Pholidobolus dolichoderes, Parra & Nunes & Torres-Carvajal, 2020

Parra, Vanessa, Nunes, Pedro M. Sales & Torres-Carvajal, Omar, 2020, Systematics of Pholidobolus lizards (Squamata, Gymnophthalmidae) from southern Ecuador, with descriptions of four new species, ZooKeys 954, pp. 109-156 : 109

Pholidobolus dolichoderes sp. nov. Figures 12 , 13 , 14 Proposed standard English name: Long-necked cuilanes Proposed standard Spanish name: Cuilanes de cuello largo

Holotype.

QCAZ 16353 (Figs 12 , 13 ), adult male, Ecuador, Provincia Azuay, San Felipe de Oña, 3.4292S, 79.2364W, WGS84, 2672 m, 16 March 2018, collected by Diego Almeida, Darwin Núñez, Eloy Nusirquia, Alex Achig and Katherine Nicolalde.

Paratypes (4).

Ecuador: Provincia Azuay: QCAZ 16349, 16352 (adult females), San Felipe de Oña, Susudel-Poetate road, 3.4322S, 79.2369W, WGS84, 2506 m, 16 March 2018; QCAZ 16350-51 (juveniles), San Felipe de Oña, 3.4275S, 79.2339W, WGS84, 2675 m, 16 March 2018, same collectors as holotype.

Diagnosis.

Pholidobolus dolichoderes is unique among its congeners in having a long neck with granular scales between the posterior corner of the orbit and the anterior edge of the tympanum, as well as an inconspicuous ventrolateral fold between fore and hindlimbs. In addition, P. ulisesi , P. dicrus , P. hillisi , and P. vertebralis differ from P. dolichoderes in having a conspicuous light vertebral stripe. The new species further differs from P. affinis in lacking ocelli on flanks, and from P. condor sp. nov., P. macbrydei , and P. montium in having prefrontal scales. Pholidobolus dolichoderes has more dorsals (35-40) and ventrals (25-27) than P. samek sp. nov. (27-29 and 19-21, respectively) and P. condor sp. nov. (26-30 and 18-20), and, unlike P. fascinatus sp. nov., it has widened medial scales on collar. In addition, P. dolichoderes has more temporals (7-9) and gulars (22-23) than P. samek sp. nov. (4-5 and 15-18, respectively), P. condor sp. nov. (4-5 and 14-16), and P. fascinatus sp. nov. (3-5 and 14-17).

Characterization.

(1) Three supraoculars, anteriormost larger than posterior ones; (2) prefrontals present; (3) femoral pores present in both sexes; (4) four to six opaque lower eyelid scales; (5) scales on dorsal surface of neck smooth, becoming slightly keeled from forelimbs to tail; (6) two or three rows of lateral granules at midbody; (7) 35-20 dorsal scales between occipital and posterior margin of hindlimb; (8) lateral body fold present but inconspicuous; (9) keeled ventrolateral scales on each side absent; (10) dorsum dark brown with a diffuse pale brown vertebral stripe that becomes grayish brown towards tail; (11) labial stripe white; (12) flanks of body gray brown; (13) white stripe along forelimb present; (14) hemipenial body cylindrical, with sulcus spermaticus originating between thick lips.

Description of holotype.

Adult male (QCAZ 16353) (Figs 12 , 13 ); SVL 41.1 mm; TL 96.3 mm; dorsal and lateral head scales imbricated, smooth; rostral hexagonal, 1.75 times as wide as high; frontonasal heptagonal, slightly wider than long, laterally in contact with nasal, similar in size to frontal; prefrontals present, in wide contact medially, and in contact with loreal and first superciliary laterally; frontal hexagonal, longer than wide, wider anteriorly, in contact with first and second supraoculars; frontoparietals hexagonal, longer than wide, slightly wider posteriorly, each in contact with second and third supraoculars, parietals and interparietal; interparietal heptagonal, lateral borders nearly parallel to each other; parietals wider than interparietal, heptagonal, and positioned anterolaterally to interparietal, each in contact with third supraocular and dorsalmost postocular; postparietals three, medial scale smaller than lateral ones; seven supralabials, fourth one the longest and below center of eye; five infralabials, fourth one below center of eye; temporals small, irregularly, smooth; supratemporal scales not well differentiated, smooth; nasal shield divided above the nostril, longer than high, in contact with rostral anteriorly, first and second supralabials ventrally, frontonasal dorsally, loreal posteriorly; loreal pentagonal, slightly wider dorsally, in contact with second and third supralabials; frenocular longer than high, in contact with loreal; three supraoculars, with the first one being the widest; four elongate superciliaries, anteriormost one enlarged, in contact with loreal; palpebral disc oval, pigmented, divided into four scales; four suboculars, two elongated and similar in size, the anteriormost and posteriormost larger than the others; three postoculars, dorsalmost wider than the others; ear opening vertically oval, without denticulate margins; tympanum recessed into a shallow auditory meatus; mental wider than long; postmental pentagonal, slightly wider than long, followed posteriorly by three pairs of genials, the anterior two pairs in contact medially and the third pair separated by postgenials; all genials in contact with infralabials; gulars imbricate, smooth, widened in two longitudinal rows; gular fold complete, posterior row of gulars (collar) with six scales, the medial two distinctly widened.

Nuchal scales slightly smaller than dorsals, except for the anteriormost that are widened; scales on sides of neck small and granular; dorsal scales elongate, juxtaposed, arranged in transverse rows; scales on dorsal surface of neck striated, becoming slightly keeled from forelimbs to tail; dorsal scales between occipital and posterior margin of hindlimbs 35; dorsal scale rows in a transverse line at midbody 32; one longitudinal row of smooth, enlarged ventrolateral scales on each side; dorsals separated from ventrals by three rows of granular scales at level of 13th row of ventrals; lateral body fold between fore and hindlimbs poorly defined; ventrals smooth, arranged in 26 transverse rows between collar fold and preanals; six ventral scales in a transverse row at midbody; subcaudals smooth; axillary region with granular scales; scales on dorsal surface of forelimb smooth, imbricate; scales on ventral surface of forelimb granular; two thick, smooth thenar scales; supradigitals (left/right) 3/0 on finger I, 7/7 on II, 9/8 on III, 10/10 on IV, 5/5 on V; supradigitals 4/4 on toe I, 7/7 on II, 11/11 on III, 12/11 on IV, 9/8 on V; subdigital lamellae of fingers I and II mostly single, III and IV paired proximally, on finger V all single; subdigital lamellae 5 on left finger I (right finger missing), 10/10 on II, 14/14 on III, 14/14 on IV, 9/9 on V; subdigital lamellae on toe I single, on toe II paired at the middle, on toe III and IV paired along proximal half, and on toe V paired proximally; subdigital lamellae 5/5 on toe I, 10/10 on II, 14/14 on III, 18/19 on IV, 11/11 on V; groin region with small, imbricate scales; scales on dorsal surface of hindlimbs striated and imbricate; scales on ventral surface of hindlimbs smooth; scales on posterior surface of hindlimbs granular; femoral pores present, three on left leg and five on right leg; preanal pores absent; cloacal plate paired, bordered by four scales anteriorly, of which the two medialmost are enlarged.

Additional measurements (mm) and proportions of the holotype: HL 9.8; HW 6.2; ShL 5.4; AGD 20.7; TL/SVL 2.4; HL/SVL 0.2; HW/SVL 0.1; ShL/SVL 0.1; AGD/SVL 0.5.

Color of holotype in life.

Dorsal background of head dark brown; diffuse pale brown vertebral stripe that becomes grayish brown towards tail; creamy white dorsolateral stripes on head extending posteriorly and fading away at midbody; white longitudinal stripe extending from first supralabial to shoulder; sides of neck brown; flanks grayish brown with diffuse dark brown marks; limbs brown; ventrolateral region of body grayish brown; throat and chest cream; belly grayish cream; base of tail gray with dark little spots (Figs 12 , 14B ).

Color of holotype in preservative.

Dorsal background uniformly brown with a diffuse light brown vertebral stripe extending from occiput onto tail, but fading at posterior end of body; dorsal and ventral surface of head brown; flanks light brown, with scattered dark brown spots; head and neck with two distinct white longitudinal stripes, the ventral one extending from first supralabial to forelimb, and the dorsal one from canthus rostralis to scapular region, posterior to which if fades into a light brown stripe; lateral aspect of neck dark brown; tail grayish brown; gular, chest and venter regions pale gray; ventral surface of tail and limbs gray.

Variations.

Measurements and scutellation data of Pholidobolus dolichoderes are presented in Table 6 . Superciliaries 4/5 (left/right) in specimen QCAZ 16350; palpebral disc divided into 5/6 scales in QCAZ 16352 and 3/5 in QCAZ 16351; frontonasal pentagonal in QCAZ 16349-52; prefrontals pentagonal in QCAZ 16349, 16350 and 16352; two rows of lateral granules at midbody in QCAZ 16439, 16350 and 16351. Usually six gular (collar) scales, eight in QCAZ 16349. Male is smaller (SVL 41.1 mm, N = 1) than females (maximum SVL 48.1 mm, N = 2).

Adult females differ from holotype in having a grayish brown vertebral stripe, fading away posteriorly, and grayish brown flanks (Fig. 14 ). Juvenile QCAZ 16350 differs from holotype in having grayish brown flanks, without scattered dark brown spots; juvenile QCAZ 16351 is unique in having white spots on flanks and over forelimbs.

Distribution and natural history.

Pholidobolus dolichoderes is known to occur between 2506-2675 m in San Felipe de Oña, southwestern Azuay province (Fig. 7 ). This area is composed of many different landscapes including small valleys, desert areas and wet paramo. Most specimens were found active at day (10h26-15h30), mostly on the ground or near spiny ground bromeliads known as achupallas ( Puya sp.).

Conservation status.

Pholidobolus dolichoderes is only known from unprotected localities around Oña. The population size of this species is unknown, but our sampling suggests low abundances. Because of the small known distribution and lack of additional data, we suggest assigning P. dolichoderes to the Data Deficient category according to IUCN (2012) guidelines.

Etymology.

The specific epithet dolichoderes derives from the Greek words dolikhós, meaning long, and derē, meaning neck, in allusion to the distinctively long neck of this species.

Figures

Figure 7. Distribution of samples of the " Pholidobolus macbrydei " species complex included in phylogenetic analyses. Circles correspond to four new species described in this paper: P. samek sp. nov. (red), P. condor sp. nov. (blue), P. dolichoderes sp. nov. (brown), and P. fascinatus sp. nov. (green). Triangles are " Pholidobolus macbrydei " clades as illustrated in the phylogenetic tree (Fig. 1): A (green) B (white) C (red) D (blue) E (black) F (turquoise). Orange diamond corresponds to type locality of P. macbrydei. This map was created in QGIS v 3.10.

Figure 12. Holotype of Pholidobolus dolichoderes sp. nov. (QCAZ 16353) in life in dorsal (A), ventral (B), and lateral (C) views. Male, SVL = 41.1 mm. Photographs by Gustavo Pazmino.

Figure 13. Head of holotype of Pholidobolus dolichoderes sp. nov. (QCAZ 16353) in lateral (A), dorsal (B), and ventral (C) views. Photographs by Valeria Chasiluisa. Scale bar: 5 mm.

Figure 14. Close-up of head and neck of Pholidobolus dolichoderes sp. nov. in life. QCAZ 16349 (A adult female); QCAZ 16353 (B male holotype). Photographs by Gustavo Pazmino.