paravertebrals adjacent to vertebral row slightly enlarged, keeled, and imbricate; paravertebrals 92; ventrals smooth, imbricate, more than twice the size of dorsals; preauricular fringe absent; antegular, antehumeral, gular, longitudinal, oblique, postauricular, supra-auricular, and transverse antegular neck folds present; ventrolateral and prefemoral folds present; dorsal scales of fore limbs imbricate, smooth; dorsal scales of hind limbs imbricate, keeled, mucronate; ventral humeral scales smooth and imbricate, about one third the size of the dorsal humeral scales; ventral scales of forearms and hind limbs imbricate, smooth; palmars and plantars imbricate, keeled; lamellae on Finger IV 26; lamellae on Toe IV 26; tail rounded; caudals strongly keeled, mucronate, imbricate; basal subcaudals smooth, imbricate; posthumeral mite pocket present as one or more vertical folds [Type 1 of Torres-Carvajal (2007 b)]; postfemoral mite pocket present as a distinct pocket slightly depth with a posteroventrally oriented slit-like opening [Type 2 of Torres-Carvajal (2007 b)]. Color in life of holotype (Fig. 3): head dorsal surface green, loreal region and snout bright green, temporal region and sides of head behind the eyes brown with white spots, thin dark brown temporal and two postocular stripes present; dorsum brown with several pale green spots, diffuse dark brown bands, and a bold black transversal band at midbody that extends ventrolaterally and is incomplete middorsally; black antehumeral collar present, incomplete middorsally; flanks brown with several white spots on sides of neck and pale green flecks on sides of trunk; tail brown; arms greenish brown and legs brown, both with dark brown flecks; throat greenish-white with diffuse brown spots, chest and belly white with brown borders and gray flecks; ventral surface of thighs and cloacal region cream. Variation. Measurements, scutellation, and other morphological characters of Stenocercus arndti are presented in Table 1. Loreals 1���5; supralabials 6���7; infralabials 5���7; second infralabials not in contact with third sublabials in 58 % of specimens; first pair of postmentals not in contact medially in 25 % of specimens. In two dissected specimens the postxiphisternal pairs of inscriptional ribs were two in one specimen (both pairs long), and in the other specimen (the first and second pair were long and the third pair short) (Pattern 2 B and 4 A of Torres-Carvajal 2004). CHARACTERS n = 13 Scales around midbody 85���100 (91.54 �� 6.32) Vertebrals 74���94 (83.23 �� 6.77) Paravertebrals 87���108 (100.23 �� 5.83) Gulars 49���60 (52.92 �� 3.65) Supraoculars 5���7 (6) Internasals 3���4 (4) Subdigitals Finger IV 22���28 (24.15 �� 2.03) Subdigitals Toe IV 23���30 (25.54 �� 1.94) Tail length/total length 0.53���0.64 (0.58 �� 0.08) Maximum SVL males 90 Maximum SVL females 77 Sexual dimorphism is evident in adult individuals. Males differ from females in having a conspicuous anthehumeral black collar and a bold black transversal band at midbody, whereas females only have several diffuse dark brown short transversal bands along the back. The other two collected adult males (CORBIDI 01681- 82 and 01688) and one subadult male (CORBIDI 01690) differ from the holotype only by having the head completely brown and the throat brown with white spots (Fig. 4: A & B). Subadult males (CORBIDI 01686- 87) and a juvenile male (CORBIDI 01691) differ from the aforementioned males in having the diagnostic mark on the dorsum dark brown, while is conspicuous black in adult males. In life, the adult female (CORBIDI 01692; Fig. 4: C & D) has a gray dorsum, while in the other collected females the dorsum was brown as in the males (Fig. 4: E & G). The throat of females was gray with white spots (CORBIDI 01692; Fig. 4 D), yellowish-green with gray spots (CORBIDI 01685; Fig. 4 F), or yellow with white spots (CORBIDI 01689; Fig. 4 H). The two referred specimens, two adult females from Ka��aris (Fig. 4: I���L), differ from the adult females from the type locality only in having the throat and chest yellow without white or gray spots on the throat (Fig. 4: J & L). Subadult females and the single hatchling collected have the transverse bands on the dorsum more evident than in the adult female, especially the anthehumeral black collar. Natural history observations. The habitat at the type locality and in the vicinity of Ka��aris are croplands with scattered large boulders and bushes. Croplands are embedded in a matrix of shorter-stature and drier forest. However, the forest has been almost removed and only some small patches of secondary forest remain close to ravines. All individuals of Stenocercus arndti were observed active on sunny days at between 10:00 and 15:00 h basking on large rocks, fallen tree trunks, and fences of rock or wood. Stenocercus arndti is sympatric with S. huancabambae and S. stigmosus in Quebrada La Iraca, and with only S. huancabambae in Quebrada Checos. Other sympatric species of squamate reptiles collected with the new species in the type locality were Chironius monticola, Dipsas peruana, Epictia teaguei, Liophis taeniurus, Micrurus peruvianus, and Pholidobolus sp. In the vicinity of Ka��aris, S. arndti was found sympatric only with Pholidobolus sp. Distribution. Stenocercus arndti is known only from two adjacent localities close to La Granja village in the interandean valley of the R��o Chotano and from one locality in the vicinity of Ka��aris village in the interandean valley of the R��o Huancabamba on the Amazonian slope of the northern portion of the Cordillera Occidental in northwestern Peru (Fig. 5). It occurs at elevations from 1997���2318 m in the regions of Cajamarca and Lambayeque. The type locality lies within the Yungas (500���2300 m) ecoregions according to Brack (1986) and Pe��aherrera del ��guila (1989). Etymology. The specific name is a patronym for Dr. Rudolf G. Arndt of Pomona, New Jersey, USA, in recognition of his financial support for the improvement of the herpetological collection of CORBIDI through the BIOPAT-Programme., Published as part of Venegas, Pablo J., Echevarria, Lourdes Y. & Alvarez, Silvana C., 2014, A new species of spiny-tailed iguanid lizard (Iguania: Stenocercus) from northwestern Peru, pp. 47-58 in Zootaxa 3753 (1) on pages 48-56, DOI: 10.11646/zootaxa.3753.1.4, http://zenodo.org/record/285378, {"references":["Cadle, J. E. (1991) Systematics of lizards of the genus Stenocercus (Iguania: Tropiduridae) from northern Peru: new species and comments on relationships and distribution patterns. Proceedings of the Academy of Natural Sciences of Philadelphia, 143, 1 - 96.","Fritts, T. H. (1972) New species of lizards of the genus Stenocercus from Peru (Sauria: Iguanidae). Occasional Papers of the Museum of Natural History, The University of Kansas, Lawrence, Kansas, 10, 1 - 21","Torres-Carvajal, O. (2004) The abdominal skeleton of tropidurid lizards (Squamata: Tropiduridae). Herpetologica, 60, 75 - 83.","Brack, A. (1986) Las Ecoregiones del Peru. Boletin de Lima, 44, 57 - 70.","Penaherrera del Aguila, C. (1989) Atlas del Peru. Instituto Geografico Nacional, Lima, Peru."]}

Pristimantis miktos sp. nov. (Figs. 2, 5) Eleutherodactylus luscombei ���In part. Univ. Kansas Sci. Bull., 55: 354. Eleutherodactylus (Eleutherodactylus) luscombei ��� Lynch and Duellman, 1997, Univ. Kansas Mus. Nat. Hist. Spec. Publ., 23: 227. Pristimantis luscombei ��� Heinicke, Duellman, and Hedges, 2007, Proc. Natl. Acad. Sci. USA, Suppl., 104: Table 2. Pristimantis luscombei ��� Hedges, Duellman, and Heinicke, 2008, Zootaxa, 1737: 128. Pristimantis luscombei ��� Duellman and Lehr, 2009, Nature und Tier Verlag: 190. Pristimantis luscombei ��� Ortega-Andrade and Valencia, 2010, Herpetology Notes, 3: 251���256. Holotype: QCAZ 53531, an adult female collected on 25 June 2012 at Juyuintza, S 2.110, W 76.190, 200 m altitude, by H. Mauricio Ortega-Andrade, Pastaza Province, Ecuador. Paratypes: ECUADOR: Orellana: Obe oriental, S 1.1, W 75.87, 212 m, collected by H. M. Ortega-Andrade on 22 August 2005, specimens DHMECN 3365 ��� 66; Parque Nacional Yasun��, S 0.99, W 76.25, 270 m, collected by J. Valencia, specimens FHGO 5252 ��� 54. Pastaza: Bufeo, S 2.19, W 76.79, 311 m, collected by H. M. Ortega-Andrade on 5 June 2007, specimen DHMECN 4448; Juyuintza, S 2.11, W 76.19, 192 m, collected by H. M. Ortega-Andrade and F. Timias on 25 June 2012, specimens QCAZ 53528 ��� 30; Lorocachi, S 1.63, W 75.97, 195 m, collected by O. Torres-Carvajal, M.C. Ter��n, X. Cisneros on 20 February 1996, specimen QCAZ 10131. PERU: Loreto: Andoas, S 2.35111, W 75.81622, 182 m, collected by V. Duran on 1 March 2008, specimens CORBIDI 4653, 4934, 4941, 4946, 4952, 4962, 5043; Curaray Paiche, Lote 66, S 2.01836, W 74.96976, 179 m, collected by G. Gagliardi, specimen GGU 663; Curaray Paiche, Lote 67, S 1.49719, W 75.39673, 190 m, collected by G. Gagliardi, specimen GGU 476, 508, 511, 536; Iquitos, S 3.75, W 73.25, 94 m, collected by L. N. Cotlow, specimen MZUNAP 989; Jibarito, S 2.73691, W 76.03007, 211 m, collected by G. Ch��vez on 12 September 2008, specimen CORBIDI 6562; Jibarito, S 2.73565, W 76.03178, 211 m, collected by G. Ch��vez on 14 July 2009, specimen CORBIDI 6588; Nanay, Lote 123, S 3.20457, W 74.71207, 154 m, collected by G. Gagliardi, specimens GGU 807-808; San Jacinto, S 2.33083, W 75.86369, 169 m, collected by A. Delgado on 1 September 2008, specimen CORBIDI 1183; 1202; Shiviyacu, S 2.48187, W 76.08565, 218 m, collected by A. Delgado on 7 July 2009, specimens CORBIDI 6429 - 30. Diagnosis. The new species is placed in the genus Pristimantis following our phylogenetic analyses (see above). Pristimantis luscombei is characterized by: (1) skin of dorsum shagreen with or without scattered low tubercles in females; males with dorsum shagreen with scattered tubercles or pustules, and a X-shaped scapular fold (sensu Duellman & Lehr 2009); dorsolateral folds absent; skin of belly areolate; discoidal fold barely evident; (2) tympanic annulus and membrane visible on skin, round, its length about two-thirds length of eye; (3) snout short, sub-acuminate in dorsal view, rounded in profile; lips not flared, canthus rostralis weakly concave in dorsal view, rounded in cross-section; (4) upper eyelid about 80 % of inter-orbital distance, bearing one small non-conical tubercle; (5) dentigerous processes of vomer narrow, oblique, bearing 2���3 small teeth; (6) males lack vocal slits, vocal sac; white nuptial excrescences are present on thumb; (7) fingers large and slender, first shorter than second; discs on outer fingers expanded, bluntly rounded, about 1.5 x the width of digit proximal to pad; supernumerary tubercles large, rounded; (8) fingers lacking lateral fringes; (9) forearm in females bearing a single low, non-conical ulnar tubercle; males bearing two or three small non-conical tubercles; (10) heel bearing a single sub-conical tubercle; outer border of tarsus shagreen in females, bearing three or four small sub-conical tubercles in males; inner border of tarsus smooth; (11) two metatarsal tubercles; inner elliptical, about 5 x the outer tubercle; supernumerary plantar tubercles absent or barely visible; (12) toes lacking lateral fringes; webbing absent; discs equal in size or slightly smaller than those on fingers; Toe V longer than Toe III; (13) in life, dorsum reddish���brown with some greenish orange stains in scapular region, with or without pale yellowish spots; snout with yellowish white reticulated blotches, upper lips with dark brown diagonal stripes; cream interorbital stripe or stains in some males. Groin yellowish-tan; anterior and posterior surfaces of thighs uniform brown. The belly is cream with minute brown flecks or brown mottling. Iris deep orange, finely reticulated with black. In preservation, all brown areas turn into dark brown to metallic green, with cream tubercles and dermal ridges. Dorsal surfaces of thighs with dark brown transversal stripes; anterior and posterior surfaces of thighs uniform brown. Snout cream, reticulated. Belly cream with minute brown flecks; (14) SVL in adult males 19.0�� 1.1 (17.7���21.3 mm); females with 27.9 �� 1.3 mm (26.7���29.2 mm). Differences with other species of Pristimantis in the upper Amazon Basin that have brownish dorsum and a differentiated tympanum are listed in Table 1. The new species is most similar to P. altamnis (Elmer & Cannatella 2008), P. d e l i us (Duellman & Mendelson 1995), P. l i br a r i us (Flores & Vigle 1994), P. luscombei (Duellman & Mendelson 1995), P. kichwarum (Elmer & Cannatella 2008), P. matidiktyo (Ortega-Andrade & Valencia 2012) and P. ockendeni (Boulenger 1912), being differentiated from all these species by having a deep orange iris, finely reticulated with black, easily distinguished in living specimens. In preserved specimens, the new species can be differentiated from P. luscombei, P. altamnis and P. kichwarum by lacking a thin W-shaped dermal ridge on scapular region. Also, from P. ki chw ar um by having a dark black canthal stripe, and by smaller size of P. miktos (females 23.6 �� 1.2 mm and males 17.9 �� 1.1 mm in P. k i ch w a r u m vs. females 28.6 �� 0.9 mm and males 19 �� 1.1 mm in P. m i k t os). Pristimantis altamnis is similar in size to P. m i k t os, but lacks ulnar tubercles and bears a weak tarsal fold. The skin texture is smooth in P. librarius and P. ockendeni, in contrast with the shagreen with scattered tubercles or pustules in the new species. Furthermore, both species have dorsal color pattern that includes inverted brown chevrons, in contrast with the reddish brown with or without pale yellowish spots or blotches in P. miktos. Also, P. ockendeni lacks tubercles on eyelids (present in the new species). Pristimantis miktos resembles P. delius and P. matidiktyo in having a shagreen dorsum, but it can be distinguished from those two species by lacking a tarsal fold and a by having a black canthal stripe. Furthermore, females of P. deli us commonly have a stripped dorsal pattern. Pristimantis miktos has tubercles or pustules on dorsum, which are absent in P. matidiktyo. Furthermore, the latter species presents a characteristic bicolored iris that is yellowish silver and is heavily reticulated with black and has a median dark reddish stripe. Other Pristimantis in the Upper Amazon Basin differ from the new species by lacking a differentiated tympanum (P. m a r t i a e, P. carvalhoi, P. quaquaversus, P. imitatrix, P. croceoinguinis, P. acuminatus, P. tantanti and P. academicus), by having contrasting colors in groin and hidden surfaces of limbs (P. diadematus, P. lythrodes, P. orcus, P. altamazonicus, P. divnae and P. eurydactylus), by having a greenish dorsum in life (P. pseudoacuminatus, and P. paululus), by having enlarged conical tubercles on eyelids (P. orphonolaimus) or by having a yellow interorbital bar and dorsolateral stripes (P. aureolineatus). Description of holotype (Fig. 5). Body slender; head wider than body; slightly longer than wide, about 40 % of SVL; snout short, sub-acuminate in dorsal view, rounded in profile; distance from nostril to corner of eye slightly shorter than diameter of eye; canthus rostralis weakly concave in dorsal view, rounded in cross-section; lips not flared; internarial area not depressed, nostrils slightly protuberant, directed anterolaterally, situated about threequarters the distance from the eyes to the tip of the snout; interorbital area flat, IOD 37 % of head width; eye large, protuberant, its diameter is about 2.5 x depth of lip below eye and about 32 % of head length; upper eyelid about 80 % of inter-orbital distance; bearing a single, small non-conical tubercle; no interocular fold; cranial crests, absent. Tympanic annulus and membrane visible on skin; prostrictal tubercles, low, barely visible in preservative; choana small, triangular, not concealed by the palatal shelf of maxillary arc; dentigerous processes of vomer narrow, oblique, bearing 2���3 small teeth, positioned posterior to level of choanae; tongue elliptical, posterior border notched, not adherent to the floor of the mouth in about 30 % of its length. Skin on dorsum shagreen with low granulated tubercles on flanks; no occipital ridges or dorsolateral folds; ventral surfaces of belly, chest, throat and thighs areolate; discoidal folds barely visible; no thoracic fold. Forearm slender; fingers large and slender, all with oval (broader than long) pads, Fingers III-IV with large pads, all fingers with large discs; pad on Finger III about 1.5 x wider than narrowest portion of penultimate phalanx; disc on Finger I distinctive smaller than those on other fingers; relative length of Fingers I Measurements (in mm) of holotype. Specimen QCAZ 53531 is an adult female with following measurements: SVL= 26.7; HL= 10.72; HW= 9.79; ED= 3.72; IOD= 3.63; EN= 3.1; TD= 1.16; TL= 12.62; FoL= 17.1. Proportions: HL/SVL= 0.4; HW/HL= 0.91; TL/SVL= 0.47; FoL/SVL= 0.64; EN/HL= 0.29; ED/HL= 0.35; IOD/ HW= 0.37. Variation. Measurements are listed on Table 2. Sexual dimorphism is evident in this species, males being smaller (19 �� 1.1 mm; 17.7���21.3 mm) than females (28.6 �� 0.9 mm; 27.9���29.2 mm). Furthermore males often have a more tuberculated dorsum, tarsus and heel, having a thick X-shaped dermal fold on scapular region (e.g. DHMECN 3365, QCAZ 53528, Fig. 6), and a pale yellow color in groin, compared with the shagreen dorsum and Measurements Pristimantis luscombei Pristimantis miktos sp. n��v. Females (N = 7) Males(N = 4) Females(N = 4) Males (N = 8) creamy tan groin in juveniles and females. Some specimens may present yellow dots on dorsum (e.g. QCAZ 53528, 53529), but the proportional occurrence of this variation is in about 6 % of the total specimens reviewed. In five paratypes, (CORBIDI 1202, 4941, 6562, 6588) the ventral coloration is paler (creamy white) with few scattered brown flecks or mottling; specimens CORBIDI 4946 and 6429 have the top of the snout and sides of the head pale brown and barely reticulated, differentiated from the well-defined cream reticulation on the snout of the other specimens. Coloration in life (Figs. 5, 6). The dorsal sides of the body are dark brown to tan with dark blotches and/or yellow spots, and with faint, dark transverse bars on the hind limbs. During the day the coloration turns darker. The dorsum in juveniles tends to be reddish brown. The snout has reticulated white blotches, and there are no canthal or postorbital stripes, but lips are barred with dark brown stripes below the eyes. A cream interorbital stripe is present in some males. The groin is pale yellow with the anterior and posterior surfaces of thighs uniform brown. The belly is cream, densely stippled with brown flecks. The iris is deep orange, finely reticulated and bordered around with black. Coloration in preservative. Dorsum dark brown to metallic green (e.g. DHMECN 4448), with cream tubercles and dermal ridges in males. Dorsal surfaces of thighs with dark brown transversal stripes; anterior and posterior surfaces of thighs uniform brown. Snout and interorbital stripe reticulated, pale cream. Belly cream with brown flecks. Darker coloration in preserved specimens could depend of the time in which each specimen was prepared as voucher. Etymology. The specific name is derived from the Greek word ��� miktos ��� (���mixed together, blended���) and refers to the fact that this species was mixed with the type specimens of another one. Natural history and distribution. Pristimantis miktos is known from five scattered localities along the Amazonian evergreen lowlands of Ecuador, in Orellana and Pastaza Provinces, and seven localities from Loreto Department in northern Peru, at elevations between 195 and 300 m a.s.l. The distribution area covers 44,196.7 km 2 in the upper Amazon basin (Fig. 4). This rare species is a forest inhabitant that can be found perching at night on leaves (0.3��� 1.5 m) such as those of heliconia or other low plants of terra firme forest. The mating call and reproductive behavior are unknown., Published as part of Ortega-Andrade, H. Mauricio & Venegas, Pablo J., 2014, A new synonym for Pristimantis luscombei (Duellman and Mendelson 1995) and the description of a new species of Pristimantis from the upper Amazon basin (Amphibia: Craugastoridae), pp. 31-57 in Zootaxa 3895 (1) on pages 35-45, DOI: 10.11646/zootaxa.3895.1.2, http://zenodo.org/record/287609, {"references":["Lynch, J. & Duellman, W. E. (1997) Frogs of Genus Eleutherodactylus (Leptodactylidae) in Western Ecuador: Systematic, ecology and biogeography. The University of Kansas Museum of Natural History. Special Publication N ° 23, Lawrence, Kansas, 236 pp.","Heinicke, M. P., Duellman, W. E. & Hedges, B. (2007) Major Caribbean and Central American frog faunas originated by ancient oceanic dispersal. PNAS, 104, 10092 - 10097. http: // dx. doi. org / 10.1073 / pnas. 0611051104","Hedges, S. B., Duellman, W. E. & Heinicke, M. P. (2008) New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 2008, 1 - 182.","Duellman, W. E. & Lehr, E. (2009) Terrestrial breeding frogs (Strabomantidae) in Peru. Nature und Tier Verlag, Munster, Germany, 382 pp.","Ortega-Andrade, H. M. & Valencia, J. (2010) First country records of Pristimantis luscombei (Duellman and Mendelson) and Syncope tridactyla (Duellman and Mendelson) in eastern lowlands of Ecuador (Amphibia: Anura: Strabomantidae, Microhylidae). Herpetology Notes, 3, 251 - 256.","Elmer, K. & Cannatella, D. (2008) Three new species of leaflitter frogs from the upper Amazon forests: cryptic diversity within Pristimantis \" ockendeni \" (Anura: Strabomantidae) in Ecuador. Zootaxa, 1784, 11 - 38.","Duellman, W. E. & Mendelson, J. (1995) Amphibians and reptiles from northern Departamento Loreto, Peru: Taxonomy and biogeography. The University of Kansas Science Bulletin, 55, 329 - 376.","Flores, G. & Vigle, G. O. (1994) A new Species of Eleutherodactylus (Anura: Leptodactylidae) from the Lowland Rainforest of Amazonian Ecuador, with notes on the Eleutherodactylus frater Assembly. Journal of Herpetology, 28, 416 - 424. http: // dx. doi. org / 10.2307 / 1564952","Ortega-Andrade, H. M. & Valencia, J. H. (2012) A new species of the Pristimantis frater Group (Anura: Strabomantidae) from the Eastern Evergreen Lowland Forests of Ecuador. Herpetologica, 68, 244 - 255. http: // dx. doi. org / 10.1655 / HERPETOLOGICA-D- 10 - 00066.1","Boulenger, G. A. (1912) Description of new Batrachians from the Andes of South America, preserved in the Brithish Museum. Annals and Magazine of Natural History, 8, 185 - 191. http: // dx. doi. org / 10.1080 / 00222931208693215"]}

Pristimantis miktos sp. nov. (Figs. 2, 5) Eleutherodactylus luscombei —In part. Univ. Kansas Sci. Bull., 55: 354. Eleutherodactylus (Eleutherodactylus) luscombei — Lynch and Duellman, 1997, Univ. Kansas Mus. Nat. Hist. Spec. Publ., 23: 227. Pristimantis luscombei — Heinicke, Duellman, and Hedges, 2007, Proc. Natl. Acad. Sci. USA, Suppl., 104: Table 2. Pristimantis luscombei — Hedges, Duellman, and Heinicke, 2008, Zootaxa, 1737: 128. Pristimantis luscombei — Duellman and Lehr, 2009, Nature und Tier Verlag: 190. Pristimantis luscombei — Ortega-Andrade and Valencia, 2010, Herpetology Notes, 3: 251–256. Holotype: QCAZ 53531, an adult female collected on 25 June 2012 at Juyuintza, S 2.110, W 76.190, 200 m altitude, by H. Mauricio Ortega-Andrade, Pastaza Province, Ecuador. Paratypes: ECUADOR: Orellana: Obe oriental, S 1.1, W 75.87, 212 m, collected by H. M. Ortega-Andrade on 22 August 2005, specimens DHMECN 3365 – 66; Parque Nacional Yasuní, S 0.99, W 76.25, 270 m, collected by J. Valencia, specimens FHGO 5252 – 54. Pastaza: Bufeo, S 2.19, W 76.79, 311 m, collected by H. M. Ortega-Andrade on 5 June 2007, specimen DHMECN 4448; Juyuintza, S 2.11, W 76.19, 192 m, collected by H. M. Ortega-Andrade and F. Timias on 25 June 2012, specimens QCAZ 53528 – 30; Lorocachi, S 1.63, W 75.97, 195 m, collected by O. Torres-Carvajal, M.C. Terán, X. Cisneros on 20 February 1996, specimen QCAZ 10131. PERU: Loreto: Andoas, S 2.35111, W 75.81622, 182 m, collected by V. Duran on 1 March 2008, specimens CORBIDI 4653, 4934, 4941, 4946, 4952, 4962, 5043; Curaray Paiche, Lote 66, S 2.01836, W 74.96976, 179 m, collected by G. Gagliardi, specimen GGU 663; Curaray Paiche, Lote 67, S 1.49719, W 75.39673, 190 m, collected by G. Gagliardi, specimen GGU 476, 508, 511, 536; Iquitos, S 3.75, W 73.25, 94 m, collected by L. N. Cotlow, specimen MZUNAP 989; Jibarito, S 2.73691, W 76.03007, 211 m, collected by G. Chávez on 12 September 2008, specimen CORBIDI 6562; Jibarito, S 2.73565, W 76.03178, 211 m, collected by G. Chávez on 14 July 2009, specimen CORBIDI 6588; Nanay, Lote 123, S 3.20457, W 74.71207, 154 m, collected by G. Gagliardi, specimens GGU 807-808; San Jacinto, S 2.33083, W 75.86369, 169 m, collected by A. Delgado on 1 September 2008, specimen CORBIDI 1183; 1202; Shiviyacu, S 2.48187, W 76.08565, 218 m, collected by A. Delgado on 7 July 2009, specimens CORBIDI 6429 - 30. Diagnosis. The new species is placed in the genus Pristimantis following our phylogenetic analyses (see above). Pristimantis luscombei is characterized by: (1) skin of dorsum shagreen with or without scattered low tubercles in females; males with dorsum shagreen with scattered tubercles or pustules, and a X-shaped scapular fold (sensu Duellman & Lehr 2009); dorsolateral folds absent; skin of belly areolate; discoidal fold barely evident; (2) tympanic annulus and membrane visible on skin, round, its length about two-thirds length of eye; (3) snout short, sub-acuminate in dorsal view, rounded in profile; lips not flared, canthus rostralis weakly concave in dorsal view, rounded in cross-section; (4) upper eyelid about 80 % of inter-orbital distance, bearing one small non-conical tubercle; (5) dentigerous processes of vomer narrow, oblique, bearing 2‒3 small teeth; (6) males lack vocal slits, vocal sac; white nuptial excrescences are present on thumb; (7) fingers large and slender, first shorter than second; discs on outer fingers expanded, bluntly rounded, about 1.5 x the width of digit proximal to pad; supernumerary tubercles large, rounded; (8) fingers lacking lateral fringes; (9) forearm in females bearing a single low, non-conical ulnar tubercle; males bearing two or three small non-conical tubercles; (10) heel bearing a single sub-conical tubercle; outer border of tarsus shagreen in females, bearing three or four small sub-conical tubercles in males; inner border of tarsus smooth; (11) two metatarsal tubercles; inner elliptical, about 5 x the outer tubercle; supernumerary plantar tubercles absent or barely visible; (12) toes lacking lateral fringes; webbing absent; discs equal in size or slightly smaller than those on fingers; Toe V longer than Toe III; (13) in life, dorsum reddish–brown with some greenish orange stains in scapular region, with or without pale yellowish spots; snout with yellowish white reticulated blotches, upper lips with dark brown diagonal stripes; cream interorbital stripe or stains in some males. Groin yellowish-tan; anterior and posterior surfaces of thighs uniform brown. The belly is cream with minute brown flecks or brown mottling. Iris deep orange, finely reticulated with black. In preservation, all brown areas turn into dark brown to metallic green, with cream tubercles and dermal ridges. Dorsal surfaces of thighs with dark brown transversal stripes; anterior and posterior surfaces of thighs uniform brown. Snout cream, reticulated. Belly cream with minute brown flecks; (14) SVL in adult males 19.0± 1.1 (17.7–21.3 mm); females with 27.9 ± 1.3 mm (26.7–29.2 mm). Differences with other species of Pristimantis in the upper Amazon Basin that have brownish dorsum and a differentiated tympanum are listed in Table 1. The new species is most similar to P. altamnis (Elmer & Cannatella 2008), P. d e l i us (Duellman & Mendelson 1995), P. l i br a r i us (Flores & Vigle 1994), P. luscombei (Duellman & Mendelson 1995), P. kichwarum (Elmer & Cannatella 2008), P. matidiktyo (Ortega-Andrade & Valencia 2012) and P. ockendeni (Boulenger 1912), being differentiated from all these species by having a deep orange iris, finely reticulated with black, easily distinguished in living specimens. In preserved specimens, the new species can be differentiated from P. luscombei, P. altamnis and P. kichwarum by lacking a thin W-shaped dermal ridge on scapular region. Also, from P. ki chw ar um by having a dark black canthal stripe, and by smaller size of P. miktos (females 23.6 ± 1.2 mm and males 17.9 ± 1.1 mm in P. k i ch w a r u m vs. females 28.6 ± 0.9 mm and males 19 ± 1.1 mm in P. m i k t os). Pristimantis altamnis is similar in size to P. m i k t os, but lacks ulnar tubercles and bears a weak tarsal fold. The skin texture is smooth in P. librarius and P. ockendeni, in contrast with the shagreen with scattered tubercles or pustules in the new species. Furthermore, both species have dorsal color pattern that includes inverted brown chevrons, in contrast with the reddish brown with or without pale yellowish spots or blotches in P. miktos. Also, P. ockendeni lacks tubercles on eyelids (present in the new species). Pristimantis miktos resembles P. delius and P. matidiktyo in having a shagreen dorsum, but it can be distinguished from those two species by lacking a tarsal fold and a by having a black canthal stripe. Furthermore, females of P. deli us commonly have a stripped dorsal pattern. Pristimantis miktos has tubercles or pustules on dorsum, which are absent in P. matidiktyo. Furthermore, the latter species presents a characteristic bicolored iris that is yellowish silver and is heavily reticulated with black and has a median dark reddish stripe. Other Pristimantis in the Upper Amazon Basin differ from the new species by lacking a differentiated tympanum (P. m a r t i a e, P. carvalhoi, P. quaquaversus, P. imitatrix, P. croceoinguinis, P. acuminatus, P. tantanti and P. academicus), by having contrasting colors in groin and hidden surfaces of limbs (P. diadematus, P. lythrodes, P. orcus, P. altamazonicus, P. divnae and P. eurydactylus), by having a greenish dorsum in life (P. pseudoacuminatus, and P. paululus), by having enlarged conical tubercles on eyelids (P. orphonolaimus) or by having a yellow interorbital bar and dorsolateral stripes (P. aureolineatus). Description of holotype (Fig. 5). Body slender; head wider than body; slightly longer than wide, about 40 % of SVL; snout short, sub-acuminate in dorsal view, rounded in profile; distance from nostril to corner of eye slightly shorter than diameter of eye; canthus rostralis weakly concave in dorsal view, rounded in cross-section; lips not flared; internarial area not depressed, nostrils slightly protuberant, directed anterolaterally, situated about threequarters the distance from the eyes to the tip of the snout; interorbital area flat, IOD 37 % of head width; eye large, protuberant, its diameter is about 2.5 x depth of lip below eye and about 32 % of head length; upper eyelid about 80 % of inter-orbital distance; bearing a single, small non-conical tubercle; no interocular fold; cranial crests, absent. Tympanic annulus and membrane visible on skin; prostrictal tubercles, low, barely visible in preservative; choana small, triangular, not concealed by the palatal shelf of maxillary arc; dentigerous processes of vomer narrow, oblique, bearing 2‒3 small teeth, positioned posterior to level of choanae; tongue elliptical, posterior border notched, not adherent to the floor of the mouth in about 30 % of its length. Skin on dorsum shagreen with low granulated tubercles on flanks; no occipital ridges or dorsolateral folds; ventral surfaces of belly, chest, throat and thighs areolate; discoidal folds barely visible; no thoracic fold. Forearm slender; fingers large and slender, all with oval (broader than long) pads, Fingers III-IV with large pads, all fingers with large discs; pad on Finger III about 1.5 x wider than narrowest portion of penultimate phalanx; disc on Finger I distinctive smaller than those on other fingers; relative length of Fingers I Hind limbs slender; tibia length in about 47 % of SVL; knee lacking tubercles; heel with a single, sub-conical tubercle; foot length in about 64 % of SVL; outer and inner border of tarsus shagreen; inner metatarsal tubercle oval, 5 x size of round outer; supernumerary tubercles barely visible, rounded, small; subarticular tubercles subconical, rounded; toes lacking lateral fringes; webbing absent between toes; pads of Toes III −V large, in all other pads and discs of toes like those of fingers; relative lengths I Measurements (in mm) of holotype. Specimen QCAZ 53531 is an adult female with following measurements: SVL= 26.7; HL= 10.72; HW= 9.79; ED= 3.72; IOD= 3.63; EN= 3.1; TD= 1.16; TL= 12.62; FoL= 17.1. Proportions: HL/SVL= 0.4; HW/HL= 0.91; TL/SVL= 0.47; FoL/SVL= 0.64; EN/HL= 0.29; ED/HL= 0.35; IOD/ HW= 0.37. Variation. Measurements are listed on Table 2. Sexual dimorphism is evident in this species, males being smaller (19 ± 1.1 mm; 17.7–21.3 mm) than females (28.6 ± 0.9 mm; 27.9–29.2 mm). Furthermore males often have a more tuberculated dorsum, tarsus and heel, having a thick X-shaped dermal fold on scapular region (e.g. DHMECN 3365, QCAZ 53528, Fig. 6), and a pale yellow color in groin, compared with the shagreen dorsum and Measurements Pristimantis luscombei Pristimantis miktos sp. nοv. Females (N = 7) Males(N = 4) Females(N = 4) Males (N = 8) creamy tan groin in juveniles and females. Some specimens may present yellow dots on dorsum (e.g. QCAZ 53528, 53529), but the proportional occurrence of this variation is in about 6 % of the total specimens reviewed. In five paratypes, (CORBIDI 1202, 4941, 6562, 6588) the ventral coloration is paler (creamy white) with few scattered brown flecks or mottling; specimens CORBIDI 4946 and 6429 have the top of the snout and sides of the head pale brown and barely reticulated, differentiated from the well-defined cream reticulation on the snout of the other specimens. Coloration in life (Figs. 5, 6). The dorsal sides of the body are dark brown to tan with dark blotches and/or yellow spots, and with faint, dark transverse bars on the hind limbs. During the day the coloration turns darker. The dorsum in juveniles tends to be reddish brown. The snout has reticulated white blotches, and there are no canthal or postorbital stripes, but lips are barred with dark brown stripes below the eyes. A cream interorbital stripe is present in some males. The groin is pale yellow with the anterior and posterior surfaces of thighs uniform brown. The belly is cream, densely stippled with brown flecks. The iris is deep orange, finely reticulated and bordered around with black. Coloration in preservative. Dorsum dark brown to metallic green (e.g. DHMECN 4448), with cream tubercles and dermal ridges in males. Dorsal surfaces of thighs with dark brown transversal stripes; anterior and posterior surfaces of thighs uniform brown. Snout and interorbital stripe reticulated, pale cream. Belly cream with brown flecks. Darker coloration in preserved specimens could depend of the time in which each specimen was prepared as voucher. Etymology. The specific name is derived from the Greek word “ miktos ” (“mixed together, blended”) and refers to the fact that this species was mixed with the type specimens of another one. Natural history and distribution. Pristimantis miktos is known from five scattered localities along the Amazonian evergreen lowlands of Ecuador, in Orellana and Pastaza Provinces, and seven localities from Loreto Department in northern Peru, at elevations between 195 and 300 m a.s.l. The distribution area covers 44,196.7 km 2 in the upper Amazon basin (Fig. 4). This rare species is a forest inhabitant that can be found perching at night on leaves (0.3– 1.5 m) such as those of heliconia or other low plants of terra firme forest. The mating call and reproductive behavior are unknown.

Gastrotheca aguaruna new species Holotype: KU 212022, an adult female, from Molinapampa, 6 ˚ 10 ' 50 "S, 77 ˚ 37 '01"W, 2400 m, Provincia Chachapoyas, Departamento Amazonas, Peru, one of a series collected by S. S. Barantes, F. M. Cuadros, W. E. Duellman, M. E. Morrison, and J. J. Wiens on 26 January 1989. Paratypes: KU 212023 –212025, adult males, 212026–212027, adult females, 212028 and 212030 – 31, adult males collected with the holotype, and KU 212021, adult male, from 5 km N Levanto, 6 ˚ 17 ' 59 "S, 77 ˚ 53 ' 55 "W, 2850 m, Provincia Chachapoyas, Departamento Amazonas, Peru, collected by W. E. Duellman on 25 January 1989. Referred specimens: CORBIDI 380, adult female, from ACP Huiquilla, 6 ˚ 22 ' 45 "S, 77 ˚ 58 ' 42 "W, 2935 m, Provincia Luya, Departamento Amazonas, Peru, collected by G. Chávez 1 February 2008. CORBIDI 592, adult female, from Camino Leimebamba–Los Chilchos, 6 ˚ 39 ' 51.2 "S, 77 ˚ 41 ' 23.9 "W, 3240 m, Provincia Mariscal Caceres, Departamento San Martín, Peru, collected by P. J. Venegas on 23 January 2008; CORBIDI 10933, adult male, from Tragadero–Las Piñas, 6 ˚ 39 ' 45.5 "S, 77 ˚ 44 ' 28.8 "W, 3235 m, Provincia San Martín, Departamento San Martín, Peru, collected by P. J. Venegas and L. Echevarría on 10 May 2012; CORBIDI 11747, 11752, 11763, 11767, 11770, adult males, 11773, juvenile, 11774, adult male, 11788 – 89, adult females, 11792 – 93, adult males, 11794, adult female from Hornillo, 6 ˚08' 30 "S, 77 ˚ 29 '04.9"W, 3308 m, Distrito de Vista Alegre, Provincia Rodríguez de Mendoza, Departamento Amazonas, Peru, collected by P. J. Venegas and V. Duran on 23 August– 2 September 2012; KU 138238 – 40, adult males, and 128241, juvenile, from Chachapoyas, 6 ° 12 ' 63 "S, 77 ° 51 ' 22 " W, 2340 m, Provincia Chachapoyas, Departamento Amazonas, Peru, collected by T. H. and P. R. Fritts on 1 May 1970; KU 181741, juvenile, from 20.5 km (by road) WSW of Leimebamba, 6 ° 55 ' 40 "S, 77 ° 54 ' 48 " W, 3220 m, Provincia Chachapoyas, Departamento Amazonas, Peru, collected by D. C. Cannatella on 6 March 1979; KU 212029, adult female (skeleton), MUSM 6116 – 21, adult males and MUSM 6292, juvenile, all from the type locality, collected with the holotype; KU 215627, juvenile, from Chachapoyas, 6 ° 12 ' 63 "S, 77 ° 51 ' 22 " W, 2360 m, Provincia Chachapoyas, Departamento Amazonas, Peru, collected by F. M. Cuadros and J. J. Wiens on 25 January 1989. Diagnosis. A moderately large species (SVL 41.6–46.8 mm in males, 49.4–50.8 mm in females) with (1) tibia length 45–54 % SVL, about same length as foot; (2) interorbital distance about equal to width of upper eyelid; (3) skin on dorsum weakly granular to smooth, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus smooth; (7) Finger I> II, with discs slightly wider than digits; (8) fingers unwebbed; (9) webbing extending maximally to antepenultimate subarticular tubercle on Toe IV and nearly to penultimate subarticular tubercle on Toe V; (10) dorsum green with or without faintly darker paravertebral marks; (11) head markings consisting of a pale labial stripe and dark canthal stripe in some; (12) pale dorsolateral stripe present or not; (13) flanks and anterior surfaces of thighs lacking dark markings, but black spots or reticulations present in groin; (14) venter cream with dark flecks or spots; (15) brood pouch single, dorsal. Gastrotheca aguaruna most closely resembles four other species in northern Peru. In contrast to the much larger G. monticola— SVL 49.0– 55.7 mm (x = 51.7 ± 2.62, n = 11 males), 48.0– 66.3 mm (x = 58.4 ± 5.11, n = 16 females)— G. aguaruna lacks a pale supracloacal stripe and black spots on the flanks and anterior surfaces of the thighs (Table 1; Fig. 3). The slightly larger G. aratia— SVL 42.8–55.7 mm (x = 48.1 ± 4.6, n = 8 males), 56.8 (n = 1 female)—differs by lacking black marks in the groin, having unpigmented nuptial excrescences, a brown vocal sac, and by having slightly more webbing on the foot; the web extends to the penultimate subarticular tubercle of Toe IV. In addition, G. aratia differs from G. aguaruna by having a tarsal fold extending the full length of the tarsus. Gastrotheca dysprosita Duellman differs from G. aguaruna by having a green dorsum with a narrow, diffuse, yellow middorsal stripe, green flanks with small yellow spots, and a granular tympanic annulus. Gastrotheca lateonota Duellman and differs from G. aguaruna by having a truncate snout in profile, smooth skin on the dorsum, and a nearly uniform grayish brown venter. Six other species of Gastrotheca in the Andes in northern Peru include G. peruana (Boulenger), G. phalarosa Duellman and Venegas, and G. phelloderma Lehr and Catenazzi, all of which have pustular skin on the dorsum. In G. ossilaginis Duellman and Venegas the skin on the skull is co-ossified with the underlying dermal bones. Gastrotheca abdita Duellman, differs by having an acuminate snout in dorsal view. Lastly, G. galeata Trueb and Duellman differs from all the others by having a spatulate labium; the latter two also produce eggs that undergo direct development. The Ecuadorian G. pseustes Duellman and Hillis differs from G. aguaruna by having an anteroventrally inclined snout in profile and a pale supracloacal stripe. Description of holotype. Adult female; body robust; SVL 50.8 mm; head as wide as long; snout rounded in dorsal view and in profile, extending slightly beyond margin of lower lip; canthus rostralis acutely rounded in section; loreal region concave; lips rounded; top of head flat; interorbital distance equal to width of upper eyelid; internarial area flat; nostrils not protuberant near terminus of canthus rostralis, posterior to level of anterior margin of lower jaw; diameter of eye much greater than its distance from nostril; tympanum round, separated from eye by distance about one and a half times length of tympanum; tympanic annulus distinct (barely evident on left side), smooth; supratympanic fold moderately heavy, extending from posterior corner of orbit to point posterior to tympanum, obscuring upper edge of tympanum. Arm robust; row of ulnar tubercles absent; hand moderately large; fingers short, unwebbed; discs small, rounded, slightly wider than digits; width of disc on Finger III about equal to half length of tympanum; relative lengths of fingers I> II II 1— 2 III 1.5— 2 IV 2 — 1 V; subarticular tubercles small, rounded; supernumerary tubercles present on proximal segments of digits. Skin on dorsum and flanks weakly granular to smooth; skin on throat, belly, and ventral surfaces of thighs granular, on other surfaces of smooth; cloacal tubercles and folds absent; pouch opening puckered; pouch filled with eggs. Dentigerous processes of the vomers inclined posteromedially, narrowly separated medially, between round choanae, bearing five teeth each. Coloration in preservative: The dorsum of the head, body, and limbs are gray. A pair of slightly darker paravertebral marks are barely discernible. The flanks are dull brown; the groin is gray with irregular black markings. A fragmented, faint pale gray dorsolateral stripe is bordered below by a fragmented, narrow black stripe; these stripes extend from the thoracic region and the groin. A dark brown canthal stripe is present; a white labial stripe borders the entire margin of the upper lip and extends posteriorly to the base of the forelimb. The posterior surfaces of the thighs are gray with faintly darker small spots or irregular markings. All ventral surfaces are dull tan with small gray spots on the throat and chest. Coloration in life: No color notes were recorded for the holotype. The coloration is most similar to the individual in Figure 4 C. Measurements (in mm): SVL 50.8, tibia length 23.3, foot length 23.3, head length 14.8, head width 16.5 interorbital distance 5.0, eyelid width 5.0, internarial distance 3.1, eye–nostril 4.9, eye diameter 5.3, tympanum diameter 4.0, orbit–jaw 2.2, nostril–jaw 4.0, thumb length 10.3, third finger length 17.3, width of disc on third finger 2.3. G. aguaruna (6 ♂, 3 ♀) G. aratia (8 ♂, 1 ♀) Character Sex Mean Range SD Mean Range SD Snout-vent ♂ 45.1 41.6–46.8 2.17 * 48.1 42.8–55.7 4.6 length ♀ 50.1 49.4–50.8 – 56.8 – – Tibia length ♂ 21.8 20.0– 22.8 1.17 * 22.4 19.7–27.7 2.72 ♀ 25.0 23.3–26.8 – 25.4 – – Foot length ♂ 22.0 20.1 –24.0 1.39 22.4 20.1–26.3 2.33 ♀ 24.4 23.3–25.8 – 25.1 – – Head length ♂ 16.5 13.9–25.4 4.41 16.3 15.0–18.0 1.09 ♀ 15.8 14.8–16.6 – 18.8 – – Head width ♂ 17.4 14.8–26.4 4.43 16.8 14.6 –19.0 1.54 ♀ 17.2 16.5 –18.0 – 19.5 – – Thumb length ♂ 8.4 7.9–9.5 0.56 * 9.0 8.0– 10.4 1.03 ♀ 9.9 9.3–10.3 – 10.7 – – Third finger ♂ 14.9 13.2–15.8 1.10 * 15.5 13.5–18.5 1.92 length ♀ 17.2 17.2–17.3 – 18.0 – – Interorbital ♂ 4.6 4.3–4.8 0.26 4.6 3.8–5.5 0.34 distance ♀ 5.1 5.0– 5.3 – 5.3 – – Internarial ♂ 3.3 2.9–3.7 0.33 3.5 2.9–3.9 0.34 distance ♀ 2.9 2.7–3.1 – 3.4 – – Eye-nostril ♂ 4.0 3.8–4.2 0.13 4.4 3.5–5.8 0.74 ♀ 4.7 4.1 –5.0 – 5.0 – – Eye diameter ♂ 4.7 4.3–5.5 0.43 *** 5.5 5.0– 5.9 0.31 ♀ 5.4 5.3–5.5 – 6.9 – – Tympanum ♂ 2.9 2.5–3.3 0.33 ** 3.6 3.1–4.5 0.49 diameter ♀ 3.3 2.8 –4.0 – 4.4 – – Eyelid width ♂ 4.4 4.1 –5.0 0.34 5.0 4.4–5.9 0.60 ♀ 5.0 4.9 –5.0 – 6.3 – – Orbit-jaw ♂ 2.1 1.9–2.3 0.15 ** 2.3 2.2–2.4 0.10 ♀ 2.4 2.2–2.6 – 2.6 – – Nostril-jaw ♂ 3.1 2.7–3.7 0.35 * 3.0 2.8–3.5 0.23 ♀ 3.9 3.8 –4.0 – 4.1 – – Disc width ♂ 1.7 1.5–1.9 0.14 *** 2.1 1.8–2.5 0.23 ♀ 2.3 2.2–2.3 – 2.2 – – * Differences between means of sexes in G. aguaruna significant (ANOVA, PG aguaruna significant and differences between means for males of the two species significant (ANOVA, P Variation. The maximum SVL in six males is 46.7 mm and 50.8 mm in three females (Table 2). The basic dorsal color is green with or without creamy yellow labial and dorsolateral stripes. The dorsum is green with or without faintly distinct darker green or brown paravertebral marks (Fig. 4). Small black flecks are present on the dorsum in some individuals. There is a faint tinge of pale blue on the posterior surfaces of the thighs and in the groin, where small black marks are present. The venter is cream with or without black spots (Fig. 4); the vocal sac in calling males is pale green. The iris is a copper color. The distinctness of a dorsal pattern of a pair of paravertebral marks is shown in figure 3, in which no pattern is evident in KU 212021 (Fig. 4 A); a faint outline is present in KU 212022 (Fig. 4 B). The paravertebral marks are faint green, slightly darker than the rest of the dorsum in KU 212024 (Fig. 4 C); and distinct brown paravertebral marks are evident in KU 212025 (Fig. 4 D). Breeding males have small brown nuptial excrescences on the medial surface of the thumb; the vocal slits extend posterolaterally from the base of the tongue. In juveniles from the type locality, the dorsum is uniform green, green with darker green or brown paravertebral marks, or tan with dark brown paravertebral marks The venter is dirty white; the iris is copper. A juvenile from Chachapoyas had a tan dorsum with pale brown blotches edged with dark brown. A subadult found at the edge of a grassy pond 20 km WSW of Leimebamba had a tan dorsum with bright green markings; the venter was cream, and the labial stripe was creamy white. Distribution and ecology: Gastrotheca aguaruna is known from several localities at elevations of 2360–3308 m in the northern part of the Cordillera Central in Peru (Fig. 5). The type locality is in a valley where the frogs were found on bushes along, and on emergent vegetation in, a sluggish slowly moving marshy stream in a pasture by day. At Chachapoyas three individuals were found on low bushes, and under a clod of dirt by day. At the locality north of Levanto one adult was found under sod in a potato field in cloud forest. Hyloxalus leucophaeus and Scinax oreites were syntopic with G. aguaruna at the type locality; the latter and Pristimantis schultei were found in bromeliads at the locality where G. aguaruna was found north of Levanto. The referred specimen CORBIDI 380 from the Área de Conservación Privada Huiquilla was in a small pond by day in humid puna above the tree line. CORBIDI 509 and 10933 from Camino Leimebamba-Los Chilchos were basking on leaves of terrestrial bromeliads by day in the humid puna. All specimens from Hornillo were in humid puna above the tree line. Gravid females (CORBIDI 11788 – 89 and 11794) were basking on the ground by day. At night, males were calling on bushes close to streams. Hornillo and Camino Leimebamba-Los Chilchos are in grassy regions dominated by bunch grass (Stipa) with scattered bushes, terrestrial bromeliads, and some scattered patches of elfin forest. In ACP Huiquilla and Camino Leimebamba-Los Chilchos, G. aguaruna was sympatric with Pristimantis melanogaster. At Hornillo, G. aguaruna was sympatric with G. phalarosa, G. phelloderma, and an unidentified species of Pristimantis of the P. orestes group. Life history: The large number of small eggs in the brood pouches of two brooding females are indicative of eggs that hatch as tadpoles that complete their development in ponds (Duellman et al. 2001). Metamorphosing young with tail stubs collected at the type locality substantiate this mode of life history. Etymology. The specific name is a noun in apposition. Aguaruna is the name of the indigenous people of the Andes of northern Peru including the Departamento Amazonas; they never were successfully conquered by the Inca. Remarks. This species was first discovered at Chachapoyas in May 1970 (KU 138238 – 41). Subsequently, a few green Gastrotheca were found throughout the geographic range of G. monticola in northern Peru. These specimens were considered to be a color variant of G. monticola until genetic data revealed that they were a separate species. This species apparently is abundant in agricultural areas, as well as in high Andean regions not suitable to agriculture. Thus, we suggest that Gastrotheca aguaruna should be categorized as Least Concern.

Stenocercus chinchaoensis sp. nov. (Figs. 1–5) Proposed standard English name: Chinchao whorltail lizard Proposed standard Spanish name: capón de Chinchao Holotype. CORBIDI 0 9024 (Figs. 1, 2), an adult male from Dos Aguas (9 ° 48´30 ´´ S, 75 ° 49´59.1 ´´ W), 1879 m, Distrito de Chinchao, Provincia de Huánuco, Región de Huánuco, Perú, collected by K. García-Burneo on 17 March 2011. Paratypes. CORBIDI 0 9023 and 0 9025, two adult males from Rinconada (9 ° 48´50.1 ´´ S 75 ° 47´18.2 ´´ W), 1766 m, Distrito de Chaglla, Provincia de Pachitea, Región de Huánuco, Peru, collected by K. García-Burneo on 29 March 2011; CORBIDI 09320- 22, an adult female, adult male, and juvenile female, respectively, collected from the same area as the holotype by P. J. Venegas and V. Duran on 8 July 2011. Diagnosis. Stenocercus chinchaoensis is distinguished from other species of Stenocercus (except S. boettgeri, S. haenschi, S. humeralis, and S. varius) by having granular scales on the posterior surface of the thighs, enlarged vertebrals, three caudal whorls per autotomic segment, a medially complete antegular fold, non-spinose caudals, and by males lacking a black transverse band on the ventral surface of the neck. In life, Stenocercus chinchaoensis is distinguished from these species by lacking yellow or pale green spots on the dorsum (Fig. 3), with the exception of S. haenschi, and with the ability to change colors between green or water green and grey. Furthermore, S. chinchaoensis has more scales around the midbody (104–107, x = 105.66) than S. boettgeri (79–104, x = 88.61), S. haenschi (57–64, x = 60.50), and S. varius (74–88, x = 82.35) (Torres-Carvajal 2007 b); S. chinchaoensis has lateral and dorsal nuchals similar in size (lateral nuchals less than half the size of dorsal nuchals in S. boettgeri, S. haenschi, and S. varius); S. chinchaoensis has the dorsal scales of the neck granular (keeled and imbricate in S. boettgeri, S. haenschi, and S. varius). In addition, males and females of S. boettgeri are larger (maximum SVL = 108 and 94 mm, respectively) than S. chinchaoensis (maximum SVL= 86 mm in males and 71 mm in females). Although S. chinchaoensis is similar in scutellation to S. humeralis, the new species can be easily distinguished from the latter by having the scales in the frontonasal region nearly equal in size to the scales in the occipitoparietal region, while in S. humeralis the scales on the frontonasal region are twice or three times longer than the scales on the occipitoparietal region. Another useful character to distinguish between S. humeralis and the new species is the type of postfemoral mite pocket: present in S. chinchaoensis as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007 b)], but present as a slitlike opening in S. humeralis [Type 2 of Torres-Carvajal (2007 b)]. Finally, S. humeralis differs in coloration from the new species by presenting black spots on the gular region in both males and females, however, the spots are diffuse gray or white in the new species. Characterization. (1) Maximum total length in males 86 mm (n= 4); (2) maximum total length in females 71 mm (n = 2); (3) vertebrals 79–90; (4) paravertebrals 117–139; (5) scales around midbody 104–107; (6) supraoculars 6; (7) internasals 4; (8) postrostrals 6; (9) loreals 3–5; (10) gulars 43–50; (11) lamellae on Finger IV 23–30; (12) lamellae on Toe IV 32 –34; (13) posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007 b)]; (14) postfemoral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007 b)]; (15) parietal eye absent; (16) occipital scales small, smooth, juxtaposed; (17) projecting angulate temporal absent; (18) enlarged supraoculars occupying most of supraocular region in one row absent; (19) scales on frontonasal region juxtaposed; (20) preauricular fringe short; (21) antegular, antehumeral, gular, longitudinal, oblique, postauricular, supra-auricular, and transverse antegular neck folds present; (22) lateral and dorsal nuchals similar in size; (23) posterior gulars cycloid, smooth, slightly inbricate, not notched; (24) lateral scales reduced in size, approximately half the size of dorsal body scales; (25) vertebrals slightly enlarged, forming inconspicuous longitudinal row between fore and hind limbs; (26) dorsolateral crests absent; (27) ventrals smooth, imbricate; (28) scales on posterior surfaces of thighs granular; (29) prefemoral fold present; (30) inguinal groove present; (31) preanals not projected; (32) tail not strongly compressed laterally in adult males; (33) tail length 61–64 % of total length; (34) three caudal whorls per autotomic segment; (35) caudals not spinose; (36) dark stripe extending anterodorsally from subocular region to supraciliaries absent; (37) color pattern of gular region in adult females with dark gray flecks and white spots, similar to ventral color pattern; (38) color pattern of gular region in adult males with gray flecks and white spots, similar to ventral color pattern; (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, turquoise or green; (43) postxiphisternal inscriptional ribs not in contact midventrally (Pattern 1 A, 1 B, and 2 C of Torres-Carvajal 2004). Description of holotype. Male (Figs. 1–2); SVL 73.2 mm; TL 130 mm; maximum head width 14.8 mm; head length 19.7 mm; head height 10.7 mm; posterior dorsal head scales small, smooth, juxtaposed; parietal eye not visible; supraoculars in six rows, smooth, slightly imbricate, with the most lateral three rows less than half the size of the medial adjacent rows; distinct circumorbitals absent; canthals two; internasals four; postrostrals six, approximately as wide as long; supralabials five; infralabials six; loreals three; lorilabials in one row; preocular divided into three scales, the dorsal-most in contact with posterior canthal; lateral temporals granular; gulars in 49 rows between tympanic openings; all gulars cycloid, smooth, slightly imbricate, not notched; second infralabial in contact with first two sublabials; first pair of postmentals in contact medially; mental without contact with first pair of infralabials but in contact with first pair of postmentals; dorsal and lateral scales of neck granular; dorsal scales of body imbricate, slightly keeled, becoming gradually granular toward flanks; scales around midbody 106; vertebrals enlarged, slightly keeled, imbricate, in 90 rows, forming distinct vertebral row; paravertebrals adjacent to vertebral row slightly enlarged, slightly keeled, and imbricate; paravertebrals 139; ventrals smooth, imbricate, more than twice the size of dorsals; preauricular fringe short, composed of three enlarged, posteriorly projected granular scales; antegular, antehumeral, gular, longitudinal, oblique, postauricular, supra-auricular, and transverse antegular neck folds present; ventrolateral and prefemoral folds present; dorsal scales of fore limbs imbricate, keeled; dorsal scales of hind limbs imbricate, strongly keeled, not mucronate; ventral humeral scales granular; ventral scales of forearms and hind limbs imbricate, smooth; palmars and plantars imbricate, keeled; lamellae on Finger IV 28; lamellae on Toe IV 33; tail rounded; caudals strongly keeled, slightly mucronate, imbricate dorsally; basal subcaudals smooth, imbricate; posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007 b)]; postfemoral mite pocket present as one or more vertical folds or ridges [Type 1 of Torres-Carvajal (2007 b)]. Color in life of holotype (based on digital photograph): Head and dorsum grey with a bold black mantle on the vertebral line becoming a zig-zag bold stripe from midbody to the base of tail; black antehumeral collar present; thin black postocular stripe present; venter white with pale gray spots in the throat. Variation. Measurements, scutellation, and other morphological characters of Stenocercus chinchaoensis are presented in Table 1. Loreals 3–5; supralabials 5–6; infralabials 5–6; second infralabials not in contact with third sublabials in all specimens; first pair of postmentals not in contact medially in 50 % of specimens. In three dissected specimens the postxiphisternal pairs of inscriptional ribs were two in two specimens (one with two long pairs and the second specimen with the first pair long and the second pair short) and three in one specimen (the first pair long and the two following pairs short) (Pattern 1 A, 1 B, and 2 C of Torres-Carvajal 2004, respectively). An adult male CORBIDI 0 9321 (Fig. 4) had the following coloration in life: head greenish brown with a thin black postocular stripe present; dorsum green, but mostly covered by wide transverse bold black marks arranged longitudinally over vertebral line; black antehumeral collar present; first third of tail sky blue; flanks and limbs green with scattered black flecks; throat white with pale grey spots; chest and belly white; ventral surface of thighs, pelvic region, vent, and base of tail brownish cream. This individual changed its coloration immediately after it was caught, replacing the green coloration by grey tones (Fig. 4 A). An adult female CORBIDI 0 9320 (Fig. 3 A–B) had the same dorsal color as the males, but with a grayish green tail dorsally. However, it had a diffuse middorsal black mantle with a pale vertebral stripe and tranverse black bars on the tail, and the flanks and dorsal surface of the limbs were sprinkled by dark and light flecks. After the change of coloration, the head of this individual turned green (Fig. 3 A). A juvenile female CORBIDI 0 9322 (Fig. 3 C–D) had a dorsal pattern of middorsal transverse bold black marks arranged longitudinally over the vertebral line and transverse black bars on the tail. Both sexes of Stenocercus chinchaoensis have the ability to change their dorsal background color from green or water green to grey. This ability was reported previously only in S. torquatus (Torres-Carvajal et al. 2005). Color change was observed immediately after capture, suggesting that it occurs as a response to stress as occurs in S. torquatus according to Torres-Carvajal et al. (2005). The green or water green coloration matches the color of the mosses than cover the tree trunks where the individuals were found (Fig. 5). Natural history. In a field trip to the type locality by PJV and VD, six individuals of Stenocercus chinchaoensis were observed active on sunny days between 1000 and 1400 h. Of these, three were found basking on house walls from 2–4 m above the ground, and three were basking on broad tree trunks between approximately 1–8 m above ground. These observations suggest that this species is mainly arboreal. The habitats at the type locality on both sides of the Huallaga River (villages of Dos Aguas and Rinconada) contain second-growth vegetation, coffee and citric plantations, and cattle pasture. S. chinchaoensis is sympatric with S. cupreus; S. prionotus occurs allopatrically at lower elevations (900 m) in the same region. Other sympatric species of squamate reptiles collected with S. chinchaoensis were Bothriopsis chloromelas, Dipsas peruana, Liophis janaleeae, Micrurus annellatus, and Philodryas sp. One adult female (CORBIDI 0 9320, SVL = 71mm) collected on 8 July contained two oviductal eggs, in the right oviduct, and four follicles. Distribution. Stenocercus chinchaoensis is known only from two adjacent localities (Dos Aguas and Rinconada) in the upper valley of the Río Huallaga on the Amazonian slope of the Andes in central Peru (Fig. 6). It occurs at elevations of 1700–1900 m in the Región Huánuco. The type locality lies within the Yungas (500–2300 m) ecoregion according to Brack (1986) and Peñaherrera del Águila (1989). Etymology. The specific name refers to the District of Chinchao in which the type locality is situated.

Venegas, Pablo J., Duran, Vilma, Garcia-Burneo, Karla (2013): A new species of arboreal iguanid lizard, genus Stenocercus (Squamata: Iguania), from central Peru. Zootaxa 3609 (3): 291-301, DOI: http://dx.doi.org/10.11646/zootaxa.3609.3.3

Ameiva concolor Ruthven 1924 Figs. 8���9 Ameiva bifrontata concolor Ruthven, Occ. Pap. Mus. Zool. 155: 3���6. ��� Terra typica: Paipoy, Rio Crisnejas, 24 km from Mara����n (elevation 1067 m), province of Cajamarca, Peru. ��� 1924 Ameiva bifrontata concolor��� Burt & Burt, Bulletin American Museum of Natural History, 61: 227���395. ��� 1931 Ameiva bifrontata concolor��� Peters & Donoso-Barros, Smithsonian Institution Press, Washington D.C. & London: 20. ��� 1970 Ameiva bifrontata concolor��� Peters & Donoso-Barros, Smithsonian Institution Press, Washington D.C. & London: 20. ��� 1986 Ameiva concolor��� Harvey et al., Zootaxa, 3459: 1���156. ��� 2012 Diagnosis. A medium-sized Ameiva that can be distinguished from all other mainland congeners by the following combination of characters: (1) maximum known SVL of 128 mm; (2) lacking longitudinal ridge on frontal scale; (3) frontal plate divided in two subequal scales; (4) postnasals separated from prefrontals by frontonasals; (5) parietal scales usually 5; (6) median gular scales scarcely enlarged; (7) enlarged median mesoptychial scales slightly larger than largest gulars; (8) gulars posterior to the interauricular crease smaller than anterior gulars; (9) nasal suture passes centrally through nostril; (10) rostral projecting beyond the nasal suture and contacting postnasal; (11) supranasals not contacting supralabials; (12) circumorbital semicircle occasionally extending to frontal suture; (13) 31���33 enlarged ventral scales between gular and vent; (14) 10���12 longitudinal rows of ventral plates, outermost often distinctly smaller; (15) 80���93 DOM; (16) 174���196 DL; (17) postbrachials not or hardly dilated; (18) 34���41 LFT; (19) 29���35 SCF; (20) 10���21 FP; (21) vertebral region in most specimens with a trace of a pale vertebral streak. Description and variation. Maximum known SVL in males 128 mm, maximum total length in males 404 mm (Holotype, UMMZ 59192); maximum SVL in females 85.8 mm, maximum total length in females 295.8 mm (ZFMK 91788). HL 0.26���0.32 (0.28 �� 0.02, n = 10) times SVL in both sexes; HH 0.11���0.14 (0.13 �� 0.1 n = 11) times SVL; HW 0.08���0.14 (0.11 �� 0.02, n = 10) times SVL; SL 0.59���0.80 (0.69 �� 0.08, n = 9) times the HL; ED 0.19���0.31 (0.24 �� 0.04, n = 9) times the HL; DSN 0.10���0.18 (0.13 �� 0.02, n = 9) times the HL; DNE 0.29���0.36 (0.31 �� 0.02, n = 9) times the HL; DEE 0.20���0.30 (0.23 �� 0.03, n = 9) times the HL. Tail round in cross section, tapering toward the tip; 2.25���2.67 (2.46 �� 0.16, n = 6) times SVL. Body cylindrical, AGL 0.40���0.54 (0.48 �� 0.04, n = 11) times SVL. Limbs well developed, FLL 0.33���0.44 (0.37 �� 0.04, n = 10) times SVL, HLL 0.63���0.87 (0.72 �� 0.02, n = 10) times SVL, TIL 0.15���0.21 (0.18 �� 0.02, n = 10) times SVL, foot 0.37���0.44 (0.4 �� 0.03, n = 10) times SVL. Snout elongate, bluntly pointed; canthus rostralis distinct. Rostral in posterior part acute-angled and bordered by supranasals, in anterior part laterally stretched, projecting beyond the nasal suture and in short contact with postnasal; about as wide as high; smooth, except for a short posterolateral suture; visible from above. Supranasals almost triangular, in short medial contact, not contacting supralabials, bordered posteriorly by rhomboidal or oval frontonasal. Postnasal almost triangular, in short contact with rostral and frontonasal and in broad contact with loreal and first and second supralabials, in some specimens even in short contact with third supralabial. Oblique nasal suture passing centrally through oval nostril. Prefrontals paired and roughly pentagonal, with a medial suture longer than that between supranasals; laterally in contact with loreal and first supraocular and in some specimens in short contact with first supraciliary. Frontal plate divided transversely in two subequal scales, the suture between both scales forming a straight line; anterior frontal pentagonal, laterally in contact with first and second supraoculars, distinctly larger than posterior frontal; posterior frontal almost quadrate or rectangular, laterally in contact with second and third supraoculars. Pair of trapezoidal or irregular pentagonal frontoparietals, longer than wide and with long medial suture, fused in one specimen (ZFMK 91790); laterally separated from third supraocular by small circumorbital scales. Parietal series composed of 3���5 scales, mostly 5 including interparietal; interparietal more or less rectangular or irregularly pentagonal, higher than wide in most specimens, as wide or slightly wider than adjacent parietals, sutures with parietals straight or slightly oblique; lateral parietals irregularly shaped divided by oblique suture. Supraoculars 3���5 (mostly four) at each side, second and third largest. Circumorbital semicircle formed by 14���20 scales at left side, 29���37 scales combining both sides, bordering fourth up to middle or anterior portion of third supraoculars, occasionally extending to frontal suture and thus contacting second supraocular and separating third supraocular from frontoparietals; bordering posterior edge of last or last two supraoculars, and separating them from parietals by 2���4 rows of circumorbital scales. Laterally, all except first supraocular entirely separated from supraciliaries by a single, occasionally double row of small scales; 21���28 combining both sides. Supraciliaries 6���10 (on one side, mostly seven), first highest, middle longest, remaining ones subequal, anterior supraciliaries normally longer than those in posterior part. Loreal very large, single, in contact with postnasal, frontonasal, prefrontal, first supraciliary, all preoculars, first subocular, third and fourth supralabials and occasionally in narrow contact with second and fifth supralabial and first supraocular. Preoculars 1���2, if two, first one much smaller; second almost similar in size or slightly larger than first supraciliary, but distinctly smaller than suboculars. Suboculars three, all in contact with supralabials, normally longer than wide, second subocular longest. A curved keel reaching from preocular through first and second subocular. Postoculars 3���5. Enlarged supralabials 5���8 (mostly seven) to below center of eye; followed to commissure of mouth by 4���9 small scales. Supratemporals almost not distinguishable from surrounding scales. Temporal region with polygonal or rounded scales, smaller and almost granular centrally. External auditory meatus large, more or less round, bordered by granular scales. Tympanum recessed. All dorsal and lateral head scales juxtaposed and smooth. Mental anteriorly ellipsoid, posteriorly straight, bordered by first infralabials and postmental. Postmental single and pentagonal, in contact with first and second infralabials, followed by 7���8 pairs of enlarged chinshields. First pair (and in few specimens also second pair) in contact with infralabials and in broad medial contact. Remaining chinshields separated from infralabials by one row of small scales, and separated medially by scales of anterior gular region. Medial chin scales moderately small, slightly convex, smooth, juxtaposed, oval or polygonal, all subequal in size. Enlarged infralabials 5���7 (mostly six) to below center of eye; followed to commissure by 4���8 smaller scales. Gular region divided into two areas: anterior region with round or polygonal and flat scales in slightly oblique rows that usually remain subequal in size, delimited posteriorly by line uniting lower margin of ear openings. Posterior gular region covered by smaller polygonal or round scales in transverse rows. Mesoptychial scales slightly enlarged, in about two indistinct rows, polygonal or hexagonal, flat, smooth, and juxtaposed. Scales on nape and sides of neck slightly smaller than dorsals. Dorsals and scales on flanks granular, round, smooth, juxtaposed; slightly larger in vertebral region; 174���196 (188 �� 6.88, n = 11) DL; 80���93 (84 �� 4.69, n = 10) DOM. Ventrals large, smooth, rectangular, wider than long, in 10���12 longitudinal and 31���33 (32 �� 0.89, n = 11) transverse rows, outermost ventrals nearly as wide as those of the adjacent row or distinctly smaller; transition between ventrals and scales on flanks sharp. Preanal shield centrally with several enlarged scales, surrounded anteriorly and laterally by smaller scales; posteriorly by much smaller scales. FP 10���21 (16 �� 4.45, n = 6), in males in a continuous row along each thigh, with short gap medially; not visible in female and juvenile specimens. Each pore surrounded by four scales. Dorsal scales on tail slightly imbricate, rectangular, smaller than subcaudals, longer than wide, with a slightly oblique keel; in transverse and oblique rows, continuous with subcaudals around tail (except first few rows); 29���35 (31 �� 2.31, n = 10) SCF. Subcaudals rectangular; smaller than ventrals; wider than long close to base, longer than wide in most parts of the tail; smooth, mostly juxtaposed. Forelimbs with row of very large, smooth, slightly imbricate, almost rectangular (distinctly wider than long) antebrachial scales on anterodorsal aspect of forearms and similar but smaller brachial scales on upper arms that extend almost to insertion of forelimbs. Antebrachials and brachials usually separated by smaller scales at elbow. Dorsoposterior, posterior, and ventral aspect of arms granular, equal in size to dorsals, except for two rows of rhomboidal scales directly adjacent to antebrachials and some moderately enlarged irregular, hexagonal scales adjacent to brachials. Legs with large, smooth, imbricate scales on anterior and ventral aspects of thighs, and ventral aspect of shanks. Ventral scales on thigh, gradually becoming smaller and irregular toward pores. On ventral aspect of shanks enlarged scales arranged in two to three rows, anterior largest, dilated, and more or less trapezoidal, posterior one or two rhomboidal, decreasing in size from anterior toward posterior row. Tibiotarsal spurs not distinguishable from other scales in most specimens. Elsewhere on hindlimbs scales similar to dorsals. Supradigital scales dilated, single and smooth. 17���20 (19 �� 0.97, n = 10) LFF; 34���41 (37 �� 2.34, n = 11) LFT. Lamellae of hand transversely enlarged, convex, and single, moderately to distinctly tubercular towards base. Lamellae of outer and inner fingers continuing to wrist and only separated by few granules, tubercular and increasing in size towards it. Lamellae of toes mostly single sometimes paired; becoming distinctly tubercular towards base of second, third and fourth toe; continuing to heel on outer toe. Coloration: No conspicuous difference in color pattern between males, females and juveniles present. In life, color on dorsum, head and dorsal surface of limbs light to moderately brown or brownish-olive; dorsum and head with or without some small dark spots; vertebral region in most specimens with a trace of a pale vertebral streak; sides of body brownish or bluish-gray and heavily mottled with tiny dark-brown or black spots, that are often merged together, resulting in dark body sides, with in most specimens a smooth transition from the lighter vertebral region; rostral cream-colored; sides of head light reddish-brown or brownish-olive, supralabials sometimes darkly edged; tail dorsally reddish-brown or moderately brown, lightening towards tip; head and body ventrally pale yellow, cream-white or grayish-white; plates of 3 outer longitudinal rows of ventrals darkly-grayish edged, some ventrals of the outermost row even almost completely dark gray; limbs and tail ventrally cream-white; tubercular lamellae of hands and feet accentuated with brown. In preservative, the general dorsal color is mainly grayish-brown or dark brownish-black; sides of the body are bluish���gray; the tail is grayish in anterior part fading posteriad into beige; head, body, limbs and tail ventrally pale yellowish, cream-white or grayish-white; edging of outer ventrals bluish-gray. Distribution and natural history. This species is endemic to the dry forest of the Canyons of the Crisnejas and Mara����n River in the Northern Peruvian Andes (Fig. 3) from the type locality Paipoy, Region of Cajamarca, 24 km from the Mara����n (1067 m, Ruthven 1924) to P��as (Laguna), Province of Tayabamba, Region of La Libertad (1720 m, Fig. 10). In addition to the locations from where we collected voucher specimens of A. concolor, we sighted this species in Vijus, Province of Pataz, Region of La Libertad, but did not collect voucher specimens. Ameiva concolor was found during daytime moving hectically on the ground in low vegetation. In P��as, the habitat was interspersed with big stones and rocks underneath which they quickly sheltered when being scared. During night time individuals were found sleeping under stones. Air temperature during the active hours of this species was between 39.3 ��C and 41.4 ��C and substrate temperature of the ground was between 33.9 ��C and 41.2 ��C. Similar to the other two Ameiva species described herein, this species was also found in the same microhabitat as Microlophus stolzmanni. Niche comparisons. The first two axes (PC 1 and PC 2) of the PCA-env explain together 91.86 % of the variation among the 19 bioclimatic variables in the analysis of A. aggerecusans versus A. concolor, 80.41 % in the analysis of A. aggerecusans versus A. nodam and 84.59 % in the analysis of A. concolor versus A. nodam. Correlation circles illustrating the contribution of each of the 19 bioclimatic variables to each PC are shown in Figure 11. For all three analysis the variables ���annual mean temperature��� (BIO 1), ���temperature annual range��� (BIO 7), ���mean temperature of the wettest quarter��� (BIO 8), ���mean temperature of the warmest quarter��� (BIO 10), ���mean temperature of the coldest quarter��� (BIO 11), ���annual precipitation��� (BIO 12), ���precipitation of the wettest month��� (BIO 13), and ���max temperature of warmest month��� (BIO 5) correlated negatively to PC 1, whereas ���temperature seasonality��� (BIO 4), ���mean temperature of the driest quarter��� (BIO 9), and ���min temperature of coldest month��� (BIO 6) correlated positively to PC 1. A. nodam seems to use the greatest environmental space of the three species and A. concolor the smallest but this observation may be related with a higher number of species records in A. nodam. There are a small differences in the climatic niches of A. aggerecusans and A. concolor in temperature related variables (BIO 1, BIO 7, BIO 8, BIO 10, BIO 11) and precipitation related variables (BIO 12, BIO 13) with A. aggerecusans occupying slightly warmer and wetter regions than A. concolor (Fig. 11 A). The differences in PC 2 between the niches of A. aggerecusans and A. concolor are only marginal. The climatic niche of A. aggerecusans is nested within the niche of A. nodam but there is a shift in the highest density of occurrences driven by precipitation and climatic seasonality (BIO 14, BIO 15, BIO 16, BIO 18, BIO 19, see appendix II for a detailed description of the variables) with A. nodam occupying more humid regions with a higher climatic seasonality (Fig. 11 B). The differences in PC 1 between the niches of A. aggerecusans and A. concolor are only marginal. The climatic niches of A. concolor and A. nodam differ in temperature related variables (BIO 1, BIO 7, BIO 8, BIO 10, BIO 11) and in precipitation related variables (BIO 12, BIO 13) with A. nodam occupying warmer and wetter regions than A. concolor (Fig. 11 C). The differences in PC 2 between the climatic niches of A. concolor and A. nodam are only marginal. Pairwise comparison revealed low niche overlap values of D = 0.034 for A. aggerecusans versus A. concolor, D = 0.026 for A. aggerecusans versus A. nodam and D = 0.008 for A. concolor versus A. nodam. The hypothesis of niche equivalency was rejected for the three lineage pairs, due to significant differences between the niches of the sister taxa (niche equivalency test: P = 0.02). The results of the niche similarity are shown in Table 1 and were significant in one of the paired comparisons. *significance value of p, Published as part of Koch, Claudia, Venegas, Pablo J., R��dder, Dennis, Flecks, Morris & B��hme, Wolfgang, 2013, Two new endemic species of Ameiva (Squamata: Teiidae) from the dry forest of northwestern Peru and additional information on Ameiva concolor Ruthven, 1924, pp. 263-295 in Zootaxa 3745 (2) on pages 280-284, DOI: 10.11646/zootaxa.3745.2.6, http://zenodo.org/record/247483

Ameiva aggerecusans sp. nov. Figs. 5���6 Diagnosis and comparison. This comparatively small Ameiva is diagnosed by the following combination of characters: (1) maximum known SVL of 99.3 mm; (2) lacking longitudinal ridge on frontal scale; (3) frontal plate divided in two subequal scales; (4) postnasals separated from prefrontals by frontonasals; (5) parietal scales 5���7; (6) median gular scales not enlarged; (7) enlarged median mesoptychial scales slightly larger than largest gulars; (8) gulars posterior to the interauricular crease smaller than anterior gulars; (9) nasal suture passes through the superior half or centrally through nostril; (10) rostral not contacting or in short contact with postnasal; (11) supranasals not contacting or in short contact with supralabials; (12) scales of circumorbital semicircle not extending to anterior margin of third supraocular; (13) 30���33 enlarged ventral scales between gular and vent; (14) 10���12 longitudinal rows of ventral plates, outermost often distinctly smaller; (15) 73���92 DOM; (16) 144���198 DL; (17) postbrachials not or hardly dilated; (18) 31���39 LFT; (19) 26���32 SCF; (20) 12���22 FP; (21) cream-colored vertebral stripe present in most females and juvenile specimens. Ameiva aggerecusans sp. nov. can be differentiated from all other mainland congeners except for A. bifrontata, A. concolor and A. nodam sp. nov. by having a transversely divided frontal plate. From A. bifrontata and A. nodam sp. nov. this species can be distinguished by lacking distinctly dilated postbrachials. It resembles A. concolor but has a lower maximum SVL and a distinctly more defined cream-colored vertebral stripe on the dorsum. Holotype. An adult male (ZFMK 85024, Figs. 5, 6 A) from Balsas, Province of Chachapoyas, Region of Amazonas, Peru (06�� 49 ��� 11.6 ������S, 78 ��00��� 12.2 ������W, 1000 m above sea level), collected on 0 8 July 2005 by P. Venegas and C. Koch. Description of holotype: An adult male with a SVL of 99.3 mm. Head 0.28 times SVL, 2.32 times longer than wide, 0.9 times as wide as high. Snout elongate, bluntly pointed; canthus rostralis distinct. Neck only slightly narrower than head, and body. Body cylindrical. Limbs well developed, forelimbs 0.36 times SVL, hindlimbs 0.71 times SVL, tibia 0.21 times SVL. Tail slightly pentagonal in cross section, tapering toward the tip; 2.11 times SVL. Rostral in posterior part acute-angled and bordered by supranasals, as wide as high; smooth, with a little suture on each side posterolaterally; visible from above. Supranasals almost triangular, in short medial contact, and in short contact with first supralabials, bordered posteriorly by hexagonal frontonasal. Postnasal almost trapezoid, not contacting rostral, in median contact with frontonasal and in broad contact with loreal and first and second supralabial. Oblique nasal suture passing through the superior half of the oval nostril. Prefrontals paired and roughly pentagonal, medial suture about twice as long as that between supranasals; laterally in contact with loreal, first supraocular and first supraciliary. Frontal plate divided in two subequal scales, the suture between both scales forming a straight line; anterior frontal pentagonal, laterally in contact with first and second supraoculars, larger than posterior frontal; posterior frontal almost squarish, laterally in contact with second and third supraoculars. Paired frontoparietals, slightly longer than wide and with long medial suture; laterally in contact with third supraocular, and small circumorbital scales bordering posterior part of third and fourth supraocular. Interparietal almost squarish, wider than adjacent parietals, sutures with parietals straight; interparietal bordered at each side by two irregular parietals divided by an oblique suture; outermost parietals slightly larger than inner parietals; parietal series composed of 5 scales including interparietal. Supraoculars four at each side. Circumorbital semicircle formed by 14 scales on each side, bordering inner edge of third supraocular and posterior edge of fourth supraocular, separating the latter from frontoparietals; fourth supraocular separated from parietals by two rows of circumorbital scales. Laterally, second and third supraocular separated from supraciliaries by a single row of small scales; 21 combining both sides. Supraciliaries seven (on left side), first highest, fourth longest, remaining ones shorter and subequal. Loreal very large, single, in contact with postnasal, frontonasal, prefrontal, first supraciliary, second preocular, first subocular, and third and fourth supralabials. Preoculars two; first granular, second distinctly larger, but explicitly smaller than suboculars. Suboculars three, first one almost squarish, second and third longer than wide, all in contact with supralabials, second subocular longest. A curved keel reaching from first through second subocular. Postoculars small, in two rows, first consisting of five and second of four scales. Enlarged supralabials eight, fifth below center of eye, third largest; followed to commissure of mouth by seven small scales. An indistinct row of seven slightly enlarged supratemporals. Temporal region with polygonal scales, slightly smaller centrally. External auditory meatus large, vertically oval, bordered by granular scales, anterior margin semicircular, posterior one straight. Tympanum recessed. All dorsal and lateral head scales juxtaposed and smooth. Mental anteriorly rounded, posteriorly straight, bordered by first infralabials and postmental. Postmental single and pentagonal, in contact with first and second infralabials, and followed by eight pairs of enlarged chinshields. First pair in broad medial contact and in contact with infralabials. Second pair of chinshields separated from infralabials by one row of small, almost granular scales, and third pair of chinshields separated by two of those granular scale rows. Remaining chinshields separated from infralabials by one or two larger scales, almost equal in size than posterior chinshields and separated medially by scales of anterior gular region. Medial chin scales moderately small, convex, smooth, juxtaposed, oval or polygonal, in slightly oblique rows, all subequal in size. Enlarged infralabials eight, fifth and sixth below center of eye; followed to commissure by 6 smaller scales. Gular region divided into two areas: anterior region with round or polygonal, juxtaposed, smooth and flat scales in slightly oblique rows, subequal in size, delimited posteriorly by line uniting lower margin of ear openings. Posterior gular region covered by smaller polygonal or round scales in transverse rows. Mesoptychial scales moderately enlarged, slightly larger than anterior gular scales, in about three rows, polygonal or hexagonal, flat, smooth, and juxtaposed. Scales on nape and sides of neck similar in size to dorsals. Dorsals and scales on flanks granular, small, round, smooth, slightly larger in vertebral region; 188 DL; 88 DOM. Ventrals large, smooth, rectangular, wider than long, in 10 longitudinal and 32 transverse rows; transition between ventrals and scales on flanks sharp. Preanal shield with three distinctly enlarged scales. Preanal plate surrounded anteriorly and laterally by smaller scales; posteriorly by much smaller scales. 19 FP in a continuous row along each thigh, first pair of pores medially separated by five scales. Each pore surrounded by four scales, anterior one distinctly larger than posterior ones. Scales on tail dorsally rectangular, smaller than subcaudals, longer than wide, slightly keeled in anterior part, stronger keeled in posterior part, slightly imbricate; arranged in transverse and nearly longitudinal rows, continuous with subcaudals around tail (except first few rows incomplete ventrally); 28 SCF. In medial verticile caudals longer and narrower than in anterior and posterior part of tail. Subcaudals rectangular, smaller than ventrals, wider than long close to base, longer than wide in posterior part of the tail, smooth, mostly juxtaposed. Forelimbs with row of very large, smooth, juxtaposed or slightly imbricate, almost rectangular (distinctly wider than long) antebrachial scales on anterodorsal aspect of forearms and similar but smaller brachial scales on upper arms that extend almost to insertion of forelimbs. Antebrachials and brachials separated by smaller scales at elbow. Dorsoposterior, posterior, and ventral aspect of arms with small almost granular scales, almost equal in size than dorsals, except for scales directly adjacent to brachials and antebrachials, which are slightly to moderately enlarged and irregular. Legs with large, smooth, imbricate scales on anterior and ventral aspects of thighs, and ventral aspect of shanks. Row of large, almost rectangular scales anteriorly on thigh, gradually becoming smaller and irregular toward pores. On ventral aspect of shanks, two rows of very large scales, anterior larger and more or less trapezoidal. Tibiotarsal spurs form a cluster of three rows, each existing of about 4���5 sharply mucronate scales which are positioned along the postaxial edge of the distal end of the shank. Elsewhere on hindlimbs scales granular and slightly smaller than dorsals. Subdigital lamellae of toes transversely enlarged and single, moderately to distinctly tuberculate towards base. Lamellae of outer toe continuing to heel. On palms lamellae of outer and inner fingers continuing to wrist and only separated by few granules, tubercular and increasing in size towards it. Supradigital scales dilated, single and smooth. 18 LFF; 35 LFT. Measurements of holotype (in mm). SVL 99.3; HL 28.43 HH 13.5; HW 12.2; SL 16.3; ED 6.6; DSN 3.0; DNE 8.9; DEE 7.1; TL 209.5; AGL 45; BHM 15.8; BWM 19.6; FLL 36.7; HLL 70.5; TIL 21.1. Coloration of holotype: In life (Fig. 6 A), the dorsal surface of the head and body of the male holotype is reddish-brown and mottled with small frayed dark-brown spots of various sizes, less mottled and slightly lighter colored towards the cloacal region. The sides of the head are cream-colored with some dark dots in the temporal region. The granules on the eyelids are white to grayish-white. Head ventrally white to grayish-white. Body laterally and outermost row of ventral grayish mottled with frayed blackish spots of various sizes. Other ventrals yellowish-white to pale yellow in thoracal region. Forelimbs dorsally striking greenish-yellow (Fig. 6 E), ventrally pale yellow. Hindlimbs and tail dorsally pale brown, ventrally off-white. In preservative, the ground color is dorsally light to moderately brown and grayish on flanks, mottled with blackish spots. Sides of the head are gray-brownish. Ventrals pale gray to yellowish-white. Forelimbs dorsally greenish-gray. Hindlimbs and tail dorsally grayish intermixed with some brownish parts. Pattern and coloration of remaining body parts as in life. Variation. Paratypes (20): Two adult males (ZFMK 85010, 85013), two subadult males (ZFMK 85009, 85011) and an adult female (ZFMK 85012) with the same data as the holotype; an adult male (ROM 16453), an adult female (ROM 16326) and a juvenile (ROM 16459) from Balsas, Province of Chachapoyas, Region of Amazonas, Peru, collected 27 June 1986 by L.D. Wilson; two adult males (CORBIDI 0 5765, ZFMK 90860), an adult female (CORBIDI 05764) and a juvenile (ZFMK 90859) from Balsas, Province of Chachapoyas, Region of Amazonas, Peru (06�� 49 ��� 11.6 ������S, 78 ��00��� 12.2 ������W, 1000 m above sea level), collected on 19 April 2009 by A. Garcia Bravo and C. Koch; three adult males (KU 134829, 134830, 134832), an adult female (KU 134831) and a juvenile (KU 134833) from Balsas, Province of Chachapoyas, Region of Amazonas, Peru, collected on 29 April 1970 by T.H. Fritts; an adult male (ZFMK 90858) from Chacanto, Province of Celendin, Region of Cajamarca (06�� 50 ' 41.5 ���S, 078��01��� 50.8 ���W, 852 m a.s.l.), collected on 16 April 2009 by A. Garcia Bravo and C. Koch; an adult female (ZFMK 90861) and a juvenile (CORBIDI 05766) from Zapatalgo, Province of Utcubamba, Region of Amazonas (06��04'S, 78 �� 29 'W, 1011���1029 m a.s.l.), collected on 0 7 December 2008 by A. Garcia Bravo and C. Koch. Description of paratypes: Maximum SVL in male paratypes 92.6 mm, maximum total length in male paratypes 327.6 mm (both ROM 16453); maximum SVL in female paratypes 71.2 mm, maximum total length in female paratypes 242.2 mm (both KU 134831). Shape of head, body, limbs as in holotype. HL 0.24���0.29 (0.28 �� 0.01, n = 20) times SVL in both sexes; HH 0.11���0.14 (0.13 �� 0.01, n = 12) times SVL; HW 0.11���0.13 (0.12 �� 0.01, n = 21) times SVL; SL 0.57���0.69 (0.63 �� 0.04, n = 12) times the HL; ED 0.23���0.36 (0.27 �� 0.04, n = 12) times the HL; DSN 0.10���0.18 (0.12 �� 0.02, n = 12) times the HL; DNE 0.27���0.35 (0.31 �� 0.02, n = 12) times the HL; DEE 0.20���0.26 (0.23 �� 0.02, n = 12) times the HL; TL 1.62���2.68 (2.30 �� 0.30, n = 18) times SVL; AGL 0.40���0.48 (0.44 �� 0.03, n = 12) times SVL; FLL 0.33���0.41 (0.37 �� 0.02, n = 12) times SVL, HLL 0.56���0.82 (0.73 �� 0.06, n = 12) times SVL, TIL 0.18���0.23 (0.2 �� 0.02, n = 12) times SVL; foot 0.36���0.43 (0.4 �� 0.02, n = 12) times SVL. Arrangement, shape and surface of scales as in holotype except for the following variations: Rostral in posterior part right- to acute-angled, in most specimens in anterior part laterally stretched, projecting beyond the nasal suture and in small contact with postnasal, in few specimens not projecting beyond the nasal suture and not contacting postnasals. Supranasals in short contact with first supralabial or not contacting supralabials. Postnasal trapezoid or almost triangular, in short contact with rostral or not contacting rostral, in contact with first and second supralabials and in some specimens also third supralabial. Oblique nasal suture passing through the superior half or centrally through nostril. Medial suture of prefrontals slightly longer or up to twice as long as medial suture between supranasals. Posterior frontal almost squarish or slightly pentagonal. Interparietal squarish, almost rectangular or irregularly pentagonal, as high or higher than wide; outermost parietals slightly smaller or larger than inner parietals; parietal series composed of 5���7 scales, mostly five, including interparietal. Supraoculars 3���5 (mostly four) at each side, the latter normally much smaller than the others. Circumorbital semicircle formed by 8��� 16 scales at left side, 16���31 scales combining both sides, bordering fourth up to middle or anterior portion of third supraoculars, occasionally extending to frontal suture and thus contacting second or even first supraocular and separating third supraocular from frontoparietals entirely (most specimens) or by parts; bordering posterior edge of last or last two supraoculars, and separating them from parietals by 2���4 rows of circumorbital scales. Laterally, second and third supraocular separated from supraciliaries by a single, occasionally double row of small scales; 22���28 combining both sides. Supraciliaries six or seven, anterior ones normally longer than those in posterior part. Loreal contacting third and fourth supralabials and occasionally in narrow contact with second and fifth supralabial and/or first supraocular, and first and/or second prefrontal. Preoculars 1���2. Postoculars in indistinct two rows, first consisting of 4���7 and second of 3���5 scales. Enlarged supralabials 5���7, fifth or sixth below center of eye; followed to commissure of mouth by 5���10 small scales. Row of slightly enlarged supratemporals distinguishable in most specimens, consisting of 4���9 scales decreasing in size posteriad. Temporal region with polygonal or rounded, slightly convex scales. External auditory meatus slightly oval or almost round. Postmental followed by 7���8 pairs of enlarged chinshields. First pair and in few specimens also second pair in contact with infralabials. Enlarged infralabials five or six (seven in one specimen), fifth or sixth below center of eye; followed to commissure by 6���10 smaller scales. Scales on nape and sides of neck similar in size or slightly smaller than dorsals. 144���198 (175 �� 14.60, n = 13) DL. 73���92 (85 �� 6.03, n = 13) DOM. Ventrals in 10���12 longitudinal and in 30���33 (32 �� 0.98, n = 21) transverse rows, outermost ventrals nearly as wide as those of the adjacent row or distinctly smaller. Preanal shield with 2���3 rows of enlarged scales. 12���22 (20 �� 2.16, n = 19) FP; not or only hardly visible in some female and juvenile specimens. Tail round or slightly pentagonal in cross section; 26���32 (29 �� 1.94, n = 13) SCF. Postbrachials not or hardly dilated. 1���2 rows of slightly to moderately enlarged irregular, hexagonal scales adjacent to brachials. On ventral aspect of shanks, two (occasionally three) rows of enlarged scales, anterior largest, dilated, and more or less trapezoidal, posterior one (or two) rhomboidal, decreasing in size from anterior toward posterior row. Tibiotarsal spurs form a cluster of 2���3 rows of about 3���6 sharply mucronate scales. Subdigital lamellae of toes mostly single, sometimes paired on inner toes. 16���20 (18 �� 1.00, n = 21) LFF; 31���39 (34 �� 1.87, n = 21) LFT. Color variation: In life, juveniles (Fig. 6 D) show a dark brown dorsal ground color on head, body and limbs; a light cream-colored or vanilla-colored vertebral stripe, 4���15 scales in width, extending from behind the head to the base of the tail or less, very striking in anterior part and fading towards posterior part in most specimens; rostral, supralabials and infralabials light yellowish-brown; lower eye granules, suboculars and lower temporals creamwhite; lateral body parts between armpit and groin cream-colored; tail dorsally reddish-brown or medium brown, lightening towards tip; venter cream-white or grayish-white, limbs and tail ventrally cream-white; tubercular lamellae of hands and feet accentuated with brown. Males dorsally reddish-brown or grayish-brown with or without a trace of a white to cream-colored vertebral stripe in anterior part, normally not reaching base of tail; sides brownish-gray or bluish-gray; dorsum of head and body and lateral body parts with or without mottling of tiny, indistinct black spots of various sizes; most specimens with a dark, fringed dorsolateral stripe, which begins faintly and discontinuously behind the eye and almost extends to the insertion of the hindlimbs, being broadest at midbody; dorsal coloration of the tail as in juveniles; head ventrally cream-white, immaculate; venter and ventral surface of limbs and tail immaculate, cream-white to pale yellow, especially frontlegs can show a yellow dorsal and ventral coloration in some males; tubercular lamellae of hands and feet as in juveniles. Females have a grayishbrown to bluish-gray or dark brown dorsal ground color, on body and limbs, head usually brown or grayish-brown; dorsum with (Fig. 6 B) or without a light vertebral stripe, with or without a dark, fringed dorsolateral stripe; lateral body parts between armpit and groin maybe bright cream-colored, dirty gray or whitish; tail dorsally as in juveniles and males; venter, limbs and tail ventrally as in juveniles as well as ventral parts of hands and feet. In pres, Published as part of Koch, Claudia, Venegas, Pablo J., R��dder, Dennis, Flecks, Morris & B��hme, Wolfgang, 2013, Two new endemic species of Ameiva (Squamata: Teiidae) from the dry forest of northwestern Peru and additional information on Ameiva concolor Ruthven, 1924, pp. 263-295 in Zootaxa 3745 (2) on pages 274-280, DOI: 10.11646/zootaxa.3745.2.6, http://zenodo.org/record/247483

Koch, Claudia, Venegas, Pablo J., Rödder, Dennis, Flecks, Morris, Böhme, Wolfgang (2013): Two new endemic species of Ameiva (Squamata: Teiidae) from the dry forest of northwestern Peru and additional information on Ameiva concolor Ruthven, 1924. Zootaxa 3745 (2): 263-295, DOI: http://dx.doi.org/10.11646/zootaxa.3745.2.6