Pseudogonatodes quihuai, Rojas-Runjaic & Koch & Castroviejo-Fisher & Prudente, 2024

Rojas-Runjaic, Fernando J. M., Koch, Claudia, Castroviejo-Fisher, Santiago & Prudente, Ana L. C., 2024, A new dwarf gecko of the genus Pseudogonatodes (Squamata: Sphaerodactylidae) from the eastern slope of the Andean Cordillera de Mérida in northern South America, Zootaxa 5418 (4), pp. 301-327 : 304-322

Pseudogonatodes quihuai sp. nov.

Figs. 2–11 , 12C–E

Holotype: MHNLS 21643 , adult female, from El Potrerito, ⁓ 2.2 km NW from Altamira de Cáceres , Bolívar municipality, Barinas state, Venezuela (8.839790°, -70.513770°; elevation 1,120 m asl; Fig. 1 ), collected on February 15, 2015, by Daniel Quihua.

Paratypes: Six specimens: an adult female ( MHNLS 22711 , field number DQ 83 ), an adult male ( MHNLS 22710 , field number DQ 82 ), and a juvenile ( MHNLS 22709 , field number DQ 81 ), from Sendero El Oso , Posada San Ramón , ⁓ 6 km N from Altamira de Cáceres , Bolívar municipality, Barinas state, Venezuela (8.878750°, - 70.497580°; elevation 1,319 m asl), collected on October 17, 2016, by Daniel Quihua, José Vieira & Dennys Mora; an adult male ( MHNLS 22712 , field number DQ 84) and a juvenile ( MHNLS 22713 , field number DQ 85), from the type locality, collected on October 20, 2016, by Daniel Quihua, José Vieira & Dennys Mora; and a juvenile ( MHNLS 21644 , without field number) with the same data as the holotype .

Definition. Pseudogonatodes quihuai is morphologically defined by the following combination of character states: (1) SVL: 27.5–30.2 mm in adult males, 31.3–33.6 mm in adult females; (2) posterodorsal margin of rostral W-shaped, with a median cleft; (3) three postrostrals; (4) median postrostral similar in size to lateral ones; (5) a single large postnasal scale; (6) five to six loreal scales; (7) 19–22 scales in a transverse row between the sutures of the 1 st and 2 nd supralabials on each side; (8) four to six supralabials; (9) four to five supraciliaries, the second being the largest; (10) four to five infralabials; (11) posterior margin of mental U-shaped, without posterior clefts; (12) four to six postmentals; (13) dorsal and lateral scales granular, juxtaposed; (14) 85–91 scales around midbody; (15) 20–22 ventrals in a transverse row at midbody; (16) 40–43 ventral scales along a midventral row, between throat and anterior border of cloaca; (17) 33–36 ventral scales along a midventral row, from anterior level of arms to anterior border of cloaca; (18) 26–29 ventral scales along a midventral row, between anterior levels of fore- and hind limbs; (19) 7–9 lamellae under fourth finger; (20) 8–10 lamellae under fourth toe; (21) third lamella under fourth toe not distinctly enlarged; (22) subcaudal pattern 1’1”.

Diagnosis. The new species differs from the other seven Pseudogonatodes species by the presence of a single large postnasal scale (vs. two or three postnasal scales; see Table 1 ). In addition, it differs from P. barbouri by having juxtaposed granular dorsal scales (vs. flat and imbricate), by lacking an expanded third lamella under fourth toe (TIV) (vs. expanded), and by its larger body size, which reaches 33.6 mm SVL in adult females (vs. up to 25.6 mm SVL; Koch et al. 2018). It is differentiated from P. lunulatus by lacking an expanded third lamella under TIV (vs. expanded), by having heterogeneous scales on the sole of the foot (vs. homogeneous), a large median postrostral, similar in size to the lateral ones (vs. median postrostral smaller than lateral ones), 5–6 loreal scales in a straight line between postnasal and orbit (vs. up to nine), 4–6 postmentals (vs. 3–4), 8–10 infradigital lamellae under TIV (vs. 6–7), and larger body size (vs. up to 29 mm SVL; Esqueda et al. 2016). The new species further differs from P. guianensis by lacking an expanded third infradigital lamella under TIV (vs. expanded), by having 33–36 ventral scales between anterior level of forelimb and cloaca (vs. 36–47 VMR-b), 8–10 lamellae under TIV (vs. 5–7), a medial row of slightly expanded subcaudal scales arranged in a 1’1” pattern (vs. in some cases slightly expanded but without a pattern), and a larger body size in adult females (vs. up to 30 mm; Avila-Pires 1995). Pseudogonatodes quihuai is distinguished from P. gasconi by having a medial cleft on the posterior margin of the rostral (vs. without cleft on posterior margin of rostral), granular dorsal scales (vs. conical to flat conical dorsals), 26–29 ventral scales between anterior levels of fore- and hind limbs (vs. 37 VMR-c), 5–6 loreal scales in a straight line between postnasal and orbit (vs. 7 loreals), a medial row of slightly expanded subcaudal scales arranged in a 1’1” pattern (vs. subcaudals unexpanded and unpatterned), and a larger body size, which ranges between 31.3–33.6 mm SVL in adult females (vs. 24 mm SVL in the only known specimen, which is an adult female; Avila-Pires & Hoogmoed 2000). It is differentiated from P. manessi by having 5–6 loreal scales (vs. 7–8), posterior margin of mental U-shaped, without posterior clefts (vs. W-shaped, with or without two posterior clefts), 4–6 postmentals (vs. 6–9), 26–29 ventral scales between anterior levels of forelimbs and hind limbs (vs. 31–35 VMR-c), and 33–36 ventral scales until border of cloaca (vs. 37–40 VMR-b), and a smaller body size (vs. up to 38 mm SVL; Avila-Pires & Hoogmoed 2000). It is distinguished from P. peruvianus by having a large median postrostral, similar in size to the lateral ones (vs. median postrostral about one-third the size of laterals), 85–91 scales around midbody (vs. 79 SAM), 33–36 ventral scales between anterior level of forelimbs and border of cloaca (vs. 42 VMR-b). Finally, P. quihuai differs from P. furvus by having 5–6 loreal scales (vs. 7–10), posterior edge of mental without clefts (vs. two median clefts), 4–6 postmentals (vs. 2–4), 27–29 ventral scales between anterior levels of forelimbs and hind limbs (vs. 34–39 VMR-c), and 33–36 ventral scales until border of cloaca (vs. 44–67 VMR-b), 7–9 lamellae under 4 th finger (vs. 11–13), 8–10 under TIV (vs. 11–15), and a smaller body size, which does not exceed 33.6 mm (vs. up to 45 mm SVL; Montes-Correa et al. 2021).

Description of the holotype. Adult female in good state of preservation ( Fig. 2 ); SVL = 33.6 mm. Head coneshaped, HL = 6.8 mm (0.2 times SVL), slightly longer than wide (HL/HW = 1.3), depressed (HD/HW = 0.7), and as wide as the neck ( Fig. 2 ). Snout 2.6 mm long (0.4 times HL), subacuminate in dorsal view, rounded in profile, gently sloping toward top of head ( Fig. 3A–C ). Neck cylindrical, slightly narrower than body ( Fig. 2 ). Body wider than high, slightly depressed; axilla-groin distance 15.3 mm. Limbs short (forelimb 0.27 times SVL; hind limbs 0.33 times SVL), robust; digits clawed, well developed, five on hands and feet ( Fig. 4A–B ). Tail complete (not regenerated; Fig. 2 ), almost as long as the body (TL = 29.4 mm; 0.9 times SVL), stout, tapering toward tip, round in cross section.

Rostral large, well visible from above ( Fig. 3A ); lateral borders vertical, almost straight, posterolaterally in contact with 1 st supralabial and anterior border of nostril of each side ( Fig. 3B–C ); posterodorsal margin of rostral W-shaped, with the medial concavity forming an angle of ~ 90º; a median cleft extending forward from posterior margin to near midpoint of the scale; rostral posteriorly in contact with three large postrostrals.All three postrostrals distinctly larger than adjacent scales on snout ( Fig. 3A ). Medial postrostral pentagonal, slightly wider than long, barely smaller than lateral ones; lateral postrostrals nearly rectangular, about twice as wide as long, posterolaterally in contact with postnasal. Nostril located at the anterior portion of nasal scale, bordered by lateral postrostral, rostral, and 1 st supralabial. A single large postnasal, nearly rounded, distinctly larger than adjacent loreal scales ( Fig. 3B–C ). Scales on dorsum of snout and loreal region juxtaposed, smooth; polygonal (nearly rounded) and flat on the anteriormost rows, gradually changing into smaller granules toward top of head ( Fig. 3A ). Nineteen scales on a transverse row on snout, between suture of 1 st and 2 nd supralabials of each side. Loreal scales 5/ 6 in row between postnasal and orbit. Scales on supraocular region granular, similar in size to adjacent scales on top of head. Four large, flattened scales aligned on anterior and upper margin of eye forming a supraciliary flap; second one the largest. Supralabials 5/5; 4 th supralabial (right side) and suture between 4 th and 5 th supralabial (left side) below center of eye. Scales on temporal region similar to those on top of head. Ear-opening small (less than half of eye diameter), roughly elliptical; 19/21 temporal scales in the shortest straight line between eye and ear-opening.

Mental scale large, U-shaped, posterior margin deeply concave, without posterior clefts. Postmentals five, polygonal, about twice the size of adjacent scales on chin, aligned forming an arc along the posterior margin of mental; the three medial postmentals nearly rounded, the two most lateral postmentals roughly rectangular. Anteriormost scales of chin nearly flat, polygonal (almost rounded), juxtaposed, gradually changing into slightly smaller granules toward center of throat; noticeably larger, flatter, and elongate those scales of chin in contact with or near to infralabials ( Fig. 3D ). Infralabials 5/5, the first much larger, 2.6–2.7 times as wide as the second; first infralabial reaching (left side) or surpassing (right side) anterior border of orbit; suture between third and fourth infralabials below center of eye ( Fig. 3B–C ). Scales on dorsal and lateral surfaces of neck granular, slightly larger than those on top of head and similar in size to dorsal body scales. Anterior scales of throat small, granular, juxtaposed; progressively larger towards the posterior throat edge, with a short transitional area at the mid-throat level; scales on posterior half of throat large, cycloid, imbricate, similar to those on ventral body surface.

Dorsal scales of body granular ( Fig. 2A ), slightly larger than scales on top of head; granules on vertebral area conical, erected, juxtaposed, somewhat misaligned; granules on dorsolateral surfaces and flanks barely larger than vertebrals, juxtaposed, arranged in diagonal rows, subtly flattened anterodorsally and slightly tilted posteriorly. A narrow lateral transitional zone between dorsals and ventrals. Ventral scales distinctly larger, distally rounded to ovate, flat, smooth, imbricate, arranged in both longitudinal and oblique rows ( Fig. 2B ); 91 scales around midbody, 22 of which are ventrals; 27 scales in a midventral row between anterior levels of fore- and hind limbs, 33 until border of cloaca, 40 between first enlarged ventral on throat to cloaca; anterior margin of cloaca bordered by one to two rows of minute scales. First 10–12 rows of dorsal scales at tail base (about five rows on lateral surfaces) similar in shape and size to dorsal body scales, followed by a short transitional area (about 4–5 scale rows) and then by large, flat, smooth, imbricate scales, similar to those on belly. A midventral row of slightly enlarged, roughly hexagonal caudal scales, starting about 5–6 scales from posterior margin of cloaca (not regenerated tail), alternately in contact latero-posteriorly with one or two smaller scales in a 1’1” pattern ( Fig. 4C ); enlarged midventral scales in latero-posterior contact with two scales (i.e., 1”) larger than those in contact with a single scale (1’).

Scales on upper and anterior surfaces of forelimbs, and on anterior and ventral surfaces of hind limbs large, flat, smooth, imbricate; scales elsewhere on limbs small, granular, conical, juxtaposed to sub-imbricate. Scales on palms and soles heterogeneous in size, flat, polygonal, imbricate ( Fig. 4A–B ). Lamellae under first (I) through fifth (V) finger (right/left): I: 4/5; II: 7/7; III: 9(1 st divided)/8; IV: 8(1 st divided)/8(1 st divided); V: 6(1 st divided)/6. Lamellae under first (I) through fifth (V) toe (right/left): I: 5(1 st –3 rd divided)/5; II: 8(1 st –2 nd divided)/7; III: 8(1 st divided)/8(1 st divided); IV: 8(1 st divided)/9(1 st –2 nd divided); V: 8/8(1 st –3 rd divided). Third lamellae under fourth toe not distinctly enlarged ( Fig. 4B ). Basal lamellae of fingers and toes larger than distal ones. Claws enclosed in ungual sheaths composed of five enlarged scales.

Color in life and preservative. In life, dorsal and lateral surfaces of head, dorsum, flanks, and limbs dark brown, with some scattered whitish dots, mainly on flanks; background color of dorsal and lateral surfaces of tail also dark brown, but darker than head and trunk. Dorsal surface of head with a lighter, poorly defined occipital band posteriorly bordered by a thin blackish brown transverse line; supralabials and infralabials mottled blackish brown, with pale vertical stripes on lateral borders of such scales; a pale diagonal postrictal band, finely bordered with blackish brown, extending posteroventrally from angle of jaws to throat, where it continues backwards flanking the throat. A pair of thin, ill-defined, pale dorsolateral stripes extending backwards from postocular corners to distal third of tail, where they become inconspicuous; both dorsolateral stripes finely bordered with blackish brown from posterior third of dorsum to base of tail; from base of tail and to its distal third, pale dorsolateral stripes become broader and slightly sinuous. Chin and throat dirty white, laterally delimited by a blackish line. Chest, belly, ventral surfaces of thighs and tail brown, slightly lighter than flanks and dorsum; all the enlarged scales on these ventral regions are pale centrally and brown on margins. Iris bronze.

In preservative (after eight years), dark brown background color has become pale brown; blackish line posteriorly bordering the pale occipital band and those blackish lines flanking pale diagonal postrictal band, more conspicuous. Pale occipital band and pale dorsolateral stripes somewhat diffuse, less conspicuous. Whitish dots scattered on dorsum and flanks have faded, almost imperceptible. Iris pale gray.

Cranial osteology. The skull of Pseudogonatodes quihuai ( Figs. 5–11 ) closely resembles that of P. barbouri ( Bauer et al. 2018) in many aspects. Therefore, we only highlight those character states that differ from P. barbouri . Unless otherwise indicated, this osteological description applies to all three specimens examined. A comparative synopsis of osteological character states among P. quihuai , P. barbouri , and P. furvus is presented in Table 2 .

Along the inner wall of the supraoccipital, prootics, and otooccipitals are large, well-developed endolymphatic sacs that project slightly posteriorly and distinctly anteriorly beyond the supraoccipital. Anteriorly, they extend along the posterior third of the parietals and are in close contact with them dorsally ( Figs. 6‒8 ). Endolymphatic sacs are also present in MHNLS 22711 but absent in MHNLS 22712. A small posteromedial gap between the parietal and the medial border of the supraoccipital is present in MHNLS 22711 ( Fig. 11E ) and MHNLS 22712 ( Fig. 11J ), but it is missing in MHNLS 21643 ( Fig. 6B ), where both bones are in firm contact. In MHNLS 22711 the gap is filled by the endolymphatic sacs.

The premaxilla contacts the nasals and maxillae in all three specimens and the septomaxilla (internally) only in MHNLS 21643. The premaxilla bears 11 (n = 3) tooth loci on the ventral portion and there are 3‒4 rows of replacement tooth buds ( Fig. 8A ). The ascending nasal process extends beyond the anterior half of the median suture of the nasal bones ( Figs. 5A , 6A , 7A , 11A, D, F, I ). The palatal process does not contact the vomer in all three specimens. The maxillae do not contact the ectopterygoid medially in MHNLS 21643 whereas the two bones are in contact in MHNLS 22711 and MHNLS 22712. The maxilla bears 22‒24 (23.33 ± 0.82; n = 6) tooth loci ( Fig. 8A ). Slightly below the facial process, the maxilla bears a series of 2‒5 (3.17 ± 0.98; n = 6) foramina ( Figs. 5B , 11B, G ). The nasals are in firm medial contact along their entire length and anteromedially both nasals together form a channel into which the ascending process of the premaxilla fits, framing it ventrally and laterally ( Figs. 5A , 9A ). Posteriorly, the nasals greatly overlap the anterior part of the frontal ( Fig. 9 ). Viewed from internally, the posterolateral margin of each nasal contacts the medial margin of the prefrontal ( Fig. 8C ). The anteromedial ventral surface of each nasal has a broad contact zone with the anteromedial dorsal surface of the septomaxilla. Viewed from internally, the medial margin of the prefrontal contacts the posterolateral margin of each nasal ( Fig. 8C ). The lacrimal foramen is oval shaped rather than triangular.

The frontal bone is only partly fused or even paired, and the suture or at least a medial longitudinal indentation is visible dorsally and ventrally along the entire structure, being more pronounced ventrally and dorsally on the posterior third ( Figs. 9 , 11 ). Anterolaterally, the frontal bone is in broad contact with the posterodorsal part of the facial process of the maxilla ( Figs. 5A , 11A, F ). This aspect is not mentioned in Bauer et al. (2018), but is also clearly visible in the figures of Pseudogonatodes barbouri . Anteriorly, the paired septomaxillae diverge, while in the posterior half they are about parallel to each other and only slightly separated without touching each other medially ( Figs. 7B , 8B ). The main body has a concave dorsal surface. The septomaxilla overlays the vomer dorsally and touches it with its posteromedial margin or remains slightly separated from it. In addition to the maxilla and vomer, the septomaxilla also contacts the nasal anteromedially and in MHNLS 21643 it even has a small contact point with the premaxilla anteriorly. The vomers are longer than wide, and become slightly broader posteriorly ( Fig. 5C ). They are paired but in firm contact in MHNLS 21643 ( Fig. 5C ) and MHNLS 22712 ( Fig. 11H ), but seem to be partially fused in the posterior region in MHNLS 22711 ( Fig. 11D ). The vomer does not contact the premaxilla anteriorly and whether it contacts the palatine posteriorly depends on the volume rendering threshold; however, if so, it is very marginal ( Fig. 7C ). A defined area for the vomerine process of the palatine is lacking. The palatines are partially very thin and broadly separated from another ( Figs. 7C , 11C, H ). The pterygoid does not contact the parabasisphenoid medially ( Figs. 7B , 8A , 11C, H ), as should be the case in P. barbouri according to its osteological description ( Bauer et al. 2018); however, this appears to be an error in the description as the two bones appear clearly separated in Bauer et al. (2018: fig. 1C). The anterior process of each ectopterygoid is broader and slightly shorter than the posterior process ( Fig. 8B ). The posterior end of the maxilla extends beyond the vertex of the ectopterygoid in ventral view. The vertex region of the ectopterygoid laterally contacts the jugal ( Fig. 8A ).

As in Pseudogonatodes barbouri ( Bauer et al. 2018) in MHNLS 21643 ( Figs. 5 , 6B , 7B , 8A ) and in MHNLS 22711 ( Fig. 11A‒C, E ) the parabasisphenoid, basioccipital, supraoccipital, prootic, and otooccipital are fused to form the braincase, although traces of a suture between the parabasisphenoid and basioccipital are visible in MHNLS 22711. In MHNLS 22712 ( Fig. 11F‒H, J ), the bones of the braincase are in close contact but not fused, and all sutures are visible. Dorsomedially the braincase contacts the posteriormedial part of the parietals in MHNLS 21643 ( Fig. 6B ), but not in MHNLS 22711 ( Fig. 11E ) and MHNLS 22712 ( Fig. 11J ). The two trabeculae between the basipterygoid processes are distinctly larger and more pronounced ( Figs. 7B , 11C, H ) than in P. barbouri .

The dentary bears 26‒30 (27.67 ± 1.63; n = 6) tooth loci ( Fig. 10A, D ). Laterally, the dentary bears 5‒7 (5.67 ± 0.82; n = 6) mental foramina, which lie in a longitudinal series between the second or third (2.83 ± 0.41; n = 6) and twentieth to twenty-fourth (22.17 ± 1.47; n = 6) tooth loci ( Fig. 10C ). The surangular is completely fused with the compound bone and not distinguishable in MHNLS 21643 ( Fig. 10 ) and MHNLS 22711, whereas a soft suture is visible in MHNLS 22712. The anterior process of the compound bone anteriorly surpasses the anteromedial process of the coronoid + splenial complex. However, this anterior extension is only visible when the dentary is (digitally) removed.

Variation in type series. Morphological variation in external phenotypic characters, related to sexual dimorphism, is only evident in the SVL, with females being slightly larger than males (31.3–33.6 mm vs. 27.5–30.2 mm). Regarding morphometry, there is very little variation in the observed body proportions of the type series. The head is little longer ( HL / SVL = 0.3) in the juvenile MHNLS 21644 (0.2 in all the others), whereas it is slender in the subadult MHNLS 22709 ( HL /HW = 1.4 vs. 1.3 in the others). The head is always depressed but the HD/HW ratio is between 0.6–0.7. Finally, the snout is slightly shorter in the adult male MHNLS 22712 ( SL /HL = 0.3 vs. 0.4 in the others). All measurements of the type series are listed in Appendix 2.

Phenotypic variation is larger in scale counts (Appendix 3) and in the shape and proportions of some head scales; although the concavity of the dorsal-posterior margin of the rostral typically forms an angle of ⁓ 90°, it is> 90° in the specimen MHNLS 22709 . The median postrostral scale can be barely smaller ( MHNLS 21643 , 22709 , and 21644), equal ( MHNLS 22710–11 , 22713 ) or barely larger ( MHNLS 22712 ) than lateral postrostrals. The loreals range between 5–6 and vary in number even between opposite sides of the same individual in four of the seven specimens of the type series. The number of supraciliaries varies between 4 and 5; however, the 2 nd is always the largest. The number of scales of the snout in a transverse line between 1 st and 2 nd supralabials ranges from 19 to 22. The number of both supralabials and infralabials varies from 4 to 5; however, the specimen MHNLS 22710 has six supralabials on the right side. The number of postmentals varies between 4–6, with one or two scales in the middle of the row. The number of lamellae under the fingers and toes varies as follow: FI: 4–5, FII: 7–8, FIII: 8–9, FIV: 7–8, FV: 6–7; TI: 5–6, TII: 7–8, TIII: 8–10, TIV: 8–10, TV: 7–10. The distal subdigital lamellae of the fingers are sometimes divided sagittally, and those of toes are frequently divided. The 1 st subdigital lamella (counted from the tip to the base) is divided in 24 of 70 fingers (34%) and in 58 of 70 toes (83%); the 2 nd lamella is divided in 4 of 70 fingers (6%) and in 23 of 70 toes (33%); the 3 rd lamella is only divided in 5 of 70 toes (7%). The number of scales around midbody is 85–91 and there are 20–22 enlarged ventral scales in a transverse line at midbody. The number of enlarged ventral scales from throat to cloaca is 40–43; and 33–36 from the anterior border of forearms to cloaca, while there are 26–29 ventrals between anterior borders of fore- and hind limbs .

The color pattern also exhibits some variation in the type series ( Fig. 12C–E ). The light occipital band may be ill-defined ( MHNLS 21643 , 22710 , and 22711), evident ( MHNLS 22713 ), or very well defined ( MHNLS 21644 , 22709 , and 22712); regarding color, this light band may be almost immaculate white ( MHNLS 22712 ), dirty white ( MHNLS 21644 , 22709 , 22713 ), light grayish brown ( MHNLS 22710–11 ), or light brown ( MHNLS 21643 ). The light occipital band also varies in form, being medially divided ( MHNLS 22709 ), medially constricted at its anterior margin ( MHNLS 22710 , 22712–13 ), or roughly uniform in its width ( MHNLS 21346 , 22711 ). The blackish-brown transverse line juxtaposed posteriorly to the pale occipital band, is wider and very conspicuous in MHNLS 22712–13 . The pale dorsolateral stripes are absent on neck and dorsum in MHNLS 22711 , and present in the remaining specimens of the type series. There are a series of small, thin, and very-spaced blackish dashes flanking pale dorsolateral stripes from neck to thighs level, in the specimens MHNLS 22709–10 , 22712–13 , which are absent in MHNLS 21643 and 22711. Finally, the juveniles MHNLS 21644 and 22713 are more densely spotted with white on the dorsum and flanks of the body. The ventral surface is quite similar in color pattern in the type series, but immature specimens ( MHNLS 21644 , 22709 , 22713 ) exhibit a thin pale stripe running longitudinally along the posteroventral surface of the thighs.

Distribution, habitat, and natural history. Pseudogonatodes quihuai is only known from two localities along the eastern slope of the Cordillera de Mérida in the Venezuelan Andes ( Figs. 1 , 12A ). Both localities are in the state of Barinas: El Potrerito ⁓ 2.2 km NW from Altamira de Cáceres (type locality); and Sendero El Oso, Posada San Ramón, ⁓ 6 km N from Altamira de Cáceres (the northernmost and highest known locality). The known altitudinal range of P. quihuai (1,120 –1,319 m asl) is occupied by montane semideciduous forests ( Fig. 12A–B ), which is mainly characterized by the loss of foliage of most of its tree species during one to three months, in the dry season at the beginning of the year; its canopy is irregular about 20–30 m high, with emergent trees reaching 40 m; the lower stratum is dominated by Araceae and Cyclanthaceae , and epiphytes are typically scarce in this phytocoenosis ( Ataroff & Sarmiento 2004). The natural history of P. quihuai remains virtually unknown; the only field notes on the type series indicate that all specimens were collected during nocturnal surveys and all of them were found on the ground, hidden in the leaf litter of the forest and shady coffee plantations. Although the specimens were collected at night (between 21:00–23:00 h), they were not observed actively foraging but were detected by leaf litter removal, which disturbed them, and caused them to move away from their shelters (D. Quihua, pers. comm.). From dissection, we noted that the specimen MHNLS 22711 has a relatively large oviductal egg, still without a calcified shell. The specimen MHNLS 22710 has a small piece of skin shedding inside its stomach. We suspect that is from itself, indicating the occurrence of shed skin eating in this species. Finally, the juvenile MHNLS 21644, the smallest of the series (15.4 mm SVL) shows a small and barely perceptible midventral umbilical scar, indicating that it had likely hatched recently.

Etymology. This species is named in honor of José Daniel Quihua Ramírez, a young self-taught naturalist and wildlife photographer, particularly passionate about the amphibians and reptiles of Venezuela. Daniel has been an outstanding collaborator of the first author during numerous field surveys and collected the type specimens of the new species.

Remarks. Avila-Pires & Hoogmoed (2000) reported on a specimen of Pseudogonatodes (MIZA 15-185 [Museo del Instituto de Zoología Agrícola, Maracay, Venezuela]) from Las Cuevas, Uribante-Caparo hydroelectric complex, Táchira state, on the southern part of the eastern slope of the Cordillera de Mérida, Venezuela (sky-blue dot closest to the white cross in Fig. 1 ). At that time, the specimen could not be identified to the species level with certainty and the authors considered two possibilities: (i) that it represented an undescribed species, or (ii) a juvenile of P. manessi . Since then, specimen MIZA 15-185 had not been reexamined and no new specimens of Pseudogonatodes have been obtained from that locality. We reexamined the scale counts reported by Avila-Pires & Hoogmoed (2000) and some additional unpublished scale counts of the same specimen (see Table 1 ), and we preliminary conclude that: 1) The specimen MIZA 15-185 likely correspond to a species of the “3 rd Group” as it presents granular juxtaposed dorsal scales and high subdigital scale counts (7 LFIV and 9 LTIV); 2) MIZA 15-185 does not appear to be conspecific with P. manessi due to having a lower number of SAM (77 vs. 85–101), VTM (19 vs. 21), VMR-b (34 vs. 37–40), and VMR-c (29 vs. 31–35); in addition, we highlight that P. manessi is only known from the Venezuelan Cordillera de la Costa, whereas MIZA 15-185 comes from the southeastern portion of the Cordillera de Mérida (see Fig. 1 ); 3) MIZA 15-185 is not conspecific with P. quihuai since it has two postnasal scales (vs. one), three posterior clefts on the mental scale (vs. mental without clefts), and lower counts of SAM (77 vs. 85–91); and 4) MIZA 15-185 likely corresponds to a still undescribed species, but new data and additional specimens will be required in order to corroborate this hypothesis.

Figures

FIGURE 1. Map of northern SouthAmerica depicting the geographicdistribution of the species of Pseudogonatodes: P.barbouri (brown), P.furvus (green), P. gasconi (black), P. guianensis (yellow), P. lunulatus (red), P.manessi (blue), P. peruvianus (purple), P. quihuai sp. nov. (white cross), and Pseudogonatodes sp. (sky blue). Dotted symbols indicate type localities. Country limits represented by white lines. Diagonal striped area: Guayana Esequiba (disputed territory). Geographic records were obtained from Roux (1927), Mechler (1968), Huey & Dixon (1970), Gorzula & Señaris (1999), Avila-Pires & Hoogmoed (2000), Molina (2001), Ribeiro-Júnior (2015), Esqueda et al. (2016), Koch et al. (2018), Montes-Correa et al. (2021), GBIF (2023), Torres- Carvajal et al. (2023), and MHNLS.

FIGURE 2. Holotype of Pseudogonatodes quihuai sp. nov. (adult female, MHNLS 21643) in preservative. Dorsal (A) and ventral (B) views of the body. Scale bar represents 10 mm. Photos: F.J.M. Rojas-Runjaic.

FIGURE 3. Holotype of Pseudogonatodes quihuai sp. nov. (adult female, MHNLS 21643) in preservative. Dorsal (A), right lateral (B), left lateral (C), and ventral (D) views of the head. Scale bars represent 1 mm. Photos: F.J.M. Rojas-Runjaic.

FIGURE 4. Holotype of Pseudogonatodes quihuai sp. nov. (adult female, MHNLS 21643) in preservative. Ventral surfaces of the left hand (A) left foot (B), and tail (C). Scale bars represent 1 mm. Photos: F.J.M. Rojas-Runjaic.

FIGURE 5. Micro-CT images of the skull of the holotype of Pseudogonatodes quihuai sp. nov. (MHNLS 21643) in dorsal (A), lateral (B), and ventral (C) views; boc: basioccipital; cb: compound bone; co: coronoid; d: dentary; ecp: ectopterygoid; epp: epipterygoid; es: endolymphatic sacs; f: frontal; j: jugal; m: maxilla; n: nasal; otoc: otooccipital; pa: parietal; pal: palatine; pref: prefrontal; pbas: parabasiphenoid; pm: premaxilla; pof: postorbitofrontal; po: prootic; pt: pterygoid; q: quadrate; sm: septomaxilla; so: sclerotic ring; soc: supraoccipital; sq: squamosal; sta: stapes; v: vomer. Scale bar represents 1 mm.

FIGURE 6. Micro-CT images of the skull of the holotype of Pseudogonatodes quihuai sp. nov. (MHNLS 21643) in anterior (A), and posterior (B) views; boc: basioccipital; cb: compound bone; co: coronoid; d: dentary; epp: epipterygoid; es: endolymphatic sacs; f: frontal; m: maxilla; n: nasal; otoc: otooccipital; pa: parietal; pm: premaxilla; pof: postorbitofrontal; po: prootic; pt: pterygoid; q: quadrate; rp: retroarticular process of compound bone; sm: septomaxilla; soc: supraoccipital; sq: squamosal. Scale bar represents 1 mm.

FIGURE 7. Micro-CT images of the skull of the holotype of Pseudogonatodes quihuai sp. nov. (MHNLS 21643) in dorsal view: mandibulae removed (A), ectopterygoids, frontal, jugals, mandibulae, maxillae, nasals, parietals, postorbitofrontals, premaxilla, andsquamosals removed(B),all previously mentionedbones plus braincase (fused parabasiphenoid, basioccipital, supraoccipital, prootic, and otooccipital), epipterygoids, prefrontals, septomaxillae, and statolithic mass removed (C); ecp: ectopterygoid; epp: epipterygoid; es: endolymphatic sacs; f: frontal; j: jugal; m: maxilla; n: nasal; otoc: otooccipital; pa: parietal; pal: palatine; pref: prefrontal; pbas: parabasiphenoid; pm: premaxilla; pof: postorbitofrontal; po: prootic; pt: pterygoid; q: quadrate; sm: septomaxilla; soc: supraoccipital; sq: squamosal; sta: stapes; v: vomer. Scale bar represents 1 mm.

FIGURE 8. Micro-CT images of the skull of the holotype of Pseudogonatodes quihuai sp. nov. (MHNLS 21643) in ventral view: mandibulae removed (A), braincase (fused parabasiphenoid, basioccipital, supraoccipital, prootic, and otooccipital), mandibulae, maxillae, premaxilla, pterygoids, and vomers removed (B), all previously mentioned bones plus ectopterygoids, endolymphatic sacs, jugals, epipterygoids, palatines, quadrates, septomaxillae, stapes, and statolithic mass removed (C); boc: basioccipital; ecp: ectopterygoid; epp: epipterygoid; es: endolymphatic sacs; f: frontal; j: jugal; m: maxilla; n: nasal; pa: parietal; pal: palatine; pref: prefrontal; pbas: parabasiphenoid; pm: premaxilla; pof: postorbitofrontal; pt: pterygoid; q: quadrate; sm: septomaxilla; sq: squamosal; sta: stapes; stm: statolithic mass; v: vomer. Scale bar represents 1 mm.

FIGURE 9. Micro-CT images of the holotype of Pseudogonatodes quihuai sp. nov. (MHNLS 21643): frontal (f) and nasals (n) in dorsal (A) and ventral (B) views, and frontal alone in dorsal (C) and ventral (D) views.

FIGURE 10. Micro-CT images of the mandibulae of the holotype of Pseudogonatodes quihuai sp. nov. (MHNLS 21643) in dorsal (A) and ventral (B) views, and right mandibula in lateral (C), and medial (D) views; cb: compound bone; co: coronoid; d: dentary; rp: retroarticular process of compound bone; sp: splenial. Scale bar represents 1 mm.

FIGURE 11. Micro-CT images of the skulls of two paratypes of Pseudogonatodes quihuai sp. nov. (MHNLS 22711: A‒E; MHNLS 22712: F‒J) in dorsal (A, F), lateral (B, G), ventral (C, H), anterior (D, I), and posterior (E, J) views.

FIGURE 12. Panoramic view of the montane semideciduous forest (A) and its understory (B) in the eastern slope of theAndean Cordillera de Mérida, where Pseudogonatodes quihuai sp. nov. inhabits. (C) Holotype (MHNLS 21643) and (D, E) paratypes (MHNLS 21644 and 22709, respectively) of P. quihuai sp. nov. in life. Photos: F.J.M. Rojas-Runjaic (A, B) and D. Calcaño (C–E).