The Namba Flat Gecko, Afroedura bogerti was described using a single male specimen collected by Harry and Alan Chapman in 1925 during the Vernay Expedition to Angola (Loveridge 1944; Branch et al. 2017b). According to Loveridge (1944), the specimen was collected at ‘Namba (Mombolo)’, even though Bogert (1940), in his list of Angolan localities, mentioned only ‘Mombolo’ and gave rough co-ordinates for it. The collectors were the sons of a famous South African farmer, William Chapman, who had hosted the scientific members of the Vernay Expedition on his properties. Within the Mombolo Region, William Chapman and his family had built or lived in three farms, Sandula (which he named Monte Victoria-Verdun and which later became Monte Verde), Chipepe (= Quipepe) and Namba (where a religious mission was subsequently established – Missão da Namba). Branch et al. (2017b) provided a detailed systematic review of A. bogerti in Angola and, noting the uncertainties regarding the exact localities of Chapman’s material ascribed to Namba/Mombolo, restricted the type locality of the nominal form of A. bogerti to a polygon encompassing his three farms and the village of Maka-Mombolo. However, this created a taxonomical problem, as the specimens from Chapman’s farms of Sandula and Namba represent two genetically-distinct clades. To resolve this issue, we re-instated the original type locality reference of ‘Namba’ and here restrict the type locality to the northern areas of Mombolo in the vicinity of Missão da Namba and, by default, the material from William Chapman’s farm of Sandula represents an undescribed species.

Afroedura karroica bogerti – Loveridge 1944:1; Afroedura bogerti – Onderstall 1984:506, Jacobsen et al. 2014:467 (part), Branch et al. 2017b:157 (part), Marques et al. 2018: 177 (part), Branch et al. 2019a: 287 (part); Afroedura bogerti (clade 3) – Branch et al. 2017a:146–147.

AMNH 47841, adult male, collected from Namba (Mombolo) (approx. -11.91417, 14.82083, 1827 m a.s.l.), Cuanza-Sul Province, Angola, by Harry and Allan (= Alan) Chapman, between September and November 1925 during the Vernay Expedition to Angola.

Females: PEM R24185*, collected from granite bedrock and outcrops 200 m south of old farm paddock, Farm Namba (-11.91417, 14.82083, 1827 m a.s.l.), Cuanza-Sul Province, Angola, by William R. Branch, Pedro Vaz Pinto and Ninda L. Baptista on 3 November 2016; PEM R24187*, collected from rocks 400 m north of Missão da Namba grounds (-11.91528, 14.84556, 1786 m a.s.l), Cuanza-Sul Province, Angola, by William R. Branch, Pedro Vaz Pinto and Ninda L. Baptista on 4 November 2016; TM 46631–34, collected from Numba (= Namba) (approx. -11.91722, 14.84417, 1808 m a.s.l.), Cuanza-Sul Province, Angola, by Wulf Haacke on 29 June 1974. Juveniles: PEM R24184*, PEM R24186*, collected from granite bedrock and outcrops 200 m south of old farm paddock, Farm Namba (-11.91417, 14.82083, 1827 m a.s.l.), Cuanza-Sul Province, Angola, by William R. Branch, Pedro Vaz Pinto and Ninda L. Baptista on 3 November 2016. *genetically confirmed.

JLRZC0015–6, collected from Namba (-11.88132, 14.76218, 1752 m a.s.l), Cuanza-Sul Province, Angola, by Pedro Vaz Pinto and Javier Lobón-Rovira on 11 February 2020.

(description based on the adult material listed above). Head and body dorsoventrally compressed; SVL 46.4–53.5 (mean 50.0) mm, HL 10.9–12.6 (mean 11.6) mm, HW 7.8–11.2 (mean 10.6) mm, broadest at posterior level of eye and 1.1–1.4 (mean 1.3) times longer than wide. Eyes large (2.1–4.3 [mean 2.8] mm wide), pupil vertical with indented margins; circumorbital scales small and smooth, elongate at upper anterior margins, upper posterior scales with small upward pointing spines. Snout rounded, 4.5–5.0 (mean 4.3) mm long, slightly longer than distance between eye and ear openings (3.6–4.3 [mean 3.6] mm). Scales on top of snout smooth, rounded, scales to the side larger than central ones, with no intervening minute granules. Scales on snout slightly subequal in size to those on back of head or nape. Scales on eyelids larger than those on the crown. Circumorbital scales are separated by a row of smaller scales from the large scales on eyelid. Nostril pierced between rostral, two to three nasal scales and the 1^st supralabial; the supranasal being much larger than the subequal postnasals and being separated from each other by one to two smaller scales. Nostrils slightly elevated. Rostral roughly rectangular, but with its upper edges elongated due to extensions into the nostril. Supralabials 8–10 (mean 8.4). Infralabials 8–9 (mean 8.3). A total of 11–14 (mean 12.4) scales across the crown at level of front of eyes. A total of 14–16 (mean 15.4) scales from front of ear to back of eye. Mid-body scale rows 69–77 (mean 73.5). Original tail slightly dorsoventrally flattened and distinctly verticillate, with obvious lateral constrictions that are not that distinct to the tip of tail; each verticil comprising 5 imbricate rows of scales dorsally and 4 imbricate scale rows ventrally and with ventral scales approximately twice the size of those on the dorsal surface. Limbs well developed, hindlimbs slightly longer than forelimbs. All digits with a large pair of distal scansors, separated by a large, curved claw and followed after a large gap 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, enlarged subdigital lamellae on 4^th toe 6–9 (mean 6.9). Precloacal pores 8.

In life (based on PEM R24187, Fig. 5A). Greyish above with six irregularly spaced darker grey crossbars from the occiput to the sacrum, central crossbars fused to form an X-shape; head with a dark grey band across the posterior edge of crown encompassing a small central pale spot; dark grey to black bar from nostril to the anterior margins of the ear opening; a vague, thin pale canthal stripe, extends on both sides from the nasal region to anterior margins of eye, continuing posteriorly of the eye to above the ear opening; upper and lower labials grey with diffuse yellow edges; lateral sides of the body with very faintly yellow-mustard scales scattered amongst more prominent darker grey scales; limbs greyish above with scattered darker grey markings with intervening faint yellow-mustard colouration; tail (regenerated) with irregular grey mottling; iris golden with a black narrow elliptic pupil with crenulated edge and black reticulation; ventre uniform greyish with scattered black specks; ventral limbs with scattered black specks, more prominent than on the ventrum. In preservative (based on PEM R24187): dorsum with six irregularly-spaced darker grey crossbars from the occiput to the sacrum with beige intervening blotches, central crossbars fused to from dark grey X-shape; ventre beige with numerous small scattered black specks on each scale, more prominent anteriorly, laterally and posteriorly. Variation: Greyish to brownish above with five to six irregularly-spaced darker grey-brown W-shaped crossbars from the occiput to the sacrum, sometimes fused in the middle to form X-shapes, limbs and tail with grey blotches; ventre uniform greyish with scattered black specks. Juveniles with more sharply-defined pattern. The original description of the holotype (Fig. 6) was based on a preserved and bleached specimen: ‘Above, greyish; back with five or six obsolescent, irregularly W-shaped brown crossbars; limbs and tail immaculate. Below, whitish, uniform’ (Loveridge 1944). Examination of images of the holotype confirm the presence of fine brown coloured specks ventrally that indicates that the ventre was not immaculate as originally described.

(Fig. 4A). An exclusively rupicolous species sheltering in crevices and under flakes of exfoliating rock amongst large granite boulders during the day. Individuals were observed at night foraging in vertical rock faces of large granite boulders. The surrounding habitat was mostly montane grasslands and some cultivated fields, but also included stunted altitude miombo and some Afromontane forest elements in steeper gorges and between boulders. It should be noted that the southern face of Namba Mountain contains the most extensive and well-preserved Afromontane forest patches in Angola, recognised as an important biodiversity reservoir and regional hotspot (Mills et al. 2013).

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 Lobón-Rovira; B, D, E. William R. Branch; C. Pedro Vaz Pinto; F. Ninda L. Baptista.

This species complex was previously considered to be widespread in granite boulders throughout south-western Angola (Branch et al. 2017b; Marques et al. 2018). Here, we show that it is restricted to the Namba Region in the southern parts of Cuanza-Sul Province, Angola (Fig. 1). Namba consists of a 25 km west-east orientated mountain range that climbs to an elevation of 2,420 m a.s.l. and towers southwards above the highland plateau of Mombolo. Our new series was collected at elevations of 1,750–1,850 m a.s.l., at the base of the south-facing slopes of the Namba Mountain chain, but the species probably occurs in suitable habitat higher up and possibly well above 2,000 m a.s.l. (Fig. 1). Considering the deep genetic divergence between this species and Afroedura present at the southern edge of Mombolo and the recognition of Namba as a biodiversity hotspot, it seems plausible for this species to have evolved in isolation on this mountain ‘island’. Nevertheless, A. bogerti may also extend its distribution further north into other poorly-studied mountain ranges in Cuanza-Sul Province (Fig. 1). The species seems to be relatively common in the Namba Mountains, but its biology remains poorly known.

Figure 5. 

Live photos of Angolan flat geckos: A. Afroedura bogerti (PEM R24187); B. Afroedura wulfhaackei sp. nov. (paratype PEM R24232); C. Afroedura praedicta sp. nov. (holotype NB 854); D. Afroedura donveae sp. nov. (holotype PEM R17937); E. Afroedura vazpintorum sp. nov. (holotype PEM R24118); F. Afroedura vazpintorum sp. nov. (NB 0745); Photos: A, B, D, E. William R. Branch; C, F. Pedro Vaz Pinto.

Figure 6. 

Holotype of Afroedura bogerti (AMNH 47841) from Namba (Mombolo), Cuanza-Sul Province, Angola. Photos: Luis M.P. Ceríaco.

The phylogenetic analysis identified four well-defined clades (Afroedura sp. 4–7) within the southern inland/highland major clade (see Results). Our data show that this southern major clade clearly represents at least one undescribed species of Afroedura. Despite comparatively-low genetic distances between the four clades recovered (Table 2), they were all retrieved separately in our phylogenetic analyses and, as our morphological examination failed to separate the four clades, we cannot assign them as separate species. Therefore, we decided to take a conservative approach and to describe only one of the clades as a new species for now and tentatively assign Afroedura sp. 5–7 to this species pending further work. Thus, we grouped the material according to these clades and selected the clade called here Afroedura sp. 4 (Fig. 2) as the name-bearing clade for our new species. Material from lower-altitude localities near Bocoio in Benguela Province groups morphologically with our new species, but needs to be tested in a phylogenetic framework.

Afroedura bogerti – Branch et al. 2017b:157 (part); Marques et al. 2018: 177 (part); Branch et al. 2019a: 287 (part); Afroedura bogerti (clade 4) – Branch et al. 2017a:147.

PEM R24234, adult male, collected 3 km north of Maka-Mombolo (-12.17056, 14.88167, 1756 m a.s.l.), Benguela Province, Angola, by William R. Branch and Pedro Vaz Pinto on 15 November 2016.

PEM R24232–3, adult females, collected 3 km north of Maka-Mombolo (-12.17056, 14.88167, 1756 m a.s.l), Benguela Province, Angola, by William R. Branch and Pedro Vaz Pinto on 15 November 2016; PEM R24236, adult male, collected above Maka-Mombolo, north-east of Balombo (-12.19833, 14.86833, 1857 m a.s.l.), Benguela Province, Angola, by William R. Branch and Pedro Vaz Pinto on 15 November 2016.

(not examined). NB 817–9, collected at Morro do Moco, camp near Canjonde (-12.42611, 15.14778, 1931 m a.s.l), Huambo Province, Angola, by Pedro Vaz Pinto on 13 November 2017 (genetic samples included in this study).

Afroedura sp. 5 (Males): PEM R24192–4, collected at William Chapman’s Farm Victoria-Verdun (Sandula), (-12.17194, 15.02667, 1834 m a.s.l.), Cuanza-Sul Province, Angola, by William R. Branch, Ninda L. Baptista and Pedro Vaz Pinto on 6 November 2016. Afroedura sp. 5 (Females): PEM R24190–1, PEM R24195–6, PEM R24199, collected at William Chapman’s Farm Victoria-Verdun (Sandula), (-12.17194, 15.02667, 1834 m a.s.l.), Cuanza-Sul Province, Angola, by William R. Branch, Ninda L. Baptista and Pedro Vaz Pinto on 6 November 2016. Afroedura sp. 5 (Juveniles): PEM R24197–8, collected at William Chapman’s Farm Victoria-Verdun (Sandula), (-12.17194, 15.02667, 1834 m a.s.l.), Cuanza-Sul Province, Angola, by William R. Branch, Ninda L. Baptista and Pedro Vaz Pinto on 6 November 2016. Afroedura sp. 6 (Мales): PEM R24201, collected near Rio Chicanda, 5 km southwest of Lepi (-12.90861, 15.36472, 1534 m a.s.l.), Huambo Province, Angola, by William R. Branch, Ninda L. Baptista and Pedro Vaz Pinto on 8 November 2016. Afroedura sp. 7 (Мales): TM 45382, TM 45387–8, TM 45392, TM 45396, collected at Candumbo Rocks, 16 km west of Vila Nova = Huambo (-12.73614, 15.97442, 1760 m a.s.l), Huambo Province, Angola, by Wulf Haacke on 11 May 1971. Afroedura sp. 7 (Females): PEM R22490–1, collected 1 km west of Candumbo on road to Boas Águas (-12.73614, 15.97442, 1760 m a.s.l), Huambo Province, Angola, by Luke Verburgt on 11 March 2016; TM 45383, TM 45390, TM 45393–4, TM 45397, collected at Candumbo Rocks, 16 km west of Vila Nova = Huambo (-12.73614, 15.97442, 1760 m a.s.l), Huambo Province, Angola, by Wulf Haacke on 11 May 1971. Afroedura sp. 7 (Juveniles): PEM R24200, collected at Candumbo Rocks Memorial, 20 km east Humana to Cuito (-12.73722, 15.97333, 1750 m a.s.l), Huambo Province, Angola, by William R. Branch, Ninda L. Baptista and Pedro Vaz Pinto on 7 November 2016; TM 45386, TM 45389, TM 45391, TM 45395, collected at Candumbo Rocks, 16 km west of Vila Nova = Huambo (-12.73614, 15.97442, 1760 m a.s.l), Huambo Province, Angola, by Wulf Haacke on 11 May 1971. Unassigned clades> (Females): TM 45374, collected 1 km south of Luimbale (-12.25367, 15.31694, 1591 m a.s.l.), Huambo Province, Angola, collected by Wulf Haacke on 10 May 1971; TM 45367–8, collected 10 km west of Soque (-12.34590, 15.01180, 1974 m a.s.l), Benguela Province, Angola, collected by Wulf Haacke on 10 May 1971; PEM R24743, collected at Morro do Pundo (-12.44389, 13.92250, 939 m a.s.l.), Benguela Province, Angola, by Pedro Vaz Pinto on 6 June 2018; TM 46587–8, TM 465890, collected 3 km west of Bocoio (-12.46605, 14.10694, 926 m a.s.l.), Benguela Province, Angola by Wulf Haacke on 25 May 1971. Unassigned clades (Males): TM 45366 collected 10 km west of Soque (-12.34590, 15.01180, 1974 m a.s.l), Benguela Province, Angola, by Wulf Haacke on 10 May 1971, TM 46589, adult male, collected 3 km west of Bocoio (-12.46605, 14.10694, 926 m a.s.l.), Benguela Province, Angola, by Wulf Haacke on 25 May 1971.

TM 45381, 45384–5, TM 45398, collected at Candumbo Rocks, about 25 km east of the city of Huambo (-12.73614, 15.97442, 1760 m a.s.l), Huambo Province, Angola, by Wulf Haacke on 11 May 1971 (placed in Afroedura sp. 7, based on same geographical area as genetically-assigned material); FKH 0239, collected at Monte Verde-Sandula (-12.17924, 15.03086, 2055 m a.s.l.), Cuanza-Sul Province, Angola, by Pedro Vaz Pinto on 29 May 2019 (genetic sample included in this study placed it in Afroedura sp. 5).

The new species is named in honour of Wulf Haacke, retired curator of the herpetology collection at the former Transvaal Museum (now Ditsong National Museum of Natural History). His herpetological expeditions to Angola in the early 1970s paved the way for this study and much of the material used in this study resulted from his expeditions. The name is constructed in the masculine singular genitive.

A member of the greater ‘transvaalica’ group in possessing two pairs of enlarged scansors per digit and a strongly verticillate and flattened tail (Jacobsen et al. 2014). It is part of the A. bogerti-group which differs from other members of the ‘transvaalica’ group by having less than 88 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 being mostly in contact ~ 68% (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 11–16 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 wulfhaackei sp. nov. differs from other members of the A. bogerti-group by a combination of the following characters (see Tables 3, 4): 76–88 (mean 79.3) mid-body scale rows (69–77 [mean 73.5] in A. bogerti, 64–78 [mean 72.8] in A. donveae sp. nov., 73–86 [mean 80.3] in A. vazpintorum sp. nov., 73–78 [mean 74.8] in A. praedicta sp. nov.); by the anterior nasals being mostly (~ 68% of the time) in contact (~ 33% of the time in contact in A. bogerti; always in contact in A. donveae sp. nov., A. vazpintorum sp. nov. and A. praedicta sp. nov.); each verticil comprising 4–5 (mean 4.0) ventral and 5–6 (mean 5.1) dorsal rows of scales (4 and 5 in A. bogerti and A. praedicta sp. nov.; 5–6 [mean 5.5] and 6–7 [mean 6.6] in A. donveae sp. nov.; 5–6 [mean 5.0] and 6–7 [mean 6.1] A. vazpintorum sp. nov.); ventral surfaces greyish with scattered small black spots (similar to A. bogerti and A. praedicta sp. nov., immaculate in A. donveae sp. nov. and A. vazpintorum sp. nov.). Afroedura wulfhaackei sp. nov. differs more specifically from its sister highland species A. bogerti in having a higher number of mid-body scale counts (76–88 [mean 79.3] versus 69–77 [mean 73.5]) and differs from A. praedicta sp. nov. in that the nasals are separated by smaller granules (versus always in contact).

Adult male; SVL 51.4 mm; tail 47.1 mm (full original tail), with a small mid-ventral incision for the removal of liver sample. Measurements and meristic characters of holotype presented in Table 5. Head and body dorsoventrally compressed; HL 11.9 mm, HW 9.0 mm, broadest at posterior level of eye and 1.32 times longer than wide. Eye large (2.3 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, slightly longer than distance between eye and ear openings (4.0 mm). Scales on top of snout smooth, rounded, scales to the edge larger than central ones, with no intervening minute granules. Scales on snout slightly subequal in size to those on the back of head or the nape. Scales on eyelids larger than those on the crown, five scales deep from circumorbital scale to crown. Circumorbital scales are separated by a row of smaller scales from the larger scales on eyelids. Nostril pierced between rostral and three nasal scales; 1^st supralabial narrowly excluded from nostril; the supranasal being much larger than the subequal postnasals and 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. Eight supralabials on each side, the labial margin flexing upwards at the rictus (approx. mid-orbital position), with 3–4 minute scales proximal to the flexure. Ten 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 two distinctly elongate postmental scales. Scales on throat much smaller than those on belly; scales touching infralabials larger. Fifteen scales across the crown at level of front of eyes; 16 scales from ear to eye; 83 scales around mid-body. Ear opening deep, oblique and roughly oval, only half as high as wide (0.3 × 0.5 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 the backbone. Original tail slightly dorsoventrally flattened and distinctly verticillate (11 verticils in total), with obvious lateral constrictions that are not that distinct to tip of tail; each verticil comprising 5 imbricate rows of scales dorsally and 4 imbricate scale rows ventrally and with ventral scales approximately twice the size of those on the dorsal surface. Limbs well developed, hindlimbs slightly longer than forelimbs, mite pockets (dermal crevices inhabited by small ectoparasitic mites) only present at anterior margin of hindlimbs. All digits with a large pair of distal scansors, separated by a large, curved claw and followed by 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; enlarged subdigital lamellae on 4^th toe 9. Precloacal pores 9.

(see Table 5 for more measurements and scale counts of type series). SVL varied from 49.1–55.7 mm; head length 1.34–1.45 times the head width; snout 1.99 times the diameter of eye. Supranasals always separated by smaller granules, usually with a single large granule in contact with the rostral between the supranasals, followed by 1–2 smaller granules in lateral contact; the first upper labial and rostral always enters 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 7–9, infralabials 8–9; scales between anterior edges of eyes 13–14; scales between nostril and anterior edge of orbit 7–9; scales between anterior edge of ear and rear margin of orbit 14–17; scales around mid-body 76–82; subdigital lamellae on 4^th toe 7–8; dorsal scales per tail verticil 5; ventral scales per tail verticil 4. Precloacal pores 9.

SVL 36.4–59.6 mm; original tail length 36.4–57.8 mm, 0.94 times SVL; head length 1.19–1.75 times head width; snout 1.91 times diameter of eye. The supranasals in contact in 22 specimens and separated by granules in eight specimens, usually with a single large granule in contact with the rostral between the supranasals, followed by 1–2 smaller granules in lateral contact; the first upper labial and rostral always enters the nostril and the width of the rostral at the lip margin is always wider than that of the mental; 2–4 postmental scales; supralabials 8–9; infralabials 8–9; scales between anterior edge of eye 12–16; scales between nostril and anterior edge of orbit 7–11; scales between anterior edge of ear and rear margin of orbit 15–18; scales around mid-body 75–88; subdigital lamellae on 4^th toe 7–9; dorsal scales per tail verticil 5 (TM 45366, TM 46588 and PEM R24743 with 6); ventral scales per tail verticil 4 (TM 46588 with 5). Precloacal pores 9–12.

In life (paratype PEM R24232, Fig. 5B): Greyish above with five irregularly-spaced darker crossbars from the occiput to the sacrum, each crossbar consisting posteriorly out of three to four black scales wide forming a W-shape; anterior to W-shape are 8–10 scales deep with a mix of dark grey and mustard colours; each dark crossbar separated by light grey to beige blotches; head with irregular dark grey-mustard blotches on the crown with intervening light grey colouration; dark mustard to dark grey bar from nostril to the anterior margins of the ear opening; a vague, thin pale grey canthal stripe, extends on both sides from the nasal region to anterior margins of eye; upper and lower labials grey with diffuse mustard edges; lateral sides of the body with a mix of dark grey and yellow-mustard colouration; limbs greyish above with scattered darker grey markings with intervening yellow-mustard colouration; tail (regenerated) with irregular grey-mustard mottling; iris golden with a black narrow elliptic pupil with crenulated edge and black reticulation with light grey intervening blotches; ventrum uniform greyish with scattered black specks; ventral limbs with scattered black specks, more prominent than on the ventrum. In preservative (holotype PEM R24234, Fig. 7): Dorsum with five irregularly-spaced dark grey 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. Variation: Greyish to brownish above with five to six irregularly-spaced darker grey-brown W-shaped crossbars from the occiput to the sacrum, limbs and tail with grey blotches; ventrum uniform greyish with scattered black specks. Juveniles have sharper patterns and colours.

Figure 7. 

Holotype of Afroedura wulfhaackei sp. nov. (PEM R24234) from Maka-Mombolo, Benguela Province, Angola. Photos: Werner Conradie.

(Fig. 4B). An exclusively rupicolous species living in crevices between rocks or under flakes of exfoliating rocks in boulders at elevations of 920–2,055 m a.s.l. Specimens were collected in cracks of relatively small- to medium-sized boulders of carbonatitic origin surrounded by montane grassland. Some individuals were also found under flakes in large granite boulders and often in steep vertical rock faces and overhangs.

This species is currently known from southern Cuanza-Sul, central Huambo and northern Benguela Provinces, Angola (Fig. 1). It appears to have a relatively large, but patchy, distribution on the Angolan highlands and may extend its range into neighbouring provinces. Although sometimes locally common, it appears to be absent from vast areas in-between, where the species would be expected to occur. Populations in isolated granite outcrops may be threatened by removal of rock flakes for construction of homes and other buildings.

NB 854, adult male, collected from Serra da Neve (-13.77354, 13.24825, 1944 m a.s.l.), Namibe Province, Angola, by Pedro Vaz Pinto, Ninda L. Baptista and Telmo António on 30 November 2017.

ZMB 91607 (NB 853), NB 855, adult males, collected from Serra da Neve (-13.77354, 13.24825, 1944 m a.s.l.), Namibe Province, Angola, by Pedro Vaz Pinto, Ninda L. Baptista and Telmo António on 30 November 2017. ZMB 91608 (NB 1053), NB 1054, adult females, collected from Serra da Neve (-13.77354, 13.24825, 1944 m a.s.l.), Namibe Province, Angola, by Pedro Vaz Pinto, Ninda L. Baptista and Telmo António on 30 November 2017. NB 1055, juvenile, collected from Serra da Neve (-13.77354, 13.24825, 1944 m a.s.l.), Namibe Province, Angola, by Pedro Vaz Pinto, Ninda L. Baptista and Telmo António on 30 November 2017.

The specific epithet reflects the earlier prediction by WRB of the potential existence of an isolated population of Afroedura at Serra da Neve. We use the specific epithet “praedicta”, the Latin participle meaning predicted or anticipated, formed in the feminine genitive to match the gender of Afroedura.

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 78 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 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 12–15 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. 2017a).

Afroedura praedicta sp. nov. differs from other members of the A. bogerti-group by a combination of the following characters (see Tables 3 and 4): 73–78 (mean 74.4 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., 73–86 [mean 80.3] in A. vazpintorum sp. nov.); by the anterior nasals always being in contact (similar to A. donveae sp. nov. and A. vazpintorum sp. nov.; ~ 33% of the time in contact in A. bogerti; ~ 68% of the time in contact in A. wulfhaackei sp. nov.); each verticil having 4 ventral and 5 dorsal rows of scales (similar to the 4 and 5 in A. bogerti, 4–5 [mean 4.0] and 5–6 [mean 5.1] in A. wulfhaackei sp. nov.; but lower than the 5–6 (mean 5.5) ventral and 6–7 (mean 6.6) in A. donveae sp. nov. and 5–6 (mean 5.0) and 6–7 (mean 6.1) in A. vazpintorum sp. nov.); ventral surfaces grey with black specks on scales (similar to A. bogerti and A. wulfhaackei sp. nov.; immaculate in A. donveae sp. nov. and A. vazpintorum sp. nov.). Afroedura praedicta sp. nov. also differs from its sister highland species A. bogerti sp. nov. and A. wulfhaackei sp. nov. in that the nasals are always in direct contact (versus mostly separated).

Adult male; SVL 51.6 mm; tail 37.0 mm (regenerated tail, except for the first verticil), with a small mid-ventral horizontal incision for the removal of liver sample. Measurements and meristic characters of holotype presented in Table 6. Head and body dorsoventrally compressed; HL 12.9 mm, HW 7.9 mm, broadest at posterior level of eye and 1.6 times longer than wide. Eye large (2.6 mm wide), pupil vertical with indented margins; circumorbital scales small and smooth, elongated at upper anterior margin, the most upper posterior scale with very small upward pointing spines. Snout rounded, 5.0 mm long, slightly larger than the distance between eye and ear openings (4.2 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, six 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; 1^st supralabial in contact with nostril; the supranasal being much larger than the subequal postnasals and are 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. Eight 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. Nine 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 elongate postmental scales. Scales on throat much smaller than those on belly; scales touching infralabials larger. Twelve scales across the crown at level of front of eye; 9 scales between nostril and front of eye; 16 scales from ear to eye; 74 scales around mid-body. Ear opening deep, oblique and roughly round, backward pointing, nearly equal as long as wide (0.7 × 0.6 mm). Scales on dorsum smooth, non-overlapping, largest at mid-body, smaller on nape and tail base. Sales on ventre flattened, not overlapping, more-or-less ovate at mid-ventrum and twice the size of lateral granules and 1.5 times those along backbone. Tail regenerated, except for the first verticil, with obvious lateral constriction before regenerated tail start; first verticil comprising 5 imbricate rows of scales dorsally and 4 imbricate scale rows ventrally and with 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 4^th toe. Precloacal pores 8.

(see Table 6 for more measurements and scale counts of type series). SVL varied from 46.0–51.6 mm; head length 1.09–1.46 times head width; snout 1.71 times diameter of eye. The supranasals always in contact; the first supralabial enters the nostril (except for ZMB 91607 and NB 855) and rostral always enters 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 8–10, infralabials 8–9; scales between anterior edge of eye 13–15; scales between nostril and anterior edge of orbit 9–11; scales between ear and eye 13–16; scales around mid-body 73–78; subdigital lamellae on 4^th toe 9–11; dorsal scales per tail verticil 5; ventral scales per tail verticil 4; precloacal pores 8.

In life (holotype NB 854, Fig. 5C). Dark grey above with random yellow-olive scattered scales, with six irregularly-spaced, darker grey to black W-shaped crossbars, first one broken up into three blotches, each crossbar anteriorly bordered by a row of light white to yellow scales; head mostly grey with scattered darker grey and yellow scales, dark black bar from nostril to posterior of eye, continuing anteriorly of the eye to ear opening; no thin pale white canthal stripe; upper and lower labials dark grey with diffuse lighter grey edges; lateral sides of the body with a mix of dark grey and yellow blotches; limbs darker grey above with scattered yellow scales; tail (regenerated) with dark black blotches and irregular white to light grey mottling; iris dark brown to golden with a narrow black elliptic pupil with crenulated edge and black reticulation; ventre uniform greyish with scattered black specks; ventral surface of limbs with scattered black specks, more prominent than on the ventrum. In preservative (holotype NB 854, Fig. 8): dorsum with five distinct, irregularly-spaced, dark grey W-shaped crossbars anteriorly, with beige intervening blotches, the posteriorly crossbar on the nape are broken up dark grey blotches; dorsally, the arms and legs are beige with irregular darker grey blotches; tail (regenerated part) with light grey to white mottling on darker grey to black background; dorsally, the head has mottled dark brown scales, dark grey bar running from the nasals through eye to anterior of the ear opening; supralabials dark brown-edged ventrally; infralabials scattered with dark brown markings dorsally; ventrum uniform greyish with scattered black specks; ventrally, limbs with scattered black specks, more prominent than on the ventrum. Variation: Similar colouration and patterning as the holotype (preserved) and paratype (life). Dorsal dark W-shaped crossbars number 5–6. Regenerated tails with fine dark brown to black mottling. Juveniles with more sharply-defined pattern and darker colouration.

Figure 8. 

Holotype of Afroedura praedicta sp. nov. (NB 854) from Serra da Neve, Namibe Province, Angola. Photos: Telmo António.

(Fig. 4C). A rupiculous species found inside deep crevices and fractured rocks in large boulders of extrusive origin. All specimens were found on one site between 1,900–2,000 m a.s.l. The surrounding vegetation included altitude dwarf miombo and montane elements. Found in syntopy with Cordylus phonolithos at Serra da Neve.

This species is known only from the alkaline mountain complex of Serra da Neve (Fig. 1).

Afroedura bogerti – Haacke 2008:6, Huntley 2009:84, Rösler 2000:57, Barts and Haacke 2010:39, Jacobsen et al. 2014:456 & 468 (part), Branch et al. 2017b:157 (part), Marques et al. 2018: 177 (part), Branch et al. 2019a: 287 (part); Afroedura cf. bogerti – Agarwal et al. 2017:649; Afroedura bogerti (clade 1) – Branch et al. 2017a:146.

PEM R17937, adult female, collected from Omauha Lodge, 15 km south of Tambor (-16.20061, 12.40183, 341 m a.s.l.), Namibe Province, Angola, by William R. Branch, Werner Conradie, Krystal Tolley and John Measey on 18 January 2009.

PEM R17936, collected from Omauha Lodge, 15 km South Tambor (-16.20061, 12.40183, 341 m a.s.l), Namibe Province, Angola, by William R. Branch, Werner Conradie, Krystal Tolley and John Measey on 18 January 2009.

Males: TM 40508, TM 40512, TM 40516, TM 40518, collected from Tambor (-16.06667, 12.43333, 355 m a.s.l.), Namibe Province, Angola, by Wulf Haacke on 1 April 1971. Females: PEM R18041–2, collected 0.5 km south of Tambor (-16.07414, 12.43328, 352 m a.s.l.), Namibe Province, Angola, by William R. Branch, Krystal Tolley and John Measey on 23 January 2009; TM 40509–11, TM 40513–5, TM 40517, TM 40536–8, collected from Tambor (-16.06667, 12.43333, 355 m a.s.l.), Namibe Province, Angola, by Wulf Haacke on 1 April 1971.

(not examined). TM 40519–20, collected from Tambor (-16.06667, 12.43333, 355 m a.s.l.), Namibe Province, Angola, by Wulf Haacke on 1 April 1971; FKH 0341–2, collected from Omauha Lodge (-16.20061, 12.40183, 338 m a.s.l.), Namibe Province, Angola, by Pedro Vaz Pinto on 3 October 2019; CAS:HERP 263012–3, collected from Omauha (-16.19858, 12.40073, 338 m a.s.l.), Namibe Province, Angola, by Luis M.P. Ceriaco, Suzana Bandeira and Isham Agarwal on 25 and 27 November 2017; CAS:HERP 248780–1, collected 0.5 km south of Tambor (-16.07414, 12.43328, 352 m a.s.l.), Namibe Province, Angola, by William R. Branch, Krystal Tolley and John Measey on 23 January 2009.

This gecko is named after Donvé Branch, WRB’s wife, with the following personal quote: “This, the most beautiful of all the Angolan flat geckos, is named for my wife, Donvé Branch (‘Dove’) who bore the long periods I was away on fieldwork, and to whose nest I returned, and surrounded me with love until the end”. The name is constructed in the feminine singular genitive.

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 78 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 donveae sp. nov. differs from other members of the A. bogerti-group by a combination of the following characters (see Tables 3, 4): 64–78 (mean 72.8) mid-body scale rows (69–77 [mean 73.5] in A. bogerti, 76–88 [mean 79.3] in A. wulfhaackei sp. nov., 73–86 [mean 80.3] in A. vazpintorum sp. nov., 73–78 [mean 74.8] in A. praedicta sp. nov.); by the anterior nasals always in contact (similar to A. vazpintorum and A. praedicta sp. nov.; in contact in ~ 33% of A. bogerti; in contact in ~ 68% of A. wulfhaackei sp. nov.); in each verticil having 5–6 (mean 5.5) ventral and 6–7 (mean 6.6) dorsal rows of scales (5–6 [mean 5.0] and 6–7 [mean 6.1] to A. vazpintorum 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. vazpintorum sp. nov.; greyish with black spots in A. bogerti, A. wulfhaackei sp. nov. and A. praedicta sp. nov.); larger average adult size 57.6 mm SVL (versus 50.0 mm in A. bogerti, 51.7 mm in A. wulfhaackei sp. nov., 51.3 mm in A. vazpintorum sp. nov.; 49.9 mm A. praedicta sp. nov.). Afroedura donveae sp. nov. differs more specifically from its sister lowland species A. vazpintorum sp. nov. in being larger (57.6 mm versus 51.3 mm average SVL) and having lower mid-body scale counts (64–78 [mean 72.8] versus 73–86 [mean 80.3]), higher numbers of precloacal pores (11–12 [mean 11.5] versus 9–11 [mean 10.2]), bolder colouration and distinct tail banding (versus duller colouration and less distinct tail banding).

Adult female: SVL 61.0 mm; tail 59.0 mm (full original tail), 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 compressed; HL 13.6 mm, HW 11.4 mm, broadest at posterior level of eye and 1.19 times longer than wide. Eyes large (3.4 mm wide), pupil vertical with indented margins; circumorbital scales small and smooth, elongated at upper anterior margin, upper three posterior scales with small upward pointing spines. Snout rounded, 5.6 mm long, longer than distance between eye and ear openings (4.5 mm). Scales on top of snout smooth, rounded, equal 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 rostral, three nasal scales; 1^st supralabial narrowly excluded from nostril; the supranasals are much larger than the subequal smaller postnasals, in broad contact with each other. Nostrils slightly elevated. Rostral roughly rectangular, but with its upper edges elongated due to extensions to the nostril. Nine supralabials on each side, the labial margin flexing upwards at the rictus (approx. mid-orbital position), with 1–2 minute scales proximal to the flexure. Ten infralabials on either side, with a small scale proximal to the flexure. At the lip, mental slightly narrower than adjacent infralabial, 63% of rostral and in contact with three distinctly elongated postmental scales. Scales on throat much smaller than those on belly, scales touching infralabials larger. Thirteen scales across the crown at level of front of eyes; 11 scales between nostril and front of eye; 13 scales from ear to eye; 74 scales around mid-body. Ear opening deep, oblique and roughly oval, only half as high as wide (0.4 × 0.9 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 the size of scales along the dorsal mid-line. Original tail slightly dorsoventrally flattened and distinctly verticillate (16 whorls in total), with obvious lateral constrictions; each verticil comprising 7 rows of imbricate scales dorsally and 6 rows of imbricate scales ventrally, with 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 on 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 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; 8 enlarged scale rows under 4^th toe.

SVL varied from 49.1–55.7 mm; original tail length 26.5–59.0 mm, 0.78 times SVL; head length 1.13–1.46 times head width; snout 1.99 times diameter of eye (see Table 7 for more measurements and scale counts of type series). Supranasals always in contact; the first upper labial and rostral always enters the nostril and the width of the rostral at the lip margin is always wider than that of the mental; 2–3 postmental scales (except PEM R18401 with only one); supralabials 8–10, infralabials 8–11; scales between anterior edges of eyes 10–13; scales between nostril and anterior edge of orbit 8–12; scales from ear to eye 11–14; scales around mid-body 64–78; subdigital lamellae on 4^th toe 6–8; dorsal scales per tail verticil 6–7 (mean 6.6); ventral scales per tail verticil 5–6 (mean 5.5); precloacal pores 11–12.

In life (holotype PEM R17937, Fig. 5D): Yellowish above, brighter anteriorly, with nine irregularly-shaped darker brown to black crossbars from the occiput to the sacrum, each separated by thinner (3–4 scales deep), light yellow crossbars; head with dark brown-black 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 on to the nape; upper and lower labials light grey with diffuse 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 anteriorly, posteriorly with black and white bars; iris dark black with irregular golden spots, a black, narrow, elliptical, crenulate-edged pupil with black reticulation; ventrum uniform beige with scattered brown specks on lateral edges only; ventrally, limbs with scattered brown spots. In preservative (holotype PEM R17937, Fig. 9): dorsum with nine irregularly spaced dark brown crossbars from the occiput to the sacrum with beige intervening blotches; dorsally, arms and legs darkly barred; tail with nine dark brown bars, bolder towards the tip; dorsally, head with mottled dark brown scales; a light beige canthal stripe from nostril to anterior corner of eyes, continuing from the posterior part of the eye to above the ear opening; dark brown bar 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 ventrally dark brown-edged; infralabials with scattered dark brown markings dorsally; ventrum is mostly immaculate, with dark brown spots ventrally on the arms and legs; ventrally, tail with nine dark brown bars, bolder posteriorly. Variation. Similar colouration and patterning as to the holotype. Dorsal crossbars are often fused to form 2–3 dark brown, X-shaped crossbars. Original tails with 6–7 broad, dark brown to black bars, separated by light beige to white bars; some specimens with fine one-scale-wide black bar separating the lighter bars. Regenerated tails with fine dark brown mottling. Juveniles with more sharply-defined patterns.

Figure 9. 

Holotype of Afroedura donveae sp. nov. (PEM R17937) from Omauha Lodge, 15 km south of Tambor, Namibe Province, Angola. Photos: Werner Conradie.

(Fig. 4D). One or two elongated white eggs were laid on limestone rock, to which they adhere in captivity; eggs measure 12.9–14.7 mm (mean 13.8 mm) × 10.5–12.1 mm (mean 11.6 mm) (n = 5) and hatch after 88‒92 days, with hatchlings measuring 30.2–30.9 mm SVL and 26.5–27.9 mm tail length (n = 5) (Barts and Haacke 2010). A rupicolous species living in crevices between rocks and under flakes of exfoliating rock amongst larger granite boulders, at elevations of 340–355 m a.s.l. in the arid Namib Desert. It appears to be associated with large granite outcrops in semi-desert shrublands, along the drier fringes of the Pro-Namib. Vegetation includes Senegalia (= Acacia) mellifera, Senegalia spp., Commiphora sp., Boscia foetida and Salvadora persica.

Currently known only from the south-western parts of Namibe Province in Angola, in granite formations around Tambor and on the right bank of the mid-Curoca River (Fig. 1). The species remains poorly known, but it is probably stable in numbers as the local habitat is currently not threatened. It likely occurs in Iona National Park.

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.

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.

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.

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.

(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.

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.

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 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).

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; 1^st 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 4^th toe. Precloacal pores 12.

(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 4^th 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.

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.

Figure 10. 

Holotype of Afroedura vazpintorum sp. nov. (PEM R24118) from 1 km east of Farm Mucungo, Namibe Province, Angola. Photos: Werner Conradie.

(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.

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.