Pholidobolus ulisesi, Venegas & Echevarría & Lobos & Sales Nunes & Abstract.-Based & The, 2016

Venegas, Pablo J., Echevarría, Lourdes Y., Lobos, Simón E., Sales Nunes, Pedro M., Abstract. - Based, Omar Torres-Carvajal & The, Peru., 2016, A new species of Andean microteiid lizard (Gymnophthalmidae: Cercosaurinae: Pholidobolus) from Peru, with comments on P. vertebralis, Amphibian & Reptile Conservation (e 121) 10 (1), pp. 21-33 : 22-27

publication ID	https://doi.org/ 10.5281/zenodo.11376950
DOI	https://doi.org/10.5281/zenodo.11377048
persistent identifier	https://treatment.plazi.org/id/0385EC6C-F366-FE1F-FC8B-FE26FC2BE9E6
treatment provided by	Felipe
scientific name	Pholidobolus ulisesi
status	sp. nov.

Treatment

Pholidobolus ulisesi View in CoL sp. nov.

urn:lsid:zoobank.org:act:283DAECE-3FD5-496D-963B-A4E8E4DC8CA7

Figs. 1–3 View Fig View Fig View Fig .

Cercosaura vertebralis — Doan and Cusi 2014 (part): 1,195 –1,200.

Pholidobolus sp. — Torres-Carvajal et al. 2015: 286.

Pholidobolus sp. — Torres-Carvajal et al. 2016: 70 ( Fig. 2 View Fig ).

Holotype: CORBIDI 12734 View Materials , an adult male from Bosque de Huamantanga (5°39’48.09’’ S, 78°56’35.8’’ W), at 2,211 m elevation, Huabal district, Jaén province, Cajamarca department, Peru, collected on 7 March 2013 by P.J. Venegas. GoogleMaps

Paratypes (17): CORBIDI 12740– 46 juveniles , CORBIDI 12735–36 View Materials , 12739 View Materials adult males , CORBIDI 12737– 38 adult females, all collected with the holotype ; CORBIDI 00871–73 View Materials , an adult female, an adult male and a juvenile, respectively, from El Chaupe (5°14´8.16” S, 79°5´56.58” W), at 2,016 m elevation, Namballe district , San Ignacio province, Cajamarca department, Peru, collected by M. Dobiey on 24 August 2008 GoogleMaps ; CORBIDI 14889 View Materials , an adult female, and CORBIDI 14896 View Materials , a juvenile, from San Felipe de Jaén (5°45’10.854” S, 79°14’19.881” W), at 2,641 m elevation, Jaén province, Cajamarca department, Peru collected by K. Garcia on 26 September 2014 GoogleMaps .

Photo voucher specimen: Cañaris (6°03´26.18´´S, 79°16´00.35´´W), at 2,318 m elevation, Ferreñafe province , Lambayeque department, Peru, captured and released by P.J. Venegas on 25 May 2007 ( Fig. 3D View Fig ) GoogleMaps .

Diagnosis: Pholidobolus affinis , P. dicrus ( Fig. 4A View Fig ), P. hillisi ( Fig. 4B View Fig ), P. prefrontalis , and P. vertebralis ( Fig. 4C View Fig ) differ from the new species in having prefrontal scales. Pholidobolus montium and P. macbrydei have striated and quadrangular dorsal scales (strongly keeled and hexagonal in P. ulisesi ), and lack the conspicuous narrow, pale brown, vertebral stripe present in P. ulisesi . In addition, the new species has fewer dorsal scales (28–31, x = 29.75) than P. affinis (45–55), P. montium (35–50), P. prefrontalis (37–46), and P. macbrydei (31–43).

Characterization: (1) Two or three supraoculars, anteriormost larger than others; (2) prefrontals absent; (3) femoral pores absent in both sexes; (4) two to six opaque lower eyelid scales; (5) scales on dorsal surface of neck striated, becoming strongly keeled between forelimbs and tail; (6) two or three rows of lateral granules at midbody; (7) lateral body fold present; (8) usually two rows of keeled ventrolateral scales on each side; (9) dorsum dark brown with a distinct pale brown middorsal stripe, slender at midbody, becoming grayish brown towards the tail; (10) labial stripe white becoming cream or pale brown along ventrolateral region; (11) sides of body dark brown; (12) cream stripe along forearm; (13) a distinct diagonal white bar with dark brown edges on each side of the mandible, extending from sixth infralabial to proximal pregular; (14) orange spots on sides of body, usually above forelimb and the base of tail in adult males.

Description of holotype: Adult male (CORBIDI 12734; Fig. 1–3A View Fig View Fig View Fig ); SVL 45. 5 mm; TL 104 mm; dorsal and lateral head scales juxtaposed, finely wrinkled; rostral hexagonal, 2.03 times as wide as high; frontonasal quadrangular, wider than long, longer than frontal, laterally in contact with nasal, loreal, and first superciliary; prefrontals absent; frontal pentagonal, longer than wide, slightly wider anteriorly, in contact with frontonasal and supraocular I on each side; frontoparietals hexagonal, longer than wide, with medial suture, each in contact laterally with supraoculars I and II; interparietal roughly heptagonal, its lateral borders parallel to each other; parietals slightly smaller than interparietal, pentagonal and positioned anterolaterally to interparietal, each in contact anteriorly with supraocular II and dorsalmost postocular; postparietals three, medial scale smaller than laterals; supralabials seven, fourth longest and below the center of eye; infralabials five, fourth below the center of eye; temporals enlarged, irregularly pentagonal or hexagonal, juxtaposed, finely wrinkled; two finely wrinkled supratemporals, dorsal conspicuously larger than ventral one; nasal divided, irregularly tetragonal, longer than wide, in contact with rostral anteriorly, first and second supralabials ventrally, frontonasal dorsally, loreal posterodorsally and frenocular posteroventrally; nostril on ventral aspect of nasal, directed lateroposteriorly, piercing nasal suture; loreal rectangular; frenocular enlarged, in contact with nasal, separating loreal from supralabials; supraoculars two, with the first being the largest; four elongate superciliaries, first one enlarged, in contact with loreal; palpebral disk divided into two pigmented scales; suboculars three, elongated and similar in size; three postoculars, ventral one smaller than the others; ear opening vertically oval, without denticulate margins; tympanum recessed into a shallow auditory meatus; mental semicircular, wider than long; postmental pentagonal, slightly wider than long, followed posteriorly by three pairs of genials, the anterior two in contact medially and the posterior one separated by postgenials; all genials in contact with infralabials; gulars imbricate, smooth, widened in two longitudinal rows; gular fold incomplete; posterior row of gulars (collar) with two enlarged scales medially, larger than the anterior gulars.

Scales on nape similar in size to dorsals, except for the anteriormost that are widened; scales on sides of neck small and granular; dorsal scales elongated, imbricate, arranged in transverse rows; dorsal scales on nape striated, becoming progressively keeled from forelimbs to tail; number of dorsal scales between occipital and posterior margin of hind limbs 30; dorsal scale rows in a transverse line at midbody 19; dorsals separated from ventrals by two longitudinal rows of large keeled scales on each side; longitudinal fold between fore and hind limbs present; ventrals smooth, wider than long, arranged in 21 transverse rows between collar fold and preanals; six ventral scales in a transverse row at midbody; subcaudals smooth; limbs overlap when adpressed against body; axillary region composed of granular scales; scales on dorsal surface of forelimb striated, imbricate; scales on ventral surface of forearm small and imbricate, those on ventral surface of arm granular; two thick, smooth thenar scales; supradigitals (left/right) 3/3 on finger I, 6/6 on II, 8/8 on III, 9/9 on IV, 6/6 on V; supradigitals 3/3 on toe I, 6/6 on II, 10/9 on III, 12/11 on IV, 8/8 on V; subdigital lamellae of forelimb single, 6/6 on finger I, 11/12 on II, 15/16 on III, 16/16 on IV, 9/8 on V; subdigital lamellae on toes I and II single, on toe III paired on the middle, on toe IV paired except for a few ones, on toe V paired at the base; number of subdigital lamellae (pairs when applicable) 6/6 on toe I, 10/11 on II, 16/17 on III, 21/21 on IV, 12/12 on V; groin region with small keeled, imbricate scales; scales on dorsal surface of hind limbs keeled and imbricate; scales on ventral surface of hind limbs smooth; scales on posterior surface of thighs granular and on shanks striated and imbricate; femoral pores absent; preanal pores absent; cloacal plate paired, bordered by two scales anteriorly, smaller than cloacal scales.

Additional measurements (mm) and proportions of the holotype: HL 9.91; HW 6.95; ShL 3.9; AGD 25.6; TL/SVL 2.05; HL/SVL 0.21; HW/SVL 0.15; ShL/SVL 0.08; AGD/SVL 0.56.

Coloration in preservative ( Figs. 1 View Fig and 2 View Fig ): Dorsum dark brown with a grayish brown vertebral stripe that is four scales broad at midbody, and extends from occiput onto tail; vertebral stripe wide anteriorly becoming slightly slender at midbody; dorsal surface of head brown, sides of head and body dark brown; two bright cream spots on each side above insertion of forelimbs; light stripe extending ventrolaterally from lips to insertion of hind limbs, white on lips and grayish brown along the body; a distinct diagonal white bar with dark edges on each side of the mandible, extending from the sixth infralabial onto the proximal pregular; dorsal surface of limbs dark brown with a cream stripe along the arms; gular region pale gray, chest and venter dark gray; ventral surface of tail dark gray.

Coloration of holotype in life ( Fig. 3A View Fig ): Similar to that in preservative, but the bright cream spots on each side above forelimbs are replaced by two black ocelli with red centers, and the sides of the base of the tail have scattered red flecks. The iris is light brown.

Variation: Variation in measurements and scutellation of Pholidobolus ulisesi is presented in Table 1 View Table 1 . Usually two supraoculars, 2/3 (left/right) in specimen CORBIDI 12742 View Materials ; superciliaries usually four, 3/ 4 in CORBIDI 12749 View Materials , 6 View Materials / 5 in CORBIDI 00873 View Materials , and 5/ 5 in CORBIDI 00872 View Materials ; little intrusive scales present on each side, in the posterior angle of frontonasal in three specimens ( CORBIDI 12735 View Materials , 12741 View Materials , 12744 View Materials ); usually seven supralabials, 7/ 6 in CORBIDI 00871 View Materials , 12738 View Materials and 6/ 6 in CORBIDI 12742–43 View Materials ; infralabials usually six, 5/ 5 in CORBIDI 12738 View Materials , 12740 View Materials , 12742 View Materials , 6 View Materials / 5 in CORBIDI 00873 View Materials , 12744 View Materials and 5/ 6 in CORBIDI 12735 View Materials , 12743 View Materials . Rows of ventrolateral keeled scales vary from two rows in nine specimens (56% of the type series), one row on each side in three specimens ( CORBIDI 00872 View Materials , 12741 View Materials , and 12745), three rows on each side in one specimen ( CORBIDI 12739 View Materials ), and absent in two adult specimens ( CORBIDI 00871 View Materials and CORBIDI 00873 View Materials ). Usually two scales on posterior cloacal plate, only two specimens ( CORBIDI 12737–38 View Materials ) have three scales, and two other specimens ( CORBIDI 00871 View Materials and 00873) have four scales.

(maximum SVL 57. 4 mm, n = 4) than males (maximum SVL 45. 5 mm, n = 5). Juvenile CORBIDI 12743 ( Fig. 3C View Fig ) differs from adults in having a fragmented dirty cream stripe along the flanks above the ventrolateral stripe.

Males can be distinguished from females by having the contacted margins of rostral and mental distinctly dark brown or black (indistinct or not contrasting in females), and by the presence of red or orange spots above the insertion of forelimbs and on the sides of the base of tail (absent in females; Fig. 3B View Fig ). Females are longer Hemipenial morphology: The left hemipenis of the holotype of Pholidobolus ulisesi ( Fig. 5 View Fig ) was everted during preservation and prepared posteriorly. The organ extends along approximately eight millimeters in length. The lobes of the organ are partially everted and the hemipenis is fully expanded. The hemipenial body is roughly conical in shape, with the basis distinctly thinner than the rest of the organ, and bears two small lobes with apical folds in the apex. The sulcus 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 fleshy fold; its branches lie on the medial region of the lobes, and end in their tips among folds. The sulcate face of the hemipenial body presents two nude areas parallel to the sulcus spermaticus that run along the entire hemipenial body.

The lateral and asulcate faces of the hemipenis are ornamented with a series of roughly equidistant flounces with calcareous spinules. Twenty-three rows of flounces extend along the body of the organ. There are four proximal rows restricted to a central position on the basal asulcate face of the hemipenis, all of them are roughly chevron-shaped. The four proximal flounces on the sides are diagonally positioned; the third to fifth flounces are separated from a complementary flounce positioned on the asulcate face and oriented in an inverse diagonal. The subsequent flounces towards the lobes cross the sides of the organ from the sulcate to the asulcate face, forming chevrons with vertices in the central region of each side pointing towards the basis of the organ. These chevron-shaped rows become reduced in size progressively towards the hemipenial apex. Similar to the description of the hemipenis of Cercosaura vertebralis by Uzzell (1973), the five distalmost lateral flounces of the hemipenis have an enlarged tooth in the vertex of the chevrons.

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, becoming Y-shaped. The region between the asulcate and lateral sides are marked by a conspicuous unevenness forming a distinctive bulge, which is also present in other species of the Macropholidus + Pholidobolus clade ( Macropholidus annectens , M. huancabambae , M. ruthveni , Pholidobolus affinis , P. hillisi , P. macbrydei , P. montium , P. prefrontalis , P. vertebralis; Nunes, 2011; Torres-Carvajal et al. 2014).

The hemipenis of the holotype of P. ulisesi described herein ( Fig. 5 View Fig ) is broadly congruent with the illustrated by Doan and Cusi (2014) for a specimen of P. ulisesi , considered by them as P. vertebralis (see “Discussion” hereafter). Although Doan and Cusi (2014) reported a reduced count of flounces ornamenting the organ (14 versus 23 in the holotype of P. ulisesi ), their Fig. 5B View Fig clearly shows at least 18 visible flounces ornamenting the hemipenis sides, plus other flounces not countable due the positioning of the organ and the lack of focus in some areas of the hemipenis photograph. Similar to the hemipenis of P. ulisesi described by Doan and Cusi (2014), but contrasting with the hemipenis of P. vertebralis illustrated by Hernández-Ruz and Bernal-Gonzalez (2011) for a specimen from Nariño, Colombia, the hemipenis of the holotype of P. ulisesi presents the four flounces in basal position at the asulcate face separated from the other flounces ornamenting the hemipenis laterally. In the drawing presented by Hernandez-Ruz (2005) for Cercosaura ampuedai (synonym of P. vertebralis according to Doan and Cusi [2014]) such flounces are not visible, probably due the distally misplaced tie made during hemipenial preparation.

Distribution and natural history observations: Pholidobolus ulisesi is known from five localities at elevations of 1,900 –2,300 m in Cajamarca and Lambayeque departments, northern Peru ( Fig. 6 View Fig ). All recorded localities lie within the Huancabamba depression, a region where the relatively low altitude of the Andean mountains causes fragmentation of montane habitats, and the northern extreme of the Central Andes at Cordillera Occidental in northern Peru. According to the terrestrial ecorregions of the world by Olson et al. (2001), P. ulisesi occurs within Eastern Cordillera real montane forest and Marañón dry forest.

Pholidobolus ulisesi was found during the day in sunny and cloudy conditions in secondary montane forest, in the edges of primary montane forest and recently opened areas for cattle ranching, as well as in small plantations of bean and coffee. In the open cattle-ranching areas, P. ulisesi was found moving on fallen trees or hiding under trunks; in secondary montane forest, the lizards were found foraging within herbaceous vegetation and running through the patches of grass. They were especially abundant in coffee and bean plantations, where they were observed running through the herbaceous vegetation and hiding in leaf litter. Sympatric squamate reptiles collected with P. ulisesi were Chironius monticola and Dipsas peruana at El Chaupe and Huamantanga, and Chironius monticola , Epictia teaguei , Erythrolamprus taeniurus , Micrurus peruvianus , Stenocercus arndti , S. huancabambae, and S. stigmosus at Quebrada La Iraca.

Etymology: The specific epithet “ ulisesi ” is a noun in the genitive case and a patronym for Ulises Gamonal Guevara, for his significant contribution to the archaeology of Cajamarca in northwestern Peru. One of his major contributions is the discovery of the>6,000-year-old Faical cave paintings in San Ignacio, declared as Cultural Patrimony of the Nation.

Remarks: In a molecular phylogeny of Cercosaura and related taxa, Torres-Carvajal et al. (2015) showed, with high support, that Pholidobolus ulisesi ( Pholidobolus sp. in their paper) and P. hillisi are sister species. Together they form a clade sister to all other species of Pholidobolus . In addition, these authors found that both “ Cercosaura ” vertebralis and “ Cercosaura ” dicra were nested within Pholidobolus , and were therefore referred to this genus ( Torres-Carvajal et al. 2015). An identical topology can be observed in a recent molecular phylogeny of the clade Cercosaurinae by Torres-Carvajal et al. (2016). Therefore, we adopt this taxonomic change in the discussion below.