An elusive new species of gymnophthalmid lizard (Cercosaurinae, Selvasaura) from the Andes of northern Peru Author Echevarria, Lourdes Y. https://orcid.org/0000-0003-0162-4408 Pontificia Universidade Catolica do Rio Grande do Sul (PUCRS), Laboratorio de Sistematica de Vertebrados. Av. Ipiranga 6681, Porto Alegre, RS 90619 - 900, Brazil & Division de Herpetologia, Centro de Ornitologia y Biodiversidad (CORBIDI), Santa Rita No. 105 Of. 202, Urb. Huertos de San Antonio, Surco, Lima, Peru lourdese.20@gmail.com Author Venegas, Pablo J. Division de Herpetologia, Centro de Ornitologia y Biodiversidad (CORBIDI), Santa Rita No. 105 Of. 202, Urb. Huertos de San Antonio, Surco, Lima, Peru & Instituto Peruano de Herpetologia (IPH), Augusto Salazar Bondy 136, Urb. Higuereta, Surco, Lima, Peru & Rainforest Partnership, 4005 Guadalupe St., Austin, Texas 78751, USA Author Garcia-Ayachi, Luis A. https://orcid.org/0000-0002-3529-1261 Division de Herpetologia, Centro de Ornitologia y Biodiversidad (CORBIDI), Santa Rita No. 105 Of. 202, Urb. Huertos de San Antonio, Surco, Lima, Peru & Instituto Peruano de Herpetologia (IPH), Augusto Salazar Bondy 136, Urb. Higuereta, Surco, Lima, Peru Author Nunes, Pedro M. Sales Universidade Federal de Pernambuco, Departamento de Zoologia, Avenida Professor Moraes Rego, 1235, 50670 - 901, Recife, PE, Brazil text Evolutionary Systematics 2021 2021-07-27 5 2 177 187 http://dx.doi.org/10.3897/evolsyst.5.68520 journal article http://dx.doi.org/10.3897/evolsyst.5.68520 2535-0730-2-177 B89E282962264A4082E6CDDDA6987C1C 4C16DD914249569A83A3F4486BDD8B4B Selvasaura evasa sp. nov. Figs 1 , 2 , 3 , 4 Proposed Standard English Name: Elusive Microtegus Proposed Standard Spanish Name: Microtegues elusivo Selvasaura Unnamed Clade 3 ( Torres-Carvajal et al. 2016 ) in part. Selvasaura sp. ( Moravec et al. 2018 ) in part. Selvasaura sp. ( Torres-Carvajal et al. in press ) in part. Holotype. Peru • ♂, adult; San Martin Department, Mariscal Caceres Province, Huicungo District, Laurel; 06°41'2"S , 77°41'44.3"W ; 2,762 m; 02 Nov. 2014; L.Y. Echevarria , A.C. Barboza and J. Briones leg.; CORBIDI 15119 (Figs 1 , 2 ). Figure 1. Dorsal ( A ), lateral ( B ) and ventral ( C ) views of the head, and dorsal ( D ) and ventral ( E ) views of the body of Selvasaura evasa sp. nov. holotype, CORBIDI 15119. Photographs by Luis A. Garcia-Ayachi . Scale bar: 5 mm ( A - C ); 10 mm ( D , E ). Figure 2. Dorsolateral ( A ) and ventral ( B ) views of Selvasaura evasa sp. nov. holotype, CORBIDI 15119, in life. Photographs by L.Y. Echevarria . Paratypes. Peru • 2 ♀ adults, 1 ♀ subadult, 2 juveniles, collected with the holotype; CORBIDI 15118 (Fig. 3A, B ), 15120, 15117, 15115-16 • 2 ♂ adults; Amazonas Department, Bongara Province, Yambrasbamba District, Copal; 05°46'36.317"S , 77°50'47.197"W ; 2,582 m; 22 Jun. 2017; P.J. Venegas and J.C. Chavez-Arribasplata leg.; CORBIDI 18900 (Fig. 3C, D ), 18901 • 1 juvenile; Amazonas Department, Chachapoyas Province, Chachapoyas District, Gajmal; 06°16'30.885"S , 77°41'41.896"W ; 2,139 m; 13 Oct. 2018; P.J. Venegas and L.A. Garcia-Ayachi leg.; CORBIDI 19955 (Fig. 3E, F ) • 1 ♂ adult; San Martin Department, Rioja Province, Pardo Miguel District, Fundo Alto Nieva 05°40'13.919"S , 77°45'45.475"W ; 1,938 m; 23 Jan. 2020; L.A. Garcia-Ayachi and J. Ormeno leg.; CORBIDI 22197. Figure 3. Paratypes of Selvasaura evasa sp. nov.: dorsolateral ( A ) and ventral ( B ) views of adult female CORBIDI 15118; dorsolateral ( C ) and ventral ( D ) views of adult male CORBIDI 18900; dorsolateral ( E ) and ventral ( F ) views of juvenile CORBIDI 19955. Photographs A, B by LYE and C-F by PJV. Characterization. A small gymnophthalmid (SVL 51.1-52.2 mm [n = 2] in females and 41.9-46.1 mm [n = 4] in males) characterized by: 1) body slender; 2) head pointed, 1.4 to 1.6 times longer than wide; 3) ear opening distinct, moderately recessed; 4) nasals separated by undivided frontonasal; 5) prefrontals, frontal, frontoparietals, parietals, postparietals and interparietal present; 6) parietals longer than wide; 7) three supraoculars; 8) superciliar series complete, consisting of four scales; 9) nasal completely or partially divided (behind the nostril); 10) loreal in contact with second supralabial; 11) supralabials usually seven; 12) two pairs of genials; 13) collar present, composed by 9-10 enlarged scales; 14) 33-38 transverse rows of dorsal scales; 15) 20-25 transverse rows of ventral scales; 16) 10-12 longitudinal rows of ventral scales; 17) 44-50 scales around midbody; 18) lateral scales at mid-body reduced or absent, present in a maximum of three rows; 19) limbs pentadactyl, all digits clawed; 20) subdigital lamellae under Finger IV 14-19, under Toe IV 17-24; 21) femoral pores 7-10 in females and 9-12 in males; 22) cloacal plate with four large scales; 23) tail 1.3-1.9 times longer than body; 24) caudals keeled dorsally and smooth ventrally; 25) lower palpebral disc undivided, pigmented or transparent; 26) dorsum brown with a yellow vertebral stripe, bordered by broad dark brown stripes in adult males; 27) cream labial stripe extending up to insertion of forelimb in males, females and juveniles; 28) sides of body brown and cream on ventrolateral region in males, females and juveniles; 29) well defined ocelli usually absent; adults and juveniles with a row of faint ocelli (black outline and cream center) followed by cream spots, usually up to midbody. Diagnosis. Selvasaura evasa sp. nov. can be distinguished from S. brava (character states in parentheses) by having 0-3 lateral rows of reduced scales at midbody (6-7); 9-12 femoral pores in males (7-9); keeled dorsal scales, usually flanked by longitudinal striae in adults and juveniles (slightly rugose in adults and slightly keeled in juveniles); ventral surface of tail orange in adult males (yellowish white); and a unilobed hemipenis (bilobed). Selvasaura evasa sp. nov. can be distinguished from S. sp. Ecuador (character states in parentheses) by having a larger size, SVL 41.9-46.1 mm, n = 3, in adult males (maximum SVL 39.7 mm); keeled dorsal scales, usually flanked by longitudinal striae (striated); 33-38 transverse rows of dorsal scales (25-32); 10-12 longitudinal rows of ventral scales at midbody (8-10); 9-10 scales on collar (7-9); a yellow vertebral stripe with broad dark brown stripes on each side in adult males (cream or gray with scattered black marks along sides). Description of holotype. Adult male (CORBIDI 15119); SVL 44.8 mm; TL 86.0 mm; dorsal and lateral head scales juxtaposed, smooth; rostral rectangular, 1.9 times as wide as high; frontonasal pentagonal, as long as wide, laterally in contact with nasals, same size than frontal; prefrontals pentagonal, longer than wide, with medial suture, laterally in contact with nasal, loreal and first superciliary; frontal hexagonal, longer than wide, slightly wider anteriorly, in contact with frontoparietals, prefrontals and supraoculars I and II; frontoparietals pentagonal, longer than wide, with medial suture, each in contact laterally with supraoculars II and III; interparietal hexagonal, lateral borders parallel to each other; parietals polygonal, positioned anterolaterally to interparietal, each in contact laterally with supraocular III and dorsalmost postocular; postparietals three, medial one smaller than laterals; 8/7 (right/left) supralabials, fourth below center of eye; six infralabials, fourth below center of eye; temporals enlarged, juxtaposed, smooth; two large, smooth supratemporal scales; nasal divided in two, subtriangular, in contact with rostral anteriorly, first and second supralabials ventrally, frontonasal and prefrontal dorsally and loreal posteriorly; nostril piercing nasal in the center, directed lateroposteriorly; loreal pentagonal; frenocular subtriangular, in contact with loreal, first subocular and third supralabial; three supraoculars, first one the largest; four superciliaries, first one the largest and dorsally extended in contact with prefrontal and loreal; lower eyelid scale single and pigmented; four suboculars, fourth one the largest; three postoculars, similar in size; ear opening round without denticulate margins; tympanum recessed into a shallow auditory meatus; mental wider than long; postmental pentagonal, slightly wider than long, followed posteriorly by two pairs of genials; gulars imbricate, smooth, in ten rows; gular fold complete; posterior row of gulars (collar) composed by enlarged scales, twice as large as anterior gulars. Scales on nape wider than dorsals; scales on sides of neck small and granular; dorsal scales elongated, imbricate, arranged in transverse rows; scales on dorsal surface of neck striated; 33 dorsal scales between occipital scales and posterior margin of hind limbs; 21 dorsal scales rows in a transverse line at midbody; ventrolateral scales smooth; dorsals separated from ventrals by one (right) or two (left) rows of small scales at the level of the 10th transverse row of ventrals; lateral body fold present; ventrals smooth, longer than wide, arranged in 22 transverse rows between collar fold and preanals; 10 ventral scales in a transverse row at midbody; subcaudals smooth; limbs overlap when adpressed against midbody; axillary region composed of granular scales; scales on dorsal surface of forelimb striated, imbricate; scales on ventral surface of forelimb granular; three thick, smooth thenar scales; supradigitals (right/left): 3/3 on finger I, 7/7 on II, 9/9 on III, 11/11 on IV, 7/7 on V; supradigitals 4/4 on toe I, 6/6 on II, 10/10 on III, 12/11 on IV, 9/9 on V; subdigital lamellae on fingers divided proximally and single distally, 7/7 on finger I, 12/13 on II, 16/16 on III, 16/19 on IV, 12/13 on V; subdigital lamellae on toes divided proximally and single distally, 7/7 on toe I, 12/10 on II, 17/16 on III, 22/22 on IV, 15/15 on V; groin region with small, imbricate scales; scales on dorsal surface of hind limbs striated, imbricate; scales on ventral surface of hind limbs smooth, imbricate; scales on posterior surface of hind limbs granular; 10 femoral pores on each thigh; preanal pores absent; two anterior and four posterior cloacal plate scales. Measurements and proportions of the holotype in mm: HL 10.6, HW 7.1, ShL 5.5, AGD 25.4; TL/SVL 1.9; HL/SVL 0.2; HW/SVL 0.2; ShL/SVL 0.1; AGD/SVL 0.6. Color of holotype in life. Dorsum brown with a broad yellow vertebral stripe bordered by a broad dark brown stripe on each side, and extending from interparietal to base of tail, where it becomes a light brown stripe. The vertebral stripe is continuous with a light brown stripe on the dorsal surface of the tail. Dorsal surface of head brown with pale yellow suffusion on frontoparietals and frontal. Lateral surface of head brown. Iris orange. Supralabials and infralabials cream with brown mottling. Cream labial stripe extending posteriorly up to insertion of forelimb. Flanks of body brown, ventrolateral region cream. A longitudinal row of small pale-yellow spots on dorsolateral region. Flanks with two faint ocelli followed by small cream spots, from forelimb insertion to base of tail. Dorsal surface of tail light brown with orange splotches on the sides. Dorsal surface of forelimbs and hind limbs brown; only fingers I and II and toes I and II yellow. Ventral surface of head cream with brown flecks. Ventral surfaces of neck, chest and belly cream with blurred brown splotches on each scale. Ventral surface of tail orange. Ventral surface of forelimbs and hind limbs yellow with brown flecks. Color of holotype in preservative. Similar to coloration in life. Suffusion on frontoparietals and frontal cream; ventral surface of forelimbs and hind limbs cream with brown spots; ventral surface of tail pale pink on anterior third and cream on the remainder. Variation. Variations in scale counts and SVL of Selvasaura evasa sp. nov. and comparisons with congeners are presented in Table 1 . Table 1. Variation in scutellation and sexual dimorphism in SVL (mm) of Selvasaura evasa sp. nov. and its congeners, S. brava and S. sp. Ecuador.

Character	Selvasaura evasa sp. nov. (n = 7)	Selvasaura brava (n = 6)	Selvasaura sp. Ecuador (n = 3)
Scales in collar	9-10	9-11	7-9
Transverse rows of dorsals	33-38	33-36	25-32
Laterals at midbody	0-2	6-7	5
Transverse rows of ventrals	20-25	22-25	20-23
Longitudinal rows of ventrals	10-12	10	8-10
Scales on cloacal plate	4-5	4	4
Lamellae under Finger IV	14-19	14-16	12-16
Lamellae under Toe IV	17-24	18-22	18-20
Femoral pores in females (one leg)	7-10	0	unknown
Femoral pores in males (one leg)	9-12	7-9	9-12
Maximum SVL in females (mm)	52.2	42.1	unknown
Maximum SVL in males (mm)	46.1	45.9	39.7

Two juvenile (CORBIDI 15116 and 19955) and two adult male (CORBIDI 18900-01) specimens have keeled dorsal scales, without the usual striae flanking the central keel. Male specimens CORBIDI 18900-01 and CORBIDI 22197 lack rows of lateral scales. Specimen CORBIDI 15118 has five superciliaries on left side. Specimen CORBIDI 15118 has six supralabials on left side, and CORBIDI 19955 has eight. The number of infralabial scales is quite variable, 5/5 (CORBIDI 15116), 6/5 (CORBIDI 15118), 6/7 (CORBIDI 15115), 7/7 (CORBIDI 15117, 18900, 19955), and 7/8 (18901). Femoral pores are absent only in one juvenile specimen (CORBIDI 19955). All adult males have a similar coloration, only CORBIDI 22197 has a light brown vertebral stripe. Adult females are similar in color to adult males. However, the vertebral stripe in females, when present, has a different color pattern. Subadult female specimen CORBIDI 15117 has a light brown vertebral stripe with scattered dark brown spots along margins. Female specimen CORBIDI 15118 (Fig. 3A, B ) lacks a vertebral stripe and has two faint black stripes along the dorsolateral margins of the tail. Females lack orange spots on the lateral surface of tail. The ventral surface of tail in females is cream with brown flecks (CORBIDI 15118) or cream with orange margins (CORBIDI 15117). Juvenile specimens CORBIDI 15115-16 are similar in color to adult males. However, the broad dark brown stripes on each side of the vertebral stripe are discontinuous. Ventrally, the brown flecks on head and the brown splotches on neck, chest and belly are more conspicuous. The tail is orange (CORBIDI 15115) or cream (CORBIDI 15116) ventrally. Specimen CORBIDI 19955 (Fig. 3E, F ) has a dark brown dorsum with abundant black flecks and a cream vertebral stripe. It is the only specimen with well-defined ocelli on the anterior portion of flanks, three and seven on right and left sides, respectively, followed by faint ocelli. One to four faint ocelli (black outline and cream center) are present on the flanks of adult and juvenile specimens. These ocelli are usually followed by cream spots, except in CORBIDI 15118 (Fig. 3A, B ) and CORBIDI 18901. Adult female CORBIDI 15117 and adult male CORBIDI 22197 lack ocelli and have a row of three cream spots. The iris is orange, except in one adult female (CORBIDI 15118) and a juvenile (CORBIDI 15115) which have a yellow iris. Hemipenial morphology. The left hemipenis of the holotype is completely everted and fully expanded, 5 mm long (~four subcaudals); organ unilobed, capitate, with lobe detached from body and bordered by the branches of the sulcus spermaticus; sulcus spermaticus broad and shallow throughout the hemipenial body, originating at the base of the hemipenis, central in position, and extending in a straight line until it divides into two branches at the half-length of the organ; sulcal branches separated by fleshy fold. Hemipenial body roughly Y-shaped with 15 pairs of transversal flounces, separated by a longitudinal nude area on asulcate face, except for the first five proximal-most flounces; all flounces ornamented with calcified spicules (Fig. 4 ). Figure 4. Left hemipenis of Selvasaura evasa sp. nov. (CORBIDI 15119 - holotype) in sulcate (left), lateral (middle), and asulcate (right) views. Photographs by PMSN. Distribution and natural history. Selvasaura evasa sp. nov. is known from four localities along the eastern slopes of the Andean Cordillera Central, in Amazonas and San Martin departments, northern Peru, at elevations from 1,938 to 2,762 m (Fig. 5 ). All localities lie in the Peruvian Yungas ecoregion ( Olson et al. 2001 ). Individuals collected at Laurel, in San Martin department, were found in the morning of a cloudy day, inside bromeliads, at ground level and up to 2 m above the ground, in a patch of poorly drained soil covered by Sphagnus mosses, with scattered shrubs and bromeliads. All specimens were lying inside the water stored in the bromeliads; only CORBIDI 15120 was found dead on the ground, beginning to decompose, missing the right hind limb and a portion of the tail, and with the head partially eaten. Specimens from Copal, in Amazonas department, were found on the border of a small lagoon, under a rock (CORBIDI 18900) and among trash (CORBIDI 18901) in a patch of bunchgrass. Specimen CORBIDI 22197 from Fundo Alto Nieva, was found crossing a road in an area of dense montane forest. Specimen CORBIDI 19955, from Quebrada Gajmal, was collected when it fell from the stick roof of a house surrounded by cattle pasture and patches of secondary montane forest. Figure 5. Distribution map of Selvasaura . Symbols with a dot in the middle correspond to type localities. The two adult females collected at Laurel were gravid, each with two eggs. Specimen CORBIDI 15118 had eggs 12.7-13.7 mm long, 6.1 mm wide and 247.3-266.8 mm3 of volume. Specimen CORBIDI 15120 had eggs 12.7-13.4 mm long, 5.6-5.9 mm wide and 208.4-244.1 mm3 of volume. Etymology. The specific epithet Selvasaura evasa is derived from the Latin adjective ēvasa meaning scaped and refers to the evasive nature of this species. The first specimens of Selvasaura evasa sp. nov. were collected in 2014 despite continuous surveys, since 2003 in 20 localities along the montane forests of Amazonas and San Martin departments, it is only known from four localities.