Euspondylus oreades, Chavez, German, Siu-Ting, Karen, Duran, Vilma & Venegas, Pablo J, 2011

Chavez, German, Siu-Ting, Karen, Duran, Vilma & Venegas, Pablo J, 2011, Two new species of Andean gymnophthalmid lizards of the genus Euspondylus (Reptilia, Squamata) from central and southern Peru, ZooKeys 109, pp. 1-17 : 8-13

Euspondylus oreades sp. n. Figs 2-3

Holotype.

(Fig 3.a-b) Adult male (CORBIDI 07219), Perú, Pasco Department, Santa Barbara (10°20'29.1"S, 75°38'27.1"W), at 3439 m of elevation, collected by Caroll Z. Landauro, Lesly Luján, Vilma Duran and Pablo J. Venegas on 23 September 2010.

Paratypes.

CORBIDI 07214, 07216-18, 07220, 07222, 07224-25 (all females), 07215, 07221, 07229 (all males), 07223 (juvenile), same data as holotype.

Diagnosis.

1) 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, four; (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 quadrangular, juxtaposed, keeled; (11) transverse dorsal count (enlarged rows at midbody) at midbody 20-26 in both sexes; (12) longitudinal dorsal count 37-43 in both sexes; (13) longitudinal ventral count 20-22 in both sexes; (14) lateral scale rows at midbody two; (15) femoral pores in males 3-8, in females 2-8; two scales between femoral pores; (16) subdigital scales on 4th finger 5-13, on 4th toe 10-19; (17) limbs overlapping, pentadactyl; digits clawed; forelimb reaching anteriorly to fourth supralabial; (18) anterior preanal plate scales paired; (19) hemipenis acapitate; flounces forming two chevrons on distal half of hemipenis whereas basal half is covered with one or two transverse flounces; 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 and divided by a small protrusion; (20) dorsum brown or pale brown with a middorsal pale stripe bordered by an discontinous dark line on neck and body; lateral ocelli usually absent; ventral surfaces white or creamy white; (21) transparent lower palpebral disc an undivided oval; (22) prefrontals usually present.

Euspondylus oreades can be distinguished from the Peruvian species of Euspondylus by the following character states (condition for Euspondylus oreades in parentheses). Euspondylus maculatus and Euspondylus 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 caideni : maximum SVL = 82.0 mm (61.0 mm), pale middorsal stripe absent (present), dorsal scales reduced in size above longitudinal band of laterals granules (not reduced), longitudinal dorsal count 41-48 (36-43). Euspondylus josyi : pale middorsal stripe absent (present), supraoculars three, exceptionally four (always four) limb overlapping 10-13 dorsal scales (10-11) and longitudinal dorsal count 29-35 (36-43). Euspondylus rahmi : anteriormost supraocular not fused with anterior most superciliar (fused), dorsal scales reduced in size above longitudinal band of lateral granules (not reduced), longitudinal dorsal count 49-54 (36-43) and maximum SVL to 71.0 mm (61.0 mm). Euspondylus simonsii : a pale line between the tympanum and shoulder present (absent), dorsal scales smooth or only faintly keeled on posterior dorsum (all dorsal scales keeled), transversal count at midbody less than 35 scales (40-46) and longitudinal dorsal count 33-39 (36-43). Euspondylus spinalis : head acuminate from dorsal and lateral view (rounded), dorsal scales reduced in size above longitudinal band of lateral granules (not reduced), subdigital lamellae of fourth toe 20-24 (11-19), longitudinal dorsal count 39-46 (32-43), femoral pores on one side in females 1-6 (4-8), supracaudal scales only faintly keeled or smooth (strongly keeled) and lateral ocelli usually present (usually absent). Euspondylus nellycarrillae : dorsal scales subhexagonal (rectangular), longitudinal dorsal count 41-49 (36-43) femoral pores of one side 12-15 in males, 12-14 in females (6-8 in males, 2-5 in females). Euspondylus chasqui : superciliar series five (four), subdigital scales on the fourth toe 17-26 (13-19), femoral pores of the one side 8-11 in males, 7-10 in females (6-8 in males, 2-5 in females) and maximum SVL = 74.0 mm (61.0 mm).

Euspondylus oreades can be distinguished from all species currently assigned to Petracola, Proctoporus , and Riama by the presence of prefrontal scales (absent in all speciesin these three genera). Euspondylus oreades can be further distinguished by the following character states (condition for Euspondylus oreades in parentheses) from all Bolivian and Peruvian species of Proctoporus except Proctoporus pachyurus and Proctoporus bolivianus : longitudinal dorsal count fewer than 36 scale rows (37-43 scale rows). Proctoporus pachyurus : longitudinal dorsal count 49-59 (37-43). Proctoporus 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 Riama columbiana : no band of granular scales along the sides of body between dorsal and ventral scales (granular scales present). Riama 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 Euspondylus 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 Euspondylus oreades ).

Description of the holotype.

Adult male (CORBIDI 07219) (Fig. 2c,d; 3a,b); SVL 55.0 mm, tail (complete) length 101.0 mm; axilla to groin distance 26.7 mm; head length 13.2 mm; head width 8.8 mm; shank length 7.2 mm. Head scales smooth, glossy; rostral scale wider (2.4 mm) than long (1.4 mm), slightly higher than adjacent supralabials, in contact with frontonasal, nasoloreal, and first supralabials posteriorly; frontonasal wider (2.8 mm) than long (2.6 mm), widest posteriorly, in contact with nasoloreal and frenocular laterally, prefrontals posteriorly; nasoloreal suture present; prefrontals present, in contact with each other medially, in contact with fused anteriormost superciliary-anteriormost supraocular, frontal posteriorly; frontal slightly longer (2.3 mm) than wide (2.2 mm), anterior suture angular with point directed anteriorly, lateral sutures straight, posterior suture angular with point directed posteriorly, in contact with second and third 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 (3.2 mm) than wide (1.7 mm), in contact with parietals laterally, postparietals posteriorly; parietals pentagonal, in contact with fourth supraocular anterolaterally, temporal scales laterally, dorsalmost postocular, postparietals posteriorly; postparietals three, lateral postparietals pentagonal, medial postparietal squarish; palpebral disc an undivided oval, unpigmented; frenocular quadrangular, in contact with nasoloreal anteriorly; postoculars three; temporals polygonal; supratympanic temporals three; supralabials eight; infralabials seven; mental wider (2.3 mm) than long (1.4 mm), in contact with first infralabials, postmental posteriorly; postmental single, pentagonal, posterior suture angular, point directed posteriorly, in contact with first and second infralabials; genials in two pairs, both pairs subquadrangular, in contact with second and third infralabials; posterior genials subpentagonal, in contact with fourth and fifth infralabials laterally; scale rows between genials and collar fold (along midventral line) 10; medialmost scales of three penultimate gular scale rows slightly enlarged; posteriormost gular row enfolded posteriorly, concealing two granular scale rows; lateral neck scales rounded, smooth.

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; midbr achial 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 Euspondylus oreades . A total of 33 individuals of Euspondylus 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 Euspondylus 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 Euspondylus 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 Euspondylus oreades we did not have such observations. However, one of us (GC) found Euspondylus chasqui running between stones when sampling, which might be a behavior that could occur in Euspondylus oreades as well.

Remarks.

Most of the Euspondylus species occur on, isolated points in the Andes ( Kizirian 1996; Köhler 2003; Köhler and Lehr 2004), in montane forests or transition areas between montane forests and grasslands. Several genera from the families Gymnophthalmidae and Iguanidae have been recorded in the high Andes ( Fritts 1972; Cadle 1991; Laurent 1992; Laurent 1998). Of these Anadia , Euspondylus, Petracola , Pholidobolus , Proctoporus and Riama ( Gymnophthalmidae ) and Stenocercus and Liolaemus ( Iguanidae ) are the only genera found above 3000 meters of elevation.

Both species described in this paper, Euspondylus chasqui and Euspondylus 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 Euspondylus spinalis (Fig. 2 e,f). Even though there is no natural history data published for Euspondylus spinalis , we have observed that the latter occurs in montane habitats, while Euspondylus oreades occurs in grasslands. We have also found that Euspondylus spinalis , was the most abundant species at two localities: Chacos Community (0°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 Euspondylus spinalis in sympatry with the iguanid lizard Stenocercus boettgeri , even though, Stenocercus 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 .