Pholidobolus odinsae, 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 odinsae sp. nov.

Type material.

Holotype. (Figs 1 , 3 ) Adult male, with genitalia in a separate microvial. Original field label: AA_7090. Museum ID: MHUA-R13907. Type locality in Colombia, Antioquia: Municipality of Jardín, Mesenia-Paramillo Nature Reserve, 5°29.76'N, 75°53.35'W, visitor centre, among pastures, 14 November 2020. Collected by Ubiel Rendón and Luis A. Mazariegos-H.

Paratypes. Six males, five females, and three juveniles. Table 2 shows field codes, localities, elevation, and geographic coordinates. Eleven specimens were collected in Colombia, Antioquia: Municipality of Jardín, Mesenia-Paramillo Nature Reserve (MPNR), between June 2018 and June 2020. Collected by Osman López, Ubiel Rendón, Jorge Jaramillo, and Luis A. Mazariegos. One from Colombia, Antioquia: Municipality of Andes, vereda Santa Rita, El Chaquiro. One from Colombia, Antioquia: Municipality of Jericó, vereda Quebradona, Finca La Aurora. The other from Colombia, Chocó: Municipality of Carmen de Atrato, vereda La Isla, Finca Gualandai.

Diagnosis.

The species can be diagnosed combining the following characters: (1) 3-4 (usually 3) supraocular scales; (2) prefrontal scales present; (3) 11-28 temporal scales; (4) dorsal scales keeled; (5) 28-32 transverse rows of dorsal scales; (6) 17-23 transverse rows of ventral scales; (7) 31-45 scales around mid-body; (8) 3-5 rows of lateral scales; (9) lateral and medial ventral scales equal in size; (10) 0-2 femoral pores; (11) no sexual dimorphism in number of femoral pores; (12) labial scales similar in colour to other head scales, crossed by a curved pale lip line, best described as two oblique white lines converging in the eye; (13) ventral head colouration homogeneous; (14) cream or white 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 four scales on the body; (15) lateral colour pattern brown, with a complete longitudinal line laterally, white and continuous from the posteroventral edge of the ear until the insertion of the hind limbs; with very few ocelli usually above the insertion of the forelimbs and absent between the limbs insertions, small; ocelli white in centre and surrounded by black scales and, beyond that, sometimes a few reddish scales; (16) venter strongly dimorphic in colouration between the sexes, uniformly pink to pale orange in females, sometimes with very few black speckles but no markings; usually glossy black and sometimes medium grey in males; (17) hemipenial body with 7-8 and 11-12 rows of spinulated flounces in the lateral columns of the sulcate and asulcate sides, respectively; (18) lateral columns of spinulated flounces connecting in the medial region of the asulcate side.

Comparisons.

Pholidobolus vertebralis differs from P. odinsae 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. odinsae sp. nov. in ventral colouration of males: reddish brown in P. paramuno , and pink to orange in P. argosi sp. nov. and P. celsiae sp. nov. (black to grey). In addition, males of P. argosi sp. nov. lack prefrontal scales (present) and have two supraocular scales (3-4). Lastly, males of P. celsiae sp. nov. are larger in size (Table 3 ), between 60.7-68.6 mm (35.4-54.7 mm).

Description of the holotype.

Adult male; snout-vent length 54.2 mm; tail length 50.0 mm; other body measurements in Table 4 . Head scales smooth, juxtaposed, glossy, with small pits organized 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 contact, touching the frontonasal, the frontal, the anterior supraocular, and the loreal. Frontal single, hexagonal, longer than wide, wider anteriorly, in contact with the prefrontals, the first supraocular and the frontoparietals. Frontoparietals pentagonal, longer than wide, narrower anteriorly, contacting one to three supraoculars laterally, and the parietal and interparietals posteriorly. Supraoculars four, the anterior most 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 uppermost 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 two in the posterior row. Nasal single, rhomboidal, 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, quadrangular in contact with the first infraocular and the second and third supralabials. Superciliaries three, the anteriormost noticeable larger than the others, contacting the uppermost preocular. Suboculars five, contacting supralabials three to five. Postoculars two, increasing in size antero-posteriorly. Temporals 26, contacting supralabials five to eight. Supralabials eight and infralabials six. Mental single, pentagonal, wider than long, contacting the first infralabial and post-mental. 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 eight, wider than long, in two longitudinal rows; collar scales 17, decreasing in size laterally. Dorsal scales longer than wide, hexagonal, keeled, imbricate, arranged in 30 transverse rows. Longitudinal rows of dorsal scales 24, 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 45. 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 62 rings, hexagonal and keeled dorsally, quadrangular and smooth ventrally.

Limbs pentadactyl with clawed fingers. Dorsal brachial and antebrachial scales lanceolate to polygonal, longer than 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, imbricate and longer than wide. Palmar scales polygonal, juxtaposed and small. Subdigital lamellae domelike with a quadrangular base, and often divided longitudinally, with six on finger I, 8 on II, 13 on III, 15 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 smooth 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. Two femoral pores per leg; preanal pores absent. Anterior and ventral crus scales polygonal and smooth. 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 five on Toe I, 9 on II, 13 on III, 15 on IV, and 7 on V.

Colouration.

In life, dorsally brown or pale brown, bisected by a mid-dorsal (i.e. vertebral) white stripe, extending from the head to the mid tail; vertebral stripe bordered with darker, usually dark brown or black, stripes; on the head, the pale stripe extends from the first supralabial to the shoulder dorsally reaching the rostral scale, and laterally including the frontonasal, prefrontal, frontal, frontoparietal, interparietal, and postparietal scales; sides of neck, flanks, and limbs predominantly brown, usually with less than five, small and white ocelli, bordered by a black stripe, and predominantly on the shoulders; white or cream lateral line from the supralabials, passing through the shoulder and extending continuously up to the insertion of the limbs, but not towards the tail; very few scattered red scales, more common around the shoulder ocelli; throat cream to pink in females, but grey to black in males; chest, belly and base of the tail pink to orange in adult females, but grey to black in males, with bare or no patterning in all cases (Figs 1 , 3 , 6 ). In preservative, brown surfaces become paler, the dorsal and lateral stripes become white, the orange chest, belly and tail of females fade to white or very pale pink, and the black chest, belly and tail of males fade into dark grey.

Etymology.

The species epithet is dedicated to the company Odinsa, for their decisive involvement in the Cartama Conservation Project, in southwestern Antioquia, aimed at restoring ecosystem services by regenerating the Andean forest along the Quebrada San Antonio basin. Together with other stakeholders, the initiative planted more than 320000 native trees during 2019-2020 alone.

Distribution, ecology, and conservation.

The species is currently known from forest edges, and open areas including pastures, crops, and around human buildings. Most specimens were seen and found amongst grass or leaf litter even hundreds of metres away from the nearest forests. They appeared clearly heliothermic: within minutes after the sun appeared, they came out of their refuges, remain exposed, and extended their ribs increasing the dorsal surface available for sun basking. Under sunny conditions, the species seems to be abundant at the known localities. Its distribution seems not to be patchy, and it is known from more than five locations. Although they encompass less than 500 km2 (Fig. 7 ), the actual distribution could arguably exceed this threshold area, given the species adaptability to disturbed habitats. Therefore, we suggest listing the new species as Data Deficient, DD ( IUCN 2012), until proper information is collected to evaluate the species conservation status. 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.