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Cordylus angolensis (Bocage, 1895) was described 128 years ago on the basis of a single specimen collected at Caconda in the west-central highlands of Angola. Additional specimens referred to this species were collected at 'Mombolo' (also in the central highlands) during the Vernay Angola Expedition in 1925. As the holotype was apparently destroyed in the fire of 1978 at the Museu Bocage in Lisbon and no additional specimens have been collected, its taxonomic status and phylogenetic relationships has remained uncertain. The species has eluded all efforts aimed at its re-discovery in the vicinity of the type locality, with a single specimen from near Condé, north of Mombolo—collected in 1970 by Wulf Haacke—the only other specimen of Cordylus known from west-central Angola. Recent field work in the Angolan highlands resulted in the collection of a series of specimens from Taqueta Mountain (west of Caconda), Monte Verde (Sandula, 'Mombolo') and Uassamba (Vondo). A phylogenetic analysis, using three mitochondrial and six nuclear genes, indicated the existence of two distinct species-level lineages in the Angolan highlands. These two species are allopatric and morphologically distinct, differing especially in terms of their colour patterns, eye colour and certain scalation characteristics. We therefore confirm that C. angolensis is a valid species and designate a neotype, and describe a new species, Cordylus momboloensis sp. nov.

Bauerius gen. nov. (Figs. 2–4) Type species. Phyllodactylus ansorgii Boulenger, 1907 Etymology. We name this new genus of gecko after Professor Aaron M. Bauer, renowned herpetologist at Villanova University (USA), in recognition of his large contributions to African herpetology, mainly to the biodiverse Gekkonidae group. The name is a conversion to the Latin nominative of the name “ Bauer ”. Definition. Bauerius gen. nov. is distinguished by the other leaf-toed geckos based on combinations of the following characters: moderate size (45 mm maximum SVL), with largely elongated head and body, and slender, rounded, semi-prehensile and non-ornamented tail. Well-ossified skull, elongated, oval (almost egg shaped), which displays no co-ossification with the overlying skin, and wider posterior snout region than postoccipital region. Parietal and fronto-parietal sutures well sealed. Frontal and nasal unpaired. Otostapes imperforate. Maxilla widely overlapping frontal, nasal, and prefrontal bones. Jugal bone reduced, almost vestigial. 14 scleral ossicles. 25–26 maxillary, 10–11 premaxillary and 31 dentary tooth loci. Surangular and compound bone fused. Splenial bone small and uniform, without alveolar foramen. 3 pairs of cervical ribs, 2 pairs of sternal ribs, and 3 pairs of uniform mesosternal ribs. Bauerius gen. nov. has 26 presacral vertebrae (paired Atlas, Axis, 5 cervical, 18 thoracic and 1 lumbar vertebrae) and one pair of well-developed cloacal bones. Digits are free and bear a single pair of dilated terminal adhesive pads (‘leaf toes’); the phalangeal formula is 2-3-4-5-3 (manus)/2-3-4-5-4 (pes); claws greatly reduced but present on all digits. Dorsal scalation consists of small and smooth granular scales, without tubercular scales. Males with 6–8 precloacal pores. One pair of postcloacal spurs. The eye bears a vertical pupil with crenate margins. Content. Bauerius ansorgii (Boulenger, 1907)

Nucras broadleyi sp. nov. Angolan Sandveld Lizard urn:lsid:zoobank.org:act: C82E3A75-96FF-4D2A-9B52-3ABF4B58BC2B (Figures 4–6) Chersonymy. Nucras tessellata var. taeniolata (Bocage 1895: 30), Nucras tessellata var. taeniolata (Boulenger 1910: 474), Nucras tessellata var. holubi (Boulenger 1917: 210), Nucras intertexta var. holubi (Boulenger 1920: 20), Nucras tessellata (Monard 1937: 73; Laurent 1964: 56), Nucras ornata (Broadley 1965: 23), Nucras tessellata (Broadley 1972: 30; Ceríaco et al. 2016: 56; Burger 2014: 171), Nucras aff. tessellata (Marques et al. 2018: 221; Branch et al. 2019: 317). Type material. The type series is comprised of the three most recently collected specimens, which are housed in PEM and TM. Holotype. A subadult male (PEM R 24005, AG 018), 10 km west of Lola, edge of Bentiaba River valley, Namibe Province, Angola (-14.29028, 13.53056, WGS 84, 802 m asl). Collected by W. R. Branch, P. Vaz Pinto, and J.S. de Almeida on 2 November 2015. Paratypes (2). a) A subadult female (PEM R 24157, AG 166), 8.8 km southwest of Farm Mucungo, Namibe Province, Angola (-14.80167, 12.41917, WGS 84, 385 m asl). Collected by W. R. Branch, P. Vaz Pinto, and J. S. de Almeida on 8 November 2015. b) An unsexed adult (TM 40392), “ 34 km S of Moçâmedes to Porto Alexandre, Angola, 1512 Ca ” (= 34 km S Namibe to Tômbwa), Namibe Province, Angola (approx. -15.48220, 12.18289). Collected by W.D. Haacke on 30 March 1971. Additional referred material: The following additional material was used to expand the description of variation within the species: a) an adult male (MD 1967, Laurent 1964), “km 34 de la route de Moçâmedes à Sa da Bandeira” (= 34 km from Namibe on Lubango road, -15.03333, 12.41667), collected 24 October 1949, b) MBL 646, 647 a, 647b (Bocage 1895: 30) from Maconjo (approx. -15.01667, 13.20000), c) BM 1970.6.29.10–11 (Boulenger 1910: 474) from Ponang Kuma (= Donguena, approx. -17.01667, 14.71667), and d) MHNC 91.0524 (Monard 1937) from Capelongo (approx. -14.88333, 15.083333), collected April 1933. Etymology. The specific epithet is a patronym in honor of Donald G. Broadley for his numerous contributions to the herpetofauna of Africa. Don (as most of us knew him) was the first to recognize the Angolan population as a separate species (Broadley 1972). The name is constructed in the masculine genitive. Diagnosis. Assigned to Nucras due to a well-defined collar (absent in Ichnotropis), toes not serrated or fringed (versus serrated or fringed in Meroles), subdigital lamellae smooth (versus keeled in Pedioplanis and Heliobolus), subocular bordering lip, the nostril is pierced between two nasals, nasal well separated from upper labial, and dorsal scales small, smooth, and juxtaposed. * anterior subocular (posterior subocular), ** total (largest number, t = truncated) The new species can be diagnosed from other Nucras species based on a combination of the following characters: series of transversely enlarged plates present under forearm (versus absent or only feebly enlarged in Nucras lalandii), a small series (0–6) of small granules present between supraciliaries and supraoculars (versus mostly absent in N. boulengeri and N. lalandii), 23–29 lamellae under 4 th toe (versus less than 22 in N. lalandii), dorsum with a series of longitudinal pale stripes (versus dark cross bands present in N. lalandii and N. scalaris or a series of pale vertebral spots, sometimes forming irregular transverse bands in N. intertexta or lack of any dorsal patterns in N. aurantiaca), four pale stripes on nape with outer stripes forming a. Bloemfontein Museum Province Province Province Province Province Province Province Province Province National Cape Cape Cape Cape Cape Cape Cape NMB – , , Northern Western, Northern,, Northern Northern, Western,, Western Kamanjab, KwaZulu-Natal Cape , Western Museum Locality South Africa Africa South South Africa South Africa South Africa GenBank Africa South South Africa Namibia, South Africa Africa South Elizabeth Port – 1 005257 951543 951540 951542 951541 632229 871208 547691 871206 871207 547718 RAG PEM HG HF HF HF HF EF DQ HF DQ DQ HF, Science 4 005232 951535 951536 751398 547725 547732 547733 547759 of ND HG — HF — HF — FR HF HF HF HF Academy 005211 951559 951556 951558 951557 080358 871152 547772 871148 871149 547800 California S 16 HG HF HF HF HF AF DQ HF DQ DQ HF CAS – : number 206725 R 18745 R 16873 R 11574 R 16872 209602 R 18376 abbreviations Museum accession CAS PEM PEM NMB PEM NA NA NMNW CAS PEM Museum. accession 5584 08 069 - 20650 21061 20687 0531 08 6001 6067 049 analysis Field ID AMB KTH MB MB MB MH GW AMB AMB SVN genetic in Samples used Species tessellata tessellata tessellata tessellata tessellata longicaudata australis australis capensis capensis suborbitalis. ) continued 2 Table (Genus Nucras Nucras Nucras Nucras Nucras Outgroup Latastia Australolacerta Australolacerta Ichnotropis Ichnotropis Meroles continuous light stripe with the outer edges of the parietals (similar to Broadley’s (1972) N. tessellata tessellata var. “ T;” differs from N. livida and N. tessellata where the outer stripes often do not form a continuous light stripe with the outer edges of the parietals; differs from N. caeiscaudata and N. ornata where there are only three longitudinal stripes present on nape and sometimes the vertebral ones are absent), well defined occipital scale separating parietals (versus reduced or absent in northern Namibia N. holubi, which is referred to as N. intertexta damarana Parker; as well as absent in N. caesicaudata), parietal foramen absent (often present in all other species except N. taeniolata), and postnasals separated (usually fused in N. taeniolata). In the phylogenetic analysis, the uncorrected p -distances show that this clade differs by>8% for 16S,>14% for ND4, and>1% for RAG 1 sequence divergence from other members of the N. tessellata clade. Description of Holotype (Fig. 4). Body relatively slender (SVL approx. 4.5 times the head length, tail truncated), with hindlimbs larger than forelimbs (femur of hind limb equal to length of tibia); head narrow and elongated (56% longer than wide) with narrow pointed but blunt snout, that is slightly longer than distance from back of eye to rear of ear opening. Rostrum protruding and visible from below. Nasals paired and in contact (0.2 mm suture length), not swollen, nostril directed backwards separating postnasals. Frontonasal single, wider than long (1.1 × 1.8 mm). Prefrontals paired and in broad median contact with one another (0.6 mm suture length), wider than long (1.1 × 1.2 mm). Frontal entire, longer than wide (2.7 × 1.9 mm). Two large rounded supraoculars, both in contact with the frontal, with anterior supraocular preceded by a single large scale in contact with prefrontal, frontonasal, and posterior loreal, with posterior supraocular bordered by a single large scale in contact with parietal and frontoparietal. Paired frontoparietal in broad contact (1.3 mm suture length), nearly as wide as long (1.7 × 1.5 mm). Parietals twice as long as wide (3.1 × 1.8 mm), fully separate by a large, pentagonal interparietal (2.5 × 1.2 mm) that is twice as long as wide, slightly shorter than frontoparietals and nearly equal to length of frontonasal and prefrontal combined. Small subtriangular occipital (0.5 × 0.7 mm). Two loreals, second much larger than first. Six supraciliaries on each side, 1 st is the longest. A single minute granule scale between supraocular and supracilliares on right side, none on left side. Four supralabials anterior to subocular and three supralabials posterior to subocular, on both sides. Subocular slightly elevated medial and bordering the lip, its lower border being shorter than the upper. Three temporal scales, first longer than others, smooth. Tympanic shield as wide as long, border of ear opening. No ear lobes. Lower eyelid with transparent brille formed by five larger scales, surrounded by numerous smaller scales. Lower eyelid separated from subocular and enlarged temporal scales by a series of 10 smaller scales. Small scale above 3 rd supralabial separating the posterior loreal and subocular. Enlarged scale bordering 1 st post subocular, supralabial, and the subocular. Six infralabials on both sides, with 3 rd being longest; four enlarged pairs of chin shields, last largest and first three in broad contact. Twenty-four gular scales in a straight line between symphysis of chin shields and median collar plate, equal in size except last 4–5 larger. Collar free, comprising seven enlarged plates (median subtriangular) and extending slightly onto side of neck as a crease, bordered by 2–3 smaller scales. Dorsal scales small, juxtaposed, granular, smooth, larger on sides toward ventrals. Midbody scales 42. Ventral plates eight longitudinal and 28 transverse rows (from collar to groin), plates of the innermost rows longer than broad, with outer row notably smaller than other rows, transverse row of ventrals across chest just behind collar longer than broad; preanal scales irregular, median ones larger. Scales on upper surface of forearm large, smooth or slightly keeled. Scales on lower surface of forearm with eight enlarged plates, at least twice the width of scales on upper forearm. Scales on upper surface of tibia rhombic, subimbricate, smooth, and much larger than dorsal scales. Tibia below with a series of large plates. Subdigital lamellae under fourth toe 23 R / 25L. Femoral pores 13 R / 15L. Dorsal scales on tail oblique, strongly keeled diagonally, and truncate behind, ventral scales on tail obtusely keeled. Coloration. Dorsum with eight pale cream to white dorsolateral longitudinal stripes, separated by dark brown to black stripes. These stripes are more boldly patterned anteriorly, fading posteriorly. No light vertebral stripe. The two pale paravertebral stripes are separated by a very narrow strip of darker scales that starts on the interparietal through the occipital scale and fades posteriorly onto body and tail. The dorsolateral stripe extending along outer borders of parietals continues onto the tail. It is followed by the upper lateral stripe extending from posterior of the eye onto the head through the mid-temporal with a brief break above the ear opening, and continues onto the tail. The lower lateral stripe starts at the subocular, through the ear opening, broken briefly above the arm, after which it continues all the way onto the tail. Ventrum white and lower limbs oblique white. Fore limbs upper surface black with scattered pale blotches. Hind limbs light brown with pale blotches. Upper surface of tail red-brown, similar to hind limbs. Scales bordering the orbit are black edged. Variation (Figs. 5–6). Meristic and escalation data are summarized in Table 1. The largest specimen examined is (BM 1907.6.29.10) 74 + 144 mm (tail regenerated). Regarding coloration, there seem to be three main variations among material examined: 1) 8–9 longitudinal stripes as in holotype (in PEM R 24005, MBL 647a, 647b, MHNC 91.0524–5), 2) 4–5 pale longitudinal stripes broken up posteriorly with flanks spotted (in BM 1970.6.29.10–11, TM 40392, MD 1967), and 3) broken paravertebral stripes, continuous dorsolateral line and barred flanks (in PEM R 24157), similar to N. intertexta. Distribution. Found only in semi-arid south-western Angola, throughout much of Namibe Province and extending onto the escarpment of southern Hu í la and Cunene Provinces (Fig. 1). Known localities include: Maconjo (Bocage 1895: 30), Ponang Kuma (=Donguena) (Boulenger 1910: 472), 34 km from Namibe on Lubango road (Laurent 1964: 56), 34 km south of Tombwa (TM 40397), 8.8 km southwest of Farm Mucungo (this study), 10 km west of Lola (this study), and Capelongo (Monard 1937: 73). The locality of Caconda (Bocage 1895) extends the species distribution further north into Huíla Province, but the specimens could not be critically evaluated by Broadley (1972) and are now presumably lost. Habitat. The species appears to be associated with mopane woodlands, dry savannas, and semi-desert shrublands (Barbosa 1970). The new material was found in sandy plains with scattered low granite outcrops, with varying degrees of short grass cover and scattered bushes. Vegetation included Colophospermum mopane, Ficus sp., Senegalia (=Acacia) mellifera, Commiphora sp., Boscia foetida, and Salvadora persica. The confirmed historical records were also obtained within the dry woodland zone, even though the possible occurrence of the species in Caconda would place the species above 1,500 m asl and well into the mesic conditions of Brachystegia habitats (Barbosa 1970)., Published as part of Branch, William R., Conradie, Werner, Pinto, Pedro Vaz & Tolley, Krystal A., 2019, Another Angolan Namib endemic species: a new Nucras Gray, 1838 (Squamata: Lacertidae) from south-western Angola, pp. 82-95 in Amphibian & Reptile Conservation 13 (2) on pages 86-91, DOI: 10.5281/zenodo.3731977, {"references":["Bocage JVB. 1895. Herpetologie d'Angola et du Congo. Ministerio da Marinha e das Colonias, Lisbonne, Portugal. 203 p., 20 pls.","Boulenger GA. 1910. 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Branch, William R., Conradie, Werner, Pinto, Pedro Vaz, Tolley, Krystal A. (2019): Another Angolan Namib endemic species: a new Nucras Gray, 1838 (Squamata: Lacertidae) from south-western Angola. Amphibian & Reptile Conservation 13 (2): 82-95, DOI: http://doi.org/10.5281/zenodo.3731977, {"references":["Aljanabi SM, Martinez I. 1997. Universal and rapid saltextraction of high quality genomic DNA for PCR based techniques. Nucleic Acids Research 25: 4,692- 4,693.","Arevalo E, Davis SK, Sites JJW. 1994. Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in central Mexico. Systematic Biology 43: 387-418.","Barbosa LAG. 1970. Carta Fitogeografica de Angola. Instituto de Investigacao Cientifica de Angola, Luanda, Angola. 323 p.","Bauer AM, Childers JL, Broeckenhoven C, Mouton PLN. 2019. A new Nucras Gray, 1838 (Squamata: Lacertidae) from the Strandveld of the Western Cape, South Africa. 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Zootaxa 4061(3): 201-226.","Tolley K, Weeber J, Maritz B, Verburgt L, Bates M, Conradie W, Hofmeyr R, Turner A, Da Silva J, Alexander G. 2019. No safe haven: protection levels show imperiled South African reptiles not sufficiently safe-guarded despite low average extinction risk. Biological Conservation 233: 61-72."]}

Cordylus meculae Branch, Rödel & Marais, 2005 African Journal of Herpetology, 54(2):132; Figs. 1–2. Holotype: PEM R16166; summit rock outcrops of Serra Mecula (12°03’11”S, 37°38’49”E; 1029 m a.s.l.), Niassa Game Reserve, Niassa Province, Mozambique; W.R. Branch, M.O. Rödel and J. Marais, 27 October 2003. Paratypes (8): a) PEM R16163 (male allotype), R16164 –5; same data as holotype. b) PEM R16180, 16184 [wrongly listed in as R 16182 in type description], Serra Mecula (12°02’25”S, 37°38’32”E; approx. 1200 m a.s.l.); 29 October 2003. c) PEM R16198; Serra Mecula (12°03’30”S, 37°38’22”E; approx. 1100 m a.s.l.); 31 October 2003. d) PEM R16202 –3; western slopes of Serra Mecula (12°02’46”S, 37°37’21”E; 643 m a.s.l.); 1 November 2003. All collected by W.R. Branch, M.O. Rödel and J. Marais. Remarks. All with ventral incisions except for the holotype and two paratypes (PEM R16180 & 16184).

Pachydactylus capensis oculatus Hewitt, 1927 Records of the Albany Museum, 3(5):394; Pl XXII, Fig. 1. Current name: Pachydactylus oculatus (Hewitt, 1927) Holotype: PEM R11953 (formerly AMG 5304); farm Cyrilhurts, about six miles from Tarkastad, Eastern Cape Province, South Africa; Mr. R. Essex, August 1925. Remarks. The description does not mention where the specimens were deposited. A specimen agreeing in appearance with the illustrated specimen in the original description is present in the PEM. The only difference is that the original label reads ‘King’s Farm, Tarkastad’ and not ‘farm Cyrilhurts about six miles from Tarkastad’. One more juvenile specimen from the same locality (PEM R11928) is present in the PEM. Hewitt specifically states the ‘type’ (holotype) to be an adult male. No further types are listed. Hewitt considered it a form between maculatus and capensis, calling it Pachydactylus maculatus oculatus in his plate. Loveridge (1947) subsequently synonymised it with P. maculatus, but it was later elevated to a full species (Branch 1988)., Published as part of Conradie, Werner, Branch, William R. & Watson, Gillian, 2019, Type specimens in the Port Elizabeth Museum, South Africa, including the historically important Albany Museum collection. Part 2: Reptiles (Squamata), pp. 1-45 in Zootaxa 4576 (1) on page 14, DOI: 10.11646/zootaxa.4576.1.1, http://zenodo.org/record/2624562, {"references":["Hewitt, J. (1927) Further descriptions of reptiles and batrachians from South Africa. Records of the Albany Museum, 3 (5), 371 - 415.","Loveridge, A. (1947) Revision of the African lizards of the Family Gekkonidae. Bulletin of the Museum of Comparative Zoology, 98, 1 - 469."]}

Tetradactylus fitzsimonsi Hewitt, 1915 Annals of the Transvaal Museum, 5(2):101. Syntype: PEM R4439; Schoemachers Kop [=Schoenmakerkop] near Port Elizabeth, Eastern Cape Province, South Africa; collector and date unknown. Remarks: Although FitzSimons (1943) lists the cotypes as being in Port Elizabeth and Albany Museum, this must have been in error as the type description clearly states two specimens loaned to Hewitt from the Port Elizabeth Museum’s director F.W. FitzSimons. However, only the larger specimen (Schoenmakerskop) is currently extant in the PEM. Both Hewitt (1915, 1937b) and subsequent authors questioned the second smaller specimen from Kroonstad and subsequently disregarded it (FitzSimons 1943; Loveridge 1942; Branch 1990) and considered the taxon to be restricted to the Port Elizabeth region.The original label gives no details for collector or the date of collection. Based on consistent morphological differences, Bates (2014) treats this as a full species., Published as part of Conradie, Werner, Branch, William R. & Watson, Gillian, 2019, Type specimens in the Port Elizabeth Museum, South Africa, including the historically important Albany Museum collection. Part 2: Reptiles (Squamata), pp. 1-45 in Zootaxa 4576 (1) on page 22, DOI: 10.11646/zootaxa.4576.1.1, http://zenodo.org/record/2624562, {"references":["Hewitt, J. (1915) Descriptions of two new South African lizards, Tetradactylus laevicauda and T. fitzsimonsi. Annals of the Transvaal Museum, 5 (2), 101 - 103.","FitzSimons, V. (1943) The Lizards of South Africa. Transvaal Museum Memoirs, 1, i-xv, 1 - 528. FitzSimons, V. (1959) Some new reptiles from southern Africa and southern Angola. Annals of the Transvaal Museum, 23, 405 - 409.","Hewitt, J. (1937 b) A Guide to the Fertebrate Fauna of the Eastern Cape Province, South Africa, Part II: Reptiles, Amphibians, and Freshwater Fishes. Albany Museum, Grahamstown, vii + 141 pp.","Loveridge, A. (1942) Revision of the African lizards of the family Gerrhosauridae. Bulletin of the Museum of Comparative Zoology, 89, 485 - 543.","Branch, W. R. (1990) The genus Tetradactylus (Sauria: Gerrhosaurinae) in the Cape Province, South Africa: new records and their taxonomic status. Journal of the Herpetological Association of Africa, 37, 13 - 16. https: // doi. org / 10.1080 / 04416651.1990.9650261"]}

Scolecoseps broadleyi Verburgt, Verburgt & Branch, 2018 African Journal of Herpetology, 67(1):5; Figs. 2–3 Paratypes (4): a) PEM R22697; from near Maganja (10°50’08.2”S, 40°33’25.4”E), Palma District, Cabo Delgado Province, Mozambique; L. Verburgt, 24 August 2013. b) PEM R22698; near Quionga, 20 km north of Palma (10°35’55.4”S, 40°29’40.8”E), Palma District, Cabo Delgado Province, Mozambique; L. Verburgt, 4 December 2013. c) PEM R22699; 15 km inland from Olumbe, 20 km S of Palma (10°59’23.8”S, 40°25’13.8”E), Palma District, Cabo Delgado Province, Mozambique; L. Verburgt, 21 February 2014. d) PEM R22696; 10 km inland from Olumbe, 20 km south of Palma (11°00’15.6”S, 40°21’47.5”E), Palma District, Cabo Delgado Province, Mozambique; L. Verburgt, 22 February 2014. Remarks. All paratypes are in good condition. The holotype is in NMZB according to the type description., Published as part of Conradie, Werner, Branch, William R. & Watson, Gillian, 2019, Type specimens in the Port Elizabeth Museum, South Africa, including the historically important Albany Museum collection. Part 2: Reptiles (Squamata), pp. 1-45 in Zootaxa 4576 (1) on page 30, DOI: 10.11646/zootaxa.4576.1.1, http://zenodo.org/record/2624562, {"references":["Engelbrecht, H. E., Branch, W. R. Greenbaum, E., Alexander, G. J., Jackson, K., Burger, M., Conradie, W. & Tolley, K. A. (2018) Diversifying into the branches: species boundaries in African green and bush snakes, Philothamnus (Serpentes: Colubridae). Molecular Phylogenetics and Evolution, 30, 357 - 365. https: // doi. org / 10.1016 / j. ympev. 2018.10.023."]}