Proctoporus bolivianus Werner, 1910

Goicoechea, NoemÍ, Padial, José M., Chaparro, Juan Carlos, Castroviejo-Fisher, Santiago & Riva, Ignacio De La, 2013, A taxonomic revision of Proctoporus bolivianus Werner (Squamata: Gymnophthalmidae) with the description of three new species and resurrection of Proctoporus lacertus Stejneger, American Museum Novitates 2013 (3786), pp. 1-32 : 7-14

Proctoporus bolivianus Werner, 1910

Figures 3A , 4

Proctoporus bolivianus Werner, 1910: 30 .

Proctoporus longicaudatus Andersson, 1914: 6 .

Proctoporus bolivianus: Dirksen and De la Riva, 1999: 203 .

Proctoporus bolivianus: Doan and Schargel, 2003: 75 .

Proctoporus subsolanus Doan et al., 2005: 330 .

Proctoporus bolivianus: Goicoechea , et al., 2012: 953.

NEOTYPE: CBF 3437 (field number MNCN 4729 ) (fig. 4) adult male, from Millipaya , 12 km S of Sorata (15°51′30.2″S / 68°37′37.6″W), Province Larecaja , Department of La Paz, Bolivia, 3743 m, collected by Ignacio De la Riva, José Manuel Padial, Rodrigo Aguayo, and Nayarit Ayllón on 1 March 2006.

DIAGNOSIS: (1) Frontonasal length longer than frontal length; (2) nasoloreal suture present in some specimens; (3) supraoculars three; (4) superciliaries 3–4, first expanded onto dorsal surface of head; (5) postoculars 1–2; (6) palpebral disc made up of a single, undivided scale; (7) four supralabials anterior to the posteroventral angle of the subocular; (8) 2–3 pairs of genials in contact; (9) dorsal body scales quadrangular, slightly keeled; (10) transverse rows of dorsals 33–37; (11) transverse ventral rows 22–24; (12) a continuous series of small lateral scales separating dorsals from ventrals; (13) posterior cloacal plate made up of six scales in both sexes; (14) anterior preanal plate scales paired; (15) femoral pores per hind limb in males 5–7; (16) preanal pores absent; (17) subdigital lamellae on toe IV 16–19; subdigital lamellae on toe V 5–11; (18) limbs not overlapping when adpressed against body in adults; (19) pentadactyl; digits clawed; (20) dorsum and lateral surface of head dark brown, lip irregularly barred with cream coloring; ventral surface of head cream with clumps of black stippling on each scale; pregular region like head but with fainter stippling.

Specimens of Proctoporus bolivianus have an undivided palpebral disc, which identifies them as members of the genus Proctoporus , as opposed to Riama and Petracola ( Uzzell, 1970; Doan and Castoe, 2005). Proctoporus bolivianus can be distinguished from its congeners by having the frontonasal longer than the frontal (subequal in all other members of the genera). It can also be distinguished from P. pachyurus by having three supraoculars (four in P. pachyurus ) and fewer transverse dorsal scale rows (47–60 in P. pachyurus ; 33–37 in P. bolivianus ); from P. sucullucu by having limbs not overlapping when adpressed; from P. unsaacae and P. guentheri by the lack of a conspicuous series of lateral ocelli; from Proctoporus carabaya by having smaller size (SVL P. carabaya = 58.1–66.5; SVL P. bolivianus = 43.4–56.5); from P. kiziriani and P. lacertus by having first superciliary fused with first supraocular; from P. iridescens by having four supralabials anterior to the posteroventral angle of the subocular (three in P. iridescens ); from P.xestus by the lack of prefrontal scales (present in P.xestus ) and having keeled dorsal scales (smooth in P. xestus ); and from P. chasqui by the lack of prefrontal scales.

DESCRIPTION OF THE NEOTYPE: Adult male, snout-vent length (SVL) 52.1 mm, regenerated tail length 88.0 mm; head scales smooth, without striations or rugosities; rostral scale wider (1.8 mm) than tall (1.0 mm), meeting supralabials on either side at above the height of supralabials, and becoming higher medially, in contact with frontonasal, nasals, and first supralabials; frontonasal longer than wide, longer in length than frontal, widest posteriorly, in contact with rostral, nasals, anteriormost supraocular, and frontal; prefrontals absent; frontal larger than wide, roughly pentagonal, not in contact with anteriormost superciliary, in contact with frontonasal, first two supraoculars, and frontoparietals; frontoparietals roughly hexagonal, in contact with frontal, second and third supraoculars, parietals, and interparietal; supraoculars three, all in contact with superciliaries, third in contact with frontoparietal, pari- etal, and postocular; interparietal longer than wide, almost hexagonal, in contact with frontoparietals anteriorly, with parietals laterally, and occipitals posteriorly; parietals polygonal, with posterior sutures in contact with occipital, lateral sutures diagonally in contact with temporals, and medial sutures in contact with interparietal; occipitals three, smaller than parietals, medial pentagonal smaller than laterals. Nasal suture present, separating nasal from loreal scale; nasal in contact with rostral, first and second supralabials, frenocular, loreal, and frontonasal; nostril situated in anterior third of nasal scale; loreal scale not in contact with supralabials; three superciliaries, first fused with first supraocular; palpebral disc made up of a single transparent scale; suboculars two; postoculars two; temporals smooth, glossy, polygonal; supratympanic temporals two; four supralabials anterior to the posteroventral angle of the subocular. Mental wider (1.8 mm) than long (1.2 mm), in contact with first infralabial and postmental posteriorly; postmental single, pentagonal, in contact with mental, first infralabials and first pair of genials; two pairs of genials in contact, anterior pair in contact with second and third infralabials; second genials in contact with third and fourth supralabials; one pair of chin shields, separated by four smaller median pregulars; gular scale rows six; collar fold slightly distinct, concealing one row of small scales; lateral neck scales round, smooth. Dorsals rectangular, longer than wide, juxtaposed, slightly keeled, in 37 transverse rows; longitudinal dorsal scale rows 23 at midbody; continuous lateral scale series, smaller than dorsals; reduced scales at limb insertion regions present; transverse ventral scale rows 22; longitudinal ventral scale rows at midbody 10; anterior preanal plate scales paired; posterior preanal plate scales six, lateralmost scales small; scales on tail rectangular, juxtaposed, smooth. Limbs pentadactyl; digits clawed; dorsal brachial scales polygonal, subequal in size, subimbricate, smooth; ventral brachial scales roundish, subimbricate, smooth; antebrachial scales polygonal, subequal in size, smooth, ventral antebrachial scales smallest; dorsal manus scales polygonal, smooth, subimbricate; palmar scales small, rounded, subimbricate, domelike; dorsal scales on fingers smooth, quadrangular, covering dorsal half of digit, overhanging subdigital scales, three on finger I, six on II, nine on III, 11 on IV, seven on V; scales on anterodorsal surface of thigh large, polygonal, smooth, subimbricate; scales on posterior surface of thigh small, rounded, juxtaposed; scales on ventral surface of thigh large, rounded, flat, smooth; femoral pores five; preanal pores absent; scales on anterior surface of crus polygonal, smooth, juxtaposed, decreasing in size distally; scales on anterodorsal surface of crus rounded, juxtaposed; scales on ventral surface of crus large, smooth, flat, subimbricate; scales on dorsal surface of toes quadrangular, smooth, overhanging supradigital lamellae, three on toe I, seven on II, 12 on III, 17 on IV, nine on V; subdigital lamellae of toes single distally, double proximally, three on toe I, four on II, 11 on III, 14 on IV, nine on V; limbs not overlapping when adpressed against the body, separated by 8–9 dorsal scale lengths.

COLORATION IN PRESERVATIVE: Dorsal and lateral surfaces of head dark brown; lip irregularly barred with cream coloring; ventral surface of head yellowish cream with clumps of black stippling on each scale; pregular region like head but with fainter stippling, medial scales lacking stippling; gular region like head but with denser stippling per scale, forming longitudinal clumps. Dorsal surface of body nearly of the same color as head with two lighter dorsolateral longitudinal stripes originating near occiput and extending to forelimb insertions; lateral surface of body of the same coloration as dorsum, fading lighter brown near venter; ventral surface of body with yellowish-cream ground color, scales with black stippling, medialmost scales lacking stippling. Color of limbs similar to body. Dorsal tail coloration like that of body, ventral surface of tail light brown with dark brown mottling anteriorly.

COLORATION IN LIFE: Similar to that in preservative but with orange coloration along flanks, forelimbs, tail, and ventral surface of the body (fig. 3A).

VARIATION: Coloration is similar among all specimens, with some of them having a lighter dorsum and more distinct longitudinal stripes. Stippling on the venter is more distinct or darker in some specimens and nearly absent in others, tending to form ventral lines (especially in younger specimens). The presence of ocelli is also variable. Variation of morphometric and meristic characters is shown in table 3.

DISTRIBUTION: This species is known from montane forests and humid puna grasslands in the eastern versant of the Andes in northern Bolivia and adjacent areas of Departamento de Puno in southern Peru (fig. 9) between 2100–3743 m, encompassing an approximate air distance of 200 km between the most distant localities, Sandia (Puno, Peru, 14°20′33.9″S / 69°27′45.8″W) and Sorata (La Paz, Bolivia, 15°51′30.2″S / 68°37′37.6″W).

HABITAT AND ECOLOGY: Individuals were found during the day under logs and stones, occupying disturbed areas and agricultural fields.

COMMENTS: Proctoporus subsolanus was described by Doan et al. (2005) from several specimens collected in Sandia (Puno, Peru: fig. 1). This species was distinguished from P. bolivianus by having frontonasal scales much longer than frontals (purportedly subequal in P. bolivianus ). Nonetheless, frontonasals are subequal to frontals only in populations from central Cusco and Puno considered as P. bolivianus by Uzzell (1970), Doan and Castoe (2003) and Doan et al. (2005) and now corresponding to Proctoporus carabaya , Proctoporus iridescens , Proctoporus kiziriani , and Proctoporus lacertus (table 2). In fact, topotypic specimens of P. bolivianus have frontonasal scales much longer than frontals, just as in P. subsolanus (table 1). Furthermore, the presence of a frontonasal larger than the frontal was also noted in the original description of P. longicaudatus , a synonym of P. bolivianus collected near the type locality of this species in Sorata Valley ( Bolivia) ( Andersson, 1914). Proctoporus subsolanus has also been reported to have two supraoculars (three in our character analyses, but the first fused with the first superciliary), which would distinguish it from P. guentheri and P.pachyurus (Doan et al., 2005) . This character state is nonetheless also observed in topotypic samples of P. bolivianus . In addition, our analysis of additional morphological characters did not reveal differences between P.subsolanus and P. bolivianus (table 1) and their color pattern is also similar. According to the original description, P. subsolanus has dorsal and lateral surfaces of the head and body dark brown, lips irregularly barred with cream, and two dorsolateral light brown longitudinal stripes originating near the occiput and extending to the forelimb insertions (Doan et al., 2005); the lateral and ventral surfaces of the body, the hind limbs and the tail were described as orange, with lateral scales having black stippling. As it is shown in figure 2A1 and 2A2, all these states are also present in topotypic specimens of P. bolivianus . Given the identical and/or broadly overlapping ranges of character states between Proctoporus subsolanus and P. bolivianus , in addition to results from molecular phylogenetic analyses, we follow Goicoechea et al. (2012) and consider P. subsolanus as a junior synonym of P. bolivianus .

Three other synonyms of P. bolivianus are currently available: Proctoporus lacertus Stejneger , Proctoporus obesus Barbour and Noble , and Proctoporus longicaudatus Andersson. We consider P. lacertus a valid species and furthermore consider P. obesus a junior synonym of P. lacertus . A rationale for these decisions is provided under the account for P. lacertus . We nonetheless retain P. longicaudatus as a junior synonym of P. bolivianus . Proctoporus longicaudatus was described from a single specimen collected at Pelechuco (La Paz, Bolivia), approximately 115 km west of the type locality of P. bolivianus (Sorata; Andersson, 1914). We have not been able to examine the holotype of this species, but the original description is quite complete. All morphological characters mentioned in the description of P. longicaudatus ( Andersson, 1914) , and those observed in specimens from Pelechuco (MNCN 43655 and MNCN 43656) and topotypic specimens of P. bolivianus examined by us have overlapping ranges of morphological character states (see also Uzzell, 1970). Given the broadly overlapping ranges of character states between this species and P. bolivianus , and the geographic proximity of their type localities, we follow Uzzell (1970) and Doan and Castoe (2005) and consider P. longicaudatus as a junior synonym of P. bolivianus .

Figures

FIG. 3. Dorsal and ventral views of living specimens of A, Proctoporus bolivianus from Millipaya (La Paz, Bolivia); B, Proctoporus iridescens from the road between Huancasarani and Limbani (Puno, Peru); and C, Proctoporus kiziriani from road between Marcapata and Tambopampa (Cusco, Peru).

FIG. 4. Neotype of Proctoporus bolivianus (CBF 3437; snout-vent length, 52.1 mm).