Afroedura vazpintorum, Branch & Schmitz & Lobón-Rovira & Baptista & António & Conradie, 2021

Branch, William R., Schmitz, Andreas, Lobon-Rovira, Javier, Baptista, Ninda L., Antonio, Telmo & Conradie, Werner, 2021, Rock island melody: A revision of the Afroedura bogerti Loveridge, 1944 group, with descriptions of four new endemic species from Angola, Zoosystematics and Evolution 97 (1), pp. 55-82 : 55

Afroedura vazpintorum sp. nov. Coastal Flat Gecko Osga-achatada da planície costeira Figures 5E, F , 10 ; Tables 3, 8

Synonym.

Afroedura bogerti - Branch et al. 2017b:157 (part); Marques et al. 2018: 177 (part); Branch et al. 2019a: 287 (part); Afroedura cf. bogerti - Butler et al. 2019:231; Afroedura bogerti (clade 2) - Branch et al. 2017a:146; Afroedura sp. - Baptista et al. 2018:400.

Holotype.

PEM R24118, adult female, collected 1 km east of Farm Mucungo (-14.78361, 12.49694, 314 m a.s.l.), Namibe Province, Angola, by William R. Branch, Ninda L. Baptista and Pedro Vaz Pinto on 7 November 2015.

Paratypes.

Males: PEM R24114-5, collected 1 km east of Farm Mucungo (-14.78361, 12.49694, 314 m a.s.l.), Namibe Province, Angola, by William R. Branch, Ninda L. Baptista and Pedro Vaz Pinto on 7 November 2015. Females: PEM R24116-7, collected 1 km east of Farm Mucungo (-14.78361, 12.49694, 314 m a.s.l.), Namibe Province, Angola, by William R. Branch, Ninda L. Baptista and Pedro Vaz Pinto on 7 November 2015.

Additional material examined.

Males: PEM R21596, collected 52 km north of Lubango-Namibe junction to Lucira (-14.65806, 12.52717, 586 m a.s.l.), Namibe Province, Angola, by William R. Branch on 8 December 2012; PEM R22489, collected from Praia do Meva (near Santa Maria) (-13.39667, 12.58972, 10 m a.s.l.), Benguela Province, Angola, by Pedro and Afonso Vaz Pinto on 28 December 2015; TM 40264-6, TM 40285, TM 40283, collected from Caraculo (-15.01667, 12.66667, 463 m a.s.l.), Namibe Province, Angola, by Wulf Haacke on 27 March 1971; TM 41137, TM 41141, TM 41144, collected from turn off to Morro do Chapéu Armado (-14.51185, 12.50190, 462 m a.s.l.), Namibe Province, Angola, by Wulf Haacke on 18 April 1971; NB 746, collected from Bimbe, Estação Zootecnica (-14.915538 13.249479, 2268 m a.s.l), Namibe Province, Angola, by Ninda L. Baptista and Pedro Vaz Pinto on 11 March 2018. Females: PEM R21595, collected 50 km east of Namibe on main tar road to Leba Pass (-15.01558, 12.55503, 516 m a.s.l.), Namibe Province, Angola, by William R. Branch on 8 December 2012; PEM R22488, collected from Praia do Meva (near Santa Maria) (-13.39667, 12.58972, 10 m a.s.l.), Benguela Province, Angola, by Pedro and Afonso Vaz Pinto on 28 December 2015; PEM R24203-4, collected 10.4 km south of Rio Mucungo on tar road to Bentiaba (-14.84194, 12.42778, 360 m a.s.l), Namibe Province, Angola, by William R. Branch and Pedro Vaz Pinto on 11 November 2016; PEM R24219, collected approx. 20 km south of Bentiaba (-14.40278, 12.44972, 426 m a.s.l.), Namibe Province, Angola, by William R. Branch and Pedro Vaz Pinto on 12 November 2016; TM 40263, TM 40267-9, TM 40280-2, TM 40284, TM 40286-7, TM 40289-90, collected from Caraculo (-15.01667, 12.66667, 463 m a.s.l.), Namibe Province, Angola, by Wulf Haacke on 27 March 1971; TM 41132-6, TM 41138-9, TM 41142, collected from turn-off to Morro do Chapéu Armado (-14.51185, 12.50190, 462 m a.s.l.), Namibe Province, Angola, by Wulf Haacke on 18 April 1971; TM 41211-4, collected from Lucira road, 5 km south of Catara River (-13.60430, 12.62890, 341 m a.s.l), Namibe Province, Angola, by Wulf Haacke on 19 April 1971; NB 744-5, collected from Bimbe, Estação Zootécnica (-14.915538 13.249479, 2268 m a.s.l), Namibe Province, Angola, by Ninda L. Baptista and Pedro Vaz Pinto on 11 March 2018. Juveniles: TM 41215-6, collected from Lucira road, 5 km south of Catara River (-13.60430, 12.62890, 341 m a.s.l), Namibe Province, Angola, by Wulf Haacke on 19 April 1971; NB 743, collected from Bimbe, Estação Zootecnica (-14.915538 13.249479, 2268 m a.s.l), Namibe Province, Angola, by Ninda L. Baptista and Pedro Vaz Pinto on 11 March 2018.

Additional referred material

(not examined). TM 24545, collected from Caraculo (-15.01667, 12.66667, 463 m a.s.l.), Namibe Province, Angola, by Charles Koch in September 1956; TM 40288, TM 40291-5, collected from Caraculo (-15.01667, 12.66667, 463 m a.s.l.), Namibe Province, Angola, by Wulf Haacke on 27 March 1971; TM 41140, TM 41143, collected from turn-off to Morro do Chapéu Armado (-14.51185, 12.50190, 462 m a.s.l.), Namibe Province, Angola, by Wulf Haacke on 18 April 1971; FKH 0248, collected from road to Praia do Furado (-14.78306, 12.41726, 403 m a.s.l.), Namibe Province, Angola, by Javier Lobón Rovira, Afonso and Pedro Vaz Pinto on 5 July 2019; P9-154, collected from Mariquita (-14.78355, 12.41783, 401 m a.s.l), Namibe Province, Angola, by Javier Lobón Rovira, Afonso and Pedro Vaz Pinto on 5 July 2019; NB 834-5, collected approx. 18 km E of Lucira (-13.90749, 12.69201, 332 m a.s.l.), Namibe Province, Angola, by Pedro Vaz Pinto, Ninda L. Baptista and Telmo António on 28 November 2017; CAS:HERP 264666-80, 26467, 264704, collected from Mocongo (= Mucungo) Farm (-14.7789, 12.48745, 309 m a.s.l.), Namibe Province, Angola, by Luis M.P. Ceriaco, Mariana Marques and Joyce Janota between 2-6 August 2018; NB 602, collected from Sta Maria (-13.49813, 12.60921, 332 m a.s.l.), Benguela Province, Angola, by Afonso and Pedro Vaz Pinto on 12 July 2017; FKH 0005-6, collected from Fazenda Carivo (-13.195467, 13.424319, 438 m a.s.l.), Benguela Province, Angola, by Pedro Vaz Pinto on 5 June 2018; CAS 263848-9, INBAC: AMB 10691, CAS 263878, collected from Tundavala (-14.82386, 13.38114, 1295 m a.s.l.), Huíla Province, Angola, by Mariana P. Marques, Luis M.P. Ceriaco, Suzana Bandeira, Matthew Heinicke, Brent Butler and Timóteo Júlio on 1 and 9 August 2017. All material referable to the new species based on geographical distribution and closeness to examined material.

Etymology.

This species is named in honour of father and son, Pedro and Afonso Vaz Pinto, two enthusiastic Angolan naturalists with whom WRB spent a great deal of time in the field, to recognise their contributions in collecting and studying Angolan herpetofauna. The name is constructed in the masculine plural genitive.

Diagnosis.

A member of the greater ' Afroedura transvaalica ' group as it possesses two pairs of enlarged scansors per digit and a strongly verticillate and flattened tail ( Jacobsen et al. 2014). Part of the A. bogerti -group which differs from other members of the ' Afroedura transvaalica ' group by having less than 86 mid-body scale rows (vs. 97-102 in A. gorongosa , 113-120 in A. loveridgei , 102-119 in A. transvaalica ); by the rostral bordering the nostril (nostril excluded from rostral in A. loveridgei ); by the anterior nasals always in contact (separated by 1-3 granules in A. gorongosa ; always in broad contact in A. loveridgei ; usually in broad contact in A. transvaalica ~ 3-18%); and in having 11-14 scales between the anterior borders of the eyes (19-22 in A. gorongosa ; 15-19 in A. loveridgei ; 15-20 in A. transvaalica ) (comparative data fide Branch et al. 2017a).

Afroedura vazpintorum sp. nov. differs from other members of the A. bogerti -group by a combination of the following characters (see Tables 3 , 4 ): 73-86 (mean 80.3) mid-body scale rows (69-77 [mean 73.5] in A. bogerti , 76-88 [mean 79.3] in A. wulfhaackei sp. nov., 64-78 [mean 72.8] in A. donveae sp. nov., 73-78 [mean 74.8] in A. praedicta sp. nov.); by the anterior nasals always in contact (similar to A. donveae and A. praedicta sp. nov.; ~ 33% of the time in contact in A. bogerti ; ~ 68% of the time in contact in A. wulfhaackei sp. nov.); in each verticil having 5-6 (mean 5.0) ventral and 6-7 (mean 6.1) dorsal rows of scales (5-6 [mean 5.5] and 6-7 [mean 6.6] in A. donveae sp. nov.; 4 and 5 in A. bogerti and A. praedicta sp. nov., 4-5 [mean 4.0] and 5-6 [mean 5.1] in A. wulfhaackei sp. nov.); ventral surfaces immaculate (similar to A. donveae sp. nov.; greyish with black spot in A. bogerti , A. wulfhaackei sp. nov. and A. praedicta sp. nov.). Afroedura vazpintorum sp. nov. differs from its sister lowland species A. donveae sp. nov. in being smaller (51.3 versus 57.6 mm average SVL), in having greater mid-body scale counts 73-86 (mean 80.3) versus 64-78 (mean 72.8), a lower number of precloacal pores (9-11 [mean 10.2] versus 11-12 [mean 11.5]), duller colouration and less distinct tail banding (versus bolder colouration and distinct tail banding).

Holotype description.

Adult male; SVL 50.6 mm; tail 44.5 mm (partly regenerated tail); with a small mid-ventral incision for the removal of liver sample. Measurements and meristic characters of holotype presented in Table 8 . Head and body dorsoventrally compressed; HL 11.1 mm, HW 7.4 mm, broadest at posterior level of eye and 1.5 times longer than wide. Eyes large (2.5 mm wide), pupil vertical with indented margins; circumorbital scales small and smooth, elongate at upper anterior margin, the most upper posterior scale with minute upward pointing spines. Snout rounded, 4.3 mm long, almost equal to distance between eye and ear openings (4.1 mm). Scales on top of snout slightly granular and elevated, rounded, mostly equal in size, with no intervening minute granules. Scales on snout slightly subequal in size to those on back of head or nape which, in turn, is irregular in size and mostly smooth. Scales on eyelids larger than those on the crown, 6 scales deep from circumorbital scales to crown. Circumorbital scales are separated by a row of smaller scales from the larger scales on eyelid. Nostril pierced between rostral, three nasal scales; 1st supralabial narrowly excluded from nostril; the supranasal being much larger than the subequal postnasals and being separated from each other by two smaller scales. Nostrils slightly elevated. Rostral roughly rectangular, but with its upper edges elongated due to extensions into the nostril. Nine supralabials on each side, the labial margin flexing upwards at the rictus (approx. mid-orbital position), with 2-3 minute scales proximal to the flexure. Eight infralabials on either side, with a small scale proximal to the flexure. At the lip, mental slightly narrower than adjacent infralabial, only three quarters the width of rostral and in contact with three distinctly elongated postmental scales. Scales on throat much smaller than those on belly; scales touching infralabials larger. Eleven scales across the crown at level of front of eye; 8 scales between nostril and front of eye; 14 scales from ear to eye; 73 scales around mid-body. Ear opening deep, oblique and roughly oval, backward pointing, much taller than wide (0.6 × 0.1 mm). Scales on dorsum smooth, non-overlapping, largest at mid-body, smaller on nape and tail base. Scales on ventrum flattened, not overlapping, more-or-less ovate at mid-ventrum, twice the size of lateral granules and 1.5 times those along backbone. Original tail slightly dorsoventrally flattened and distinctly verticillate (7 whorls in total), with obvious lateral constrictions; each verticil comprising 6 imbricate rows of scales dorsally and 5 imbricate scale rows ventrally and ventral scales approximately twice the size of those on the dorsal surface. Limbs well developed, hindlimbs slightly longer than forelimbs, both without obvious mite pockets at posterior margin of limb insertions, mite pockets present at anterior margin of hindlimbs. All digits with a large pair of distal scansors, separated by a large, curved claw and followed after a large gap (twice the length of terminal scansor) by a smaller pair of scansors; infero-median row of digital scales enlarged transversely, particularly towards the scansors, where the terminal scale adjoining the first pair of scansors may be medially constricted, swollen and scansor-like; 9 enlarged scale rows under 4th toe. Precloacal pores 12.

Paratypes and additional material variation

(see Table 8 for more measurements and scale counts of type series). SVL 43.9-58.8 mm; original tail length 42.3-67.0 mm, 0.94 times SVL; head length 1.07-1.61 times head width; snout 1.75 times diameter of eye. The supranasals are always in contact; the first upper labial and rostral always enter the nostril and the rostral at the lip margin is always wider than the mental; 2-3 postmental scales; supralabials 8-10, infralabials 8-9; scales between anterior edges of eyes 11-15; scales between nostril and anterior edge of orbit 7-11; scales between anterior edge of ear and rear margin of orbit 7-11; scales around mid-body 73-86; subdigital lamellae on 4th toe 6-10; dorsal scales per tail verticil 6-7(mean 6.1); ventral scales per tail verticil 5-6(mean 5.0); precloacal pores 9-12.

Colouration.

In life (paratype PEM R24118, Fig. 5E ). Light grey to yellow-olive above with six irregularly-spaced darker, brown to black, W-shaped crossbars from the occiput to the sacrum, central two broken up to form irregularly-shaped crossbars; each crossbar separated by yellow-olive to light grey blotches; head mostly light grey with scattered darker grey scales; dark brown band from nostril across the upper margins of the ear opening connecting with dark brown lateral bar on to the neck; a thin pale yellow canthal stripe extends on both sides from the nasal region to anterior margins of eye, continuing on to the nape; upper and lower labials light grey with diffuse dark brown edges; lateral sides of the body with a mix of dark grey and yellow blotches; limbs yellowish above with scattered darker grey markings; tail (original) with irregular dark brown bars separated by yellow bars; tail (regenerated) with black mottling on light grey background; iris dark brown with a black narrow elliptic pupil with crenulated edge and black reticulation; ventrum uniform beige with scattered brown specks on lateral edges of ventrum alone; ventrally, limbs with scattered brown spots. In preservative (holotype PEM R24118, Fig. 10 ): dorsum with three distinct, irregularly-spaced, dark brown W-shaped crossbars anteriorly, with beige intervening blotches, the posterior crossbars are broken up and irregularly spaced; dorsally, arms and legs darkly barred; tail (original part) with dark brown bars that are not well defined; tail (regenerated part) with dark brown mottling on beige background; dorsal head with mottled dark brown scales, light beige canthal stripe from nostril to anterior corner of eyes, continuing posterior to the eye to above the ear opening; dark brown band running from the nasals, above the ear opening, to the neck; light beige stripe (one-scale-wide) from above the supralabials to the ear opening (two-scales-wide); supralabials dark brown edged ventrally; infralabials scattered with dark brown markings dorsally; ventrum mostly immaculate with dark brown spots ventrally on arms and legs. Variation: Similar colouration and patterning to the holotype (preserved) and paratype (as above, in life). Dorsal crossbars are often fused to form an X-shape or are irregular. When present, crossbars number 5-6. Coastal material is often very light in colouration and not as boldly patterned or brightly coloured compared to highland material. Original tails with 6-7 broad, dark brown to black bars, separated by light beige to white bars, some specimens with a fine one-scale-wide black bar splitting the lighter bars. Regenerated tails with fine dark brown to black mottling. Juveniles with more sharply defined patterns.

Natural history and habitat

(Fig. 4E, F ). This species was frequently observed at night in coastal areas hunting in small branches, bushes and small trees growing amongst rock crevices between large granite boulders. When disturbed, it quickly jumps to the rock faces and finds shelter under granite flakes. Mainly rupiculous, it is strongly associated with large granite boulders present throughout the semi-arid ecosystems of the Angolan Kaokoveld. Usually found in rocky habitats with succulent plants and dominated by acacia savannah ( Senegalia spp.), preferring more wooded and less arid environments when compared to A. donveae sp. nov. Some specimens, referable to A. vazpintorum sp. nov., were found at high altitude near Bimbe, in crevices in pre-cambrian limestone formations, a rocky habitat framed by stunted Afromontane forest elements and surrounded by extensive montane grassland.

Distribution and conservation.

Known from various localities in the coastal lowlands of Namibe and Benguela Provinces of Angola, stretching for about 250 km along the coast and up to 60 km inland, which is probably a fair reflection of the species’ global range (Fig. 1 ). Occurs from near sea level to 500 m elevation. An apparently isolated population occurs on the Angolan highlands along the Humpata Plateau. This population occurs at a much higher elevation (above 2,000 m a.s.l.) and may warrant further phylogenetic analysis employing nuclear markers. The species seems common and widely distributed, mostly in relatively undisturbed habitat and, therefore, it is likely not threatened.

Figures

Figure 1. Digital elevation map showing the predicted distribution of the five newly-recognised species of Afroedura occurring in Angola (data for A. wulfhaackei sp. nov. also contain the morphologically-indistinguishable Afroedura sp. 5 - 7 clades). Confirmed locality records are indicated by solid circles, with type localities indicated by stars. Coloured polygons indicate potential distribution areas with more than 90 % of climatic habitat suitability.

Figure 4. Typical habitat of Angolan Afroedura. A. Namba area (A. bogerti); B. Mt Sandula (Afroedura sp. 5); C. Serra da Neve (A. praedicta sp. nov.); D. Omauha Lodge (A. donveae sp. nov.); E. Farm Mucungo (A. vazpintorum sp. nov.); F. Bimbe (A. vazpintorum sp. nov.). Photos: A. Javier Lobon-Rovira; B, D, E. William R. Branch; C. Pedro Vaz Pinto; F. Ninda L. Baptista.

Figure 5. Live photos of Angolan flat geckos: A. Afroedura bogerti (PEM R 24187); B. Afroedura wulfhaackei sp. nov. (paratype PEM R 24232); C. Afroedura praedicta sp. nov. (holotype NB 854); D. Afroedura donveae sp. nov. (holotype PEM R 17937); E. Afroedura vazpintorum sp. nov. (holotype PEM R 24118); F. Afroedura vazpintorum sp. nov. (NB 0745); Photos: A, B, D, E. William R. Branch; C, F. Pedro Vaz Pinto.

Figure 10. Holotype of Afroedura vazpintorum sp. nov. (PEM R 24118) from 1 km east of Farm Mucungo, Namibe Province, Angola. Photos: Werner Conradie.