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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)

Schismaderma branchi sp. nov. (Figs. 6���10) urn:lsid:zoobank.org:pub: 5C42BEA8-4D3E-4659-9A16-B60C19012B7B Holotype. ZMB 91630, adult male, road between Mussolo and Dumba Cambango, 10��06���40������ S, 17��19���58������ E, elevation ~ 1100 m a.s.l., Malanje Province, Angola, small depression near road, 26September 2016, coll. Pedro Vaz Pinto. Paratypes. Female. PEM A14246, a few kilometres south of Mussolo on a road between Cangandala National Park and Luando Strict Reserve, 10��08���56������ S, 17��21���00������ E, elevation ~ 1100 m a.s.l., Malanje Province, Angola, small pond near road, 27 October 2015, coll. Pedro Vaz Pinto (Fig. 7). Males. CHL 0338, road between Mussolo and Dumba Cambango, 10��08���51������ S, 17��20���59������ E, elevation ~ 1100 m a.s.l., Malanje Province, Angola, small pond near road, 26 September 2016, coll. Pedro Vaz Pinto; ZMB 91628, road between Mussolo and Dumba Cambango, 10��08���51������ S, 17��20���59������ E, elevation ~ 1100 m a.s.l., Malanje Province, Angola, small pond near road, 26September 2016, coll. Pedro Vaz Pinto; ZMB 91629, road between Mussolo and Dumba Cambango, 10��08���51������ S, 17��20���59������ E, elevation ~ 1100 m a.s.l., Malanje Province, Angola, small pond near road, 26 September 2016, coll. Pedro Vaz Pinto. Additional material. Tadpoles. PEM A14307, road between Mussolo and Dumba Cambango, 10��06���08������ S, 17��19���50������ E, elevation ~ 1100 m a.s.l., Malanje Province, Angola, pond near the road, 17 November 2017, coll. Pedro Vaz Pinto (Fig. 9). Etymology. This new species is named after William Roy (Bill) Branch (1946���2018) as a tribute to this inspiring and passionate scientist, in recognition of his crucial role in African herpetology. He correctly suggested from an early stage that we were dealing with an undescribed Angolan Schismaderma, and strongly encouraged the collection of additional material and the investigative work leading to the current paper. Bill tragically passed away in October 2018, but his efforts and dedication during the last few years of his career had a major impact on the renascence of Angolan herpetology. His legacy will continue to inspire future generations. The species epithet is used as an invariable noun in apposition to the generic name. We suggest Angolan Red Toad and Sapo-vermelho de Angola as English and Portuguese common names, respectively. Diagnosis. The molecular results confirm that the new species is a member of the family Bufonidae and the genus Schismaderma. This is also shown by the presence of granular ventral skin, pupil horizontally elongated in bright light, a conspicuous tarsal fold, inner metatarsal tubercle small and not flanged, no adhesive terminal discs on fingers and toes, and no hard claws on toes on adults (Du Preez & Carruthers 2017). The genus Schismaderma is characterized by moderate-size, presence of external tympanum and tarsal fold, absence of externally visible parotoid glands, a prominent glandular fold running from above the tympanum laterally to near the groin and demarcating dorsal and ventral surfaces, dorsal skin granular, with reddish colouration and flat warts, ventral colouration white with bold black reticulations. This combination of features distinguishes Schismaderma from all other southern African bufonid genera and all other toad species in Angola. The back is usually uniformly coloured, sometimes carrying a pair of mid-dorsal dark blotches. The venter is heavily mottled in black, the gular region in males suffused with dark grey. The soles of hand and feet are black, with white single subarticular tubercles. The tympanum is clearly discernible, very large and rounded, the horizontal diameter larger than distance between eye and nostril. The tadpoles are also characteristic of Schismaderma, with a horse-shoe shaped fold of vascularised skin on the back (Channing et al. 2012; Viertel & Channing 2017). Schismaderma branchi sp. nov. can be distinguished from the only other congener, S. carens, by its smaller size; ventral skin of hind limbs, belly, chest and throat heavily mottled in black (Figs. 6B, 7B) (lightly speckled or immaculate in S. carens); fewer and smoother tubercles on the palm of hands and sole of feet (Fig. 8E���H) (prominent, well-developed, and more numerous in S. carens); and larger inner metatarsal tubercle in relation to shortest toe. Males of S. branchi sp. nov. have shorter NL, TLB, TE, UEW, TypH and IMT, and longer EL, ED, TypW and FTL than males of S. carens. Description of the holotype. ZMB 91630 (Fig. 6) is a moderate-sized adult male (SUL 66.7 mm, with robust limbs (Fig. 6). Head triangular, longer than wide (24.9 mm x 23.4 mm); short snout (8.1 mm), projecting slightly beyond upper jaw; rostral tip slightly rounded in dorsal, ventral and lateral views. Loreal region concave, canthus rostralis rounded. Internarial region flat, small nostrils, directed dorsolaterally, much closer to the snout tip than to eyes. Tympanum and tympanic annulus clearly discernible. Tympanum diameter (6.9 mm) approximately 1.1 times eye diameter (6.1 mm). Eye diameter approximately 1.3 times eye-nostril distance (4.7 mm); eye diameter 0.6 times interorbital distance (10 mm). Interorbital distance approximately 2.8 times internarial distance (3.6 mm). Upper eyelid width 5.3 mm. Prominent eyes; not projecting laterally out of margin of head in dorsal view, projecting about 50% above margin of head in lateral view, not visible ventrally. Eye pupil ellipsoid oriented horizontally. Well-developed limbs and digits. Thigh length 25.9 mm. Crus (tibiofibula) length 24.0 mm. Foot length 22.9 mm. First toe length 5.2 mm. Hand length 14.7 mm. Forearm length 14.9 mm. Tarsal fold present. No webbing on hands. Fingers with no terminal bulbs or discs. Nuptial pads on fingers absent. Relative length of fingers IV>III>I>II. Fingers and toes with rounded tips, and rounded, not very prominent single subarticular tubercles. A small round thenar tubercle and a large flat palmar tubercle on right hand (Fig. 6D). Relative length of toes IV>III=V>II>I. Three phalanges of toe IV free of webbing; a well-developed margin of web reaching distal phalanges. Toes with small and single subarticular tubercles. Prominent oval inner metatarsal tubercle (2.8 x 1.9 mm), more than half the size of shortest toe, and around two times the size of outer metatarsal tubercle (1.7 x 1.3 mm) (Fig. 8H). Both feet have an oval abrasion of the skin on the heel, between the inner and outer metatarsal tubercles (Fig. 6B,E). Dorsal granular skin with flat warts and with numerous, prominent conical asperities (Fig. 6A) with a pale edge and a very small dark tip, only visible with magnifying glass. Parotoid glands absent or not visible externally. Lateral glandular ridge extending from above tympanum to near leg insertion. Skin of ventral surfaces and flanks granular (Fig. 6A���B). Colouration: After four years preserved in formalin, dorsal ground colour greyish-brown; plain dorsum with no markings; small darker blotches in fore and hind limbs. Lateral ridge paler dorsally and darker ventrally, flanks greyish-brown, darker than dorsum, mottled in black (Fig. 6A). Tympanum with colour similar to dorsum, centre of tympanum with a paler blotch. Dorsal snout similar in colour and pattern to dorsum. Dorsal surface of forelimbs and fingers white with black blotches similar to ventral colouration. Lateral surface dark red speckled with black markings becoming more evident towards the belly. Dorsal surface of hind limbs greyish-brown with dark black markings. One dark blotch in middle of thigh. Base colour of dorsal foot greyish-brown; with scattered dark blotches, extending across toes. Region surrounding cloaca white heavily mottled in black. Margin of upper jaw greyish-brown with small dark markings posterior and anterior to eye. Vocal sac diffuse dark grey. Throat, belly, and ventral and anterior surfaces of fore and hind limbs white, heavily mottled in black; mottling slightly thinner in posterior part of the ventral region (Fig. 6B). Soles of hands and feet black with white tubercles. Webbing of feet black with thin white edge (Fig. 8H). The holotype was collected in the breeding season (as all other collected adults, see below). It is unclear if the colour might be duller during other parts of the year, as observed in S. carens (see Fig. 3). Variation of paratypes. In life, dorsal ground colour bright red. Iris golden with black pupil. Little variation was observed in the four paratypes. Three paratypes (CHL 0338; ZMB 91628; ZMB 91629) are smaller than the holotype (SUL S. carens (Channing & R��del 2019). Tadpole morphology. Tadpoles at Gosner stage 26 (Gosner 1960) have a morphology and behaviour typical of Schismaderma tadpoles, i.e. concerning body shape, presence of a horse-shoe shaped fold of vascularised skin on the back (Fig. 9B), and congregating in compact schools (Fig. 11F) (Channing et al. 2012). Lateral view (Fig. 9A): body round to ovoid. Head fold is present with the posterior edge free from the main body. Body width nearly equal to the body height. Snout oblique. Mouth directed ventrally. Spiracle sinistral, tubular, small, joined to body wall, positioned laterally and situated midbody. Dorsal view (Fig. 9B): snout rounded. Nasals rounded, large (0.2 mm), positioned laterally, closer to eyes than to pupils. Eyes moderately sized (ED 8.6% of body length), positioned and directed dorsally; central parietal eye clearly visible. Ventral view (Fig. 9C): internal collided intestine clearly visible through dermal skin. LTRF: 2/3. Size: Total length 13.7���17.8 mm and BL 5.3���7.8 mm. Tail 135% of the BL. Table 9 summarizes all measurements. Colouration in preservative: Body dark brown-black. Ventrum paler with scattered darker pigmentation and transparent anterior dermal skin. Tail musculature more heavily pigmented than fins. Advertisement call. The species was first found while calling in large, loud choruses from small ponds. The toads were observed calling while floating in the water, on two independent occasions in the early rainy season (see below). The call is a characteristic deep and loud ���whoob, whoob.��� Temporal and spectral parameters of the calls of S. branchi sp. nov. are provided in Table 10 and Fig. 10B. Calls of S. branchi sp. nov. consist of a single note that is repeated in time, lasting 0.70 to 0.80 s (average 0.76 �� 0.04 s; n = 3), and inter-call intervals longer than the calls varying between 1.81 and 2.73 (average 2.19 �� 0.48 s). Average call repetition rate 24.57 times/minute (n = 24); average pulses per note 61���64 (average 62.33 �� 1.25; n = 3); pulse repetition rate 76.05���87.98 pulses per second (average 82.60 �� 4.94; n = 3); pulse duration 11.37���13.15 ms (average 12.15 �� 0.74 ms; n = 3). Dominant frequency of 375.0 (375.0 Hz; n = 3), mostly constant throughout the call; 90% frequency bandwidth 281.25���562.5 Hz. Habitat and life history notes. All specimens of S. branchi sp. nov. were found during the early rains, the species��� peak breeding season. Adults were breeding during the day in temporary pans that consisted of fairly small (2 to 5 m across) but relatively deep (1 to 2 m at the deepest section), and irregularly distributed depressions filled with murky water, recently accumulated after storms at the onset of the rainy season. These small pans were associated with small streams connected to large inland lakes within the Luando River sub-basin (Fig. 1). They had little vegetation apart from flooded grass (Fig. 11F), and some were artificial, having originated from abandoned road construction works. The first time the species was found, in 27 October 2015, it was breeding in two of these temporary ponds. On a second visit to the site, in 26 September 2016, red toads were found breeding in other additional temporary ponds near the original site. Breeding was observed after heavy rains at the beginning of the rainy season in two consecutive years. On the first occasion, between 100 and 200 toads were seen in three ponds over a distance of around 5 km. On the second visit, around 50 toads were observed in two ponds. No red toads were found in the nearby streams��� main channels nor in larger perennial ponds. Observed breeding behaviour was typical of Schismaderma toads (Balinsky 1969; Tandy & Keith 1972), with males calling loudly and incessantly throughout the day in shallow water at the edges of the pond, during a mating frenzy. Egg strings were observed floating in the water and attached to vegetation (Fig. 11E). The eggs were arranged in one row; the number of eggs was not counted.Although S. carens is reported to have eggs in double strings (Channing 2001; Du Preez & Carruthers 2017), the single strings we observed were very likely from S. branchi sp. nov., as no other amphibian species was recorded reproducing in the same ponds. Further observations are necessary to confirm if egg strings differ between the two species or not. Tadpoles moved through the pools in large dense schools, this schooling behaviour being characteristic of Schismaderma (Fig. 11F). Although only found in pans while mating, S. branchi sp. nov. is expected to be terrestrial outside the breeding season like S. carens (Balinsky 1969). Habitat in the surrounding areas, where they likely spend most of the year, consisted of mesic woodlands and savannas within the Angolan Miombo woodlands ecoregion (Burgess et al. 2004; Huntley 2019). Just a few kilometres to the east, there were extensive floodplains with papyrus reed beds surrounding large inland lakes. The species was found in sympatry with Afrixalus wittei, Hyperolius angolensis, H. benguellensis, H. cinnamomeoventris, Kassina senegalensis, Leptopelis bocagii, Ptychadena anchietae, Sclerophrys gutturalis, and S. pusilla. Distribution. So far, S. branchi sp. nov. was only found in four nearby sites spanning 5 km along a road where it crosses small streams flowing to Lussechi and Loge rivers, both tributaries of the larger Luando River, part of the Cuanza River basin (Fig. 1). Just east of these sites are two large lakes, Capalanga and Quipemba surrounded by extensive swamps and floodplains and connected via the Loge River. Even though many sites have been surveyed within a 50 km radius, mostly to the north, west, and south, the species has not yet been recorded elsewhere. It is likely for S. branchi sp. nov. to have a wider distribution, which may extend into the neighbouring Cuango River drainage, but for the moment it appears to be restricted to the Luando River sub-basin, bordering the Cuanza-Congo watershed. The region where the species was found constitutes a fairly large inland depression stretching for about 100 km, roughly between the towns of Mussolo and Quirima. Conservation status. This species is currently known from a very small area in central Angola and might be endemic to that region. It is expected to occur in a wider area in the surroundings of the type locality, but it has not yet been recorded in two formally protected areas nearby that were proclaimed primarily for the conservation of the endemic and threatened giant sable antelope (Hippotragus niger variani): Cangandala National Park and Luando Strict Nature Reserve, both within 50 km radius to the northwest and southwest from the type locality, respectively (Fig. 1). Its habitat does not appear to be locally threatened and the species probably tolerates disturbance, similar to S. carens (Poynton 1996; IUCN 2013). Following IUCN criteria, uncertainties regarding abundance and distribution recommends it to be classified as Data Deficient (IUCN 2012)., Published as part of Baptista, Ninda L., Pinto, Pedro Vaz, Keates, Chad, Edwards, Shelley, R��del, Mark-Oliver & Conradie, Werner, 2021, A new species of red toad, Schismaderma Smith, 1849 (Anura: Bufonidae), from central Angola, pp. 301-332 in Zootaxa 5081 (3) on pages 320-325, DOI: 10.11646/zootaxa.5081.3.1, http://zenodo.org/record/5775710, {"references":["Du Preez, L. & Carruthers, V. (2017) Frogs of Southern Africa, a complete guide. Struik Nature, Cape Town, 519 pp.","Channing, A., Rodel, M. - O. & Channing, J. (2012) Tadpoles of Africa: the biology and identification of all known tadpoles in sub-Saharan Africa. Edition Chimaira, Frankfurt am Main, 402 pp.","Viertel, B. & Channing, A. (2017) The larva of Schismaderma carens (Smith, 1849) (Anura: Bufonidae) - a redescription. Alytes, 33, 38 - 46.","Channing, A. & Rodel, M. - O. (2019) Field guide to the frogs and other amphibians of Africa. Struik Nature, Cape Town, 408 pp.","Gosner, K. L. (1960) A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica, 16 (3), 183 - 190.","Balinsky, B. I. (1969) The reproductive ecology of amphibians of the Transvaal highveld. African Zoology, 4 (1), 37 - 93. https: // doi. org / 10.1080 / 00445096.1969.11447365","Tandy, M. & Keith, R. (1972) Bufo of Africa. In: Blair, W. F. (Ed.), Evolution in the Genus Bufo. University of Texas Press, Austin, Texas and London, pp. 119 - 170.","Channing, A. (2001) Amphibians of Central and Southern Africa. Cornell University Press, New York, New York, 470 pp. https: // doi. org / 10.7591 / 9781501733697","Burgess, N., Hales, J. A., Underwood, E., Dinerstein, E., Olson, D., Itoua, I., Schipper, J., Ricketts, T. & Newman, K. (2004) Terrestrial ecoregions of Africa and Madagascar: a conservation assessment. Island Press, Washington, 497 pp.","Huntley, B. J. (2019) Angola in outline: physiography, climate and patterns of biodiversity. In: Huntley, B. J., Russo, V., Lages, F. & Ferrand, N. (Eds.), Biodiversity of Angola. Science & conservation: a modern synthesis. Springer, Cham, pp. 15 - 42. https: // doi. org / 10.1007 / 978 - 3 - 030 - 03083 - 4 _ 2","Poynton, J. C. (1996) Diversity and conservation of African bufonids (Anura): some preliminary findings. African Journal of Herpetology, 45, 1 - 7. https: // doi. org / 10.1080 / 21564574.1996.9649958","IUCN SSC Amphibian Specialist Group (2013) Schismaderma carens. The IUCN Red List of Threatened Species, 2013, e. T 54885 A 3021025. https: // doi. org / 10.2305 / IUCN. UK. 2013 - 2. RLTS. T 54885 A 3021025. en","IUCN (2012) IUCN Red List Categories and Criteria: Version 3.1. 2 nd Edition. IUCN, Gland and Cambridge, 32 pp."]}

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. A revised list of South African reptiles and batrachians, with synoptic tables, special reference of specimens in the South African Museum, and descriptions of new species. Annals of the South African Museum 5: 455 - 538.","Boulenger GA. 1917. A revision of the lizards of the genus Nucras, Gray. Annals of the South African Museum 13: 195 - 216.","Boulenger GA. 1920. Monograph of the Lacertidae. Volume 1. Trustees of the British Museum (Natural History), London, United Kingdom. x + 352 p.","Monard A. 1937. Contribution a l'herpetologie d'Angola. Arquivos do Museu Bocage 8: 19 - 154.","Laurent RF. 1964. Reptiles et amphibiens de l'Angola (troiseme contribution). Publicacoes Culturais Com- panhia de Diamantes de Angola 67: 1 - 165.","Broadley DG. 1965. Some problems presented by sand lizards of the Nucras tessellata group. Journal of the Herpetological Association of Africa 1: 18 - 23.","Broadley DG. 1972. A review of the Nucras tessellata group (Sauria: Lacertidae). Arnoldia 20: 1 - 35.","Ceriaco LMP, de Sa SC, Bandeira S, Valerio H, Stanley EL, Kuhn AL, Marques M, Vindum JV, Blackburn DC, Bauer AM. 2016. Herpetological survey of Iona National Park and Namibe Regional Natural Park, with a synoptic list of the amphibians and reptiles of Namibe Province, Southwestern Angola. Proceedings of the California Academy of Sciences 63 (2): 15 - 61.","Burger M. 2014. Nucras tessellata (A. Smith, 1838). Pp. 171 - 172 In: Atlas and Red List of the Reptiles of South Africa, Lesotho, and Swaziland. Editors, Bates MF, Branch WR, Bauer AM, Burger M, Marais J, Alexander GJ, de Villiers MS. Suricata 1. South African National Biodiversity Institute, Pretoria, South Africa. 485 p.","Marques MP, Ceriaco LMP, Blackburn DC, Bauer AM 2018. Diversity and distribution of the amphibians and terrestrial reptiles of Angola: atlas of historical and bibliographic records (1840 - 2017). Proceedings of the California Academy of Sciences Series 4, 65 (Supplement II): 1 - 501.","Branch WR, Baptista N, Vaz Pinto P, Conradie W. 2019. The reptiles of Angola - History, updated checklists, endemism, hot spots, and future directions for research. Pp. 283 - 326 In: Biodiversity of Angola. Science and Conservation: A Modern Synthesis. Editors, Huntley BJ, Ferrand N, Russo V, Lages F. Springer Open, Cham, Switzerland. 552 p.","Barbosa LAG. 1970. Carta Fitogeografica de Angola. Instituto de Investigacao Cientifica de Angola, Luanda, Angola. 323 p."]}

1. We developed a biogeographical regionalization of Angolan mammals based on data collected before major declines occurred during the civil war (1975–2002). In terms of its biodiversity, Angola is one of the least known of all African countries., 2. We used 9880 grid records of 140 species (rodents, ungulates and carnivores) collected mainly in 1930–80, at a quarter degree cell resolution. Biogeographical regions were identified by using cluster analysis, based on βsim dissimilarity matrices and a hierarchical classification using Ward's method. An indicator value analysis was used to identify species characterizing each region. Distance-based redundancy analysis was used to investigate the environmental correlates of mammalian assemblages., 3. Four biogeographical subdivisions emerged from ungulate distributions, while rodent and carnivore data were largely uninformative. In the north, the Zaire-Lunda-Cuanza region was mainly characterized by ungulate species associated with Congolian forests. In the south, the Namibe and Cunene-Cuando Cubango regions were mainly characterized by ungulates widespread in south-western and southern Africa. In between these regions, the Central Plateau region was mainly characterized by a few widespread ungulate species that are relatively common in dense miombo woodlands., 4. Biogeographical patterns were significantly associated with a dominant north–south gradient of decreasing humidity and increasing temperature, and with a concurrent gradient from dense forests and woodlands to open savannas, grasslands and deserts., 5. The biogeographical regions we identified in Angola were largely consistent with other bioregionalizations developed using various taxonomic groups at larger spatial scales. Biogeographical patterns reflected the southward penetration of Congolian forest species in the north, and the northward penetration of southern African desert/grassland species in the south-west and of open savanna species in the south. These processes seem to be controlled by the distribution of vegetation types, which in turn are associated with climatic gradients and soil types. The stronger patterns observed for ungulates than for other mammals may reflect the close association of ungulates to specific vegetation types.