Afroedura wulfhaackei, 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 wulfhaackei sp. nov. Angolan Flat Gecko Osga-achatada de Angola Figures 5B , 7 ; Tables 3, 5

Note.

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.

Synonym.

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.

Holotype.

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.

Paratypes.

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 .

Additional referred material

(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).

Tentative referred additional material examined.

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.

Tentative referred additional material not examined.

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).

Etymology.

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.

Diagnosis.

A member of the greater ' Afroedura 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 ' Afroedura 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).

Holotype description.

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; 1st 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 4th toe 9. Precloacal pores 9.

Paratype variation

(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 4th toe 7-8; dorsal scales per tail verticil 5; ventral scales per tail verticil 4. Precloacal pores 9.

Additional material variation.

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 4th 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.

Colouration.

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.

Natural history and habitat

(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.

Distribution and conservation.

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.

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 2. Phylogenetic tree topology for the 16 S gene (identical for Bayes and ML) using Afroedura loveridgei as outgroup. Support values for posterior probability and bootstraps are indicated by circles (ML:> 70 % / PP:> 0.95).

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 7. Holotype of Afroedura wulfhaackei sp. nov. (PEM R 24234) from Maka-Mombolo, Benguela Province, Angola. Photos: Werner Conradie.