Pholidobolus celsiae, Amézquita & Mazariegos-H & Cañaveral & Orejuela & Barragán-Contreras & Daza, 2023

Amezquita, Adolfo, Mazariegos-H, Luis A., Canaveral, Santiago, Orejuela, Catalina, Barragan-Contreras, Leidy Alejandra & Daza, Juan M., 2023, Species richness under a vertebral stripe: integrative taxonomy uncovers three additional species of Pholidobolus lizards (Sauria, Squamata, Gymnophthalmidae) from the north-western Colombian Andes, ZooKeys 1141, pp. 119-148 : 119

Pholidobolus celsiae sp. nov.

Type material.

Holotype. Adult male, with genitalia in a separate microvial. Original label: AA_7061. Museum ID: MHUA-R13906. Type locality in Colombia, Risaralda: Municipality of Mistrató, 5°28.01'N, 75°53.44'W, secondary forest, under rocks, 7 October 2020. Collected by Ubiel Rendón and Luis A. Mazariegos-H.

Paratypes. Eleven males, two females, and one juvenile. Table 2 shows field codes, localities, elevation, and geographic coordinates. Twelve specimens were collected in Colombia, Risaralda: Mistrató, Mesenia-Paramillo Nature Reserve (MPNR), May 2018, June 2019, and October 2020. Collected by Ubiel Rendón, Luis A. Mazariegos-H., Jorge Jaramillo, and Osman López. One from Colombia: Risaralda, Municipality of Mistrató, Mampay village. Collected by Juan P. Hurtado. The other from Colombia, Risaralda: Municipality of Pereira, vereda La Suiza, Santuario de Fauna y Flora Otún Quimbaya. Collected by Melisa Galeano.

Diagnosis.

The species can be diagnosed combining the following characters: (1) three supraocular scales; (2) prefrontal scales present; (3) 14-28 temporal scales; (4) dorsal scales keeled; (5) 28-32 transverse rows of dorsal scales; (6) 18-21 transverse rows of ventral scales; (7) 36-44 scales around mid-body; (8) 1-3 rows of lateral scales; (9) lateral and medial ventral scales equal in size; (10) 5-6 femoral pores; (11) no sexual dimorphism in number of femoral pores; (12) labial scales pale, often with black markings; (13) ventral head colouration homogeneous in females; with irregular orange or black markings, and paler towards the anterior half in males; (14) white to cream vertebral stripe bordered by two black stripes, 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 on the body; (15) lateral colour pattern brown and dark orange to red, with numerous ocelli, usually more than seven between the limbs insertions, white in centre and surrounded by black scales, with a longitudinal pale line laterally, continuous and white in the head, pale and discontinuous towards the body; (16) venter pink to pale orange, or brown, with darker marking towards the edge of scales in females; vivid orange to red, with scattered black markings towards the edge of scales in males; (17) subcylindrical and bilobed hemipenial body with 4-5 and 7-9 rows of spinulated flounces in the lateral columns of the sulcate and asulcate sides, respectively; (18) lateral columns of spinulated flounces connecting in the distal region of the asulcate side.

Comparisons.

Pholidobolus vertebralis differs from P. celsiae sp. nov. (character states in parenthesis) in having the lateral ventral scales smaller than the medial ventrals (lateral and medial ventral scales equal in size). The other species from the north-western and central Colombian Andes (Fig. 7 ) differ from P. celsiae sp. nov. in exhibiting smaller adult body size in males (Table 2 ), between 35.4-54.7 mm in P. odinsae sp. nov., and 42.6-57.9 mm in P. argosi sp. nov. (60.7-68.6 mm). In addition, males of P. argosi sp. nov. lack prefrontal scales (present) and have two supraocular scales (3-4). Lastly, males of P. odinsae sp. nov. exhibit black to grey and males of P. paramuno reddish brown ventral coloration (orange).

Description of the holotype.

Adult male; snout-vent length 68.2 mm; tail length 79.0 mm; other body measurements in Table 4 . Head scales smooth, juxtaposed, glossy, with small pits organised mainly around their margins. Rostral single, hexagonal, wider than high, dorsally in broad contact with the internasal and laterally in contact with the first supralabial and the nasal. Frontonasal single, wider than long, pentagonal, in contact with the nasal, loreal and prefrontals. Prefrontals two, wider laterally and narrower medially, in wide contact with the first superciliary, the frontal and the anterior supraocular. Frontal single, hexagonal, longer than wide, wider anteriorly, in contact with the prefrontals, the first supraocular and the frontoparietals. Frontoparietals two, pentagonal, longer than wide, narrower anteriorly, contacting the first two supraoculars laterally, and the parietal and interparietals posteriorly. Supraoculars three, the anteriormost nearly as wide as long and the other two wider than long, decreasing in size antero-posteriorly, contacting the superciliaries laterally and the parietal and postocular posteriorly. Interparietal single, heptagonal, longer than wide, narrower than the parietals and contacting laterally the parietals and posteriorly the postparietals. Parietals two, hexagonal, wider than long, slightly shorter and wider than the interparietal, contacting the temporals laterally and the postparietals posteriorly. Postparietals in two rows, three in the anterior row and four in the posterior row. Nasal single, wider than high, contacting the first and second supralabials, the loreal and frenocular. Loreal single, quadrangular, over the frenocular, in contact with first superciliary dorsally. Frenocular single, triangular, in contact with the first infraocular and the second and third supralabials. Superciliaries four, the anteriormost noticeable larger than the others, contacting the uppermost preocular. Suboculars five contacting supralabials three to five. Postoculars two, ventral larger than dorsal. Temporals 17 contacting supralabials five to seven. Supralabials seven and infralabials five. Mental single, pentagonal, wider than long, contacting the first infralabial and postmental. Postmental single, pentagonal, contacting the first two infralabials and the anterior genials. Genials in three pairs, the anterior one quadrangular and the posterior two pentagonal. The anterior two in contact medially and the posterior one separated by postgenials; contacting infralabials two, three, and four. Pregulars two. Gular scales seven, wider than long, in two longitudinal rows; collar scales 13 decreasing in size laterally. Dorsal scales longer than wide, hexagonal, keeled, imbricate, arranged in 29 transverse rows. Longitudinal rows of dorsal scales 23, the first two rows in each side weakly keeled and rounded. Lateral row scales at mid-body one, smooth, at least half the size of adjacent scales. Scales around mid-body 39. Longitudinal rows of ventrals six, quadrangular. Transverse rows of ventrals 20. Cloacal plates in two rows of two quadrangular scales each, the posterior row larger than the anterior one, in contact with two small scales laterally. Tail scales arranged in 54 rings, hexagonal and keeled dorsally, quadrangular and smooth ventrally.

Limbs pentadactyl with clawed fingers. Dorsal brachial and antebrachial scales lanceolate to polygonal, almost as long as wide, imbricate and smooth. Ventral brachial and antebrachial scales lanceolate to polygonal, almost as long as wide, juxtaposed, much smaller than the dorsal ones. Dorsal hand scales hexagonal, wider but shorter than the dorsal antebrachial scales. Finger length formula IV = III> II> V> I. Supradigital scales quadrangular and imbricate. Palmar scales polygonal, juxtaposed and small. Subdigital lamellae domelike with a quadrangular base, and often divided longitudinally, with four on finger I, 8 on II, 12 on III, 13 on IV, and 7 on V. Thigh scales on the dorsal, anterior and ventral surfaces lanceolate to rhomboidal, longer than wide, those in the dorsal surface keeled and the others smooth and imbricate. Thigh scales on the posterior surface of the legs rounded, smooth, juxtaposed and much smaller than those of the anterior and dorsal surfaces. Five femoral pores per leg; preanal pores absent. Anterior and ventral crus scales polygonal and keeled. Lateral and posterior crus scales rounded, small and subimbricate. Toe length formula IV> III> II> IV> I. Supradigital scales quadrangular, imbricate and longer than wide. Plantar scales polygonal, juxtaposed and small. Subdigital lamellae domelike with a quadrangular base, and often divided longitudinally, with four on Toe I, 8 on II, 13 on III, 15 on IV, and 9 on V. Thigh scales on the dorsal, anterior and ventral surfaces lanceolate to rhomboidal, longer than wide, those in the dorsal surface keeled and the others smooth and imbricate. Thigh scales on the posterior surface of the legs rounded, smooth, juxtaposed and much smaller than those of the anterior and dorsal surfaces. Five femoral pores per leg; preanal pores absent. Anterior and ventral crus scales polygonal and keeled. Lateral and posterior crus scales rounded, small and subimbricate. Toe length formula IV> III> V> II> I. Supradigital scales quadrangular, imbricate and longer than wide. Plantar scales polygonal, juxtaposed and small. Subdigital lamellae domelike with a quadrangular base, and often divided longitudinally, with seven on I, nine on II, 13 on III, 18 on IV, and 10 on V.

Colouration.

In life, dorsally dark brown, bisected by a mid-dorsal (i.e. vertebral) cream, or white stripe, extending from the head to the base of the tail; vertebral stripe bordered with darker, usually black, stripes; on the head, the pale stripe extends from the first supralabial to the shoulder dorsally reaching the rostral scale, and laterally not in contact with the supraocular and parietal scales; sides of neck, flanks, and limbs predominantly brown; neck, flanks and tail base usually with more than 10 white ocelli, bordered by a black stripe; white or cream lateral line from the supralabials to the shoulder; cream and interrupted lateral stripe, running between the insertions of fore and hind limbs, not extending towards the tail; many red scales, more common in males and grouped above the shoulder and along the lateral surface of the tail; throat cream to pale brown in males, paler towards the anterior extreme; throat pink in females; chest, belly and base of the tail cream to pink in females, but orange in males, often with black blotches, apparently more common in adult males (Figs 1 , 3 , 6 ). In preservative, brown surfaces become paler, the dorsal and lateral stripes become white, and the red surfaces on the flanks, chest, belly and tail fade to white or very pale pink.

Etymology.

The species epithet is dedicated to the Celsia Foundation, for their voluntary contribution to the restoration of cloud and dry forests in the tropical Andes, through their reforestation program Reverde-C, which already planted more than one million trees. In addition, their program for children education in rural areas, already benefited more than 16000 students in terms of school infrastructure, teacher training, and further logistic support during the Covid pandemic. We believe their commitment contributes to the well-being and education of direct neighbours and thereby stakeholders of Colombian protected nature.

Distribution, ecology, and conservation.

The specimens were mostly collected in open areas with secondary vegetation, at the edge of a cloud forest. Groups of up to nine eggs were found together with adult individuals under a rock, suggesting communal nesting. Also, the observed specimens appeared clearly heliothermic: within minutes after the sun appeared, they came out of their refuges, remained exposed, and extended their ribs increasing the dorsal surface available for sun basking. The species is currently known from three localities, two of them within protected areas: the Mesenia-Paramillo Nature Reserve, and the Santuario de Flora y Fauna (SFF) Otún-Quimbaya. Further explorations are needed to ascertain the species distribution. In the meantime, we suggest listing the new species as Endangered EN B1ab(iii), B2ac(iii), under the IUCN criteria ( IUCN 2012). Many individuals showed signs of a regenerated tail.

Figures

Figure 1. Molecular phylogenetic hypothesis. Recovered relationship between species of the leaf-litter lizards in the genus Pholidobolus, including P. vertebralis near its type locality in Ecuador. The three recovered clades from the Colombian Western Andes are outlined in blue (clade A), orange (clade B) and yellow (clade D). A fourth clade of the Colombian Central Andes is outlined in green (clade C). Green dots indicate nodal support of at least 95 %. See Methods for further details on the phylogenetic analysis and Suppl. material 2 for the detailed tree with individuals as terminal nodes.

Figure 3. Key morphological features of the three new species on the holotypes. External morphological traits allowing unambiguous diagnosis of the three north-western Colombian clades of Pholidobolus lizards recovered in our phylogenetic analysis. Pholidobolus argosi sp. nov. lacks one supraocular (numbered) and the prefrontal (filled in red) scales, both of which are present in the other two species. Males of P. odinsae sp. nov. exhibit predominantly black and dark grey ventral colouration, which is red with black markings in males of the two other species, as further shown by the image segmentation analyses: the stacked areas denote the proportion of pixels with each summarised hue. Lastly, P. celsiae sp. nov. reaches larger adult body size than the other two species, as evidenced by blue (clade A in Fig. 1), orange (B), and yellow (D) bars, which denote range of variation, and the black points, that represent the actual body size of the shown holotypes.

Figure 6. Variation in lizard ventral colouration. Among-species, among-sexes, and among-individuals variation in ventral colouration of three lizard species in the genus Pholidobolus. Pictures were taken immediately after euthanisation to reflect colour in life. To depict eventual covariation with body size, individuals are sorted from the smallest (extremes) to the largest (middle) one.

Figure 7. Geographic and altitudinal distribution. Geographic (A) and altitudinal (B) distribution of five species of Pholidobolus lizards across the north of the Western and Central Colombian Andes. The cladogram summarises the recovered phylogenetic relationship among them (Fig. 1). Each dot represents an individual deposited in the MHUA-R collection at the Universidad de Antioquia.