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Pholidobolus paramuno HURTADO-GÓMEZ, ARREDONDO, SALES-NUNES & DAZA, 2018

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Higher TaxaGymnophthalmidae (Cercosaurinae), Sauria, Gymnophthalmoidea, Squamata (lizards)
Subspecies 
Common Names 
SynonymPholidobolus paramuno HURTADO-GÓMEZ, ARREDONDO, SALES-NUNES & DAZA 2018
Prionodactylus vertebralis — UZZELL 1973: 6 (part)
Cercosaura vertebralis — HERNANDEZ-RUZ & BERNAL 2011 (part)
Cercosaura vertebralis — DOAN & CUSI 2014: 1197 (part) 
DistributionColombia (Antioquia), elevation 2700–3200 m

Type locality: near the Congo site (06°45′57.24′′N, 75°43′37.92′′W, 3,200 m) in the Santa Rita paramo between the municipalities of San José de la Montaña and Liborina, Antioquia Department, Colombia  
Reproductionoviparous (manual imputation, fide Zimin et al. 2022) 
TypesHolotype. MHUA R 12481, an adult male. Collected on November 5, 2011 by Felipe Duarte. Paratypes. (n = 11): MHUA R 12454 adult male, MHUA R 12455–12456 adult females, and MHUA R 12480 juvenile of indeterminate sex, collected with the holotype; MHUA R 10475–10476, CSJ-h 3594 adult males, MHUA R 10477, 12451–12452 adult females, and MHUA R 12453 juvenile of indeterminate sex, collected in the Belmira paramo (06°39′16.56′′N, 75°40′18.84′′W, 3,226 m), Belmira municipality, Antioquia, Colombia. 
DiagnosisDiagnosis and comparison. The species is assigned to the genus Pholidobolus on the basis of the molecular phylogenetic analysis (Fig. 1) and the presence of a palpebral area divided in several scales and a lateral fold between fore and hindlimbs (Figs. 4–5; Torres-Carvajal and Mafla-Endara, 2013). The new species can be diagnosed by the combination of the following characters: (1) 2–3 (usually 2) supraocular scales; (2) prefrontal scales present; (3) 8–13 temporal scales; (4) all dorsal scales keeled; (5) 28–33 transversal rows of dorsal scales; (6) 19–23 transversal rows of ventral scales; (7) 28–34 scales around mid-body; (8) 0–2 (usually 1) rows of lateral scales; (9) lateral and medial ventral scales equal in size; (10) 0–4 femoral pores; (11) no sexual dimorphism in number of femoral pores; (12) labial scales pale; (13) ventral head coloration homogeneous; (14) single golden vertebral stripe originating on the rostral scale, completely covering the dorsal region of the head and the vertebral region of the body, reaching only the anterior portion of the tail, with maximum width of two scales; (15) lateral color pattern brown with scattered black blotches, with a longitudinal pale line laterally; (16) venter red to reddish brown, with scattered black blotches; (17) hemipenial body with 19–20 and 21–22 rows of spinulated flounces in the lateral columns of the sulcate and asulcate sides, respectively; (18) lateral columns of spinulated flounces connecting in the proximal region of the asulcate side.
Pholidobolus vertebralis differs from P. paramuno sp. nov. (character states in parentheses) by having the lateral ventral scales smaller than the medial ventrals (lateral and medial ventral scales equal in size); golden vertebral stripe not covering the supraocular scales (vertebral stripe covers supraoculars), and extending from snout to tip of tail (vertebral stripe extends from snout to anterior portion of tail); and 23–27 and 25–29 rows of spinulated flounces in the lateral columns of the sulcate and asulcate sides of the hemipenial body, respectively (19–20 and 21–22). Pholidobolus hillisi Torres-Carvajal et al., 2014 may be distinguished from P. paramuno sp. nov. by having a hemipenis with 28–30 flounces (19–22); one diagonal white stripe on each side of the chin (none, homogeneous pigmentation), and by having a vertebral stripe extending to the posterior portion of the tail (vertebral stripe extends from snout to anterior portion of tail). Pholidobolus dicrus (Uzzell, 1973) differs from P. paramuno sp. nov. by having four supraoculars (2–3, usually two); 38–43 scales around mid-body (28– 34); 3–5 lateral scales (0–2, usually one); a higher number femoral pores and sexual dimorphism, with males having 20–25, and females 7–11 (not sexually dimorphic, 0–4); and two pale dorsal stripes that begin in the rostral, go parallel through the head and fuse posteriorly forming a vertebral pale stripe (dorsum with a single golden stripe). Pholidobolus ulisesi Venegas et al., 2016 is distinguished from P. paramuno sp. nov. by having the prefrontals absent (two), and by having the ventral coloration of the head with a diagonal white bar with dark brown edges (ventral coloration of the head homogeneous, without bars). Pholidobolus affinis, P. anomalus Müller, 1923, P.montium,P.macbrydei,andP.prefrontalisdifferfromthe new species by having the vertebral region of the dorsum cream, gray or brown, homogeneously colored (dorsum with a golden stripe bordered by two black lines). Pholi‐ dobolus anomalus has the dorsal scales smooth anteriorly and weakly keeled posteriorly (all dorsal scales keeled). Pholidobolus affinis, P. macbrydei, and P. montium have the venter yellow or orange-red (venter reddish brown or red, with disperse black blotches). Pholidobolus macbrydei and P. montium have several pale and dark dorsolateral stripes (one pale lateral stripe on each side). Pholidobolus prefrontalis has the venter gray to dark blue or black posteriorly in males (males with reddish or reddish brown, with disperse black blotches). Interspecific variation in scutellation among species of Pholidobolus is presented in Table 4. 
CommentHabitat: under rocks, inside dead leaves of the frailejon Espeletia occidentalis.

Behavior: Most specimens were found active during the day, usually in sunny conditions.

Sympatry: Riama sp. and Atractus sp. 
EtymologyThe specific epithet is an adjective referring to the word paramo, a high-altitude ecosystem restricted to the northern Andes of South America, where all specimens of the new species were found. 
References
  • Doan, Tiffany M.; Juan C. Cusi 2014. Geographic distribution of Cercosaura vertebralis O’Shaughnessy, 1879 (Reptilia: Squamata: Gymnophthalmidae) and the status of Cercosaura ampuedai (Lancini, 1968). Check List 10 (5): 1195-1200 - get paper here
  • Hernández‐Ruz E.J, Bernal C.A. 2011. Variación morfológica en Cercosaura vertebralis (Sauria: Gymnophthalmidae) en Colombia. Ingenierías & Amazonia 4:48–57
  • Hurtado-Gómez, Juan P.; Juan C. Arredondo, Pedro M. Sales-Nunes and Juan M. Daza 2018. A New Species of Pholidobolus (Squamata: Gymnophthalmidae) from the Paramo Ecosystem in the Northern Andes of Colombia. South American Journal of Herpetology 13 (3): 271–286 - get paper here
  • Uzzell, T. 1973. A revision of lizards of the genus Prionodactylus with a new genus for P. leucostictus and notes on the genus Euspondylus (Sauria, Teiidae). Postilla 159: 1-67 - get paper here
  • Zimin, A., Zimin, S. V., Shine, R., Avila, L., Bauer, A., Böhm, M., Brown, R., Barki, G., de Oliveira Caetano, G. H., Castro Herrera, F., Chapple, D. G., Chirio, L., Colli, G. R., Doan, T. M., Glaw, F., Grismer, L. L., Itescu, Y., Kraus, F., LeBreton 2022. A global analysis of viviparity in squamates highlights its prevalence in cold climates. Global Ecology and Biogeography, 00, 1–16 - get paper here
 
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