Euspondylus excelsum, Chávez & Catenazzi & Venegas, 2017

Chávez, Germán, Catenazzi, Alessandro & Venegas, Pablo J., 2017, A new species of arboreal microteiid lizard of the genus Euspondylus (Gymnophtalmidae: Cercosaurinae) from the Andean slopes of central Peru with comments on Peruvian Euspondylus, Zootaxa 4350 (2), pp. 301-316 : 303-314

Euspondylus excelsum sp. nov.

( Figs. 1–4 )

Unnamed Clade 1—Torres-Carvajal et al. 2016: 70 ( Fig. 2 )

Holotype. ( Figs. 1–2 ) CORBIDI 16457 , adult male, from Quebrada Tambo (-9.70483° Lat , -75.81383° Long, WGS 84, 1147 m above sea level): Huánuco Province, Huánuco Department, Peru; collected by Lesly Luján on 15 October 2015 .

Paratopotype. CORBIDI 16450 subadult female; collected by Lesly Luján on 12 October 2015.

Paratypes. ( Figs. 3–4 ) Two males, five females and three juveniles, all from Huánuco Department, Peru. CORBIDI 14666 , adult male, from Zaria, Pachitea Province (-9.68905° Lat, -75.84300° Long , WGS 84, 1023 m above sea level); collected by Vilma Durán on 16 July 2014 . CORBIDI 15573 , adult female, from Campamento La garganta de la Bella, Tingo Maria National Park, Leoncio Prado Province (-9.33841° Lat, -76.00206° Long , WGS 84, 1095 m above sea level); collected by Diego Vásquez and Germán Chávez on 21 November 2014 . CORBIDI 16396 , female, from Zaria, Huanuco Province (-9.71467° Lat; -75.81592° Long , WGS 84, 1157 m above sea level); collected by Diego Vásquez on 7 September 2015 . CORBIDI 16459 , juvenile female from Santa Elena, Pachitea Province (-9.73027° Lat, 75.80503° Long , WGS 84, 1144 m above sea level); collected by Lesly Luján on 18 October 2015 . CORBIDI 16463 , adult female from Shincay, Huanuco Province (-9.72563° Lat, -75.81497° Long , WGS 84, 1149 m.a.s.l); collected by Lesly Luján on 27 October 2015 . CORBIDI 16464 , adult female from Zaria, Huanuco Province (-9.71830° Lat ; -75.81575° LongWGS 84, 1159 m above sea level); collected by Lesly Luján on 28 October 2015 . CORBIDI 16465 adult male, 16466 adult female, 16467- 68 juveniles, from Zaria, Huanuco province (-9.71897° Lat, 75.8144° Long , WGS 84 1137 m above sea level); collected by Lesly Luján on 5 November 2015 .

Generic placement. We assign tentatively the new species to Euspondylus on the basis of morphological comparisons. The presence of a tongue homogeneously covered with imbricate-like papillae, of well-developed eyelids, a divided lower palpebral disc, elongated nose, striated and subimbricate dorsal scales, lateral scales reduced when are closer to ventrals, limbs clawed and well-developed, and a compact body (all this characters in accordance with the type species of the genus, E. maculatus ) leads us to allocate the new species to the genus Euspondylus (see Table 1 for comparisons with types of E. maculatus ). Additionally, a recent phylogeny (Torres- Carvajal et al. 2016) that includes paratype CORBIDI 15573 (referred as Cercosaurini CORBIDI 15573 into Unnamed clade 1) shows that this taxon is closely related to Pholidobolus and Macropholidus , and it is nested within the paraphyletic clade of Anadia .

Diagnosis. The new species can be identified by the following combination of characters: 1) body robust, moderate size, maximum SVL 83.7 mm in males, 81.4 mm in females; 2) head flat, elongated, about 1.6–1.9 times longer than wide; 3) ear openings distinct, slightly recessed; 6) nasals separated by frontonasal; frontonasal undivided; 7) prefrontals, frontal, frontoparietals, parietals, postparietals and interparietal present; 8) parietals longer than wide; 9) supraoculars four, anteriormost fused with anteriormost superciliar; 10) superciliar series complete, five, occasionally four; 11) uncomplete suture from narina to posterodorsal suture of nasal present; 12) loreal in large contact with second supralabial; 13) supralabials six; infralabials five; 14) genials in three pairs, first and second pair always in contact, third pair usually in contact; 15) collar fold present; 16) dorsals in 32–34 rows in both sexes, rectangular, about three times longer than wide, striated, subimbricate; 17) ventrals in 21–23 rows in both sexes, quadrangular, smooth, juxtaposed; 18) scales around mid-body 40–42; 19) lateral scales at mid-body reduced in 6–9 lines; 20) limbs pentadactyl, all digits clawed, forelimb reaching anteriorly to sixth supralabial; 21) subdigital lamellae under finger IV 15–18, under toe IV 20–24; 22) femoral pores in males 10–12, in females 4–7, four scales between femoral pores; 23) preanal plate scales seven; 24) tail up to 2.5 times longer than body; 25) caudals subimbricate, striated dorsally, smooth ventrally; 26) lower palpebral disc transparent, divided in four sections; 27) coloration in life golden-brown with scattered black spots or blotches on head and dorsum, with a black stripe running from the nasal plate to supratympanic temporals, only interrupted by the eye; dorsum with dorsolateral dark-bordered stripes, red-orange in males, brown in females; males with 7–14 pairs of lateral ocelli, from anterior part of the insertion of forelimbs to posterior part of the insertion of hindlimbs; limbs golden-brown; dorsal scales of tail golden-brown with pale flecks arranged dorsolaterally; venter, ventral surfaces of limbs and tail creamy white with black spots; iris copper-red.

Comparisons. Euspondylus excelsum sp. nov. resembles in scutellation and measurements to E. guentheri and E. maculatus , from which it can be easily differentiated by the strong fusion of anteriormost supraocular with anteriormost superciliar (fusion absent in E. guentheri and E. maculatus ) (See Fig. 5 ). Furthermore, E. excelsum can be distinguished from both species by the following characters (conditions for E. guentheri and E. maculatus in parenthesis): body robust (vs. slender in E. guentheri ), the presence of small rounded irregular spots (vs. large, transversal black spots on dorsum in E. guentheri ), by having striated dorsal scales (vs. smooth in E. guentheri ); third pair of chin shields usually in contact (vs. separated in E. guentheri and E. maculatus ); head 1.6–1.9 longer than wide (vs. 1.5 in E. maculatus ); eye–nose distance is 0.3–0.4 times of head length (vs. 0.5 in E. guentheri ); longitudinal dorsal count 32–34 (vs. 38 in E. maculatus ) ventral scales rows 21–23 (vs. 25 in E. maculatus ); 10–12 femoral pores in adult males (vs. 17 in E. guentheri ); and 7–8 preanal scales (vs. 5 in E. guentheri and E. maculatus ).

Furthermore, E. excelsum sp. nov. can be differentiated from other congeners (those with the same condition on the lower palpebral disc) by having a longer body with a maximum SVL of 83.7 mm (vs. 53.5 mm in E. acutirostris , 59.6 mm in E. auyanensis , 46.0 mm in E. monsfumus ); head 1.6–1.9 longer than wide (vs. 1.5 in E. sp. 1, 1.7 in E. acutirostris and E. guentheri , 1.8 in E. monsfumus ); first supraocular fused with first superciliar (vs. not fused in E. acutirostris , E. monsfumus ); most dorsal temporal 3–3.5 times longer than small lateral temporals (vs. 4–4.5 times longer than small lateral temporals in E. sp. 2); a lower number of postparietals having 2–3 (vs. 3 in E. auyanensis , 3–4 in E. sp. 1); 2–3 pairs of genials in contact (vs. 2 pairs in contact in E. acutirostris and E. sp. 2); a lower number of ventral scales rows having 21–23 (vs. 26–33 in E. acutirostris , 24 in E. auyanensis , 28 in E. monsfumus ); a higher number of scales around mid-body 40–42 (vs. 34–40 in E. acutirostris , 32 in E. auyanensis , 36 in E. monsfumus ); a lower number of femoral pores having 3–7 on each side (vs. 8–11 in E. acutirostris , 15–18 in E. auyanensis , 9–10 in E. monsfumus ); a higher number of preanal scales 7–8 (vs. 5 in E. sp. 1).

Description of the holotype ( Figs 1–2 ). Adult male (CORBIDI 16457); SVL = 69.4 mm, tail (regenerated) length = 55.6 mm; axilla to groin distance = 32.6 mm; head length = 18.1; head width = 11.2 mm; shank length = 11.0 mm.

Head scales smooth, glossy; rostral scale wider (2.7 mm) than long (1.5 mm), slightly higher than adjacent supralabials, in contact with frontonasal, nasals, and first supralabials posteriorly; frontonasal pentagonal, slightly wider (2.8 mm) than long (2.6 mm), posterior suture angular with point directed posteriorly, in contact with nasal and laterally, prefrontals posteriorly, nasoloreal suture uncomplete; prefrontals present, in contact with each other medially, in contact with fused anteriormost superciliary-anteriormost supraocular, frontal posteriorly; frontal pentagonal, longer (3.6 mm) than wide (2.7 mm), anterior suture angular with point directed anteriorly, lateral sutures straight, posterior suture angular with point slightly directed posteriorly, in contact with second supraocular laterally, frontoparietals posteriorly; frontoparietals pentagonal, in contact with third and fourth supraocular, parietals and interparietal posteriorly; supraoculars four, none in contact with ciliaries; superciliary series complete, anteriormost superciliary fused with anteriormost supraocular; interparietal pentagonal, longer (3.6 mm) than wide (2.8 mm), lateral sutures concave, in contact with parietals laterally, postparietals posteriorly; parietals polygonal, in contact with fourth supraocular anterolaterally, temporal scales laterally, dorsalmost postocular, postparietals posteriorly; postparietals four, polygonal; palpebral disc oval, unpigmented, divided in four sections; frenocular triangular, in contact with loreal anteriorly; postoculars three; temporals polygonal; supratympanic temporals two; supralabials six; infralabials five; mental wider (2.8 mm) than long (1.7 mm), in contact with first infralabials, postmental posteriorly; postmental single, pentagonal, posterior suture angular, point slightly directed posteriorly, in contact with first and second infralabials; genials in three pairs, anterior pair subquadrangular, in contact with second and third infralabials, medial pair of genials subcuadrangular, in contact with third and fourth infralabials laterally, posterior pair of genials trapezoid, in contact with fourth and fifth infralabials laterally; scale rows between genials and collar fold (along midventral line) 15; medialmost scales of three penultimate gular scale rows cuadrangular; posteriormost gular row enfolded posteriorly, concealing two granular scale rows; lateral neck scales rounded, smooth.

Dorsal scales rectangular, longer than wide, juxtaposed, striated, 33 in a longitudinal count, homogeneously arranged; transverse dorsal count (enlarged rows at mid-body) at fifth transverse ventral scale row 10, at 10th transverse ventral scale row 12, at 15th transverse ventral scale row 14; lateral scale rows at fifth transverse ventral scale row 13, at 10th transverse ventral scale row 9, at 15th transverse ventral scale row 10; lateral scales on body near insertion of forelimb small to granular; ventrals cuadrangular and juxtaposed; complete longitudinal ventral count 21; longitudinal ventral scale rows at mid-body 10; 41 scales around mid-body; anterior preanal plate scales two; posterior preanal plate scales seven, all the scales at the same size; scales on tail rectangular and juxtaposed, striated; mid-ventral subcaudals squarish.

Limbs pentadactyl; digits clawed; forelimb reaching anteriorly to sixth supralabial; dorsal brachial scales polygonal, of twice the size of anterodorsal midbrachial scale, imbricate, smooth; midbrachial anterodorsal scale small, at least twice shorter than adjacent scales, smooth; anteroventral, ventral, and posteroventral scales roundish, imbricate, smooth; antebrachial scales polygonal, at least as large as midbrachial anterodorsal scale; medial antebrachial scales larger, polygonal, smooth; dorsal manus scales polygonal, subimbricate; palmar scales granular, small, oval; dorsal scales on fingers smooth, quadrangular, covering dorsal half of digit, overhanging supradigital scales, four on I, seven on II, nine on III, 11/10 on IV, six on V; subdigital scales seven on I, 14/13 on II, seventeen on III, 16/17 on IV, eleven on V; anterodorsal thigh scales polygonal, at least three times as large as adjacent scales, becoming smaller ventrally, smooth; posterodorsal thigh scales small, rounded, arranged irregularly; anterior and anteromedial shank scales polygonal, subimbricate, smooth, anteriormost scales many times larger than lateral, posterolateral, and posteromedial shank scales; lateral, posterolateral, and posteromedial shank scales polygonal or roundish, subimbricate, smooth; dorsal pes scales polygonal, subimbricate, smooth; scales on dorsal surface of digits single, quadrangular, smooth, overhanging supradigital scales, four on I, seven on II, thirteen on III, 16/15 on IV, eleven on V; subdigital scales single, seven on I, 14/13 on II, nineteen on III, twentyfour on IV, 16/17 on V; femoral pores series not continued, 5/7; four scales between medialmost femoral pores.

Hemipenial morphology. ( Fig. 6 ) The right hemipenis was everted during preservation and prepared posteriorly. The organ extends along approximately 11 mm. The hemipenis is completely expanded and the lobes of the organ are partially everted. Hemipenial body in light bulb shape, with the basis distinctly narrower than the rest of the organ, and ending in two small lobes with apical folds in the apex. The s ulcus spermaticus is central in position, originating at the base of the organ, and proceeding in a straight line towards the lobes. The sulcus is broader in the region of the lobular crotch, where it is divided by a small bulky fold; branches running on the medial region of the lobes, and ending in their tip among folds. The sulcate face of the hemipenial body presents two fleshy nude areas, stained with blue, parallel to the sulcus spermaticus that run through all the hemipenial body extension.

Laterals and asulcate face of the hemipenis are ornamented with series of roughly equidistant flounces with calcareous spinules, all of them stained in red with Alizarin. Twenty-four rows of flounces extend along the body of the organ. There are three proximal rows restricted to a central position on the basal asulcate face of the hemipenis, all of them are roughly chevron-shaped. The five proximalmost flounces in the laterals are diagonally positioned, with the second to fifth flounces separated from a complementary flounce positioned in the asulcate face and orientated in an inverse diagonal. The nude area between the proximalmost fifth flounce is narrower than the nude area between the other four proximal flounces and its complementary ones. The subsequent flounces towards the lobes cross the laterals of the organ from the sulcate to the asulcate face, in chevron-shape with vertexes in the central region of each lateral pointed towards the basis of the organ. These chevron-shaped rows reduce in size progressively towards the hemipenial apex.

The lateral flounces are separated in two groups by a nude area in the central asulcate face that increases in size in the apical region acquiring a Y-shape. The regions between the asulcate face and the laterals in this species are marked by a conspicuous unevenness forming a bulge shared by other related species such as Macropholidus annectens , M. huancabambae , M. ruthveni , Pholidobolus affinis , P. macbrydei , P. montium , P. prefrontalis ( Nunes, 2011) , P. hillisi ( Torres-Carvajal et al. 2014) , and P. ulisesi (Venegas et al. 2016) .

Color of holotype in life. ( Fig. 2 ) Head golden-brown with scattered black spots, down to inferior level of supralabials, infralabials creamy white with black spots; a dark stripe running from the nasal plate to the anterior angle of the eye and from the posterior angle of the eye to the supratympanic temporals, iris copper-red. Dorsum golden-brown with scattered black spots, a dorsolateral dark-bordered red-orange stripe from nuchals to the base of the tail, dark border turning paler and fading entirely towards the base of tail, seven pairs of lateral ocelli. Dorsal surface of tail golden brown with pale dorsolateral spots. Throat, chest, belly and ventral surfaces of limbs creamy white, with scattered black spots or blotches. Precloacal and cloacal scales creamy white with scattered black spots; ventral surface of tail creamy white with scattered black blotches.

Variation and sexual dimorphism. ( Fig. 3 , 4 ) Variation in measurements and scutellation is summarized in Table 1. The largest male is CORBIDI 16465 (81.7 mm), the largest female is CORBIDI 16466 (81.4 mm). Females CORBIDI 15573 and 16466 have two genials in contact, not three as the rest of type series. Six specimens have the supraocular-superciliar scale fusion divided by a diagonal, weakly defined, complete or uncomplete suture (CORBIDI 16396, 16459, 16463, 16465–66, 16468); also a juvenile (CORBIDI 16468) has six superciliars. Dorsolateral red-orange dark bordered stripe is extended to 10th verticile of the tail in male CORBIDI 16465. Juveniles CORBIDI 16467–68 have yellow coloration on belly and ventral surfaces of hindlimbs and tail. The sex can be distinguished by a higher number of femoral pores in males (10–12) than females (4–6), only juvenile male CORBIDI 16459 has 6 femoral pores, furthermore males can be differentiated by the presence of 7–14 pairs of lateral ocelli (absent in females), and by the presence of black scattered spots or blotches on ventral surface of tail in males, instead black spots or blotches arranged ventrolaterally on tails of females. Additionally, male CORBIDI 16465 has an atypical high number of irregular black spots on belly, remarkably higher than the rest of males of the type series.

Etymology. The specific epithet “ excelsum ” is from Latin and means “tall”, in reference to the habitat of the new species in the canopy of the tropical rainforest.

Distribution and natural History. Euspondylus excelsum sp. nov. is known from localities on both sides of the Huallaga river, including within Tingo Maria National Park ( Fig. 7 , 8 ), at distances as far as 48.5 km between the northernmost and southernmost collecting sites. Specimens were collected in the canopy of primary and secondary forests. Specimens CORBIDI 14666 and 15573 were observed during diurnal Visual Encounter Surveys climbing trunks at 1.5 m above leaf litter. The holotype and specimens CORBIDI 16396, 16450, 16459, 16463-65, were collected in the canopy of Couma spp . at night; specimens CORBIDI 16466–68 were collected under leaves of bromeliads gathered on crags. Male CORBIDI 16465 started swimming underwater during an escape attempt prior to capture. After dissection for laboratory analyses, two eggs were observed into the ventral cavity of three females of type series collected at November 2014 and October-November 2015, eggs lengths were (measurements of egg in right oviduct is presented first): 18.0 mm and 18.0 mm in CORBIDI 15573 (23.2% of the SVL), 16.6 mm and 16.5 mm in CORBIDI 16463 (20.2% and 20.4% of the SVL respectively), 15.3 mm and 16.1 mm in CORBIDI 16466 (19.7% and 18.7% of the SVL respectively).

Euspondylus Euspondylus Euspondylus Euspondylus sp 1 Euspondylus sp 2 excelsum sp. nov. maculatus guentheri (n=2; m=1, f=1) (n=1; m=1, f=0) (n=12; m=5, f=7) (n=2; m=1, f=1) (n=1; m=1, f=0)

Max SVL (mm) Male 46–83.7 59 (59 ± 0) 92 86.1 76.3 (62.8 ± 15.8)

Female 59.9–81.8 92.6 (70.3 ± 11)

Tail Lenght (mm) Male 43.1–149.2 56 (56 ± 0) 102 106 120 (91.5 ± 50)

Female 32.9–175.2 148 (119.1 ± 47.9)

Head length (mm) Male 12.9–21.4 15.5 (15.5 ± 0) 20 23.1 17.9 (16.5 ± 3.8)

Female 13.8–19.2 20.7 (17 ± 2.1)

Head width (mm) Male 7.6–14.3 9.5 (9.5 ± 0) 11.7 15.1 11.2 (10.2 ± 2.7)

Female 8–11.9 13 (10.2 ± 1.5)

Head length/SVL Male 0.2–0.3 0.2 (0.2 ± 0) 0.2 0.18 0.2

(mm) (0.2 ± 0)

Female 0.2–0.3 0.2 (0.2 ± 0)

Head length/Head Male 1.6–1.7 1.6 (1.6 ± 0) 1.7 1.5 1.6

width (mm) (1.6 ± 0)

Female 1.6–1.9 1.6 (1.6 ± 0)

Eye-Nose distance Male 4.4–7.9 - 9.4 8.4 6.7

(mm) (5.9 ± 1.4)

Female 4.9–6.9 - 8.2 (6 ± 0.8)

Eye-Nose distance/ Male 0.3–0.4 - 0.5 0.3 0.4

Head length (mm) (0.4 ± 0)

Female 0.3–0.4 - 0.4 (0.3 ± 0)

Number of Femoral Male 6–12 10 (10 ± 0) 17 8 6

pores (9.6 ± 2.1)

Female 4–6 2 (2 ± 0) 2 (5.1 ± 1)

......continued on the next page Euspondylus Euspondylus Euspondylus Euspondylus sp 1 Euspondylus sp 2 excelsum sp. nov. maculatus guentheri (n=2; m=1, f=1) (n=1; m=1, f=0) (n=12; m=5, f=7) (n=2; m=1, f=1) (n=1; m=1, f=0)

The external appearance of Euspondylus lizards throughout their distribution ranges resembles those of other Cercosaurini, specifically to Anadia , in “long-snouted” species from Venezuela, Colombia, Ecuador and Peru ( Goicoechea et al. 2012, Mijares-Urrutia et al. 2001, Uzzell 1973), or Petracola , Proctoporus and Riama in“shortsnouted” forms distributed in Peru and Bolivia ( Goicoechea et al. 2012). Strikingly, all species recently transferred to Proctoporus ( Goicoechea et al. 2012; Torres-Carvajal et al. 2016) are similar in appearance to the short-snouted forms. In contrast, reviewed specimens of Euspondylus maculatus (type species of the genus) resemble to the long snouted species, as well as to the new species described herein. We note that an important diagnostic character is the condition of the lower palpebral disc, which is divided in vertical sections in “long-snouted” species and not divided in “short-snouted”. Euspondylus species having this character are “long-snouted” E. acutirostris , E. auyanensis , E. excelsum sp. nov., E. guentheri , E. maculatus and E. monsfumus . Therefore, although is still necessary to confirm our results with a phylogenetic analysis, we tentatively propose that Euspondylus can be differentiated in scutellation from all other Cercosaurini by having smooth, striated or slightly keeled dorsals scales (vs. keeled or striated in Cercosaura and Pholidoboluls, strongly keeled and tuberculate in Echinosaura, Gelanesaurus , Neusticurus and Potamites ), subimbricated or juxtaposed (vs. imbricated in Cercosaura , Echinosaura, Gelanesaurus , Pholidobolus , Potamites ), homogeneous (vs. heterogeneous in Echinosaura, Gelanesaurus , Neusticurus and Potamites ); lateral scales rows adjacent to ventrals reduced (vs. lateral scales rows adjacent to ventrals not reduced in Anadia ); lateral body fold absent (vs. present in Pholidobolus ); prefrontal scales present (vs. absent in Petracola and Riama ); a discontinuous series of femoral pores in males (vs. continuous in Petracola ); and a lower transparent palpebral disc divided (vs. not divided in Proctoporus and Macropholidus , opaque in Pholidobolus ).

The presence of prefrontals scales had formerly been used to differentiate “short-snouted” “ Euspondylus ” lizards from Proctoporus lizards (i.e. Köhler 2003). As previously mentioned, phylogenetic analyses dismiss the validity of this character ( Goicoechea et al., 2012; Torres-Carvajal et al., 2016), because all “short-snouted” Euspondylus that were included in these analyses have been transferred to Proctoporus . Five other “short-snouted” species lacked molecular data and are still assigned to Euspondylus ( E. caideni , E. josyi , E. nellycarrillae , E. paxcorpus , and E. simonsii ). However, Torres-Carvajal et al. (2016) suggested that four of them could be Proctoporus , we must emphasize that all of them bear an undivided transparent lower palpebral disc, in contrast to E. maculatus and the other “long-snouted” Euspondylus . According to the scutellation and measurements given in the description of the recently discovered E. paxcorpus , this species also belongs to the “short-snouted” group. On the basis of our comparisons, we support Torres-Carvajal et al. 2016 in suggesting that these five taxa should be placed into Proctoporus , but we await phylogenetic analyses to confirm their formal placement into this genus.

We assign the former records of E. maculatus originating from the Chanchamayo valley in Junín department, and the Mina Inca in Puno department, to Euspondylus sp. 1 and Euspondylus sp. 2 respectively. Despite variation in measurements and scutellation (See Table 1), the lack of genetic evidence prevents comparison of specimens from Chanchamayo (FMNH 40584, 45474) to specimens of E. maculatus , E. excelsum sp. nov., or E. guentheri . These specimens from Chanchamayo share with E. excelsum sp. nov. (and not with other species) the presence of a fusion of the anteriormost supraocular with the anteriormost superciliar scale, but differ from the new species in other characters (i.e., number of postparietals, number of preanal scales). The specimen from Mina Inca is most similar to E. excelsum sp. nov., especially in body measurements, and external appearance, however it also disagrees in scutellation (i.e. number of genials in contact, longitude of temporals), therefore we consider this specimen as Euspondylus sp. 2; collection of new specimens and genetic analyses to clarify the taxonomic status of specimens of both localities should be priorities for future research. Consequently, we consider the distribution range of E. maculatus to be restricted to the type locality “vicinity of Moyobamba” ( Tschudi, 1845) in the eastern slopes of the northern Peruvian Andes.

We note that the origin of specimen from Mina Inca , Puno department (AMNH 1704) is cited erroneously “from Juliaca, Puno department” in previous articles ( Köhler 2003, Köhler & Lehr 2004). We consider this is most likely erroneous because Juliaca is in the high altiplano at 3800 m of elevation (much higher than all other Euspondylus records, including the new one described herein) and in puna grasslands (very different from the montane forests where all other Euspondylus live). The collector of the “Juliaca” specimens, Dr. H. H. Keays clarified the confusion by sending a letter to AMNH mammologist Joel A. Allen asking to correct the error because the collection site was the montane forests around the Inca Mine, in a valley of a tributary of the Inambari River (Madre de Dios watershed), in the Cordillera de Carabaya of Puno at about 1800 m.a.s.l. ( Myers et al. 2009).

Although O´ Shaugnessy (1881) did not mention the sex of the type specimen of E. guentheri (BMNH 1946.8.8.99) in its description, Uzzell (1973) cited this specimen as a female despite Boulenger (1885) considering the same specimen as a male. We have reviewed the same material, and although this specimen does not have everted hemipenes or skin dissections around the base of the tail, the high number of femoral pores (9 on the right thigh and 8 on the left, more than any other Peruvian species) lead us to believe that the specimen is indeed a male, as originally suggested by Boulenger. Additionally, E. guentheri has previously been listed as occurring in Peru (i.e., Köhler & Lehr 2004), nevertheless we were unable to find voucher specimens to corroborate these records. Available specimens correspond to individuals caught in Ecuador, therefore we cannot confirm its presence in Peru and suggest its distribution to be restricted to Ecuadorian territory.

Among Cercosaurini, some species of the genus Anadia from Colombia, Costa Rica, and Venezuela are known for their arboreal habits ( Oftedal 1974, Rivas et al. 2012). Peruvian Gymnophtalmid lizard are usually known to inhabit leaf litter, in contrast to all specimens of E. excelsum sp. nov. captured in the canopy. Although we ignore whether “long snouted” forms share similar arboreal habits, the paucity of historic records of other Peruvian Euspondylus (no more than ten specimens collected since 1845, excluding E. excelsum sp. nov.) may suggest that species in this genus are highly arboreal and frequently missed by traditional, terrestrial-focused surveys. Therefore, we advocate for more frequent use of arboreal surveys sampling canopy habitats. For example, only two individuals of the type series of E. excelsum sp. nov. (male CORBIDI 14666 and female CORBIDI 15573), were collected using traditional Visual Encounter Surveys (both specimens were climbing a tree). All other type specimens were caught during navigation of areas flooded by a hydroelectric project, which allowed us to access to the canopy of the trees of the area. The ten individuals were observed on branches or bromeliads in the canopy that was still above water, suggesting the importance of canopy survey method to properly sample these arboreal lizards and other canopy dwelling herpetofauna.

Despite our present contribution, we consider that herpetofaunal diversity in the Huallaga drainage remains poorly documented ( Venegas et al. 2013; Chávez & Catenazzi, 2016), which motivates us to continue with future research to document its exceptional biodiversity and to contribute to its conservation.

Figures

FIGURE 1. Lateral (A), dorsal (B), and ventral (C) views of the head of the holotype of Euspondylus excelsum sp. nov., male CORBIDI 16457. Photographs by Germán Chávez.

FIGURE 2. Views of the holotype (CORBIDI 16457; SVL = 69.4 mm); A) coloration in life; B) dorsal view of the coloration in preservative; C) ventral view of the coloration in preservative. Photographs by Lesly Luján (A), and Germán Chávez (B, C).

FIGURE 3. Views of the dorsum (left) and venter (right) of the paratypes in life, A – B) Adult male (CORBIDI 16465, SVL = 83.7 mm); C – D) Adult female (CORBIDI 16463, SVL = 81.4); E – F) Juvenile male (CORBIDI 16468, SVL = 47.7). Photographs by Lesly Luján.

FIGURE 4. Sexual dimorphism in Euspondylus excelsum sp. nov., A) ventral view of the tail of a male (CORBIDI 16465) showing black blotches on the entire surface; B) Close up of the ventral side of the thigh of an adult male (CORBIDI 16465) showing the femoral pores (pointed with red arrows); C) ventral view of the tail of a female (CORBIDI 16463) showing black blotches arranged dorsolaterally; D) Close up of the ventral side of the thigh of an adult female (CORBIDI 16463) showing the absence of femoral pores. Photographs by Germán Chávez.

FIGURE 5. Dorsal views of the head of species of Euspondylus, showing the condition of the first supraocular and first superciliar, pointed with red arrows: A) Euspondylus guentheri (BMNH 1946.8.8.99); B) Euspondylus maculatus (MHNN 2105 - 1); C) Euspondylus excelsum sp. nov. (CORBIDI 16457); D) Euspondylus sp 1 (FMNH 45474); E) Euspondylus sp 2 (AMNH 1704). Photographs by Patrick Campbell (A), Alessandro Catenazzi (B, D, E), and Lesly Luján (C).

FIGURE 6. Views of the hemipenes of the holotype of Euspondylus excelsum sp. nov., A) ventral or sulcate side; B) lateral side; C) dorsal or asulcate side. Photographs by Germán Chávez.

FIGURE 7. Condition of the habitat of Euspondylus excelsum sp. nov., A – B) foothills forests flooded by an hydroelectric project, only with the canopy protruding from the water; C) Individual (pointed with a red arrow) perched on a leaf at the level of water surface in a flooded forest; D) Bromeliad inhabited by an individual of E. excelsum as a shelter. Photographs by Lesly Luján.

FIGURE 8. Type locality (yellow star) and localities for the paratypes (yellow triangle) of Euspondylus excelsum sp. nov.; type locality of E. maculatus (green circle); Chanchamayo, Junin department (blue circle), where specimens of Euspondylus sp. 1 were collected, and Inca Mine, Puno department (red circle) where Euspondylus sp. 2 was collected. Map by Caterina H. Cosmópolis.