QCAZ 14955 (Figs 4, 5), adult male, Ecuador, Provincia Zamora-Chinchipe, Cerro Plateado Biological Reserve, Cerro Plateado plateau, 4.6159S, 78.7870W, WGS84, 2844 m, 23 September 2016, collected by Diego Almeida, Eloy Nusirquia, Fernando Ayala, Javier Pinto, Alex Achig and Malki Bustos.

Figure 4. 

Holotype of Pholidobolus samek sp. nov. (QCAZ 14955) in dorsal (A), ventral (B), and lateral (C) views. Male, SVL = 46.7 mm. (A, B): preserved specimen; (C): live specimen. Photographs by Darwin Nuñez and Valeria Chasiluisa.

Ecuador: Provincia Zamora-Chinchipe: QCAZ 14954 (adult female), same data as holotype; QCAZ 14956 (adult female), Cerro Plateado Biological Reserve, 4.6050S, 78.8167W, WGS84, 2320 m, 28 September 2016; QCAZ 14969–70, 14976–77(hatchlings) Cerro Plateado Biological Reserve, 4.6179S, 78.7838W, WGS84, 2873 m, 24 September 2016, same collectors as holotype.

Figure 5. 

Head of holotype of Pholidobolus samek sp. nov. (QCAZ 14955) in lateral (A), dorsal (B), and ventral (C) views. Photographs by Valeria Chasiluisa. Scale bar: 5 mm.

Pholidobolus samek is unique among its congeners, except P. condor sp. nov., in having green dorsolateral stripes on the head. However, adult males of P. samek differ from those of P. condor sp. nov. in having brighter dorsolateral head stripes and lacking a reddish venter. In addition, P. affinis, P. prefrontalis, P. macbrydei, P. dolichoderes sp. nov., and P. montium differ from P. samek (character states of P. samek in parentheses) in having a loreal scale frequently in contact with the supralabials (loreal scale not in contact with supralabials), and dorsal scales finely wrinkled (slightly keeled). Pholidobolus ulisesi and P. hillisi differ from P. samek in having a diagonal white bar along the rictal region (white rictal bar absent). Pholidobolus samek can be distinguished from P. dicrus by lacking a bifurcating vertebral stripe at midbody. Pholidobolus affinis, P. prefrontalis, P. dicrus, P. hillisi, and P. vertebralis further differ from P. samek in having well defined prefrontal scales (if present, prefrontal scales poorly differentiated). Additionally, P. samek has fewer dorsal scales (27−29) than P. affinis (45−55), P. montium (35−50), P. prefrontalis (37−46), P. macbrydei (31−43), P. fascinatus sp. nov. (32−37), and P. dolichoderes sp. nov. (35−40). Pholidobolus samek can be further distinguished from P. fascinatus by having widened medial scales on collar, and from P. dolichoderes sp. nov. by having fewer temporals (4–5 and 7–9, respectively), fewer ventrals (19–21 and 25–27), and fewer gulars (15–18 and 22–23).

(1) Two (rarely three) supraoculars, anteriormost slightly larger than posterior one; (2) prefrontals present or absent; (3) femoral pores absent in both sexes; (4) four to five opaque lower eyelid scales; (5) scales on dorsal surface of neck striated, becoming slightly keeled from forelimbs to tail; (6) two or three rows of lateral granules at midbody; (7) 27–29 dorsal scales between occipital and posterior margin of hindlimb; (8) lateral body fold present; (9) keeled ventrolateral scales on each side absent; (10) dorsum grayish brown with a distinct golden gray middorsal stripe, slender at midbody, becoming pale gray towards tail; (11) labial stripe white or orange; (12) flanks of body dark brown; (13) conical hemipenial body, with sulcus spermaticus originating between thick lips.

Adult male (QCAZ 14955) (Figs 4, 5); SVL 46.7 mm; TL 80.9 mm; dorsal and lateral head scales juxtaposed, finely wrinkled; rostral hexagonal, 2.06 times as wide as high; frontonasal irregularly quadrangular, wider than long, laterally in contact with nasal, loreal and first superciliary, slightly bigger than frontal; prefrontal scales absent; frontal longer than wide, in contact with one supraocular on the left side, and two on the right side; frontoparietals pentagonal, longer than wide, slightly wider posteriorly, each in contact laterally with supraocular II; interparietal roughly heptagonal; parietals slightly bigger than interparietal, hexagonal, and positioned anterolaterally to interparietal, each in contact anteriorly with supraocular II (and supraocular III on right side) and dorsalmost postocular; postparietals three, medial scale smaller than laterals; seven supralabials, fourth one longest and below center of eye; six infralabials, fourth one shortest and below center of eye; temporals enlarged, irregularly hexagonal, juxtaposed, smooth; two large supratemporal scales, smooth; nasal slightly divided, irregularly pentagonal, longer than high, 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; loreal rectangular, wider dorsally; frenocular higher than long, in contact with nasal, separating loreal from supralabials; two supraoculars on left side, three on right side (posteriormost much smaller), with the first one being the largest; four elongate superciliaries, first one enlarged, in contact with loreal; palpebral disc divided into four enlarged, pigmented scales; suboculars three (on the left side the medial subocular is fragmented), elongated and homogeneous in size; two postoculars, the dorsalmost wider than the other; 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, posteriorly widened in two longitudinal rows; posterior row of gulars (collar) with six scales, the medial two widened.

Nuchal scales similar in size to dorsals, except for the anteriormost that are widened; scales on sides of neck small and granular; dorsal scales hexagonal, elongate, imbricate, arranged in transverse rows; scales on dorsal surface of neck striated, becoming progressively keeled from forelimbs to tail; number of dorsal scales between occipital and posterior margin of hindlimbs 27; dorsal scale rows in a transverse line at midbody 26; one longitudinal row of smooth, enlarged ventrolateral scales on each side; dorsals separated from ventrals by two rows of small scales at level of 13^th row of ventrals; lateral body fold between fore and hindlimbs present; ventrals smooth, wider than long, arranged in 20 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 striated, imbricate; scales on ventral surface of forelimb granular; two thick, smooth thenar scales; supradigitals (left/right) 3/3 on finger I, 6/7 on II, 8/8 on III, 9/9 on IV, 6/6 on V; supradigitals 3/4 on toe I, 6/6 on II, 10/9 on III, 11/12 on IV, 7/7 on V; subdigital lamellae of fingers I and II single, paired on III (except the four distalmost), paired at base on IV, on finger V all single; subdigital lamellae 5/5 on finger I, 11/12 on II, 15/16 on III, 17/16 on IV, 9/10 on V; subdigital lamellae on toes I and II single, on toe III, IV and V all paired, except for the three distalmost subdigitals; subdigital lamellae 6/6 on toe I, 11/10 on II, 16/15 on III, 21/21 on IV, 14/14 on V; groin region with small, imbricate scales; scales on dorsal surface of hindlimbs smooth and imbricate; scales on ventral surface of hindlimbs smooth; scales on posterior surface of hindlimbs granular; femoral pores absent; 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 11.4; HW 7.4; ShL 7.0; AGD 23.9; TL/SVL 1.5; HL/SVL 0.2; HW/SVL 0.2; ShL/SVL 0.1; AGD/SVL 0.5.

Dorsal background from head to base of tail grayish brown, with a golden light brown vertebral stripe extending from occiput to tail; bright green dorsolateral stripes on head; cream white longitudinal stripe extending from first supralabial to shoulder; sides of neck, flanks and limbs dark brown; reddish brown narrow stripe extending from tympanum to arm insertion; ventrolateral region of body grayish brown; throat cream; chest, belly and base of tail cream orange (Figs 4C, 6B).

Figure 6. 

Pholidobolus samek sp. nov. in life. A Adult female, paratype (QCAZ 14954) B adult male, holotype (QCAZ 14955).

Dorsal background uniformly grayish brown, with a golden-gray vertebral stripe extending from occiput to tail; vertebral stripe wider anteriorly, becoming slightly slender at most posterior part of body; dorsal and lateral surfaces of head brown (rostral, frontonasal, frontal, frontoparietals, and supraoculars); bluish white longitudinal stripe extending from first supralabial to shoulder and fading on flanks; ventrolateral aspect of neck dark brown with a dorsolateral light brown stripe extending posteriorly along flanks to hindlimbs; forelimbs with scattered ocelli (black with white center); flanks grayish brown with two dorsolateral stripes on each side, the dorsal one dark brown and the most ventral one brown diffuse with dark brown spots; tail brown dorsally; ventral surface of head gray, chest and venter dark gray, ventral surface of tail slightly gray, with scattered dark brown marks.

Measurements and scale counts of Pholidobolus samek are presented in Table 6. Supralabials 8/7 (left/right) and temporals five in specimen QCAZ 14956; small and separated prefrontals on both sides in QCAZ 14954 and one prefrontal on right side in QCAZ 14956; little intrusive scales between parietal and postparietal in QCAZ 14954; frontal hexagonal in QCAZ 14956; roughly decagonal interparietal in QCAZ 14954. Usually two scales on posterior cloacal plate, four in QCAZ 14954 and 14956. Male is larger (SVL 46.7 mm, N = 1) than females (maximum SVL 45.4 mm, N = 2). Hatchlings (QCAZ 14969, 14970, 14976) with eight or seven (QCAZ 14976) posterior gular (collar) scales. Unlike the male holotype, females have an orange-brown longitudinal stripe extending from third supralabial to shoulder and fading on the flanks (Fig. 6).

Pholidobolus samek inhabits cloud forests in Cordillera del Cóndor, southeastern Ecuador at elevations between 2324−2844 m (Fig. 7). The new species is known only from Zamora-Chinchipe province, on the sandstone plateaus of Cerro Plateado Biological Reserve. The ground at the type locality is covered with mosses, roots, and bromeliads. Such ground cover is locally known as bamba. All specimens were found active at 11h30–17h00 under stones or terrestrial bromeliads (Fig. 8). Four eggs, collected under flat stones on 24-09-2016, were incubated in sphagnum and perlite in captivity for two months approximately. They were 14.0–14.1 mm long, 8.0–8.5 mm wide, and weighted 0.4 g on average. Hatchlings (QCAZ 14969–70, 14976–77) weighted 0.3 g and were 24.7 mm in SVL on average.

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 v3.10.

Pholidobolus samek is only known from Cordillera del Cóndor. The population size for this species is unknown, but our sampling suggests low abundances. Because of the small known distribution, as well as habitat destruction through mining activities nearby (Van Teijlingen 2016), we suggest assigning P. samek to the Critically Endangered category under criteria B1a, b(iii); C1; D, according to IUCN (2012) guidelines.

Figure 8. 

Habitat of Pholidobolus samek sp. nov. A Vegetation around type locality, Cerro Plateado Biological Reserve, Ecuador B habitat where holotype was found. Photographs by Álvaro Pérez.

The specific epithet samek means green in the Shuar language, in allusion to the green dorsolateral head stripes distinguishing the new species from other congeners. The type locality of Pholidobolus samek lies within territory of Shuar indigenous people, who inhabit the Amazonian rainforest in Ecuador and Peru.

Pholidobolus samek sp. nov. is very similar morphologically and genetically to P. condor sp. nov. These species can be easily distinguished from each other by coloration in adult males, although we recognize that our sample size is small (N = 7 and 4, respectively) and includes only one adult male per species. However, further evidence supports recognition of P. samek and P. condor as different species. First, they are reciprocally monophyletic and they are not sister taxa, with P. samek being sister to “P. macbrydei” Clade A (Fig. 1), which is very different in color patterns from either P. samek or P. condor (V. Parra and O. Torres-Carvajal, personal observation). Second, unlike the 12S gene (the less variable gene in this study), genetic distances between P. samek and P. condor for 16S and ND4 are not the lowest (Tables 2 and 4, respectively) within Pholidobolus. For example, the 16S distance between P. samek and P. condor (3%) is the same as the distance between the well-recognized species P. paramuno and P. affinis. In addition, genetic exchange among P. samek, P. condor and Clade A is very unlikely as they are isolated from each other on mountaintops above 2000 m (Fig. 7).

QCAZ 15844 (Figs 9, 10), adult male, Ecuador, Provincia Morona Santiago, buffer zone of El Quimi Biological Reserve, plateau on the eastern side of El Quimi river valley, 3.51892S, 78.3690W, WGS84, 2209 m, 11 July 2017, collected by Diego Almeida, Darwin Núñez, Eloy Nusirquia, Alex Achig and Ricardo Gavilanes.

Figure 9. 

Holotype of Pholidobolus condor sp. nov. (QCAZ 15844) in dorsal (A), ventral (B), and lateral (C) views. Male, SVL = 42.7 mm. Preserved specimen (A); live specimen (B, C). Photographs by Malki Bustos.

Ecuador: Provincia Morona Santiago: QCAZ 16790 (hatchling), El Quimi Biological Reserve, base camp towards old heliport (high zone), 3.51894S, 78.36897W, WGS84, 2226 m, 17 April 2018; QCAZ 16788–89 (hatchlings), El Quimi Biological Reserve, near base camp, 3.5182S, 78.3913W, WGS84, 1994 m, 12 April 2018, collected by Diego Almeida, Darwin Núñez, Eloy Nusirquia, Alex Achig and María del Mar Moretta.

Figure 10. 

Head of holotype of Pholidobolus condor sp. nov. (QCAZ 15844) in lateral (A), dorsal (B), and ventral (C) views. Photographs by Valeria Chasiluisa. Scale bar: 5 mm.

Pholidobolus condor is unique among its congeners, except P. samek sp. nov., in having green dorsolateral stripes on the head. However, adult males of P. condor differ from those of P. samek sp. nov. in having lighter dorsolateral head stripes, and reddish flanks and venter. In addition, P. ulisesi, P. dicrus, P. hillisi, and P. vertebralis differ from P. condor (character states of P. condor in parentheses) in having a conspicuous light vertebral stripe (light vertebral stripe absent). Pholidobolus affinis, P. prefrontalis, P. dicrus, P. hillisi, and P. vertebralis further differ from P. condor in having prefrontal scales (prefrontal scales absent). Additionally, P. condor sp. nov. has fewer dorsal scales (26−30) than P. affinis (45−55), P. montium (35−50), P. prefrontalis (37−46), P. macbrydei (31−43), and P. dolichoderes sp. nov. (35−40). Pholidobolus condor can be further distinguished from P. fascinatus sp. nov. by having widened medial scales on collar, and from P. dolichoderes sp. nov. by having fewer temporals (7–9 and 4–5, respectively), fewer ventrals (18–20 and 25–27), and fewer gulars (14–16 and 22–23).

(1) Two (rarely three) supraoculars, anteriormost larger than posterior one; (2) prefrontals absent; (3) femoral pores absent; (4) four opaque lower eyelid scales; (5) scales on dorsal surface of neck striated or smooth, progressively striated from forelimbs to tail; (6) two rows of lateral granules at midbody; (7) 27−31 dorsal scales between occipital and posterior margin of hindlimb; (8) lateral body fold present; (9) keeled ventrolateral scales on each side absent; (10) dorsum dark brown with a narrow, pale brown stripe; (11) labial stripe white; (12) flanks of body dark brown or gray; (13) hemipenial body cylindrical with distinctive capitular groove.

Adult male (QCAZ 15844) (Figs 9, 10); SVL 42.7 mm; TL 74.8 mm; dorsal and lateral head scales juxtaposed, finely wrinkled; rostral hexagonal, 1.67 times as wide as high; frontonasal quadrangular, slightly bigger than frontal, laterally in contact with nasal, loreal and first superciliary; prefrontal scales absent; frontal pentagonal, longer than wide, wider anteriorly, in contact with first superciliary and supraocular; frontoparietals hexagonal, longer than wide, slightly wider in the middle, each in contact laterally with supraocular II; interparietal octagonal, with a short medial suture posteriorly, lateral borders nearly parallel to each other; parietals larger than interparietal, hexagonal and positioned anterolaterally to interparietal, each in contact laterally with supraocular II and dorsalmost postocular; postparietals three, medial scale smaller than lateral ones; eight supralabials, fourth one longest and below center of eye; six infralabials, fourth one below center of eye; temporals enlarged, irregularly hexagonal, smooth; two large and smooth supratemporals; nasal shield slightly divided above nostril, irregularly pentagonal, longer than high, in contact with rostral anteriorly, first and second supralabials ventrally, frontonasal dorsally, loreal posterodorsally and frenocular posteroventrally; nostril on ventral aspect of nasal, directed laterally; loreal quadrangular, slightly wider dorsally, not in contact with supralabials; frenocular higher than long, in contact with nasal; nasal separating loreal from supralabials; two supraoculars, anteriormost one the widest; four elongate superciliaries, anteriormost enlarged, in contact with loreal; palpebral disc divided into five pigmented scales; four suboculars, anteriormost three elongated and homogeneous in size, posteriormost widest; two postoculars, the dorsalmost wider than the other; 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; posterior row of gulars (collar) with nine scales, the medial three slightly 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, imbricate, arranged in transverse rows; scales on dorsal surface of neck striated, becoming progressively keeled from forelimbs to tail; dorsal scales between occipital and posterior margin of hindlimbs 27; dorsal scale rows in a transverse line at midbody 27; one longitudinal row of smooth, enlarged ventrolateral scales on each side; dorsals separated from ventrals by two rows of small scales at the level of 13^th row of ventrals; lateral body fold between fore and hindlimbs present; ventrals smooth, wider than long, arranged in 20 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 striated, imbricate; scales on ventral surface of forelimb 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, 9/9 on III, 12/12 on IV, 7/7 on V; subdigital lamellae of finger I, II, III, and V single, on finger IV few scales in the middle paired; subdigital lamellae 6/6 on finger I, 11/11 on II, 15/15 on III, 17/16 on IV, 10/10 on V; subdigital lamellae on toes I and II single, on toes III, IV and V paired, except for two or three distalmost subdigitals; subdigital lamellae 7/6 on toe I, 12/12 on II, 15/16 on III, 22/22 on IV, 12/12 on V; groin region with small, juxtaposed 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 absent; 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 11.0; HW 6.6; ShL 5.8; AGD 20.4; TL/SVL 1.7; HL/SVL 0.3; HW/SVL 0.2; ShL/SVL 0.1; AGD/SVL 0.5.

Dorsal background from head to base of tail dark brown, with a golden brown vertebral stripe extending from occiput to tail; greenish cream dorsolateral stripes on head, becoming light brown on posterior part of body; white longitudinal stripe extending from first supralabial to shoulder; sides of neck, flanks and limbs dark brown; chocolate brown narrow stripe extending from tympanum to arm insertion; ventrolateral region of body grayish brown; throat reddish cream; chest, belly, base of tail and lateral region of tail bright orange, with brown marks in some scales; ventral surface of hind limbs with orange diffuse marks (Fig. 9B,C).

Dorsal background uniformly dark brown with a grayish brown middorsal stripe extending from occiput onto tail; dorsolateral stripe distinct, pale gray, extending from snout to near base of tail; head brown dorsally (rostral, frontonasal, frontal, frontoparietals and supraoculars) and dark brown laterally; white longitudinal stripe extending from first supralabial to forelimb; lateral aspect of neck dark brown with a dorsolateral light brown stripe extending posteriorly along flanks to hindlimbs; flanks grayish brown; tail dark brown dorsally and bronze laterally; ventral surface of head gray, with dirty cream genials and scattered black marks; chest, belly and ventral surface of tail light gray with light red spots; ventral surface of limbs dark gray (Fig. 9A).

Measurements and scale counts of Pholidobolus condor are presented in Table 6. Supraoculars three on left side in specimen QCAZ 16789; supralabials six in QCAZ 16789 and 16790, and seven in QCAZ 16788; two quadrangular frontonasals in QCAZ 16788; transverse rows of ventral scales between collar fold and preanals 18 in QCAZ 16788 and 19 in QCAZ 16790. Hatchlings with eight (QCAZ 16788–89) or six (QCAZ 16790) posterior gular (collar) scales. Unlike the adult male, hatchlings lack reddish color on tail.

Pholidobolus condor occurs in Cordillera del Cóndor in southeastern Ecuador at elevations between 1994–2226 m. The new species is known from El Quimi Biological Reserve in Morona Santiago province (Fig. 7). The holotype was found active at 21h14 at the base of a bromeliad on a sandstone plateau of shrub vegetation (Fig. 11).

Figure 11. 

Habitat of Pholidobolus condor sp. nov. at El Quimi Biological Reserve, Ecuador. Photographs by Álvaro Pérez.

Several eggs were found within a bromeliad, suggesting that females of P. condor lay their eggs in communal nests. Four eggs that were found on the ground at the base of bromeliads and under a trunk were incubated in sphagnum and perlite in captivity for approximately three months. On average, hatchlings weighted 0.4 g and were 23.7 mm in SVL.

Pholidobolus condor is only known from Cordillera del Cóndor in southeastern Ecuador. This area is currently threatened by mining activities (Ron et al. 2018; Valencia et al. 2017; Van Teijlingen 2016). Habitat destruction and fragmentation is evident at a distance of ~11 km from the collection sites (Mazabanda et al. 2018). Because of the small known distribution and habitat disturbance, we suggest assigning P. condor to the Critically Endangered category under criteria B1a, b(iii); C1; D, according to IUCN (2012) guidelines.

The specific epithet condor refers to Cordillera del Cóndor, where the new species was discovered. The Cordillera del Cóndor is an eastern outlier of the main Andean chain, where a significant number of species have been discovered in the last decade (Brito et al. 2017; Huamantupa-Chuquimaco and Neill 2018; Ron et al. 2018; Torres-Carvajal et al. 2009; Valencia et al. 2017).

See remarks on Pholidobolus samek sp. nov. above.

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.

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 Pazmiño.

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.

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.

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).

(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.

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 13^th 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.

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).

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 Pazmiño.

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.

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.

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.).

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.

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.

QCAZ 15120 (Figs 15, 16), adult male, Ecuador, Provincia El Oro, Lake Chillacocha, 3.4984S, 79.6188W, WGS84, 3382 m, 20 November 2016, collected by Diego Almeida, Darwin Núñez, Eloy Nusirquia, Santiago Guamán and Guadalupe Calle.

Figure 15. 

Holotype of Pholidobolus fascinatus sp. nov. (QCAZ 15120) in life in dorsal (A), ventral (B), and lateral (C) views. Male, SVL = 52.5 mm. Photographs by Diego Quirola.

Ecuador: Provincia El Oro: QCAZ 15122 (adult male), QCAZ 15121 (adult female), QCAZ 15169−73, 15177−78, 15180, 15193, 15221, 15243−44, 15396−15405 (juveniles), same data as holotype; QCAZ 15118 (adult female), Lake Chillacocha, 3.4986S, 79.6187W, WGS84, 3348 m, 17 November 2016, same collectors as holotype.

Figure 16. 

Head of holotype of Pholidobolus fascinatus sp. nov. (QCAZ 15120) in lateral (A), dorsal (B), and ventral (C) views. Photographs by Valeria Chasiluisa. Scale bar: 5 mm.

Pholidobolus fascinatus is unique among its congeners in lacking widened medial scales on collar (posterior row of gulars). In addition, P. fascinatus differs from P. affinis, P. prefrontalis, P. macbrydei, P. dolichoderes sp. nov., and P. montium in having a loreal scale frequently in contact with the supralabials (loreal scale, if present, not in contact with supralabials in the other species). Pholidobolus ulisesi, P. dicrus, P. hillisi, P. paramuno, and P. vertebralis differ from P. fascinatus in having a conspicuous light vertebral stripe. Pholidobolus samek sp. nov. and P. condor sp. nov. differ from P. fascinatus in having bright green dorsolateral stripes on the head. In addition, P. fascinatus has more dorsals (32–37) and ventrals (21–25) than P. samek sp. nov. (27–29 and 19–21, respectively) and P. condor sp. nov. (26–30 and 18–20); and it has fewer temporals (3–5) and gulars (14–17) than P. dolichoderes sp. nov. (7–9 and 22–23, respectively).

(1) Two (rarely three) supraoculars, anteriormost larger than posterior one; (2) prefrontals present or absent; (3) femoral pores absent in both sexes; (4) four to six opaque lower eyelid scales; (5) scales on dorsal surface of neck smooth, becoming striated from forelimbs to tail; (6) one row of lateral granules at midbody; (7) 32–37 dorsal scales between occipital and posterior margin of hindlimb; (8) lateral body fold present; (9) dorsum brown with a diffused chocolate brown middorsal stripe that fades away towards tail; (11) labial stripe white or cream; (12) flanks of body brown; (13) conical hemipenial body, with sulcus spermaticus originating between distinctly thick lips; (14) 22 flounces extending along hemipenial body.

Adult male (QCAZ 15120) (Figs 15, 16); SVL 52.5 mm; TL 37.6 mm; dorsal and lateral head scales juxtaposed, finely wrinkled; rostral hexagonal, 2.27 times as wide as high; frontonasal hexagonal, wider than long, in contact with nasal laterally, slightly larger than frontal; prefrontal scales irregularly pentagonal; frontal heptagonal, longer than wide, slightly wider anteriorly, in contact with prefrontals and frontonasal anteriorly, two supraoculars laterally, and frontoparietals posteriorly; frontoparietals pentagonal, longer than wide, slightly wider posteriorly, each in contact laterally with supraocular II; interparietal heptagonal, lateral borders nearly parallel to each other; parietals hexagonal, each in contact laterally with supraocular II and dorsalmost postocular; postparietals four, with medial scales less than half the size of lateral ones; eight supralabials, fourth one the longest and below center of eye; eight infralabials, third and fourth one below center of eye; temporals enlarged, irregularly hexagonal, juxtaposed, smooth; two large, smooth supratemporal scales; nasal divided, irregularly pentagonal, longer than high, in contact with rostral anteriorly, first and second supralabials ventrally, frontonasal dorsally, loreal posterodorsally and frenocular posteroventrally; nostril in center of nasal, directed lateroposteriorly; loreal rectangular, wider ventrally; frenocular longer than high, higher anteriorly, in contact with nasal, separating loreal from supralabials; two supraoculars, homogeneous in size; four superciliaries, anteriormost enlarged and in contact with loreal; palpebral disc divided into five pigmented scales; suboculars elongated, four on right side and three on left side; two postoculars, dorsalmost wider than the other; ear opening vertically oval, without denticulate margins; tympanum recessed into a shallow auditory meatus; mental semicircular, longer than wide; postmental pentagonal, slightly longer than wide, 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; posterior row of gulars (collar) with 11 scales that are similar in size.

Nuchal scales similar in size to dorsals, except for the anteriormost that are widened; scales on sides of neck small and slightly granular; dorsal scales hexagonal, elongate, imbricate, arranged in transverse rows; scales on dorsal surface of neck smooth, becoming progressively striated from forelimbs to tail; dorsal scales between occipital and posterior margin of hindlimbs 33; dorsal scale rows in a transverse line at midbody 25; dorsals separated from ventrals by one row of small scales at level of 13^th row of ventrals; lateral body fold between fore and hindlimbs present; ventrals smooth, wider than long, arranged in 25 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 striated, imbricate; scales on ventral surface of forelimb granular; two thick, smooth thenar scales; supradigitals (left/right) 3/3 on finger I, 7/6 on II, 8/8 on III, 10/10 on IV, 5/5 on V; supradigitals 3/3 on toe I, 6/6 on II, 8/9 on III, 11/11 on IV, 8/8 on V; subdigital lamellae of finger I single, on finger II all paired, except by the three distalmost, on finger III (proximal half) paired, on finger IV slightly paired at the middle, on finger V all single in right finger and three paired in left finger; subdigital lamellae 5/5 on finger I, 9/9 on II, 13/13 on III, 14/15 on IV, 9/9 on V; subdigital lamellae on toes I and II paired proximally and single distally, on toes III, IV and V paired, except for the three to five distalmost subdigitals; subdigital lamellae 5/5 on toe I, 10/10 on II, 14/13 on III, 18/18 on IV, 11/12 on V; groin region with small, imbricate scales; scales on dorsal surface of hindlimbs smooth and imbricate; scales on ventral surface of hindlimbs smooth; scales on posterior surface of hindlimbs granular; femoral pores absent; preanal pores absent; cloacal plate paired, bordered anteriorly by two enlarged scales.

Additional measurements (mm) and proportions of the holotype: HL 12.3; HW 9.2; ShL 6.7; AGD 26.5; TL/SVL 0.7; HL/SVL 0.2; HW/SVL 0.2; ShL/SVL 0.1; AGD/SVL 0.5.

Dorsal background from head to base of tail brown, with a diffuse chocolate-brown middorsal stripe that fades away towards tail; light brown dorsolateral stripes on head extending posteriorly and fading away at midbody; white longitudinal stripe extending from third supralabial to shoulder; sides of neck, flanks, and limbs brown; reddish brown narrow stripe extending from tympanum to arm insertion; ventrolateral region of body grayish brown; throat and chest gray; belly background gray with conspicuous orange marks; tail orange anteriorly and laterally (Figs 15, 17A).

Figure 17. 

Color variation in live specimens of Pholidobolus fascinatus sp. nov. A male holotype (QCAZ 15120, SVL = 52.5 mm) B male paratype (QCAZ 15122, SVL = 42.6 mm) C female paratype (QCAZ 15118, SVL = 46.7 mm).

Dorsal background uniformly brown with a cream brown vertebral stripe extending from head onto tail; vertebral stripe slender anteriorly, becoming slightly wider posteriorly; head light brown with black dots dorsally (rostral, frontonasal, frontal, frontoparietals and supraoculars) and brown laterally; cream longitudinal stripe extending from third supralabial to shoulder; ventrolateral aspect of neck brown; forelimbs with scattered black dots; flanks brown; tail brown dorsally; ventral surface of head light gray, chest and venter dark gray, ventral surface of tail slightly brown, with scattered dark brown marks.

Measurements and scale counts of Pholidobolus fascinatus are presented in Table 6. Supralabials 9/9 (left/right) in specimens QCAZ 15118 and 15122, and supraoculars 3/3 in QCAZ 15118; loreal scale absent in QCAZ 15118; prefrontals absent in QCAZ 15122 and 15173; little intrusive scales between postparietal and frontoparietals in QCAZ 15118, 15121 and 15122; frontonasal quadrangular in QCAZ 15122; frontal nonagonal and pentagonal in QCAZ 15118 and 15173, respectively; interparietal hexagonal in QCAZ 15122; parietal pentagonal in QCAZ 15170. Four posterior cloacal scales in QCAZ 15118. Males are slightly smaller (SVL 47.6 mm, N = 2) than female (maximum SVL 48.2 mm, N = 2). Adult male QCAZ 15122 differs from holotype in having sides of tail and chest dark brown without gray spots. Adult female QCAZ 15118 differs from holotype in having a light gray chest, a dark gray ventral surface of tail, dark brown sides of tail, and in lacking orange or red brown color on sides of neck (Fig. 17).

Pholidobolus fascinatus inhabits wet paramo in the western slopes of the Andes of southern Ecuador (Fig. 7). The new species is known only from El Oro province, at 3348−3382 m. All specimens were found active at 14h00–17h00 mostly under stones.

We found 41 eggs (17 as fragmented eggshells) in a communal nest next to male QCAZ 15120. We incubated the 24 unhatched eggs in soil and perlite in captivity. They were 11.9–13.2 mm long, 5.5–9.2 mm wide, and weighted 0.5 g on average. Hatchlings (N = 20) weighted 0.4 g and were 26.2 mm in SVL on average.

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

The species epithet fascinatus is a Latin word meaning enchanted, in allusion to Lake Chillacocha, also known as the Enchanted Lake. According to local belief, this lake is enchanted and has healing powers.