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Bothriechis nubestris new species Talamancan Palm-Pitviper Figures 4–5. Bothriechis nigroviridis Peters 1859: 278. [In part.] Holotype. UTA R- 9637 (Campbell and Lamar, 2004: fig. 97 B), an adult female, from San Isidro de El General, Province of San José, COSTA RICA; approximately 3000 m; collected in October 1973 by Peter Seigfried. Paratypes. All from province of San José, COSTA RICA: UTA R- 2801, 2808, 2850, 7327, 7463, 9635, 9636, 10433, all from San Isidro de El General District; UCR 5727, 5728, 5758, 11151, 15420, 15423, 15424, 15432, all from San Gerardo de Dota; UCR 12356, from División, Páramo District. Referred specimens. From province of San José, COSTA RICA: UCR 5757, from San Gerardo de Dota; UTA R- 6799, 9364, 9365, 10432, all from San Isidro de El General District; CM 145874, 148063, 148064, LACM 154553, Cerro de la Muerte near San Gerardo; UCR 16726, Providencia; UCR 15422, 15429, Alto Palma, Parque Nacional Braulio Carrillo, Bajo la Hondura (locality possibly in error); UCR 15428, Las Nubes (locality possibly in error); UCR 3951, Naranjo, Río Naranjo; UMMZ 117734, no precise locality data. From province of Cartago: MVZ 24228. From unknown Costa Rican locality: UCR 15439. From province of Limón: UCR 22428, Talamanca, Telire, Cerro Uthyum. Diagnosis. (1) a medium-sized slender arboreal pitviper; (2) dorsum green with heavy black mottling; (3) iris blackish; (4) superciliary scales absent; (5) interrictals 22–29; (6) supraoculars thin, usually kidney-shaped; (7) intersupraoculars 6–10; (8) partial rows or two rows of irregular scales between suboculars and supralabials; (9) infralabials 9–12; (10) first dorsals usually 21 (75.9 %); (11) second dorsals usually 21 (58.6 %); (12) third dorsals usually 17 (85.7 %); (13) ventrals 150–160; (14) subcaudals 52–64; (15) tail prehensile. Specimens of Bothriechis nubestris differ from B. schlegelii and B. supraciliaris by lacking superciliary scales (present in B. schlegelii and B. supraciliaris). Bothriechis nubestris differs from all other Bothriechis species except B. nigroviridis by having green dorsal coloration with heavy black mottling and a blackish iris. Bothriechis nubestris differs from B. nigroviridis (see Table 1 for summary) by the combination of having 150–160 ventral scales (136–149 in B. nigroviridis), thin, often kidney-shaped supraoculars with a wide intersupraocular space (B. nigroviridis usually have wider supraoculars, never kidney-shaped, with narrow intersupraocular space), and higher average counts of interrictals, dorsals, and subcaudal scales than B. nigroviridis. Description of holotype. An adult female; SVL 779 mm; tail length 132.2 mm, comprising 14.51 % of total; head length 38.9 mm; maximum head width 23.8 mm; rostral scute broader than high (4.2 X 3.2 mm); nasal divided above and below nostril, nasal fused with first supralabial on right side; loreal 1 / 1, contacting canthal, upper preocular, supralacunal, prelacunal, and two prefoveals; prefoveals 6 / 8; subfoveals absent; postfoveals 3 / 4; prelacunal large, contacting third supralabial on right side, second and third supralabials on left side; sublacunals 2 / 2; postlacunal absent because supralacunal and second sublacunal in contact; preoculars 3 / 3, upper large, middle about half size of upper, lower small and rounded; suboculars 2 / 2, anterior scale long; postoculars 3 / 3; loreal pit large, directed anteriorly, center of pit located slightly below line drawn from center of eye to naris and approximately halfway between center of eye and naris; supralabials 11 / 11 (including fusion of nasal with first supralabial on right side); infralabials 10 / 11, first pair meet posteriorly. Mental broader than long (3.6 X 2.9 mm); three pairs of chin shields flanking mental groove, first two pairs contacting infralabials; five gulars between chin shields and preventrals; three preventrals. Internasals 3 anteriorly; canthals 2 / 2, anterior above nasal, posterior contacting loreal and upper preocular; 1 / 2 small scales between posterior canthal and supraocular; three scales between anterior canthal pair; four scales between posterior canthal pair; superciliaries absent; supraoculars narrow, approximately three times longer than broad; intersupraoculars nine; scales in parietal region small and keeled; interrictals 23. Dorsal scale rows 21 - 21 - 17; ventrals 156; cloacal entire; subcaudals 53, first divided; tail spine short, blunt, 1 ½ times as long as preceding subcaudal scale, covered by scales; tail prehensile. Coloration in preservative. Dorsum of head and body black and pale green mottled; dorsal surface of tail less mottled, grey-green speckled with black, last third yellowish brown lightly speckled with black; postocular stripe extending to rictus, above stripe faint black longitudinal striping, below stripe yellowish with some black blotches on upper lip; ventral surface of head and body yellow, becoming more yellowish green posteriorly. Subcaudal region grey-green with black speckling fading to brown with faint black speckling near tip. Coloration in life of holotype unknown. Variation. Dorsals were most commonly arranged in 21 – 21 – 17 (42.9 % of specimens), whereas each of the following combinations were also observed: 23 – 23 –17, 23– 21 –17, 21– 21 –15, 21– 20 –17, 21– 19 –17, 21– 19 –15, 19– 21 –17, 19– 19 –17, 19– 17 – 17, and 19 – 17 – 15. Ventral scales range 151–157 (mean = 153.7) in males and 152–160 in females (mean = 155.4). Subcaudal scales range 53–62 (mean = 58.6) in males and 52–64 (mean = 56.2) in females. Tail length comprises 14–17 % (mean = 15.2 %) of total length in males; tail comprises 14–16 % (mean = 15.0%) of total length in females. Head scale variation is as follows: prefoveals 4–7; subfoveals 0–1 (present in one specimen only, all others lack subfoveals); postfoveals 1–4; suboculars 1–4; postoculars 2–3; supralabials 9–11; infralabials 9–12; internasals 2–4; intercanthals 4–7; interrictals 22–29 (Table 1). Coloration in preservative varied among specimens and consisted of two rather distinct morphs, with few specimens showing intermediate coloration. Twenty-six percent of the specimens had nearly identical coloration to the holotype. Sixty percent of the specimens displayed a darker color morph with dorsal green being replaced by dark grey and the yellowish venter being replaced by beige, fading to grey-green on the tail. The same ventral pattern is present is both morphs but the yellow is replaced by beige in the darker morph. In two specimens, the dorsum is the “darker morph” but the venter is like the holotype’s with yellow. Recently-born juveniles had brown and black dorsums with beige-brown venters and the tail tips were completely brownish yellow. Coloration in life of three known specimens is nearly identical to the holotype coloration in preservative. Hemipenes. Paratype UCR 11151 had fully everted hemipenes. The total length of the right hemipenis was equal to the length of the first five subcaudal scales. The two lobes of the organ bifurcate at an approximate length of two subcaudals and the sulcus spermaticus divides at a distance of approximately ½ the length of a subcaudal. The base of the organ is spinous, whereas the distal portion is covered in calyces. Spines are dense and evenly dispersed; some spines are quite large. Calyces begin approximately 1 ½ subcaudal scale lengths from the tip of the organ. Distribution and natural history. Bothriechis nubestris is known from the northern and central portions of the Cordillera de Talamanca of Costa Rica in the provinces of San José, Cartago, and Limón (see Fig. 6). Three specimens (UCR 15422, 15428, 15429) have locality data that indicate they are from the Cordillera Central, but they were snakes donated by locals to the Instituto Clodomiro Picado and it is possible that the locality data were recorded in error. We exclude these localities from consideration here, but acknowledge that if these localities are correct, the species also occurs in the Cordillera Central. The species has been recorded from 2400 m on Cerro de la Muerte to over 3000 m in San Gerardo de Dota. Bothriechis nubestris is an arboreal species usually found in the transition zone between cloud forest and montane rainforest. Etymology. The specific epithet means ‘belonging to the clouds’. It is derived from the Latin noun nubes, –is, meaning cloud, and the Latin suffix –estris, meaning belonging to. This name alludes to the fact that this species inhabits cloud forests. The common name Talamancan Palm-Pitviper refers to its range in the Cordillera de Talamanca.

Euspondylus paxcorpus new species Figures 2���3. Holotype. UTA R- 62329, an adult male, from Ulcuran, district of Ulcumayo, province of Jun��n, region of Jun��n, Peru; 10 �� 57.954 ��� S, 75 �� 49.753 ���W; 3341 m (Fig. 1); collected on 8 November 2013 by Esther Ulloa Ribera. Paratypes. Twelve paratypes (7 male, 2 female, 2 juvenile) are from the Ulcumayo district, province Jun��n, region Jun��n, Peru. UTA R- 62308, an adult female, from Villac, 3741 m, collected by Franklin Jeferson Malpartida Ramos, 19 September 2013; UTA R- 62309, an adult male, from Yapacmarka, 3754 m, by Frank Hever Malpartida Ramos, 20 September 2013; UTA R- 62310, a subadult male, from Hachulan, 3581 m, by Herlis Malpartida Alderete, 5 November 2013; UTA R- 62312, a subadult male, and UTA R- 62313, a juvenile, from Huaylla, 3615 m, by Herlis Malpartida Alderete, 5 November 2013; UTA R- 62315, a female, and UTA R- 62317, a male, from La Libertad, 3523 m, by Heraclio Ulloa Condor, 6 November 2013; UTA R- 62321, a male, from Matamayo, 3508 m, by Herlis Malpartida Alderete, 6 November 2013; UTA R- 62325, a male, from Yananauwi, 3280 m, by Anibel Malpartida Arias, 7 November 2013; UTA R- 62328, a juvenile male, from Ulcuran, 3341 m, by Esther Ulloa Ribera, 8 November 2013; UTA R- 62330, a subadult male, from Ulcumayo, 3610 m, by Josben Chuco Carhuapoma, 9 November 2013. Referred specimens. MHNC TMD 1479, TMD 1491 ��� TMD 1492, TMD 1494, TMD 1496 ��� TMD 1497, TMD 1 499 ��� TMD 1500, TMD 1503 ��� TMD 1505, TMD 1508 ��� TMD 1510, TMD 1513, TMD 1515, TMD 1517 ��� TMD 1521, TMD 1523, TMD 1525 ��� TMD 1527, TMD 1530, TMD 1533 ��� TMD 1534, TMD 1536, TMD 1538 ��� TMD 1539, TMD 1541 ��� TMD 1544; UTA R- 62311, 62314, 62316, 62318���62320, 62322���62324, 62326���62327, 62331��� 62333. Diagnosis. (1) prefrontals present with long suture (Doan & Cusi 2014); (2) supraoculars four, with first supraocular/first supraciliary fusion; (3) occipitals 5���7, usually irregularly shaped; (4) palpebral eye-disc made up of a single, undivided, transparent scale; (5) supralabials five; (6) infralabials five; (7) dorsal body scales rectangular with rounded edges and low rounded keel, in transverse rows only; (8) dorsal scale rows in a longitudinal count 40���45; (9) dorsal rows in a transverse count at midbody 22���28; (10) a continuous series of small lateral scales separating dorsals from ventrals, 2���6 scales wide; (11) gular scale rows eight; (12) ventral rows in a longitudinal count 23���25; (13) ventral rows in a transverse count 12; (14) femoral pores per leg in males 8���12, femoral pores per leg in females 5���9; (15) preanal pores absent; (16) subdigital lamellae on Toe IV 19���22; (17) in preservative, dorsal and lateral surfaces dark brown with no stripes or ocelli; (18) in life, three indistinct dorsal stripes and a continuous series of white spots (ocelli) present on the lateral surface of the body. Euspondylus paxcorpus may be distinguished from all species of Petracola Doan & Castoe, Proctoporus, and Riama Gray (except Proctoporus chasqui and P. x es t u s (Uzzell)) by the presence of prefrontal scales. It may be distinguished from other species in the genus Euspondylus by the following character states (condition for E. paxcorpus in parentheses). Euspondylus acutirostris (Peters) and E. monsfumus Mijares-Urrutia, Se��aris & Arends: longitudinal ventral count 26���33 (23���25). Euspondylus auyanensis Myers, Rivas & Jadin: palpebral eye disc divided (entire) and dorsal coloration of squarish markings covering entire dorsum (plain in preservative, with dorsolateral stripes in life). Euspondylus caideni K��hler, E. nellycarrilae K��hler & Lehr, and E. spinalis (Boulenger): hexagonal or subhexagonal dorsal scales (rectangular). Euspondylus nellycarrilae: venter dirty white in preservative (uniform dark blue). Euspondylus spinalis: three supraocular scales (four). Euspondylus guentheri (O���Shaughnessy) and E. maculatus Tschudi: palpebral eye disc divided (entire), dorsal scales smooth or wrinkled (usually keeled), and longitudinal dorsal count 32���37 (40���45). Euspondylus josyi K��hler: longitudinal dorsal count 29���35 (40���45) and longitudinal ventral count 18 (23���25). Euspondylus rahmi (De Grijs): anteriormost supraocular not fused with anteriormost supraciliary (fused) and longitudinal dorsal count 49���54 (40���45). Euspondylus simonsii Boulenger: longitudinal dorsal count 33���39 (40���45) and no enlarged thenar scales (two enlarged thenar scales). Euspondylus oreades Ch��vez, Siu-Ting, Duran & Venegas: longitudinal ventral count 20���22 (23���25), sexual dimorphism in dorsal and ventral coloration (no sexual dimorphism in coloration), and in life males with black flecks on each dorsal scale and distinct black dorsolateral stripes separating lighter from darker regions (no black flecks on dorsal scales and dorsolateral stripes, if present, not distinct and located inside lighter scale region). Proctoporus chasqui: yellowish or reddish venter in life (white or bluish venter) and ventral scales in a longitudinal count 19���22 (23���25). Proctoporus xestus: a single large elongate subocular scale (several small subocular scales) and smooth dorsal scales (usually keeled). Description of holotype. Adult male, snout-vent length (SVL) 51.24 mm, tail 85.11 mm, partially regenerated, length of regenerated portion 23.05 mm; axilla to groin distance 27.27 mm; head length 10.96 mm, head width 7.71 mm. Head scales smooth, glossy, without striations or rugosities; rostral scale wider than tall, meeting supralabials on either side at above the height of supralabials, in contact with frontonasal, nasals, and first supralabials; frontonasal wider than long, pentagonal with all sutures rounded, in contact with nasals, loreals, prefrontals, approximately equal length as frontal; prefrontals paired, trapezoidal, in contact with frontal, fused supraciliary/ supraocular, second supraocular, and loreals; frontal longer than wide, hexagonal, in contact with first, second, and third pairs of supraoculars and frontoparietals; frontoparietals hexagonal, in contact with third and fourth pairs of supraoculars, parietals, and interparietal; supraoculars four, first supraocular fused with first supraciliary, second supraocular in contact with supraciliaries 1, 2, and 3, third supraocular not in contact with supraciliaries, fourth supraocular in contact with fourth supraciliary, postocular, one temporal, and parietal; interparietal longer than wide, heptagonal, in contact with parietals and occipitals; parietals hexagonal, in contact on right side with one postocular, three temporals, and one occipital, on the left the parietal is fused with a temporal and contacts only two temporals, a postocular, and one occipital; occipitals seven, lateralmost occipitals pentagonal, next most lateral occipitals polygonal, next most lateral smaller and triagonal, median occipital quadrangular. Nasal triangular, longer than high, partially divided, nostril situated in anterior third of scale, in contact with first and second supralabials, and loreal; loreal quadrangular, on left side in contact with supralabial, frenocular, preocular, and first supraciliary, loreal on right side not in contact with supralabials, in contact with frenocular and supraciliary only; frenocular wedge-shaped, in contact with first two preoculars, first subocular, and supralabials 2 and 3, on right side also in contact with first supraciliary; three preoculars, first in contact with first and second supraciliaries, second in contact with first supraciliary, third in contact with first subocular; four supraciliaries, first expanded onto dorsal surface of head and fused with first supraocular; palpebral eye-disc made up of a single transparent scale; suboculars three; postoculars two; temporals smooth, glossy, polygonal; supratympanic temporals three, first largest; ear opening oblong, tympanum recessed; supralabials five, first four to angle of jaw; infralabials five. Mental wider than long, in contact with first infralabial and postmental posteriorly; postmental single, pentagonal, posterior suture angular with point directed posteriorly, in contact with first and second infralabials and first pair of genials; two pairs of genials, anterior pair in contact with second and third infralabials, second genials in contact with third and fourth infralabials; two pairs of chin shields, separated by irregular pregulars; gular scale rows eight; collar fold distinct; lateral neck scales round, smooth. Dorsals rectangular, longer than wide, approximately twice the width as length at midbody, with anterior and posterior margins slightly rounded, juxtaposed, with single very low rounded keel, in transverse rows only, 44 in a longitudinal count at midbody, some paravertebral scales irregularly arranged; transverse dorsal count at fifth transverse ventral scale row 15, at tenth transverse ventral scale row 26, at fifteenth transverse ventral scale row 20, at twentieth transverse ventral scale row 15; dorsalmost lateral scales half the size of dorsals, rounded squarish, region approximately 3 scales wide, remaining lateral scales smaller, round, and forming a continuous series 2-4 scales wide, more numerous lateral scales at limb insertion regions, up to 11 scales wide; no lateral fold. Ventral scales smooth, in transverse and longitudinal rows, squarish, median scales wider than long, lateral ventral scales longer than wide, imbricate, 23 scales in a longitudinal count; transverse ventral count at midbody 12; scale row anterior to preanal plate three scales wide; anterior preanal plate scales 4; posterior preanal plate scales 6. Scales on tail like those on body; dorsal and dorsolateral caudal scales rectangular with low keel basally, more distal caudal scales lack keel; ventral and ventrolateral caudal scales square, smooth. Limbs pentadactyl; digits clawed; dorsal brachial scales polygonal, subequal in size, imbricate, smooth; ventral brachial scales much smaller than dorsal scales, polygonal, subimbricate, smooth; dorsal and ventral antebrachial scales polygonal, subequal in size, imbricate, smooth. Scales on dorsal surface of manus polygonal, smooth, imbricate; scales on palmar surface of manus small, rounded, subimbricate, domelike; thenar scales two, raised into triangular domes, subimbricate; finger length formula IV> III> II> V> I; scales on dorsal surfaces of fingers smooth, quadrangular, covering dorsal and lateral portion of digit, overhanging supradigital lamellae 3 on I, 6 on II, 8 on III, 8 on IV, 5 on V; subdigital lamellae 6 on I, 9 / 10 on II, 10 / 12 on III, 12 / 14 on IV, 9 / 10 on V. Scales on anterodorsal surface of thigh large, polygonal, smooth, imbricate; scales on posterior surface of thigh small, rounded, domelike, subimbricate; scales on ventral surface of thigh large, flat, smooth, polygonal, imbricate; femoral pores 9 / 8 in an arc, large, in center of scale, continuous series on right side, on left side scale lacking pore in between fifth and sixth pore and in between sixth and seventh pore; preanal pores absent; scales on anterior surface of crus polygonal, large, smooth, imbricate; scales on anterodorsal surface of crus rounded, domelike subimbricate; scales on ventral surface of crus large, polygonal, smooth, flat, imbricate. Scales on dorsal surface of pes polygonal, smooth, imbricate; scales on palmar surface of pes small, rounded, subimbricate, domelike; toe length formula IV> III> V> II> I; scales on dorsal surface of digits single, quadrangular, smooth, of varying sizes, overhanging supradigital lamellae 4 on I, 7 on II, 9 / 10 on III, 11 / 12 on IV, 9 on V; subdigital lamellae single distally, double proximally, 7 / 8 on I, 10 / 11 on II, 15 on III, 19 / 20 on IV, 12 on V; limbs not overlapping when adpressed against the body. Coloration in preservative. Dorsal and lateral surfaces of head, body, tail, forelimbs, and hindlimbs uniform dark brown (Fig. 3). No lateral ocelli visible. Ventral surface of head, gular region, body, forelimbs, and hindlimbs uniform dark blue. Ventral surface of tail the same blue color of dorsal body transitioning to brown distally; regenerated portion of tail dark brown with light brown mottling. Coloration in life. Base color of the dorsal region of the head, body, and tail dark brown, as in preservative. However, middorsal region of approximately 12 dorsals wide and the top of the head are light brown (Fig. 3). Lateralmost portion of the light region is even lighter, almost a beige color and slightly darker pigmentation separates the beige region from the light brown, creating indistinct darker stripes. In addition, a darker indistinct stripe appears paravertebrally. The light brown color does not extend laterally. The light brown head scales include the rostral, frontonasal, prefrontals, frontal, supraciliaries, supraoculars, first postocular, frontal, frontoparietals, interparietal, and occipitals. The light brown color continues onto the tail, but does not extend to the distal region. A series of lateral white spots begins at the neck region and continues to just before hindlimb insertion. The first two of these spots are surrounded by black and may be considered ocelli. More posterior spots are not surrounded by black. Ventral coloration in life is much lighter than in preservative, a pale whitish-blue. Femoral pores are a dark orangish-yellow. Variation. Adult males SVL 45.34���54.78 mm (n = 7), adult females SVL 54.47���56.87 mm (n = 4), juveniles SVL 33.28���37.80 mm (n = 2); complete original tail of juvenile 74.80 mm, no adults with complete original tails known. The paratypes and referred specimens are similar to the holotype with the following minor exceptions. In two males (UTA R- 62309 and R- 62321) the right frontoparietal is divided anomalously into two scales approximately midway through the scale. In a female (UTA R- 62308) both frontoparietals are divided in the posterior third of the scales. In a male (UTA R- 62330) the left third supraocular contacts the third supraciliary and in another male (UTA R- 62321) the right third supraocular makes contact with the third supraciliary. In a subadult male (UTA R- 62310) an extra tiny scale fills the space between the third supraocular and the third supraciliary on both sides. In a female (UTA R- 62308) some of the dorsal scales lack keels. In two males (UTA R- 62317 and R- 62321) anterior/posterior preanal plate scale rows are 2 / 6; in a male (UTA R- 62330) they are 4 / 4. In most specimens the femoral pores are small and indistinct, but in two males (UTA R- 62317 and R- 62321) the pores are much larger and fill one-third of the scale. In most the femoral pores are in a straight line along the leg, but in other specimens they proceed in an arc, as in the holotype. Meristic variation in the paratypes and referred specimens is very minimal and includes: dorsal rows in a longitudinal count 40���45; dorsal rows in a transverse count at midbody 22���28; ventral rows in a longitudinal count 23���25; subdigital lamellae on Toe IV 19���22; femoral pores per leg in males 8���12, femoral pores per leg in females 5���9; occipitals 5���7, usually irregularly shaped. Sexual dimorphism is not apparent in any of the meristic characters, except for higher averages of femoral pores in males. Coloration is quite similar among the specimens with no sexual dimorphism. In life the middorsal light region ranges from 10���12 dorsals wide. The dorsolateral and paravertebral ���stripes��� are more or less distinct in various specimens. The ���stripes��� are most distinct in all juveniles. In most specimens the femoral pores are grey. In some specimens, the lateral white spots are less distinct (but still visible upon close inspection); in other specimens they are very distinct and continue along the entire lateral surface of the body and tail. In preservative a minority of the adults and all of the juveniles and subadults have mottling on the ventral surface of the head and throat regions. The middorsal light region, stripes, and spots/ocelli are not visible in any preserved individuals. Distribution and natural history. Euspondylus paxcorpus is endemic to the eastern slope of the Cordillera Oriental of the central Peruvian Andes. All specimens were collected by day associated with groups of rocks near small farms between 3500���3800 masl in the district of Ulcumayo, region of Jun��n, Peru. The terrain is montane and andenes (agricultural terraces) constructed by the local pre-hispanic culture dominate the area and have been maintained for the continued cultivation of potatoes (Solanum spp.), olluco (Ullucus tuberosus), oca (Oxalis tuberosa), mashua (Trapaeolum tuberosum), and fava beans (Vicia faba). The climate is defined as boreal (Dwb, K��ppen���Geiger climate classification system) and is cold and dry with a marked difference between day and night (approximately 7 to 15 ˚C difference) stereotypical of the Suni and Quechua natural regions (Monge Miguel et al., 1996). An intense rainy season occurs between the months of December and March. The flora is mainly composed of small herbaceous plants and grasses including Jarava ichu. The only other reptile species found in the area was the dipsadid snake Tachymenis peruviana Wiegmann. Interestingly, in the district of Ulcumayo and surrounding areas, E. paxcorpus is used for traditional medicinal purposes; specimens are flayed and tied to extremities to heal broken bones. The use of reptiles for medicinal means is not limited to E. paxcorpus in Ulcumayo, as snakes are commonly left to steep in sugar cane alcohol called ca��aso and considered a curative. In Quechua these lizards are referred to as shakurhuay. Etymology. The specific epithet paxcorpus is a Latin noun that honors the Peace Corps, or Cuerpo de Paz in Spanish. The lizards were discovered and collected by a Peace Corps Volunteer during his service in Peru to promote community-based environmental management., Published as part of Doan, Tiffany M. & Adams, Grant, 2015, A novel species of Euspondylus (Squamata: Gymnophthalmidae) from the Andes Mountains of central Peru, pp. 129-136 in Zootaxa 4033 (1) on pages 131-135, DOI: 10.11646/zootaxa.4033.1.7, http://zenodo.org/record/240255, {"references":["Doan, T. M. & Cusi, J. C. (2014) Geographic distribution of Cercosaura vertebralis O'Shaughnessy, 1879 (Reptilia: Squamata: Gymnophthalmidae) and the status of Cercosaura ampuedai (Lancini, 1968). Check List, 10, 1195 - 1200. http: // dx. doi. org / 10.15560 / 10.5.10.5.1195","Monge Miguel, R. W., Leon Lecaros, W. R. & Chacon Abad, N. I. (1996) Geologia de los cuadrangulos de Chuchurras, Ulcumayo, Oxapampa y La Merced. Boletin Instituto Geologico Minero y Metalurgico, 78 (A), 1 - 179."]}

Cercosaura hypnoides new species Figures 1���3. Holotype. UTA R- 60249 (Figs. 1, 2), an adult male, from Vereda de Portachuelo, in the vicinity of Manzanares, Municipality of Acac��as, Department of Meta, Colombia; 4.1 ��N, 73.8 ��W; 1640 m (Fig. 4); collected on 17 August 1979 by William W. Lamar and Eduardo Thierry. Paratypes. All paratypes from Vereda de Portachuelo, 1640 m, collected by William W. Lamar and Eduardo Thierry. UTA R- 60250, was found as an egg that hatched upon touch on 0 8 July 1980. All five other paratypes were collected as eggs that subsequently hatched in captivity. UTA R- 60248 (Fig. 3), collected on 0 8 July 1979; UTA R- 60251���60254, collected on 0 9 July 1980. Diagnosis. (1) supraoculars three; (2) superciliaries 4���5, first expanded onto dorsal surface of head; (3) palpebral eye-disc made up of a single, undivided scale; (4) supralabials five; (5) infralabials five; (6) dorsal body scales hexagonal, with high rounded keel; (7) dorsal scales in transverse and oblique rows; (8) transverse rows of dorsals 28���31; (9) a continuous series of small lateral scales separating dorsals from ventrals, 4���7 scales wide; (10) transverse ventral rows 16���20; (11) longitudinal ventral rows six; (12) femoral pores per hind limb 8���12; (13) preanal pores absent; (14) subdigital lamellae on Toe IV 18���22; (15) limbs overlapping when adpressed against body; (16) dorsolateral stripe passing through dorsal surface of the eye and lateral stripe beginning on the supralabials, no vertebral stripe. Cercosaura hypnoides is most similar to the species that were formerly considered to belong to the genus Pantodactylus (Ruibal, 1952), but may be distinguished from each of them by having lateral scales that are much smaller than the dorsal scales (versus dorsals and laterals approximately equal in size in C. parkeri (Ruibal), C. quadrilineata Boettger, C. schreibersii Wiegmann, and C. steyeri). Cercosaura hypnoides may also be distinguished from C. quadrilineata by having six longitudinal rows of ventral scales (versus 4). Cercosaura hypnoides can be further distinguished from C. parkeri (Ruibal) by the high femoral pore count of 8���12 (versus 2���6 for C. parkeri). The new species can be distinguished from C. schreibersii Wiegmann by having three postocular scales (versus two) and by having 8���12 femoral pores versus 3���5. Cercosaura hypnoides can be distinguished from C. steyeri by having six longitudinal rows of ventral scales (versus 4 in C. steyeri). The new species can be distinguished from additional species of the genus Cercosaura by possessing hexagonal dorsal scales arranged in transverse and oblique series (versus quadrangular dorsals in transverse and longitudinal series for C. ocellata Wagler and by having only transverse series in C. argulus Peters, C. dicra (Uzzell), C. eigenmanni (Griffin), C. manicata O���Shaughnessy, C. nigroventris (Gorzula and Se��aris), C. phelpsorum (Lancini), and C. vertebralis O���Shaughnessy). The new species can be further separated from C. dicra and C. vertebralis by having three supraocular scales (versus four). Cercosaura hypnoides can be distinguished from C. dicra (8), C. manicata (8), and C. nigroventris (8) by having six longitudinal rows of ventral scales. The new species additionally differs from C. argulus, C. eigenmanni, C. ocellata, C. phelpsorum and C. vertebralis in its pattern of dorsolateral and lateral stripes. Description of holotype. Adult male, with hemipenes everted, snout-vent length (SVL) 38.60 mm, tail 48.12 mm, partially regenerated, length of regenerated portion 8.03 mm; head scales smooth, glossy, without striations or rugosities; rostral scale wider than tall, meeting supralabials on either side at above the height of supralabials, in contact with frontonasal, nasals, and first supralabials; frontonasal wider than long, pentagonal with all corners rounded, in contact with nasals, prefrontals, and in point contact with loreal, shorter than frontal; prefrontals paired, pentagonal, in contact with frontal, anteriormost supraoculars, loreals, and anteriormost superciliaries; frontal longer than wide, hexagonal, in contact with first two pairs of supraoculars and frontoparietals; frontoparietals hexagonal, in contact with second and third pairs of supraoculars, parietals and interparietal; supraoculars three, first supraocular in contact with first three superciliaries, second supraocular in contact with supraciliaries 3 and 4, third supraocular in contact with fourth supraciliary, one postocular, and parietal; interparietal longer than wide, heptagonal, in contact with parietals and occipitals; parietals hexagonal, in contact with one postocular, a subequally large supratemporal, and one occipital; occipitals three, smaller than parietals, median smallest, extending further posteriorly than two lateral occipitals. Nasal longer than high, nostril situated in anterior third of scale, in contact with first and second supralabials, loreal, and frenocular; loreal quadrangular, not in contact with supralabials, contacting frenocular and first superciliary; frenocular quadrangular, dorsalmost corner in contact with first superciliary, in contact with second and third supralabials, preocular, and first subocular; four superciliaries, first expanded onto dorsal surface of head; one preocular, in contact with first superciliary and first subocular; palpebral eye-disc made up of a single transparent scale; suboculars five; postoculars three; temporals smooth, glossy, polygonal; supratemporals two, first largest; ear opening oblong, tympanum recessed; supralabials five, first four supralabials to angle of jaw; infralabials five. Mental wider than long, in contact with first infralabial and postmental posteriorly; postmental single, pentagonal, posterior suture angular with point directed posteriorly, in contact with first and second infralabials and first pair of genials; two pairs of genials, anterior pair in contact with second and third infralabials, second genials in contact with third and fourth infralabials; two pairs of chin shields, separated by irregular pregulars; gular scale rows eight; collar fold indistinct; lateral neck scales round, smooth. Dorsals hexagonal, longer than wide, with posterior margins slightly curved, imbricate, with single high rounded keel, in 30 transverse rows, oblique rows also present; some paravertebral scales irregularly arranged; longitudinal dorsal scale rows 10 at fifth transverse ventral scale row, 11 at tenth transverse ventral scale row, 10 at fifteenth transverse ventral scale row; wide continuous lateral scale series, five to six scales wide, approximately half the size of dorsals, rounded squarish, smaller and more numerous lateral scales at limb insertion regions; no lateral fold. Ventral scales smooth, squarish with rounded posterior margins, imbricate, in 18 transverse rows; longitudinal ventral scale rows at midbody 6; anterior and posterior preanal plate scales paired. Scales on tail like those on body; dorsal and dorsolateral caudal scales hexagonal with keel, ventral and ventrolateral caudal scales square, smooth. Limbs pentadactyl; digits clawed; dorsal brachial scales polygonal, subequal in size, subimbricate, smooth; ventral brachial scales much smaller than dorsal scales, round, juxtaposed, smooth; dorsal and ventral antebrachial scales polygonal, subequal in size, smooth. Scales on dorsal surface of manus polygonal, smooth, subimbricate; scales on palmar surface of manus small, rounded, subimbricate, domelike; thenar scales two, smooth laterally, raised into domes medially; finger length formula IV> III> II> V> I; scales on dorsal surfaces of fingers smooth, quadrangular, covering dorsal half of digit, overhanging supradigital lamellae 3 / 4 on I, 6 on II, 8 on III, 10 on IV, 5 on V; subdigital lamellae 6 on I, 9 / 10 on II, 13 / 14 on III, 14 / 15 on IV, 8 / 9 on V. Scales on anterodorsal surface of thigh large, polygonal, smooth, subimbricate; scales on posterior surface of thigh small, rounded, subimbricate; scales on ventral surface of thigh large, rounded, flat, smooth; femoral pores 9 / 11; preanal pores absent; scales on anterior surface of crus polygonal, smooth, imbricate, decreasing in size distally; scales on anterodorsal surface of crus polygonal, subimbricate, some scales with a slight keel; scales on ventral surface of crus large, polygonal, smooth, flat, imbricate. Scales on dorsal surface of pes polygonal, smooth, subimbricate; scales on palmar surface of pes small, rounded, subimbricate, domelike; toe length formula IV> III> V> II> I; scales on dorsal surface of digits single, quadrangular, smooth, of varying sizes, overhanging supradigital lamellae 3 on I, 5 on II, 8 / 9 on III, 10 on IV, 8 on V; subdigital lamellae single distally, double proximally, 5 / 6 on I, 10 / 11 on II, 15 / 16 on III, 18 / 20 on IV, 14 on V; limbs overlapping when adpressed against the body. Coloration in preservative. Dorsal surface of head dark brown. Lateral surface of head like dorsal surface, a distinct cream stripe initiates on the ventralmost portion of the first supralabial scale and continues through all supralabials onto body, after the first supralabial the stripe bends dorsally to the tops of supralabials 3 and 4, covering approximately half of the final subocular, then the stripe bends ventrally through the middle of the final supraocular in a straight line to the bottom of the auricular opening, onto the body; a second, fainter cream stripe originates at loreal and continues posteriorly through superciliaries, dorsalmost postocular, and supratemporals, continuing onto body. Ventral surface of head yellowish cream with small brown stippling on the chin shields and lateralmost pregulars. Gular region like head but posteriormost gular scales with denser brown stippling per scale, forming clumps. Dorsal surface of body same color as head, keels tipped in light cream; the faint dorsolateral cream stripes continue longitudinally onto tail, the narrowest distance between the stripes is six dorsal scales wide near the forelimb insertions, the widest distance between the stripes is seven scales wide, approximately two-thirds of SVL. Lateral surface of body same coloration as dorsum, lateral stripe from head continues over the forelimb insertion, briefly stopping at the hindlimb insertion and continuing at the same height posterior to the hindlimb insertion onto the tail, the stripe becomes less distinct after the forelimb insertion. Ventral surface of body slightly paler brown than the dorsal and lateral surfaces, with cream coloring on the posteriormost surface of each scale, medialmost ventrals are slightly paler in color than more lateral scales. Dorsal surface of forelimbs with mottling the same brown color of the dorsal body, ground color orangish cream, cream large irregular blotches; dorsal surfaces of manus yellowish cream with small brown stippling; ventral surfaces of forelimbs cream with dense brown mottling on lateral surfaces of antebrachial and diffuse brown mottling on manus. Dorsal surfaces of hindlimbs orangish cream with diffuse brown mottling; dorsal surfaces of pes similar but with less mottling; ventral surfaces hindlimbs similar to that of forelimbs. Dorsal tail coloration made of three distinct stripes, dorsolateral stripes orangish-cream, much more distinct and wider than on body, dorsolateral stripes begin to converge and dark brown coloration fades to stippling, which disappears altogether at approximately one-third of the tail length, rendering the entire dorsal surface of the tail orangish-cream. Lateral surfaces of tail defined by three stripes, the top and bottom stripes dark brown and the middle stripe yellowish-cream; at approximately one-third of the tail length the bottom stripe disappears. Ventral surface of tail yellowish cream. All surfaces of the regenerated portion of the tail a slightly darker brownish-cream. Coloration in life. Dorsally a bronzy tan with sooty highlights on head, becoming paler on body at middorsal region; an intense creamy white stripe begins on eyelid and passes posteriorly to approximately midbody where it fades to nearly the ground color, this stripe is bordered dorsally by a narrow black stripe which begins just distal to eye and becomes faded and broken at midbody, the white stripe is bordered ventrally by a wide, shiny black stripe which begins at snout and extends posteriorly, passing just above extremities and narrowing at tail but continuing to tail tip, this wide stripe is, in turn, bordered ventrally by another intense pearly white stripe which begins just distal to rostral and passes through upper labials, just superior to insertion of forelimb, and gradually fading until its termination at insertion of hindlimb. This is bordered ventrally by another black stripe which begins at the first infralabial and passes squarely through the rictus, extending posteriorly through the forelimb and fading at the insertion of the hindlimb. Ventrally a greenish pearly white beneath the head and neck, black edging on the infralabials; ventral ground color gradually changes to a soft peach orange beneath the abdomen and it intensifies to a pinkish orange on hindlimbs, vent, and tail. Forelimb dark with pale spot on upper arm, elbow, and wrist; soft orange ventrally. Hindlimb brindled orange. Tail intense pinkish orange with black lateral stripe and another poorly defined broken black stripe from just distal to hindlimb, fading at mid-tail; a faint black dorsal line is evident in the sacral region. Iris deep brown, pupil ringed in gold. Tongue pink with dark gray forks. Hemipenis. The organ was almost completely everted during preservation. It is broad and weakly symmetrically bilobed. The sulcus spermaticus runs a medial course until it bifurcates and runs onto the apical surfaces of the lobes. The organ has 14 oblique plicae (flounces), 12 of which occur on the lobes. The plicae bear comblike rows of spicules. Variation. All paratypes are juveniles SVL 22.54���23.65 mm; complete original tails of juveniles 35.88���41.50 mm. The paratypes are very similar to the holotype with the following minor exceptions. In some specimens the frontal only makes contact with the first supraocular (versus the first two supraoculars in the holotype); in such cases the frontoparietal makes contact with all three supraoculars; in one specimen the frontal is irregularly octagonal. In UTA R- 60252 the frenocular does not make contact with the first superciliary. In UTA R- 60251 on the right side the loreal is much larger, making contact with the supralabial and preocular, separating the frenocular from the nasal and first superciliary; there are five superciliaries on the right side. The left side of this specimen resembles the holotype except the loreal makes contact with the preocular. In UTA R- 60250 there is one pair of genials in contact with the first three infralabials. In UTA R- 60250 and UTA R- 60251 the posterior cloacal plate is made up of four scales. Meristic variation in the paratypes includes: gular scale rows 5���7; transverse dorsal rows 28���31; longitudinal dorsal rows at 5 th ventral row 8���11, at 10 th ventral row 11���14, at 15 th ventral row 10���14; lateral scale row 4���7 scales wide; transverse ventral rows 16���20; supradigital lamellae on Finger IV 9���11; subdigital lamellae on Finger IV 14���17; femoral pores 8���12; supradigital lamellae on Toe IV 10���12; subdigital lamellae on Toe IV 18���22. Coloration in the juveniles in preservative is quite similar to the holotype. Overall they have paler pigmentation. The dorsal surface is orangish-brown instead of dark brown. The lateralmost stripe is much wider and covers almost the entire surface of all supraoculars, the stripe is thickest beneath the eye; the second stripe is more distinct than in the holotype, especially posterior to the eye. Ventral surface of the head is like that of holotype but with no stippling; stippling is present in the gular region in some specimens and on the ventral body in all specimens, but not as dense as in the holotype. The dorsal body stripes are much more distinct throughout length of body and tail; there are four to six scales between stripes. There is color information for two of the paratypes in life (UTA R- 60248 and 60250). In general the paratypes are more brightly colored than the holotype. However, in UTA R- 60250 the dorsal ground color is dark gray on head and back, with black on sides. In both paratypes the dorsolateral stripe is pale yellow, becoming more orange distally. The lateral stripe is pale yellow. In UTA R- 60248 the stripes are bordered in black and ventral and limb surfaces are similar to the holotype. In UTA R- 60250 the tongue is dark gray with white tips whereas in UTA R- 60248 the tongue is pink with yellow tips and a blue patch at the union of the two forks. In UTA R- 60248 the umbilical scar is plainly visible. Distribution and natural history. Cercosaura hypnoides is known only from the type locality on the Vereda Portachuelo (Fig. 5). Vereda Portachuelo is a ridge of the Cordillera Oriental of the Andes Mountains at 1560���1800 m elevation, at Kms 13���15 along the road between Guayabetal and Manzanares, Municipality of Acac��as, Department of Meta, Colombia. Type locality is 1640 m. The habitat is premontane pluvial forest (Espinal, 1977), receiving over 4000 mm annual rainfall. The annual temperature range of the type locality is 18���24 ˚C. There is steep, rugged terrain with numerous rocky streams, and relatively tall undisturbed forest that has an abundance of mosses, lichens, aroids, orchids, bromeliads, and vines, creating dense vegetation. Plant associations include trees and shrubs of the genera Vismia (lanzo, carate), Albizia (pisqu��n), Cecropia (yarumo), Trema (zurrumbo), Heliocarpus (balso blanco), Solanum (lulo), Cedrella (cedro), Ochroma (balso), Miconia (tunos), Saurauia (dulumoco), Insertia, Inga (guano), and Pollalesta. The area contains many man-made clearings and small pastures. Owing to the high precipitation, runoff is fairly constant and soils are washed regularly. Nighttime temperatures drop rather sharply, causing fog; high humidity produces a significant chill factor. Syntopic amphibians included Allobates sp., Atelopus minutulus Ruiz-Carranza, Hern��ndez-Camacho & Ardila-Robayo, Centrolene (undescribed), Hyalinobatrachium orientale (Rivero), Hyloscirtus denticulentus (Duellman), Gastrotheca nicefori Gaige, Rhaebo glaberrimus (G��nther), Rheobates palmatus (Werner), Rhinella sp., and various Pristimantis species. Reptiles included Anolis lamari Williams, Pseudogonatodes peruvianus Huey & Dixon, Ptychoglossus bicolor (Werner), Bothrops atrox (Linneaus), Chironius monticola Roze, Dipsas (undescribed), Liophis taeniurus (Tschudi), Mastigodryas boddaertii (Sentzen), Micrurus (undescribed), Ninia atrata (Hallowell), Oxyrhopus leucomelas (Werner), and Synophis lasallei (Maria). The type, Published as part of Doan, Tiffany M. & Lamar, William W., 2012, A new montane species of Cercosaura (Squamata: Gymnophthalmidae) from Colombia, with notes on the distribution of the genus, pp. 44-54 in Zootaxa 3565 on pages 45-53, DOI: 10.5281/zenodo.208567, {"references":["Ruibal, R. (1952) Revisionary studies of some South American Teiidae. Bulletin of the Museum of Comparative Zoology, Harvard, 106, 477 - 529.","Espinal T., S. (1977) Zonas de Vida o Formaciones Vegetales de Colombia: Memoria Explicativa Sobre el Mapa Ecologico. Instituto Geografico \" Augustin Codazzi \", Bogota, 238 pp."]}

RIAMA GRAY COMB. NOV. Type species: Riama unicolor Gray, 1858: 446. Definition: Tongue with imbricate, scalelike papillae. Head scales smooth without striations or rugosities: single frontonasal, frontal, and interparietal; paired frontoparietals and parietals; prefrontals absent; nostril pierced in a single nasal; nasals not in contact. Eyelids developed, lower with a translucent disc divided into several scales. Posterior gulars squarish. Limbs pentadactyl; digits clawed. Dorsal scales quadrangular or hexagonal, elongate, smooth, striate, rugose, or keeled, juxtaposed, forming transverse series only. Ventrals large, smooth, quadrangular, juxtaposed, forming regular longitudinal and transverse series. Femoral pores usually present in males and females; preanal pores present or absent in males or females. Tail cylindrical. Diagnosis: Riama differs from members of Alopoglossinae by having its tongue covered in imbricate, scale-like papillae instead of oblique plicae (Harris, 1994), from Gymnophthalminae by having moveable eyelids, from Rhachisaurinae by having external ear openings, and from Ecpelopinae by lacking prefrontal scales. Within Cercosaurinae (sensu Castoe et al., 2004), Riama differs from all genera except Pholidobolus, Proctoporus, and Petracola by lacking prefrontal scales. It differs from Proctoporus by having a divided palpebral disc, from Pholidobolus by lacking two medial rows of widened gulars, and from Petracola by lacking the combination of smooth dorsal scales and femoral pores per hind limb in both sexes less than six (exclusive of preanal pores). KEY TO THE SPECIES OF NEUSTICURUS DUM��RIL & BIBRON 1a. Single frontonasal...............................................................................2 1b. Paired frontonasals..............................................................................4 2a. Uniformly small dorsal scales.....................................................................3 2b. Heterogeneous dorsal scales: large keeled scales intermixed with small flat scales.................... N. rudis 3a. Deeply recessed tympanum............................................................... N. medemi 3b. Slightly recessed tympanum................................................................. N. tatei 4a. Dorsal scales in longitudinal rows only................................................... N. bicarinatus 4b. Dorsal scales in transverse and oblique rows only............................................. N. racenisi KEY TO THE SPECIES OF PETRACOLA GEN. NOV. 1a. Three supraoculars, supralabial-subocular fusion absent................................ P. ventrimaculatus 1b. Four supraoculars, supralabial-subocular fusion present.................................... P. labioocularis Content: Riama includes 24 species (Table 1). Distribution: Riama occurs throughout the Andes of central Peru, Ecuador, Colombia, and Venezuela, the Cordillera de la Costa of Venezuela, and the northern range of the island of Trinidad. Remarks: Gray���s (1858) description of Riama included some erroneous characters, such as a lateral longitudinal fold that he used as evidence to state that the genus was related to amphisbaenians and Bachia. Although we have not been able to examine the type specimen of this genus, it appears from his detailed drawing that the type specimen was average and did not actually have a lateral fold, but a very thin row of reduced lateral scales. Preservation of a specimen that had not recently eaten or had been dehydrated before or after fixation may have led to the overlapping of dorsal and ventral scales, producing what appeared to be a lateral fold. KEY TO THE SPECIES OF NEUSTICURUS DUM��RIL & BIBRON 1a. Single frontonasal...............................................................................2 1b. Paired frontonasals..............................................................................4 2a. Uniformly small dorsal scales.....................................................................3 2b. Heterogeneous dorsal scales: large keeled scales intermixed with small flat scales.................... N. rudis 3a. Deeply recessed tympanum............................................................... N. medemi 3b. Slightly recessed tympanum................................................................. N. tatei 4a. Dorsal scales in longitudinal rows only................................................... N. bicarinatus 4b. Dorsal scales in transverse and oblique rows only............................................. N. racenisi KEY TO THE SPECIES OF PETRACOLA GEN. NOV. 1a. Three supraoculars, supralabial-subocular fusion absent................................ P. ventrimaculatus 1b. Four supraoculars, supralabial-subocular fusion present.................................... P. labioocularis, Published as part of Doan, Tiffany M. & Castoe, Todd A., 2005, Phylogenetic taxonomy of the Cercosaurini (Squamata: Gymnophthalmidae), with new genera for species of Neusticurus and Proctoporus, pp. 405-416 in Zoological Journal of the Linnean Society 143 (3) on pages 411-412, DOI: 10.1111/j.1096-3642.2005.00145.x, http://zenodo.org/record/4634632, {"references":["Gray JE. 1858. Description of Riama, a new genus of lizards, forming a distinct family. Proceedings of the Zoological Society of London 1858: 444 - 446.","Harris DM. 1994. Review of the teiid lizard genus Ptychoglossus. Herpetological Monographs 8: 226 - 275.","Castoe TA, Doan TM, Parkinson CL. 2004. Data partitions and complex models in Bayesian analysis: the phylogeny of gymnophthalmid lizards. Systematic Biology 53: 448 - 469."]}

POTAMITES GEN. NOV. Type species: Euspondylus strangulatus Cope, 1868: 99. Neusticurus strangulatus group: Uzzell, 1966: 311. Etymology: Potamites is a masculine Greek noun, meaning water finder. It refers to the fact that most members of this genus are semiaquatic, walking on the bottom of streams and often diving into streams to escape predation. Definition: Tongue with imbricate, scale-like papillae. Nostril pierced in a single nasal; nasals separated usually by paired or single frontonasals, occasionally by irregular scales; prefrontals paired or irregular; interparietal usually bordered by a pair of parietals laterally, by a series of two to eight smaller scales posteriorly; occipital and temporal scales differentiated or not; rostral large, mental and postmental single, followed by several paired chin shields; gular crease feeble or absent. Collar fold well developed. Lower eyelid developed, with a palpebral disc undivided or divided into two to seven scales, transparent or pigmented. Tympanum at surface of head or slightly recessed, overhung by surrounding scales of surface of head. Dorsal scales heterogeneous, imbricate, with large, keeled tubercles intermixed with small flat scales, in transverse or longitudinal rows. Ventral scales wider than dorsals, usually flat, rectangular or slightly rounded posteriorly, subimbricate, in transverse and 6���10 longitudinal rows; lateral rows raised or keeled in some species. Limbs pentadactyl, digits clawed; forefoot with enlarged, plate-like scales along inner margin between thumb and wrist; under side of third and fourth toes with paired scales proximally, inner scale a rounded tubercle. Tail slightly compressed; a double caudal crest, feebly to strongly developed. Total femoral and preanal pores: 10���59 in males; 0���29 in females. Preanal plate in 2���3 rows, posterior row 2���5 scales in both sexes. Males and females with or without conspicuous, white- or browncentred, black-bordered ocelli on their lateral body surfaces. Hemipenis without basal hooks; flounces with minute calcareous spinules. Diagnosis: Potamites differs from members of subfamily Alopoglossinae by having its tongue covered in imbricate, scale-like papillae instead of oblique plicae (Harris, 1994), from subfamily Gymnophthalminae by having moveable eyelids, from subfamily Rhachisaurinae by having external ear openings and from subfamily Ecpleopinae by having heterogeneous dorsal scalation. Within subfamily Cercosaurinae (sensu Castoe et al., 2004), Potamites differs from all genera except Echinosaura, Neusticurus, and Teuchocercus by having heterogeneous dorsal scalation. It differs from Echinosaura by lacking basal spines on the hemipenes, from Teuchocercus by lacking conical scales on the tail, and from Neusticurus (character states in parentheses) by having a slightly compressed tail (strongly compressed), subimbricate ventral scales (imbricate), and calcareous spinules on flounces of hemipenes (no spinules). Content: Potamites includes six species (Table 1). Distribution: Potamites occurs throughout Amazonia in Colombia, Ecuador, Peru, Brazil, and Bolivia and in the San Jos�� Province of Costa Rica., Published as part of Doan, Tiffany M. & Castoe, Todd A., 2005, Phylogenetic taxonomy of the Cercosaurini (Squamata: Gymnophthalmidae), with new genera for species of Neusticurus and Proctoporus, pp. 405-416 in Zoological Journal of the Linnean Society 143 (3) on pages 408-410, DOI: 10.1111/j.1096-3642.2005.00145.x, http://zenodo.org/record/4634632, {"references":["Cope ED. 1868. An examination of the Reptilia and Batrachia obtained by the Orton Expedition to Equador and the upper Amazon, with notes on other species. Proceedings of the Academy of Natural Sciences of Philadelphia 1868: 96 - 140.","Uzzell TM. 1966. Teiid lizards of the genus Neusticurus (Reptilia, Sauria). Bulletin of the American Museum of Natural History 132: 277 - 328.","Linnaeus C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Stockholm.","Dixon JR, Lamar WW. 1981. A new species of microteiid lizard (genus Neusticurus) from Colombia. Journal of Herpetology 15: 309 - 314.","Boulenger GA. 1900. Descriptions of new Batrachians and Reptiles collected by Mr. P. O. Simons in Peru. Annals and Magazine of Natural History, Series 7 (6): 181 - 186.","Burt CE, Burt MD. 1931. South American lizards in the collection of the American Museum of Natural History. Bulletin of the American Museum of Natural History 61: 227 - 395.","Kohler G, Lehr E. 2004. Comments on Euspondylus and Proctoporus (Squamata: Gymnophthalmidae) from Peru with the description of three new species and a key to the Peruvian species. Herpetologica 60: 501 - 518.","Avila-Pires TCS, Vitt LJ. 1998. A new species of Neusticurus (Reptilia: Gymnophthalmidae) from the Rio Jurua, Acre, Brazil. Herpetologica 54: 235 - 245.","Boettger O. 1891. Reptilien und Batrachier aus Bolivia. Zoologischer Anzeiger 14: 343 - 347.","Tschudi JJ. 1845. Reptilium conspectus quae in Republica Peruana reperiunter et pleraque observata vel collecta sunt in itinere a Dr. J. J. de Tschudi. Archiv for Naturgeschichte 11: 150 - 170.","Doan TM, Castoe TA. 2003. Using morphological and molecular evidence to infer species boundaries within Proctoporus bolivianus Werner (Squamata: Gymnophthalmidae). Herpetologica 59: 433 - 450.","Uzzell TM. 1958. Teiid lizards related to Proctoporus luctuosus, with the description of a new species from Venezuela. Occasional Papers of the Museum of Zoology, University of Michigan 597: 1 - 15.","Kizirian DA. 1996. A review of Ecuadorian Proctoporus (Squamata: Gymnophthalmidae) with descriptions of nine new species. Herpetological Monographs 10: 85 - 155.","Kizirian DA, Coloma LA. 1991. A new species of Proctoporus (Squamata: Gymnophthalmidae) from Ecuador. Herpetologica 47: 420 - 429.","Andersson LG. 1914. A new Telmatobius and new teiidoid lizards from South America. Arkiv for Zoologi 9: 1 - 12.","Boulenger GA. 1902. Descriptions of new batrachians and reptiles from the Andes of Peru and Bolivia. Annals and Magazine of Natural History, Series 7 (6): 394 - 402.","Doan TM, Schargel WE. 2003. Bridging the gap in Proctoporus distribution: a new species (Squamata: Gymnophthalmidae) from the Andes of Venezuela. Herpetologica 59: 68 - 75.","Boulenger GA. 1908. Descriptions of new batrachians and reptiles discovered by Mr. M. G. Palmer in south-western Colombia. Annals and Magazine of Natural History, Series 8 (2): 515 - 522.","Peters W. 1862. Uber Cercosaura und die mit dieser Gattung verwandten Eidechsen aus Sudamerica. Abhandlungen der Akademie der Wissenschaften Berlin 1862: 165 - 225.","Boulenger GA. 1885. Catalogue of the Lizards in the British Museum (Natural History) I - III. London: British Museum (Natural History).","O'Shaughnessy AWE. 1879. Descriptions of new species of lizards in the collection of the British Museum. Annals and Magazine of Natural History, Series 5 (4): 295 - 303.","Kizirian DA. 1995. A new species of Proctoporus (Squamata: Gymnophthalmidae) from the Andean Cordillera Oriental of northeastern Ecuador. Journal of Herpetology 29: 66 - 72.","Gray JE. 1858. Description of Riama, a new genus of lizards, forming a distinct family. Proceedings of the Zoological Society of London 1858: 444 - 446.","Harris DM. 1994. Review of the teiid lizard genus Ptychoglossus. Herpetological Monographs 8: 226 - 275.","Castoe TA, Doan TM, Parkinson CL. 2004. Data partitions and complex models in Bayesian analysis: the phylogeny of gymnophthalmid lizards. Systematic Biology 53: 448 - 469."]}

KEY TO THE SPECIES OF RIAMA GRAY 1a. Limbs overlapping when adpressed against body.....................................................2 1b. Limbs not overlapping when adpressed............................................................10 2a. Hexagonal dorsal scales.........................................................................3 2b. Quadrangular dorsal scales.......................................................................4 3a. Dorsal scales obtusely hexagonal, preanal pores in males 2................................... R. achlyens 3b. Dorsal scales acutely hexagonal, preanal pores in males 4���6................................... R. shrevei 4a Keeled, striated, or rugose dorsal scales............................................................5 4b Smooth dorsal scales..................................................................... R. laevis 5a. Lateral scale rows 0���6...........................................................................6 5b. Lateral scale rows 7���12..........................................................................9 6a. Transverse rows of dorsal scales 25���35.............................................................7 6b. Transverse rows of dorsal scales 39���43................................................... R. balneator 7a. Lateral scale rows 2���6...........................................................................8 7b. Lateral scale rows 0���1.................................................................... R. striata 8a. Rugose dorsal scales...................................................................... R. vieta 8b. Keeled or striated dorsal scales......................................................... R. hyposticta 9a. Femoral pores on one leg 0���6............................................................. R. oculata 9b. Femoral pores on one leg 9���14........................................................... R. luctuosa 10a. Venter immaculate except near lateral surfaces.....................................................11 10a. Ventral scales with dark pigmentation............................................................12 11a. Preanal pores present in both sexes........................................................ R. inanis 11b. Preanal pores absent.................................................................. R. vespertina 12a. Keeled, striated, or rugose dorsal scales...........................................................13 12b. Smooth dorsal scales.................................................................. R. meleagris 13a. Keeled or striated dorsal scales, dorsal stripes in some individuals, preanal pores present or absent, three or four supraoculars.......................................................14 13b. Keeled dorsal scales, no dorsal stripes, preanal pores present in males and females, four supraoculars............................................................ R. labionis 14a. Transverse rows of dorsal scales greater than 31...................................................15 14b. Transverse rows of dorsals scales 27���31................................................... R. petrorum 15a. Three or four supraoculars, lateral ocelli present or absent, lip barring variable..........................17 15b. Three supraoculars, no lateral ocelli, no barring on lip...............................................16 16a. Supralabial-subocular fusion absent........................................................ R. raneyi 16b. Supralabial-subocular fusion present................................................... R. laudahnae 17a. Superciliary series complete or incomplete, transverse rows of ventrals 19���27, anterodorsal thigh scales smooth, rugose, or striate.................................................18 17b. Superciliary series incomplete, transverse rows of ventral scales 18���20, anterodorsal thigh scales smooth.................................................... R. colomaromani 18a. Keeled or striated dorsal scales, longitudinal rows of dorsal scales 20���34, preanal pores present or absent, femoral pores on one leg of males 0���11................................19 18b. Keeled dorsal scales, longitudinal rows of dorsal scales 21���22, preanal pores absent, femoral pores of males on one leg 3���5........................................... R. columbiana 19a. Keeled or striate dorsal scales, subdigital lamellae on fourth toe 10���18, superciliary series complete or incomplete.........................................................20 19b. Keeled dorsal scales, subdigital lamellae on fourth toe 15���18, superciliary series complete........................................................................... R. orcesi 20a. Transverse rows of ventral scales 19���27, hemipenis with or without small or large spines.................21 20b. Transverse rows of ventral scales 19���20, hemipenis with small spines.......................... R. simotera 21a. Femoral pores of males 0���10, preanal pores of males 0���4, femoral pores of females 0���9, preanal pores of females 0���4....................................................................22 21b. Femoral pores of males 8���9, preanal pores of males 2���4, females lack femoral and preanal pores....................................................................... R. stigmatoral 22a. Hemipenis with small or absent spines, superciliary series usually incomplete...........................23 22b. Hemipenis with large spines, superciliary series complete................................... R. anatoloros 23a. Preanal pores present in both sexes...................................................... R. unicolor 23b. Preanal pores absent................................................................. R. cashcaensis KEY TO THE SPECIES OF PROCTOPORUS TSCHUDI 1a. Two to three supraoculars........................................................................ 2 1b. Four supraoculars...................................................................... P. pachyurus 2a. Venter uniformly dark or with dark stippling or mottling near lateral scale rows.......................... 3 2b. Venter clear yellow or orange without dark mottling......................................... P. guentheri 3a. No continuous series of lateral ocelli............................................................... 4 3b. Continuous series of lateral ocelli......................................................... P. unsaacae 4a. Frontonasal scale longer than frontal scale................................................. P. sucullucu 4b. Frontonasal scale equal in length to frontal scale........................................... P. bolivianus KEY TO THE SPECIES OF PROCTOPORUS TSCHUDI 1a. Two to three supraoculars........................................................................ 2 1b. Four supraoculars...................................................................... P. pachyurus 2a. Venter uniformly dark or with dark stippling or mottling near lateral scale rows.......................... 3 2b. Venter clear yellow or orange without dark mottling......................................... P. guentheri 3a. No continuous series of lateral ocelli............................................................... 4 3b. Continuous series of lateral ocelli......................................................... P. unsaacae 4a. Frontonasal scale longer than frontal scale................................................. P. sucullucu 4b. Frontonasal scale equal in length to frontal scale........................................... P. bolivianus, Published as part of Doan, Tiffany M. & Castoe, Todd A., 2005, Phylogenetic taxonomy of the Cercosaurini (Squamata: Gymnophthalmidae), with new genera for species of Neusticurus and Proctoporus, pp. 405-416 in Zoological Journal of the Linnean Society 143 (3) on pages 413-414, DOI: 10.1111/j.1096-3642.2005.00145.x, http://zenodo.org/record/4634632

NEUSTICURUS DUMÉRIL & BIBRON Type species: Lacerta bicarinata Linnaeus, 1758: 201. Neusticurus bicarinatus group: Uzzell, 1966: 311. Definition: Tongue with imbricate, scale-like papillae. Nostril pierced in a single nasal; nasals separated usually by paired or single frontonasals; prefrontals paired or irregular, sometimes with a diamond shaped scale between frontonasal and prefrontals; interparietal usually bordered by a pair of parietals laterally, by a series of 3-9 smaller scales posteriorly; occipitals differentiated or not; temporal scales differentiated or not, irregular, sometimes keeled; rostral large, mental and postmental single, followed by several paired chin shields; gular crease feeble or absent. Collar fold moderately to well developed. Lower eyelid developed, with a palpebral disc divided into 3-6 scales, sometimes pigmented. Tympanum slightly to deeply recessed, overhung by surrounding scales of surface of head. Dorsal scales uniformly small, or heterogeneous, imbricate, with keeled tuberculate scales, sometimes intermixed with small flat or convex scales, in longitudinal, oblique, or transverse and oblique rows. Ventral scales usually flat, rounded posteriorly, imbricate, in transverse and 8–10 longitudinal rows, lateralmost rows convex or keeled in some species. Limbs pentadactyl, digits clawed; forefoot with enlarged, plate-like scales along inner margin between thumb and wrist; under side of third and fourth toes with paired scales proximally, inner scale a rounded tubercle. Tail strongly compressed; a double caudal crest, feebly to strongly developed. Femoral and preanal pores form an unbroken series: 32–72 in males; 1-18 in females. Preanal plate in 2-3 rows, posteriormost row with 5-7 scales in both sexes. Males and females with or without conspicuous, white-centred, black-bordered ocelli on their lateral body surfaces. Hemipenis without basal hooks; flounces without minute calcareous spinules. Diagnosis: Neusticurus differs from members of Alopoglossinae by having its tongue covered in imbricate, scale-like papillae instead of oblique plicae (Harris, 1994), from Gymnophthalminae by having moveable eyelids, from Rhachisaurinae by having external ear openings, and from Ecpleopinae by having heterogeneous dorsal scalation. Within Cercosaurinae (sensu Castoe et al., 2004), Neusticurus differs from all genera except Echinosaura, Potamites and Teuchocercus by having heterogeneous dorsal scales. KEY TO THE SPECIES OF POTAMITES GEN. NOV. 1a. Dorsal scales in longitudinal rows of large, keeled scales with smaller scales on either side of rows.................................................................................... 2 1b. No longitudinal rows of large, keeled dorsal scales....................................... P. strangulatus 2a. Single frontonasal scale.......................................................................... 3 2b. Frontonasal scale replaced by a series of irregular scales..................................... P. apodemus 3a. Tail annuli in complete rings of enlarged keeled scales................................................ 4 3b. Tail annuli rings not complete, large keeled scales interrupted by small flat scales......................... 5 4a. Tympanum at surface of the head......................................................... P. ocellatus 4b. Tympanum slightly recessed............................................................. P. cochranae 5a. Two to three reduced scales transversely between two paravertebral tubercular rows at mid-dorsum................................................................... P. juruazensis 5b. Four to six reduced scales transversely between two paravertebral tubercular rows at mid-dorsum.......................................................... P. ecpleopus It differs from Echinosaura by lacking basal spines on the hemipenes, from Teuchocercus by lacking conical scales on the tail, and from Potamites (character states for Potamites in parentheses) by having a strongly compressed tail (slightly compressed), ventral scales imbricate (subimbricate ventral scales), and hemipenes lacking calcareous spinules on flounces (calcareous spinules on hemipenes). Content: Neusticurus includes five species (Table 1). Distribution: Neusticurus occurs throughout the Guianan Shield region of Venezuela, Guyana, Suriname, French Guiana, and northern Brazil. KEY TO THE SPECIES OF POTAMITES GEN. NOV. 1a. Dorsal scales in longitudinal rows of large, keeled scales with smaller scales on either side of rows.................................................................................... 2 1b. No longitudinal rows of large, keeled dorsal scales....................................... P. strangulatus 2a. Single frontonasal scale.......................................................................... 3 2b. Frontonasal scale replaced by a series of irregular scales..................................... P. apodemus 3a. Tail annuli in complete rings of enlarged keeled scales................................................ 4 3b. Tail annuli rings not complete, large keeled scales interrupted by small flat scales......................... 5 4a. Tympanum at surface of the head......................................................... P. ocellatus 4b. Tympanum slightly recessed............................................................. P. cochranae 5a. Two to three reduced scales transversely between two paravertebral tubercular rows at mid-dorsum................................................................... P. juruazensis 5b. Four to six reduced scales transversely between two paravertebral tubercular rows at mid-dorsum.......................................................... P. ecpleopus

PROCTOPORUS TSCHUDI Type species: Proctoporus pachyurus Tschudi, 1845: 161. Proctoporus pachyurus group: Uzzell, 1970: 4. Definition: Tongue with imbricate, scalelike papillae. Head scales smooth without striations or rugosities: single frontonasal, frontal, and interparietal; paired frontoparietals and parietals; prefrontals absent; nostril pierced in a single nasal; nasals not in contact. Eyelids developed, lower with an undivided translucent disc. Posterior gulars squarish. Limbs pentadactyl; digits clawed. Dorsal scales quadrangular, elongate, keeled, juxtaposed, forming transverse series only. Ventrals large, smooth, quadrangular, juxtaposed, forming regular longitudinal and transverse series. Femoral pores present in males, sometimes absent in females; preanal pores absent in both sexes. Tail cylindrical. Diagnosis: Proctoporus differs from members of subfamily Alopoglossinae by having its tongue covered in imbricate, scale-like papillae instead of oblique plicae (Harris, 1994), from Gymnophthalminae by having moveable eyelids, from Rhachisaurinae by having external ear openings and from Ecpelopinae by lacking prefrontal scales. Within Cercosaurinae (sensu Castoe et al., 2004), Proctoporus differs from all genera except Pholidobolus, Riama, and Petracola by lacking prefrontal scales. It differs from Riama by having an entire palpebral eye disc, from Petracola by having keeled dorsal scales and from Pholidobolus by having juxtaposed dorsal scales. Content: Proctoporus s.s. includes five species (Table 1). Distribution: Proctoporus occurs in the Andes of the Peruvian departments of Apurimac, Ayacucho, Cusco, Jun��n, and Puno, and the Bolivian departments of Cochabamba, La Paz, and Santa Cruz. Remarks: Recent field-work recovered specimens and tissues (for molecular analysis) of P. pachyurus, for which no such material had previously been available. Preliminary molecular sampling using the mitochondrial gene ND4 revealed that Proctoporus s.s. as described here is a monophyletic group that includes P. pachyurus and an undescribed lineage (T. M. Doan, T. A. Castoe, & Ariz��bal, unpubl. data). Based on phylogenetic analyses of morphological characters, Doan (2003a) stated that, due to the lack of monophyly of the P. pachyurus group, the character of an undivided palpebral disc should not be considered a synapomorphy and was likely a pleisiomorphic character. In contrast to that study, molecular evidence strongly supports the monophyly of this group and we believe that the undivided palpebral disc is a valid synapomorphy for the genus Proctoporus s.s., despite the fact that a single specimen of P. bolivianus (AMNH R-150695) was found with a divided palpebral disc (Wiens & Servedio, 2000). Although Tschudi (1845) did not provide an etymology for the name Proctoporus, the name presumably refers to the preanal pores that he reports for the type species P. pachyurus. However, neither the type species, nor any other members of the genus Proctoporus s.s. have preanal pores (Uzzell, 1973; pers. obs.). Thus, the name erroneously describes a character that these species do not possess., Published as part of Doan, Tiffany M. & Castoe, Todd A., 2005, Phylogenetic taxonomy of the Cercosaurini (Squamata: Gymnophthalmidae), with new genera for species of Neusticurus and Proctoporus, pp. 405-416 in Zoological Journal of the Linnean Society 143 (3) on page 412, DOI: 10.1111/j.1096-3642.2005.00145.x, http://zenodo.org/record/4634632, {"references":["Tschudi JJ. 1845. Reptilium conspectus quae in Republica Peruana reperiunter et pleraque observata vel collecta sunt in itinere a Dr. J. J. de Tschudi. Archiv for Naturgeschichte 11: 150 - 170.","Uzzell T. 1970. Teiid lizards of the genus Proctoporus from Bolivia and Peru. Postilla 142: 1 - 39.","Harris DM. 1994. Review of the teiid lizard genus Ptychoglossus. Herpetological Monographs 8: 226 - 275.","Castoe TA, Doan TM, Parkinson CL. 2004. Data partitions and complex models in Bayesian analysis: the phylogeny of gymnophthalmid lizards. Systematic Biology 53: 448 - 469.","Doan TM. 2003 a. A south-to-north biogeographic hypothesis for Andean speciation: evidence from the lizard genus Proctoporus (Reptilia, Gymnophthalmidae). Journal of Biogeography 30: 361 - 374.","Wiens JJ, Servedio MR. 2000. Species delimitation in systematics: inferring diagnostic differences between species. Proceedings of the Royal Society of London B 267: 631 - 636.","Uzzell TM. 1973. A revision of lizards of the genus Prionodactylus, with a new genus for P. leucostictus and notes on the genus Euspondylus (Sauria, Teiidae). Postilla 159: 1 - 67."]}