According to Branch et al. (2021), historical material from near Bocoio in Benguela Province, Angola clustered morphologically with A. wulfhaackei. However, due to the occurrence at lower elevations and being isolated from other known populations of Afroedura it was suggested that the status of this population required further investigation (Branch et al. 2021). Newly-collected material allowed for its re-assessment within a wider phylogenetic framework, and it was determined that it represented a novel lineage, related to A. bogerti and not A. wulfhaackei, as initially hypothesised. It is therefore described below as a new species.

Afroedura bogerti – Branch et al. 2017: 162; Marques et al. 2018: 178 (in part).

Afroedura wulfhaackei – Branch et al. 2021: 66 (in part).

PEM R24743, adult female, collected at Morro do Pundo, about 25 km west of Bocoio (-12.44389, 13.92250; 946 m a.s.l.), Benguela Province, Angola by Pedro Vaz Pinto on 6 June 2018.

(six specimens). *TM 46587–8, TM 465890, adult females, collected 30 km W of Sousa Lara [= Bocoio] (approx. -12.40689, 13.90400; 670 m a.s.l.), Benguela Province, Angola by Wulf Haacke on 28 May 1974; *TM 46589, adult male, collected 30 km W of Sousa Lara [= Bocoio] (approx. -12.40689, 13.90400; 670 m a.s.l.), Benguela Province, Angola, by Wulf Haacke on 28 May 1974; FKH 0688, FKH 0689, adult females, collected from Alto Pundo – Bocoio (-12.44367, 13.92072, 920 m a.s.l.), Benguela Province, Angola by Pedro Vaz Pinto and Afonso Vaz Pinto on 2 September 2021. *Note the locality data presented as ‘3 km west of Bocoio, Benguela Province (12°28'58.0"S, 14°06'24.8"E)’ in Branch et al. (2017, 2021) is in error and we update it according to the original specimen labels and catalogue museum register.

The new species is named in reference to the area where it was found. The region lies on top of a ridge known as Morro do Pundo that translates to the ‘Hills’ or ‘Mountain’ of the Baboons. The name thus comprises two parts: pundo (= baboon) and montana (= mountain).

A member of the greater ‘transvaalica’ group, possessing two pairs of enlarged scansors per digit, and a strongly verticillate and flattened tail (Jacobsen et al. 2014). As part of the A. bogerti group it differs from other members of the ‘transvaalica’ group by having 78–82 midbody scale rows (versus 97–102 in A. gorongosa, 113–120 in A. loveridgei, 102–119 in A. transvaalica); and rostral excluded from the nostril (in contact in A. gorongosa) [Note: in Branch et al. (2021) it was incorrectly recorded that the rostral is in contact with the nostril in the A. bogerti-group]; with the supranasals always being 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 13–15 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. 2017, 2021).

Afroedura pundomontana sp. nov. differs from other members of the A. bogerti group by a combination of the following characteristics (see Tables 5–6): midbody scale rows 78–82 (mean 79.5) (71–72 [mean 71.5] in A. otjihipa sp. nov., 65–67 [mean 66.0] in A. donveae, 69–77 [mean 73.5] in A. bogerti, 73–78 [mean 74.8] in A. praedicta, 73–88 [mean 79.5] in A. wulfhaackei, 73–86 [mean 80.3] in A. vazpintorum); by the supranasals always being in contact (~33% of the time in A. bogerti; ~57% in A. wulfhaackei; always in contact in A. donveae, A. vazpintorum, A. praedicta and A. otjihipa sp. nov.); each tail verticil comprising 4–5 (mean 4.4) ventral and 5–6 (mean 5.6) dorsal rows of scales (mean 4 ventral and 5 dorsal in A. bogerti, A. praedicta and A. wulfhaackei; 5–6 [mean 5.5] ventral and 6–7 [mean 6.6] dorsal in A. donveae; 5–6 [mean 5.0] ventral and 6–7 [mean 6.1] dorsal in A. vazpintorum; 5 ventral and 6 dorsal in A. otjihipa sp. nov.); ventral surfaces greyish with scattered small black spots (similar to A. bogerti, A. praedicta and A. wulfhaackei, immaculate in A. donveae, A. vazpintorum and A. otjihipa sp. nov.). Afroedura pundomontana sp. nov. differs from its sister highland species A. bogerti in having higher numbers of midbody scale counts (78–82 [mean 79.5] versus 73–78 [mean 74.8]), supranasals always in contact (versus ~33% of the time), and the posterior scales of the dorsal W-shapes crossbars dark black (versus same colour as cross bands; Fig. 3); it differs from A. wulfhaackei in that the supranasals are always in contact (versus ~57%).

Figure 3. 

Live specimens of: A–B. Afroedura bogerti (A. P1-286, not vouchered; B. JLRZC0015, not vouchered) from Serra da Namba, Cuanza Sul Province, Angola; C–D. Afroedura pundomontana sp. nov. (FKH0689) from Morro do Pundo, Benguela Province, Angola. Photos: A, C, D. Pedro Vaz Pinto; B. Javier Lobón-Rovira.

Adult male; SVL 46.0 mm; tail 42.3 mm (detached full original tail). Small mid-ventral incision for removal of liver sample. Measurements and meristic characters of holotype are presented in Table 6. Head and body dorsoventrally compressed; HL 12.5 mm, HW 8.3 mm, broadest at posterior level of eye; head 1.51 times longer than wide. Eye large (2.6 mm wide), pupil vertical with indented margins; circumorbital scales small and smooth, elongate at upper anterior margin, upper three posterior scales with small upward pointing spines. Snout rounded, 4.9 mm long, longer than distance between eye and ear openings (3.8 mm). Scales on top of snout smooth, rounded; scales at the edge larger than central ones, with no intervening minute granules. Scales on snout slightly larger in size to those on the back of head or the nape. Scales on eyelids larger than those on the crown, six scales deep from circumorbital scale to crown. Circumorbital scales separated from the larger scales on the eyelids by two rows of smaller scales. Nostril pierced between first supralabial and three nasal scales; rostral excluded from nostril; 1^st supralabial narrowly excluded from nostril; supranasal much larger than the postnasals (which are about equal in size) and in broad contact. Nostrils slightly elevated. Rostral roughly rectangular but with the upper edges slightly elongated due to extensions to the supranasals. Eight supralabials on either side, the labial margin flexing upwards at the rictus (approx. midorbital position), with 3–4 minute scales proximal to the flexure. Nine infralabials on either side, with a small scale proximal to the flexure. At the lip, mental slightly narrower than adjacent infralabial; mental only two thirds the width of rostral (1.1 mm versus 1.8 mm respectively), and in contact with three rounded postmental scales. Scales on throat notably smaller than those on belly, but the scales touching the infralabials are larger. Fourteen scales across the crown at level of front of eyes; 18 scales from ear to eye; 83 scales around midbody. Ear opening deep, oblique and more-or-less round, nearly symmetrical (0.7 × 0.8 mm). Scales on dorsum smooth, closely set but juxtaposed, largest at mid-body, smaller on nape and tail base. Scales on venter flattened, not overlapping, more-or-less ovate at mid-ventrum, about twice the size of lateral granules and about 1.5 times larger than the scales along the backbone. Original tail slightly dorsoventrally flattened and distinctly verticillate (10 distinct verticils in total), with obvious lateral constrictions that become less distinct towards the tip of the tail; each verticil comprising 6 rows of imbricate scales dorsally and 4 rows of imbricate ventrally, with ventral scales approximately twice the size of those on the dorsal surface. Limbs well-developed, hindlimbs slightly longer than forelimbs, no notable mite pockets (dermal crevices inhabited by small ectoparasitic mites) at anterior or posterior margin of hind limbs. 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; seven enlarged subdigital lamellae on 4^th toe.

(see Table 6 for more measurements and scale counts of type series). SVL 43.4–57.8 mm; head length 1.19–1.50 times head width; snout 1.20–1.93 times the diameter of eye. Supranasals always in contact; the first upper labial and rostral always enter the nostril, and the width of the rostral at the lip margin is always wider than that of the mental; 2–3 postmental scales; supralabials 9, infralabials 9; scales between anterior edges of eyes 16–19; scales between nostril and anterior edge of orbit 10–13; scales between anterior edge of ear and rear margin of orbit 16–19; scales around mid-body 78–83; subdigital lamellae of 4^th toe 7–9; dorsal scale rows per tail verticil 5–6; ventral scale rows per tail verticil 4–5. Precloacal pores 12 (single male).

In life (holotype PEM R24743 [similar to Fig. 3C–D]): Greyish above with five evenly spaced darker crossbars from the occiput to the sacrum, each crossbar consisting of 9–12 dark scales forming a distinct W-shape, that consist anteriorly of a mix of grey and mustard scales and posteriorly by more prominent dark grey to black scales; each dark crossbar is separated by a mix of lighter grey scales; head with irregular dark grey blotches on the crown with intervening pale grey and mustard colouration; dark grey bar from nostril to the anterior margin of the ear opening; a vague, thin grey canthal stripe, extends on both sides from the nasal region to anterior margins of eye; upper and lower labials light grey anteriorly and beige posteriorly with fine black specks; lateral sides of the body with a mix of light grey and light cream-yellow; limbs light greyish above with scattered darker grey markings interspersed with cream-yellow; tail with eight dark brown to black crossbands, becoming increasingly more bold towards the tip; iris gold in colour with a narrow black elliptic pupil with crenulated edge, and black reticulation with light grey intervening blotches; venter uniform greyish with scattered black specks; ventral part of limbs with scattered black specks, more prominent than on the underparts. In preservative (Fig. 4): Dorsum with five evenly spaced dark brown W-shaped crossbars from the occiput to the sacrum with beige intervening blotches; ventrum is beige with numerous small scattered black specks on each scale, more prominent posteriorly; tail with eight dark brown to black cross bands. Paratype colouration variation: greyish above with five to six evenly or irregularly spaced darker grey-black W-shaped crossbars from the occiput to the sacrum, anterior part of these crossbars much darker than the posterior part, which is scattered with mustard coloured scales; lateral sides of body with a mixture of darker grey and mustard coloured scales; limbs and tail with grey blotches, with scattered mustard coloured scales; ventrum uniform greyish with scattered black specks.

Figure 4. 

Holotype of Afroedura pundomontana sp. nov. (PEM R24743) from Morro do Pundo, 25 km west of Bocoio, Benguela Province, Angola. Scale bar: 1 cm. Photos: Werner Conradie.

(Fig. 5A–B). A rupicolous species found in rugged landscape between 600 to 1,000 m a.s.l. No details are available regarding the conditions under which the historical material was collected, but the new material was collected during the day, underneath vertical flakes in large granite boulders. On both occasions, several individuals were sheltering under the same flake. They were found in rocky outcrops in anthropogenically disturbed mixed escarpment woodland, characteristic of the ecotone between the arid coastal plain and the inland mesic Angolan plateau. The presence of shrubs and small trees surrounding the granite outcrops suggests that these geckos might forage at night in the vegetation as reported for other Angolan species (Branch et al. 2021).

Figure 5. 

Habitat photos of: A–B. Afroedura pundomontana sp. nov.: Morro do Pundo, 25 km west of Bocoio, Benguela Province, Angola; C–D. Afroedura otjihipa sp. nov.: Otjihipa Middleberg, Kunene Region, Namibia. Photos: A–B. Pedro Vaz Pinto; C–D. Francois Becker.

This species is currently known only from central Benguela Province, Angola (Fig. 2). It was collected at three localities in close proximity to one another, on the first elevation step of the Angolan escarpment, inland from the town of Lobito. The species may be more widely distributed as our predicted mapping indicates but, so far, surveys conducted on the coastal plain and elsewhere along the escarpment did not produce additional material. Even around the type locality, the species proved to be uncommon and quite difficult to find, partly due to the inaccessible topography, but apparently also due to scarcity of granite flakes. Populations in isolated granite outcrops may be threatened by removal of rock flakes for construction of homes and other buildings. In accordance with IUCN Red List Guidelines (IUCN 2022) we propose this species to be classified as Data Deficient (DD) at this stage.

Afroedura cf. bogerti – Branch 1998: 232; Griffin 2002: 20, 2003:10; Herrman and Branch 2013: 5.

NMNW R11253, adult female, collected from Otjihipa Middleberg (-17.28314, 12.66506, 1,900 m a.s.l.), Kunene Region, Namibia, by Morgan Hauptfleisch, Francois Becker, Vera De Cauwer, Wessel Swanepoel and Ernst van Jaarsveld on 23 April 2021.

NMNW R11245, adult male (paired with female NMNW R11253 in same rock crack). Same collection details as holotype.

The new species is named in reference to the area it was collected, namely Otjihipa Mountains in northern Namibia.

A member of the greater ‘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 ‘transvaalica’ group by having less than 72 mid-body scale rows (vs. 97–102 in A. gorongosa, 113–120 in A. loveridgei, 102–119 in A. transvaalica); rostral excluded from the nostril (in contact in A. gorongosa); supranasals 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 15–16 scales between anterior borders of the eyes (19–22 in A. gorongosa, 15–19 in A. loveridgei, 15–20 in A. transvaalica) (comparative data from Branch et al. 2017, 2021).

Afroedura otjihipa sp. nov. differs from other members of the A. bogerti group by a combination of the following characteristics (see Tables 5 and 7): 65–67 (mean 66.0) mid-body scale rows (64–78 [mean 72.8] in A. donveae, 69–77 [mean 73.5] in A. bogerti, 73–78 [mean 74.8] in A. praedicta, 78–82 (mean 79.5) in A. pundomontana sp. nov.; 76–88 [mean 79.3] in A. wulfhaackei, 73–86 [mean 80.3] in A. vazpintorum); supranasals always in contact (similar to A. donveae, A. vazpintorum, A. praedicta and A. pundomontana sp. nov.; in contact in ~ 33% of A. bogerti; in contact in ~ 57% of A. wulfhaackei); each tail verticil comprises 5 ventral and 6 dorsal rows of scales (mean 4 ventral and 5 dorsal in A. bogerti, A. praedicta and A. wulfhaackei; 4–5 (mean 4.4) ventral and 5–6 (mean 5.6) dorsal in A. pundomontana sp. nov.; 5–6 [mean 5.5] ventral and 6–7 [mean 6.6] dorsal in A. donveae; 5–6 [mean 5.0] ventral and 6–7 [mean 6.1] dorsal A. vazpintorum); ventral surfaces light cream and almost immaculate, with some scattered dark spots near lateral edges (similar to A. donveae and A. vazpintorum; greyish with black spots in A. bogerti, A. wulfhaackei, A. praedicta and A. pundomontana sp. nov.); larger average adult size 58.2 mm SVL (versus 57.6 mm in A. donveae, 51.7 mm in A. wulfhaackei, 51.3 mm in A. vazpintorum; 50.3 mm in A. pundomontana sp. nov., 50.0 mm in A. bogerti, 49.9 mm A. praedicta), and by having very distinct black-and-white tail banding (similar to A. donveae). Afroedura otjihipa sp. nov. differs from its sister lowland species A. donveae in having a brown or copper coloured (versus black) iris, a relatively broader head (mean HL/HW 1.1 versus 1.3), and in dorsal colour pattern (Fig. 6): in A. otjihipa sp. nov. it is dominantly dark brown, the yellow appearing as small asymmetrical, irregular patches, and as irregular borders of four paired, asymmetrical, irregular, roughly triangular brown blotches, which merge at the scapular and sacral regions to form two additional bands (versus roughly symmetrical brown patterns on a mostly yellow background in A. donveae).

Figure 6. 

Live specimens of: A–B. Afroedura donveae from Omahua, Namibe Province, Angola (not sampled); C–D. Afroedura otjihipa sp. nov. (holotype female, NMNW R11253) from Otjihipa Middleberg, Kunene Region, Namibia. Photos: A–B. Javier Lobón-Rovira; C–D. Francois Becker.

Adult female: SVL 57.9 mm; tail regenerated, with a small mid-ventral incision for the removal of liver sample. Measurements and meristic characters of holotype presented in Table 7. Head and body dorsoventrally depressed; HL 13.6 mm, HW 13.2 mm, head broadest posterior level of eye and 1.02 times longer than wide. Eyes large (3.2 mm wide), pupil vertical with indented margins; circumorbital scales small and smooth, bottom posterior scales with small upward pointing spines. Snout rounded, 5.7 mm long, longer than distance between eye and ear openings (4.8 mm). Scales on top of snout smooth, rounded, similar in size, with no intervening minute granules. Scales on snout slightly larger than those on back of head or nape. Scales on eyelids larger than those on the crown, 5 scales deep from circumorbital scale to crown. Nostril pierced between first supralabial and three nasal scales; rostral narrowly excluded from nostril; supranasals much larger than the smaller postnasals, ventral postnasal being about half the size of its dorsal counterpart, and all in broad contact with one other. Nostrils very slightly elevated. Rostral roughly rectangular, but with its upper edges elongated due to extensions toward the nostril, and the central point extends between the nasals. Seven supralabials on each side, the labial margin flexing upwards at the rictus (approx. mid-orbital position), with 1–2 elongate scales proximal to the flexure and several minute scales along the flexure proximal to these. Seven infralabials on either side. At the lip, mental scale slightly narrower than adjacent infralabial, mental only two thirds the width of rostral (1.1 mm versus 1.8 mm respectively) and in contact with three postmental scales; mental similar in size and shape to the surrounding gular scales, the central one of which is distinctly smaller. Scales on throat much smaller than those on belly, scales touching infralabials larger. Fourteen scales across the crown at level of front of eyes; 10 scales between nostril and front of eye; 12 scales from ear to eye; 67 scales around mid-body. Ear opening deep, oblique and roughly oval, less than half as high as wide (0.42 × 0.95 mm respectively). Scales on dorsum smooth, non-overlapping, largest at mid-body, smaller on nape and tail base. Scales on ventrum flattened, not overlapping, roughly twice the size of lateral granules and 1.4 times the size of scales along the dorsal mid-line. Regenerated tail dorsoventrally flattened, roughly as broad as the neck, with ventral scales larger than those on the dorsal surface. Limbs well-developed, hindlimbs slightly longer than forelimbs; all limbs without obvious mite pockets at posterior or anterior margin of limb insertions. All digits with a large pair of distal scansors, separated by a curved claw, notably smaller on the fingers than toes, and followed after a gap (about the width of terminal scansor) by a smaller pair of scansors; infero-median row of digital scales slightly enlarged transversely, the distal two rows being paired in both digits and toes, where the terminal scale adjoining the first pair of scansors may be swollen and scansor-like; 6 enlarged central and two paired distal scale rows under 3^rd toe, while other toes have paired scale rows, 8 under the 4^th toe.

SVL 59.9 mm adult male, tail truncated, precloacal pores 12. Measurements and meristic characters of paratype are presented in Table 7. The paratype is very similar to the holotype with regard to scalation.

In life (holotype NMNW R11253, Fig. 7C–D): dark brown with yellowish patterns, fading to whitish on limbs and top of head; yellow patterns are irregular, asymmetrical patches and spots along the body, symmetrical paired spots around the nape and near the tail base; there is a thin, irregular, broken or continuous yellow bar on the nape; another broken, irregular yellow bar across the scapular region to the shoulders; three asymmetrical yellow double-bars which may present as pairs of medially-angled triangles posteriorly, across the back, each with an irregular dark brown core; another broken yellow bar or collection of symmetrical spots around the sacrum; head dark brown with yellow blotches on the crown with intervening pale yellow colouration; dark brown bar from nostril across the upper margins of the ear opening, connecting with dark brown lateral bar on the neck; a thin pale yellow canthal stripe extends on both sides from the nasal region to anterior margins of eye, continuing posteriorly from the eye onto the nape; skin above eyes copper blue with dark brown spots; upper and lower labials light grey with dense brown speckling, denser anteriorly and on supralabials; lateral sides of the body with a mix of dark brown and yellowish blotches, as a continuation of the dorsal patterns; limbs dark brown above with scattered light grey markings; tail (regenerated) with an asymmetrical chequered pattern of dark brown and light grey; iris copper in colour with a narrow black elliptic pupil with crenulated edge, and black reticulation; venter uniform beige with scattered brown specks mostly on lateral edges; ventrally, limbs with scattered brown spots, mostly near lateral surfaces. In preservative: yellowish patterns faded to light grey, dark brown to grey-brown, and eyes faded to bluish grey, with original colouration of pupils and iris no longer evident. Paratype colouration: Similar colouration and patterning as to the holotype, but the yellow bands and patterns are more clearly defined: the bar on the nape is nearly continuous, that on the scapular region has a clear dark brown core, and three pairs of asymmetrical, medially-pointing, dark brown, triangular blobs are clearly outlined by irregular yellow lines; no clear bar near the tail base, but a collection of symmetrical spots. The original tails are not present on the preserved specimen, but were observed briefly in life before capture. The original tails of another pair of individuals in a nearby rock crack (not caught) were also observed. Tail bars could not be counted, but bold black-and-white banding was clearly visible.

Figure 7. 

Holotype of Afroedura otjihipa sp. nov. (NMNW R11253) from Otjihipa Middleberg, Kunene Region, Namibia. Scale bar: 1 cm. Photos: Francois Becker.

A rupicolous species living in narrow rock crevices in relatively small sandstone outcrops in arid woodland savannah (Fig. 5C–D), at elevations of 1,800–1,900 m a.s.l. in the Otjihipa Mountains. It was not found in the dolomite formations near the type locality, despite greater search time dedicated to those areas. The rock cracks where they were found were smaller than is typical for this group and were similar throughout this surface formation. Congeners in the A. bogerti group are normally found only in deep rock cracks in and amongst large boulders. Such habitat features were present in the surveyed area, but only in dolomite formations. The much less crevice-rich sandstone formation, with thin, straight cracks formed between the sandstone strata, appeared to be favoured syntopically by A. otjihipa sp. nov. and Cordylus namakuiyus.

Currently known from a single sandstone ridge on Otjihipa Middleberg in the extreme north-west of the Kunene Region, Namibia (Fig. 2). The species remains poorly known, but it is probably stable in numbers as the local habitat is currently not threatened and is topographically unsuitable for human habitation. It likely occurs more broadly across the Otjihipa Mountain range. In accordance with IUCN Red List Guidelines (IUCN 2022) we propose this species to be classified as Data Deficient (DD) at this stage, but due to the remoteness of the locality and because no notifiable threats exist, it could be listed as Least Concern.