Two new species of Andean gymnophthalmid lizards of the genus Euspondylus (Reptilia, Squamata) from central and southern Peru

Abstract Two new species of lizards assigned to the genus Euspondylus from the montane forests of the Peruvian Andes in the Pasco Department (central Peru) and Ayacucho Department (southern Peru) both at elevations of 2550 and 3450 m, respectively, are described. The new species are distinguishable from all other Peruvian and Ecuadorian species of Euspondylus by a unique combination of morphometric, scalation and color pattern characteristics. Natural history data for the new species and for Euspondylus spinalis are also provided.

Recent herpetological surveys in the central and southern Andes of Peru resulted in the discovery of two new species of gymnophthalmids, which are described and tentatively assigned to the genus Euspondylus.

Materials and methods
The format for the description of the new species generally follows that of Köhler and Lehr (2004). For the comparisons only Ecuadorian, Peruvian, and Bolivian species of Euspondylus, Opipeuter, Petracola, Proctoporus and Riama were included because our purpose was to distinguish the two new species from any that could occur in sympatry or be similar. Nomenclature of scale characters follows that of Köhler and Lehr (2004). Scale sizes were measured using precision calipers and were rounded to the nearest 0.1 mm. For characters recorded on both sides, the condition on the right side is presented first. Everted hemipenes were fixed with formalin 10%. Abbreviations for museum collections are as follows: CORBIDI (Centro de Ornitología y Biodiversidad) and MUSM (Museo de Historia Natural Universidad Nacional Mayor de San Marcos, Lima, Peru) and GPS coordinates were taken using the geodetic datum WGS84. Diagnosis. 1) Head rounded in dorsal and lateral view, frontonasal length usually equal or slightly larger than frontal length; (2) nasoloreal suture present; (3) supraoculars four, anteriormost supraocular fused with anteriormost superciliary, all supraoculars separated from ciliaries; (4) superciliary series complete, five; (5) supralabial-subocular fusion absent; (6) postoculars three; (7) postparietals three; (8) supratympanic temporals three; (9) genials in two pairs, transverse sutures perpendicular with respect to midline of body; (10) dorsal scales rectangular, juxtaposed, keeled; (11) transverse dorsal count (enlarged rows at midbody) at midbody 20-28 in both sexes; (12) longitudinal dorsal count 35-43 in both sexes; (13) longitudinal ventral count 19-22 in both sexes; (14) lateral scale rows at midbody two or three; (15) femoral pores in males 8-11, in females 7-10; two scales between femoral pores; (16) subdigital scales on 4th finger 10-16, on 4th toe 17-26; (17) limbs overlapping, pentadactyl; digits clawed; forelimb reaching anteriorly to fourth supralabial; (18) anterior preanal plate scales paired; (19) hemipenis acapitate; flounces lacking calcified spines and forming two chevrons on distal half of hemipenis whereas basal half is covered with three transverse flounces; some asulcate flounces separated by a small expansion pleat; sulcate flounces about as wide as asulcate flounces; sulcus spermaticus single, flanked by a broad naked expansion pleat widened distally; (20) dorsum olive green, brown, or reddish brown with a middorsal pale stripe bordered by a discontinous dark line on neck and body more prominent in females than males; lateral ocelli present; ventral surfaces yellowish or reddish white; (21) transparent lower palpebral disc an undivided oval; (22) prefrontals present.
Euspondylus chasqui can be distinguished from all species currently assigned to Petracola, Proctoporus and Riama by the presence of prefrontal scales (absent in all species in these three genera). E. chasqui can be further distinguished by the following character states (condition for E. chasqui in parentheses). All Bolivian and Peruvian species of Proctoporus except P. pachyurus and P. bolivianus: longitudinal dorsal count fewer than 36 scale rows (35-43 scale rows). P. pachyurus: longitudinal dorsal count 49-59 scale rows (35-43 scale rows). P. bolivianus: 4-8 femoral pores in males (7-11 femoral pores). All Petracola and Riama species: lower palpebral disc with vertical sections (palpebral disc an undivided oval). Northern Ecuador species of Riama excluding R. columbiana: no band of granular scales along the sides of body between dorsal and ventral scales (granular scales present). R. columbiana: limbs not overlapping when adpressed against body in adults (limbs overlapping), superciliary series incomplete (complete), and some supraoculars in contact with ciliaries (all supraoculars separated from ciliaries).
Euspondylus chasqui can be distinguished from Opipeuter xestus (condition for E. chasqui in parentheses): smooth dorsal scales (keeled); a single large elongate subocular (several small subocular scales); and in hemipenis morphology, large spines at the base of the sulcus spermaticus (no such spines present in E. chasqui).
Dorsal scales rectangular, longer than wider, juxtaposed, keeled, 40 in a longitudinal count; some middorsal scales irregularly arranged; transverse dorsal count (enlarged rows at midbody) at fifth transverse ventral scale row 16, at 10th transverse ventral scale row 29, at 15th transverse ventral scale row 26; lateral scale rows at fifth transverse ventral scale row 14/ 16, at 10th transverse ventral scale row 4/4, at 15th transverse ventral scale row 3/3; lateral scales on body near insertion of forelimb small to granular; ventrals rectangular and juxtaposed; complete longitudinal ventral count 21; longitudinal ventral scale rows at midbody 12; 49 scales around midbody; anterior preanal plate scales two; posterior preanal plate scales four, all the scales at the same size; scales on tail rectangular and juxtaposed, keeled; at midventral subcaudals squarish.
The completely everted hemipenis is an acapitate organ without a medial welt; apex with two large protrusions separated by the distal end of the sulcus spermaticus; sulcus spermaticus single, flounces lacking calcified spines and forming two chevrons on distal half of hemipenis; sulcate flounces about as wide as asulcate flounces; asulcate flounces becoming shorter distally, three in the basal half and eleven in each protrusion, distal chevrons separated by a small expansion pleat; sulcus spermaticus single, flanked by a broad naked expansion pleat widened distally.
Coloration in preservative. Dorsal surface of head brown, dorsal surface of body and tail bluish brown with a middorsal dark bordered pale stripe on neck and body; lateral ocelli absent; ventral surfaces dirty white suffused with pale blue.
Coloration in life (Fig. 1a, d). Dorsal surface of head olive green; lateral surface of head, around the labial region yellowish orange with dark spots in each labial scale; ventral surface of head, pregular and gular region yellowish orange with dark grey spots on the genials and pregular scales. Dorsal surface of body same color as head, but with black spots in each scale around middorsal region, that form two indistinct and discontinous lines that extend from occiput to posterior hind limbs forming a dark bordered middorsal pale stripe; lateral surface of body same coloration as dorsum with one indistinct ocellus on both sides above insertion of forelimbs, some lateral scales bearing black or small orange spots; ventral surface of body reddish cream (resembling clay). Limbs similar to body, ventral surface of arms olive cream, ventral surface of legs cream. Coloration of dorsal and ventral surfaces of tail like that of body.
Variation (Fig. 1b-d). In the type series, the distinctness of the pale middorsal stripe is more noticeable in females than males, whereas the lateral stripes are obscure in some. Lateral ocelli are present forming a series from three to five ocelli on each side in females, and usually one on each side in males, only one male of the type series (CORBIDI 06967) has three ocelli on each side. Sexual dimorphism is evident in the size of the femoral pores, males have bigger femoral pores than females, but not in their number (8-11 in males versus 7-10 in females), however the main differences between females and males is the SVL (maximum SVL in females = 61.0 mm, maximum SVL in males = 74.0 mm). See Table 1 for variation in selected morphometric and squamation characters in the specimens examined.
Etymology. The specific epithet is based on the Quechua word "chasqui", which refers to the messengers of the Incan empire, men who, on foot, carried the messages throughout the imperial territory in the Cordillera de los Andes where these lizards are found.
Distribution and natural history. Euspondylus chasqui is known from two localities within a studied area of approximately 12 km² in the Río Apurímac valley (Fig 4). It inhabits secondary forests and human settlements. The individuals observed were mostly found at midday under the rocks or foraging between stones, always near medium-sized rocks that they use for hiding. The soil under these rocks is generally more damp compared to the rest of the soil around. A clutch with two eggs was found Holotype. (Fig 3.a- Paratypes. CORBIDI 07214, 07216-18, 07220, 07222, 07224-25 (all females), 07215, 07221, 07229 (all males), 07223 (juvenile), same data as holotype.
Euspondylus oreades can be distinguished from the Peruvian species of Euspondylus by the following character states (condition for E. oreades in parentheses). E. maculatus and E. guentheri: lower palpebral disc with vertical sections (palpebral disc an undivided oval), dorsal scales smooth or wrinkled (keeled), and longitudinal dorsal count 32-37 (36-43). Euspondylus oreades can be distinguished from all species currently assigned to Petracola, Proctoporus, and Riama by the presence of prefrontal scales (absent in all species in these three genera). E. oreades can be further distinguished by the following character states (condition for E. oreades in parentheses) from all Bolivian and Peruvian species of Proctoporus except P. pachyurus and P. bolivianus: longitudinal dorsal count fewer than 36 scale rows (37-43 scale rows). P. pachyurus: longitudinal dorsal count 49-59 (37-43). P. bolivianus: four or five supralabials (six or seven). All Petracola and Riama species: lower palpebral disc with vertical sections (palpebral disc an undivided oval). All northern Ecuadorian Riama species except R. columbiana: no band of granular scales along the sides of body between dorsal and ventral scales (granular scales present). R. columbiana: limbs not overlapping when adpressed against body in adults (limbs overlapping), superciliary series incomplete (complete), and some supraoculars in contact with ciliaries (all supraoculars separated from ciliaries).
Euspondylus oreades can be distinguished from Opipeuter xestus (condition for E. oreades in parentheses): smooth dorsal scales (keeled); having a single large elongate subocular scale (several small subocular scales); and in hemipenis morphology, large spines at the base of the sulcus spermaticus (no such spines present in E. oreades).
Dorsal scales quadrangular, longer than wide, juxtaposed, keeled, 42 in a longitudinal count; some middorsal scales irregularly arranged; transverse dorsal count (enlarged rows at midbody) at fifth transverse ventral scale row 8, at 10th transverse ventral scale row 11, at 15th transverse ventral scale row 11; lateral scale rows at fifth transverse ventral scale row 13/12, at 10th transverse ventral scale row two, at 15th transverse ventral scale row two; lateral scales on body near insertion of forelimb small to granular; ventrals rectangular and juxtaposed; one complete longitudinal at ventral count 22; longitudinal ventral scale rows at midbody 12; 29 scales around midbody; anterior preanal plate scales six; posterior preanal plate scales four (third one not totally developed), all the scales at the same size; scales on tail rectangular and juxtaposed, keeled; midventral subcaudals squarish.
Limbs pentadactyl; digits clawed; forelimb reaching anteriorly to fourth supralabial; dorsal brachial scales polygonal, of varying sizes, subimbricate, smooth; midbra-chial anterodorsal scale at least twice as large as adjacent scales, smooth; anteroventral, ventral, and posteroventral scales roundish, imbricate, smooth; antebrachial scales polygonal, of various sizes; medial antebrachial scales small, polygonal, smooth; dorsal manus scales polygonal, subimbricate; palmar scales small, oval, domelike; dorsal scales on fingers smooth, quadrangular, covering dorsal half of digit, overhanging supradigital scales, two on I, five on II, five on III, six on IV, four on V; subdigital scales four on I, seven on II, 9/10 on III, 7/8 on IV, seven on V; anterodorsal thigh scales polygonal, at least two times as large as adjacent scales, becoming smaller ventrally, smooth; posterodorsal thigh scales small, rounded, arranged irregularly; anterior and anteromedial shank scales roundish, subimbricate, smooth, anteriormost scales many times shorter than lateral, posterolateral, and posteromedial shank scales; lateral, posterolateral, and posteromedial shank scales polygonal or roundish, juxtaposed, smooth; dorsal pes scales polygonal, subimbricate, smooth; scales on dorsal surface of digits single, quadrangular, smooth, overhanging supradigital scales, two on I, 4/5 on II, eight on III, ten on IV,seven on V; subdigital scales single or double, four on I, nine on II, 12/13 on III, seventeen on IV, nine on V; femoral pores seven or eight; two scales between medial most femoral pores.
The completely everted hemipenis is an acapitate organ without a medial welt; apex with two large protrusions separated by the distal end of the sulcus spermaticus; sulcus spermaticus single, flounces lacking calcified spines and forming two chevrons on distal half of hemipenis; sulcate flounces about as wide as asulcate flounces; asulcate flounces becoming shorter distally, two in the basal half and ten, in each protrusion, distal chevrons separated by a small expansion pleat; sulcus spermaticus single, flanked by a broad naked expansion pleat widened distally.
Coloration in preservative. Dorsal surfaces of head, body, and tail brown with a middorsal dark bordered pale stripe on neck; bearing three lateral ocelli in both sides; ventral surface bluish white with black blotches in each scale, ventral surface of the hind limbs and forelimbs pale brown.
Coloration in life (Fig. 3a,b). Dorsal surface of head pale brown; lateral surface of head dark brown, dark and white spots in each labial scale forming labial bars; ventral surface of head creamy white with irregular line on postmental, genials and post genials scales, pregular and gular region creamy white. Dorsal surface of body, on the middorsal region same color as head, but with black blotches in each middorsal scale bordering the middorsal region and forming a dark bordered pale stripe from the occipital region to the posterior insertion of the hind limbs; lateral surface of body dark brown with three ocelli above insertion forelimbs on both sides, some lateral granular scales white and some keeled lateral scales bearing black blotches; ventral surface of body creamy with with black blotches. Limbs similar to body, ventral surface of arms cream with dark spots, ventral surface of legs cream with black blotches. Coloration of dorsal and ventral surfaces of tail like that of body.
Variation. In the type series, the distinctness of the pale middorsal stripes varies. However, at least the bordered stripes are visible in all specimens, whereas the lateral stripes are obscure in some, only CORBIDI 07216, is lacking the bordered stripe (Fig.  3c,d). Lateral ocelli are not present in most of the specimens except CORBIDI 07219 (three ocelli on both sides), 07221 and 07229 (both specimens with a row of ocelli on both sides). Sexual dimorphism is evident in the size of the femoral pores (bigger in males than in females) and in their number (2-5 in females versus 6-8 in males). See Table 1 for variation in selected morphometric and squamation characters in the specimens examined.
Etymology. The specific name oreades refers to the Oreades, nymphs of Greek mythology. These feminine spirits lived and protected isolated mountains and caves, places that recall the type locality where this species was found.
Distribution and natural history. Euspondylus oreades is known only from the type locality, an isolated hill at an elevation of 3400 m in the Cordillera Oriental in central Peru, inside Yanachaga-Chemillen National Park (Oxapampa Bioesphere Reserve) (Fig. 4). Individuals were found in grassland (Puna habitat) under rocks, fallen trunks, moss, and under the base of terrestrial spiny bromeliad (Puya sp.) by the day. Only the marsupial frog Gastrotheca griswoldi was found sympatric with E. oreades. A total of 33 individuals of E. oreades were found in seven hours of survey by four herpetologists. Four nests of the new species of Euspondylus were found under the rocks and the number of eggs found per nest vary from two to 15. Two eggs of E. oreades hatched during the surveys and the new hatchlings ran to hide immediately after leaving the egg shell. One of these hatchlings was collected (CORBIDI 07223) and measured (SVL = 23.0 mm). Six females were collected, five of them contained eggs in their oviducts, only CORBIDI 07216 contained one egg, the rest of mature females contained two eggs, SVL range of these specimens was 53.0-61.0 mm. Egg length ranged from 3.2-13.2 mm (x-= 8.6 mm, n= 9) and width ranged from 3.6-5.7 mm (x-= 3.7 mm, n=9). The ornithological team in Santa Barbara collected one Variable Hawk Buteo polyosoma (CORBIDI/FHC 245) that contained three whole E. oreades individuals in the crop and the remains of three other individuals in the stomach. Furthermore, the team collected one Andean Caracara Phalcoboenus megalopterus (CORBIDI/WV 315) with remains of an unassigned Euspondylus species in its stomach. These findings could suggest that these lizards can be found in the open, but for E. oreades we did not have such observations. However, one of us (GC) found E. chasqui running between stones when sampling, which might be a behavior that could occur in E. oreades as well.
Both species described in this paper, Euspondylus chasqui and E. oreades, are abundant at their respective type localities and, where surveyed, are not sympatric with any other lizard species. The only species that is distributed close to Euspondylus oreades is E. spinalis (Fig. 2 e,f ). Even though there is no natural history data published for E. spi-nalis, we have observed that the latter occurs in montane habitats, while E. oreades occurs in grasslands. We have also found that E. spinalis, was the most abundant species at two localities: Chacos Community (10°35'24.2" S, 75°16'24.4" W, 1986 m) and Oso Playa Road (10°19'21.5" S, 75°35'03.1" W, 2000 m), in the Pasco Department (Fig. 4), with 14 individuals found in two hours by one surveyer and 87 individuals in two hours by two surveyers, respectively. Only in Chacos did we find E. spinalis in sympatry with the iguanid lizard Stenocercus boettgeri, even though, S. boettgeri was not abundant (only three individuals recorded). Given the high abundances observed of these gymnophthalmid lizards, it is likely that they play an important role in the lizard community composition, and apparently, in the trophic chain of certain major predators, as evidenced by the records of Euspondylus lizards found in the stomach contents of Buteo polyosoma and Phalcoboenus megalopterus.