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Acanthocercus ceriacoi sp. nov. (Figs. 2–8) Stellio nigricollis: Bocage (1866: 43) [lapsus calami; see Bocage (1895) and Wagner et al. (2018)] Stellio atricollis: [part] Peters (1881: 147), Bocage (1895: 22) Agama atricollis: [part] Boulenger (1885: 256; 1905: 110), Ferreira (1903: 15), Monard (1937: 58) Agama cyanogaster: [part] Loveridge (1957: 195) Acanthocercus cyanocephalus: [part] Ceríaco et al. (2014: 670; 2016: 75; 2018b: 422), Marques et al. (2018: 27), Wagner et al. (2018: 27), Branch et al. (2019b: 313) Acanthocercus sp. Malanje: Wagner et al. (2021) Holotype. CAS 258430 (field number JVV 9152; Figs. 2–3), adult male from Cangandala National Park headquarters [- 9.819417° N, 16.653861° E, 1089 m a.s.l.], Malanje Province, Angola, collected by Mariana P. Marques, Luis M.P. Ceríaco, Suzana A. Bandeira, Edward L. Stanley and Jens V. Vindum on 14 September 2015. Paratypes. Five specimens, all from Angola: CAS 258429 (field number JVV 9099) adult male from the vicinity of Cangandala National Park headquarters [- 9.818917° N, 16.6541° E, 1106 m a.s.l.], Malanje Province, same collectors as the holotype on 13 September 2015; CAS 258431 (field number JVV 9460) adult female with the same collecting data as the holotype; CAS 258433 (field number JVV 9261) and CAS 258434 (field number JVV 9262), two adult males, respectively, both from the vicinity of Cangandala National Park headquarters [- 9.818583° N, 16.654028° E, 1089 m a.s.l.], Malanje Province, same collectors as the holotype on 18 September 2015; MHNC- UP / REP 860 (field number GJ 04564, Figs. 4–5), an adult female from near Zulumongo village [- 7.207669° N, 15.167342° E, 1184 m a.s.l.], Uíge Province, collected by Luis M.P. Ceríaco, Gregorius Jongsma, Stuart Nielsen and Ilola Jorge on 28 November 2019. Additional material. 42 specimens: Malanje Province: CAS 258428, 258432 258435 (field number JVV 9098, 9218, 9440) three adult males from the same locality and collectors as holotype, on 13, 17 and 22 September 2015, respectively; CAS 258436 (field number JVV 9298), an unsexed juvenile from 13.67 km southwest by road of Cangandala National Park headquarters [- 9.819222° N, 16.654139° E, 1086 m a.s.l.], collected by the same collectors as the holotype on 20 September 2015; NHMUK 1866.6.11. 1 adult male from Duque de Bragança (currently Kalandula) [- 9.1° N, 15.95° E, 1010 m a.s.l.], collected by Francisco Pinheiro Bayão in 1866 [specimen noted by Bocage (1866) and Boulenger (1885)]; NHMUK 1904.5.2. 26 adult male from Pungo Andongo [- 9.66667° N, 15.58333° E, 1220 m a.s.l.], collected by William John Ansorge in June/ July 1903 [specimen noted by Boulenger (1905)]; NHMUK 1904.5.2. 25 adult male from Bange N’Gola [- 8.433300° N, 16.56667° E, 723 m a.s.l.], collected by William John Ansorge in the end of 1903 [specimen noted by Boulenger (1905)]; NHMUK 1904.5.2.17–24 eight specimens from Quanza River (most likely Duque de Bragança), collected by William John Ansorge at the end of 1903 [specimen noted by Boulenger (1905)]; ZMB 10027 adult male from Malanje [- 9.55° N, 16.35° E, 1147 m a.s.l.], collected by Friedrich Wilhelm Alexander von Mechow between 1879 and 1881 [specimen noted by Peters (1881)]; MNHNL /Rep/A/Ag 108–126, 25 specimens from Capanda Dam area [- 9.72841° N, 15.34585° E, 859 m a.s.l.], collected by Ana Lavres between January and April 2003 [specimens noted by Ceríaco et al. (2014)]. Bengo Province: AMNH R 48179 adult female from Dande [- 8.47279° N, 13.361224° E, 45 m a.s.l.], collected by Rudyerd Boulton and Herbert Lang (Vernay-Lang expedition to Angola) on 4 August 1925 [specimen noted by Wagner et al. (2018)]. The records from Cassange [- 9.583330° N, 17.866670° E, 955 m a.s.l.], Malanje Province, noted by Bocage (1895), as well as those from Duque de Bragança [- 9.1° N, 15.95° E, 1010 m a.s.l.], Malanje Province, noted by Ferreira (1903) cannot be confirmed, as these specimens were destroyed in the fire that engulfed the Lisbon Museum in 1978. However, due to their geographic proximity and similar habitat, they are here tentatively identified as the newly described species. Diagnosis. Segmentation of the tail into distinct whorls places the new species in the genus Acanthocercus. Within this genus, heterogeneous body scalation refers it to the cyanogaster / atricollis group and the high density of enlarged trunk scales, in combination with a black patch on the shoulder, place it in the A. atricollis complex. Acanthocercus ceriacoi sp. nov. is a medium sized lizard (mean SVL 122.5 mm [99–143 mm]; mean total length 285 mm [224–330 mm]) for its genus and overall is very similar in morphology to A. cyanocephalus. However, it may be distinguished from all other species of the complex by its meristic characters. The new species has lower scale counts when compared to other species of the A. atricollis complex group (mean scale rows around midbody 108, mean dorsal scale rows 64, mean ventral scale rows 85). In life, displaying males are characterized by a blue coloration of the head; neck, shoulders, body and limbs brownish with a pattern of white and black stripes and speckles; and a pale vertebral band from the neck to the tail, distinct anteriorly between the flanks, becoming more indistinct along a brown background posteriorly. Comparison with other close related Acanthocercus species. Acanthocercus ceriacoi sp. nov. can be distinguished from other taxa of the A. atricollis complex by the following characters (see Table 2): (a) Acanthocercus atricollis has a larger SVL (mean 129 mm vs. 123 mm) and total length (mean 302 mm vs. 285 mm). It has higher mean counts of scale rows around midbody (112 vs. 108), dorsal scales (67 vs. 63), and ventral scales (87 vs. 85) than A. ceriacoi sp. nov.. The mean number of precloacal pores is higher in A. atricollis than in A. ceriacoi sp. nov. (22 vs 18). Acanthocercus atricollis presents a blue or green head and ventral surface, in some individuals extending to the body, while in A. ceriacoi sp. nov. the blue is limited to the head. (b) Acanthocercus branchi has a smaller SVL (mean 109 mm vs. 123 mm) and total length (mean 262 mm vs. 285 mm). It has higher mean counts of scale rows around midbody (117 vs. 108), dorsal scales (73 vs. 63), and ventral scales (88 vs. 85) than A. ceriacoi sp. nov.. The ranges of these characters largely overlap in both species, although they are higher in A. branchi than in A. ceriacoi sp. nov.. The mean number of precloacal pores is higher in A. branchi than in A. ceriacoi sp. nov. (20 vs 18). (c) Acanthocercus cyanocephalus has a larger SVL (mean 138 mm vs. 123 mm) and total length (mean 334 mm vs. 285 mm). It is morphologically very similar to A. ceriacoi sp. nov. and its meristic differences are subtle. Acanthocercus cyanocephalus has a higher mean dorsal scale count (68 vs. 63), and a lower mean number of ventral scales (80 vs. 85) than A. ceriacoi sp. nov.. The mean number of precloacal pores is higher in A. cyanocephalus than in A. ceriacoi sp. nov. (22 vs 18). The major difference between these two species is the coloration of displaying males, which in A. cyanocephalus is characterized by the blue coloration of the head extending to the neck and shoulders, whereas in A. ceriacoi sp. nov. the blue is limited to the head. (d) Acanthocercus cyanogaster has a smaller SVL (mean 101 mm vs. 123 mm) and total length (mean 255 mm vs. 285 mm). It has lower mean counts of ventral scales (76 vs. 85) than A. ceriacoi sp. nov.. The ranges of this character slightly overlap in the two species, although they are higher in A. ceriacoi sp. nov. than in A. cyanogaster. The mean number of precloacal pores is much higher in A. cyanogaster than in A. ceriacoi sp. nov. (38 vs 18). [A. cyanogaster is not a part of A. atricollis group but is included here due to its morphological similarity and because of the historical use of the name to describe this population]. (e) Acanthocercus gregorii has a smaller SVL (mean 119 mm vs. 123 mm) but greater total length (mean 291 mm vs. 285 mm). A. gregorii has lower mean counts of scale rows around midbody (104 vs. 108) than A. ceriacoi sp. nov.. The ranges of these characters are higher in A. ceriacoi sp. nov. than in A. gregorii. The mean number of precloacal pores is higher in A. gregorii than in A. ceriacoi sp. nov. (22 vs 18). In displaying adult males, A. gregorii are characterized by their bluish-green head and a uniform greenish-blue throat, with a uniform blue body (Klausewitz 1957; Wagner et al. 2018), whereas in A. ceriacoi sp. nov. the blue of the male is limited to the head. (f) Acanthocercus kiwuensis has a smaller SVL (mean 111 mm vs. 123 mm) but greater total length (mean 290 mm vs. 285 mm). A. kiwuensis has lower mean counts of scale rows around midbody (87 vs. 108) and ventral scales (67 vs. 85) than A. ceriacoi sp. nov.. The ranges of this character largely overlap in the two species, although they are higher in A. ceriacoi sp. nov. than in A. kiwuensis. Moreover A. kiwuensis does not possess transverse rows of enlarged scales (Wagner et al. 2018). The mean number of precloacal pores is lower in A. kiwuensis than in A. ceriacoi sp. nov. (10 vs 18). (g) Acanthocercus margaritae has a smaller SVL (mean 112 mm vs. 123 mm) and total length (mean 258 mm vs. 285 mm). It has higher mean counts of scale rows around midbody (136 vs. 108), dorsal scales (103 vs. 63), and ventral scales (101 vs. 85) than A. ceriacoi sp. nov.. The ranges of these characters are lower in A. ceriacoi sp. nov. than in A. margaritae. The mean number of precloacal pores is lower in A. margaritae than in A. ceriacoi sp. nov. (14 vs 18). (h) Acanthocercus minutus has a smaller SVL (mean 101 mm vs. 123 mm) and total length (mean 234 mm vs. 285 mm). It has higher mean counts of dorsal scales (79 vs. 63), and lower mean counts of ventral scales (78 vs. 85) than A. ceriacoi sp. nov.. The mean number of precloacal pores is higher in A. minutus than in A. ceriacoi sp. nov. (26 vs 18). (i) Acanthocercus ugandaensis has a smaller SVL (mean 103 mm vs. 123 mm) and total length (mean 275 mm vs. 285 mm). A. ugandaensis has lower mean counts of scale rows around midbody (94 vs. 108) and ventral scales (73 vs. 85) than A. ceriacoi sp. nov.. The ranges of these characters are higher in A. ceriacoi sp. nov. than in A. ugandaensis. The mean number of precloacal pores is higher in A. ugandaensis than in A. ceriacoi sp. nov. (24 vs 18). Description of the holotype. Adult male (Fig. 2). Measurements: snout–vent length (SVL) 143 mm, tail length (TL) 187 mm, head length (HL) 48 mm, head width (HW) 38 mm, head height (HH) 34 mm. Habitus stout, with a large triangular head distinct from body; nasal scale flat to faintly convex, smooth, oval and situated slightly below canthus rostralis, pierced by round nostril in posterior part of scale, directed laterally. Scales on anterior, lateral, and central parts of head large, but smaller from behind ear opening, half the size of large head scales; head scales unequal in size, not oriented in the same direction, usually smooth, sporadically heavily keeled or with a rough surface, with free anterior margins and only sporadically with sensory pits. Fourteen supralabial scales and twelve infralabial scales on both sides; supraocular scales smooth, twelve scales along the canthus rostralis from nasal to behind the eye; six scales from nasal to anterior margin of eye; parietal shield quadrangular; pineal organ small, visible in posterior part of parietal shield. Ear opening large, but smaller than eye, posterior margin lacking semicircle of spinose mucronate scales; tympanum superficial. No nuchal crest present. Gular scales smooth, flat to faintly convex in the anterior part of the gular area; mucronate and spinose in the posterior part, becoming more spiny and smaller towards gular fold. Dorsal body scales a mix of small and smooth matrix scales, sometimes giving the impression of granular scales, and scattered, pale white-edged, enlarged scales. Large scales keeled, sometimes mucronate or spinose, not arranged in clusters, but rather in seven to eight, very indistinct, transverse rows between limbs. Sixty-four dorsal scales along vertebral column from midpoint of pectoral region to midpoint of pelvic region. Vertebral region covered by a mix of small and large scales, forming a pattern distinct from rest of lateral parts of body, with a large set of vertebral scales strongly keeled to smooth or feebly keeled. Ventral body scales smooth, slightly imbricate at posterior margins, in 89 rows from midpoint of pectoral region to precloacal pores. One hundred twenty-six scale rows around body behind forelimbs; 105 around midbody and 80 around body in front of hind limbs. Precloacal scales in two rows (the more anterior weakly developed), twelve pores in the posterior row and seven pores in the row above. Scales on upper side of forelimb unequal in size and strongly keeled, smooth on underside, upper arm scales somewhat larger than largest dorsal body scales. Relative length of manual digits 4=3>2>5>1, subdigital lamellae keeled and mucronate, 21 under left 4 th finger. Scales on upper side of hind limb keeled to smooth, becoming completely smooth on underside; scales on upper thighs unequal in size, a mix of small and scattered enlarged scales that are as large as the enlarged dorsal body scales. Fourth toe longest, relative length of pedal digits 4>3>2>5>1, subdigital lamellae keeled and mucronate, 23 under left 4 th toe. Tail with indistinct whorls of five scale rings (one ring consists of distinctly smaller scales) in its basal portion, caudal scales keeled and mucronate. First third of tail slightly swollen, scales larger than in other parts of the body, strongly robust and keeled. Distal portion of the tail much thinner and slightly depressed. After formalin fixation and six years of preservation in ethanol (Fig. 2) head dark with some brown to orange scales. Gular region bluish and dark from the mental through the throat, whitish in lateral parts. Black patch on shoulders. Ground color of dorsum grayish to brownish, enlarged scales on upper and lateral parts of body and limbs speckled dirty white. Four to five brownish areola-like markings delimited by white pale circles are visible through the vertebral region. Ventral side of body, limbs and tail dirty grayish, speckled with dark spots. Enlarged scales on upper side of tail base distinctly whitish. Scales becoming more brownish-dirty white striped towards tail tip. Coloration. In displaying males (see Fig. 3), head vibrantly blue; neck, shoulders, body and limbs brownish with a pattern of white and black stripes and speckles; a pale vertebral band from the neck to the tail is present, distinct anteriorly between the flanks, becoming more indistinct and on a brown background posteriorly; enlarged dorsal scales of the lower part of the body yellowish, matrix scales greyish becoming brownish on the hind limbs; first half of the tail yellowish, second half of the tail with dark and lighter blue bands. Ventral parts of the body, limbs and tail dirty whitish with a brown patterning coloration on females in life (see Fig. 5). Shoulders brownish with a black patch, some individuals present a pale blue head. Four dark brown, broad transverse bands between limbs, each interspersed by transverse bands composed of a single row of enlarged yellow scales. Upper and lateral parts of tail with brown and whitish/yellowish bands. Ventral parts of the body, limbs and tail dirty whitish with an extensive dark brown pattern. Gular region of a vibrant blue. Regarding its Angolan congeners, the coloration of displaying males (Fig. 3) is overall more similar to A. cyanocephalus than to A. margaritae. However, in A. cyanocephalus, the blue coloration of the head extends to the neck and shoulders, while in A. ceriacoi sp. nov. the blue is limited to the head. The new species is clearly distinct from A. margaritae, in that the latter species has a blue coloration of the head, forelimbs, flanks of the upper part of the body and second half of the tail (not banded as in A. ceriacoi sp. nov.). Variation. Some variations of the type series are shown in Table 2. Snout-vent length 99–143 mm (mean 123 mm); Tail length 125–187 mm (mean 162 mm), with the ratio TL/SVL 1.26–1.31 (mean 1.32); head length 28–48 mm (mean 39 mm); supralabial scales on the left side 12–14 (mean 13); supraocular scales on the left side 6; temporal scales on the left side 7; scale rows around fore body 121–128 (mean 126); scale rows around hind body 74–87 (mean 80); subdigital lamellae under Fi4 21–23 (mean 21); subdigital lamellae under TOE4 21–23 (mean 22); precloacal pores 17–20 (mean 18). Distribution. Currently, Acanthocercus ceriacoi sp. nov. is only known from the central and northwestern regions of Angola, from central Malanje to Bengo Province in the west and Uíge Province in the north (Fig. 6). It is likely that the species may also be present in the neighboring provinces of Kwanza-Norte and Zaire, as well as in southern Democratic Republic of the Congo. Habitat and Natural History notes. In Cangandala National Park and surrounding areas, the species is commonly found dwelling in trees, several meters above ground. The park lies at an approximate elevation of 1000 m and its climate is similar to that of the Angolan central plateau, with vegetation mostly dominated by miombo woodlands (Fig. 7), with Brachystegia wangermeeana, B. floribunda, Julbernardia paniculata, Erythrophleum africanum, Combretum spp., and Rhus spp. growing on the red clay soils of the park (Grandvaux-Barbosa 1970). Some small riverine gallery forests also occur in the park. In Uíge Province, the species was found basking on the ground in open-savannah habitat, in an area particularly devoid of trees (Fig. 8). The habitat of this region mainly comprises a mosaic of Zambezian savannah and coffee plantation forests, dominated by the plant genera Hyparrhenia, Celtis, Albizia, Morus and Ficus (Grandvaux-Barbosa 1970). The female paratype (MHNCUP/REP 860) from Uíge Province, collected in November, was active on the ground around midday. After euthanasia and while preparing the specimen for fixing, three well-developed eggs were found in the abdominal cavity of the specimen.

Cordylus phonolithos sp. nov. (Tables 1���2; Fig. 4���8) lsid: urn:lsid:zoobank.org:act: 813340B4-8299-4B1B-A786-5552BDC0C722 C. namakuiyus [part]: Stanley et al. (2016: 209) Holotype. A subadult male (CAS 263581, field number AMB 10296; Figs. 4���5A) collected within a crevice in a granite boulder in the vicinity of N���Dolondolo (Figs. 2, 9), Namibe Province, Angola [- 13.80678��N, 13.13507��E, 752 m elevation], collected by L. Cer��aco, S. Bandeira and I. Agarwal, on 21 November 2016. Paratype. A juvenile specimen (INBAC: AMB 10272 [same as field number]; Fig. 5B), collected at the same locality as the holotype, under a small granite rock on a larger rock, by the same collectors, on 20 November 2016. Diagnosis. A medium sized Cordylus species, identified to genus by the following combination of characters: fully limbed, strongly depressed triangular head and body, osteoderms present, rhomboidal, imbricate and keeled dorsal scales present, occipitals non-spinose, and spinose caudal and limb scales enlarged (Branch 1998; Broadley & Branch 2002; Stanley et al. 2011). Cordylus phonolithos sp. nov. differs from all other species in the genus except for C. vittifer (Reichenow, 1887), C. machadoi and C. namakuiyus, by the presence (versus absence) of a transverse row of elongated dorsal scales immediately posterior to occipitals (Fig. 6). It is distinguished from C. vittifer by possessing an incomplete row of pre-occipital scales between posterior parietal and occipital scales (versus complete), and by having infralabials that are moderately deeply ridged (versus usually smooth). It is distinguished from C. machadoi by having a large keyhole-shaped interparietal in contact with frontoparietals and separating the anterior parietals (versus small, diamond-shaped interparietal not in contact with frontoparietal and never completely separating the anterior parietals in C. machadoi, see Fig. 6); having the intrusion of a scale in broad contact with the interparietal and occipitals, thereby separating posterior parietals (versus absence of this scale and posterior parietals in full contact in C. machadoi, see Fig. 6); a higher number of supralabials (6 in C. phonolithos sp. nov. versus 5 in C. machadoi); males with fewer caudal scales at the 15 th tail whorl (8 in C. phonolithos sp. nov. versus 10���11 in C. machadoi); an orange-brown dorsal body coloration (versus darker brown to black in C. machadoi), the absence of dark speckles on throat and ventral body surfaces (versus presence in C. machadoi) and a reduced, widely separated posteromedial parietal process (similar to C. angolensis and unlike C. machadoi, in which it is extensive and forked, and C. namakuiyus, in which it is extended and unbifurcated). It is distinguished from C. namakuiyus by having the intrusion of a scale in broad contact with interparietal and occipitals, thereby separating posterior parietals (versus absence of this scale, posterior parietals in full contact, see Fig. 6); a higher number of supralabials (6 in C. phonolithos sp. nov. versus 4���5 in C. namakuiyus); a higher number of femoral pores (7 in C. phonolithos sp. nov. versus 4-6 in C. namakuiyus); a higher number of generation glands (16���17 in C. phonolithos sp. nov. versus 12 in C. namakuiyus); males with fewer caudal scales at the 15 th tail whorl (8 in C. phonolithos sp. nov. versus 10 in C. namakuiyus); absence of osteoderms on throat and ventral surfaces (versus presence), and significantly thicker caudal osteoderms than dorsal osteoderms (resembling C. machadoi and C. angolensis); temporal scales are weakly keeled (versus strongly keeled). In coloration C. phonolithos sp. nov. is quite similar to C. namakuiyus, although the new species has a more vibrant coloration (orange-brown versus light brown). It is distinguished from C. angolensis by having a large keyhole-shaped interparietal in contact with frontoparietals, thereby separating anterior parietals, with an intrusion of a scale in broad contact with interparietal and occipitals (versus small, diamond-shaped interparietal not in contact with frontoparietal, thereby never completely separating anterior parietals, posterior parietals in broad contact, see Fig. 6); fewer ventral transverse scale rows (23 versus 27), orange-brown dorsal body coloration (versus brown with blackish speckles over paler dorsal ground coloration), and by the absence of a longitudinal series of whitish speckles along dorsal surface (versus presence of two longitudinal series of small whitish speckles along dorsum), and presence (versus absence) of a loreal. Description of holotype. SVL 71.4 mm. Head and body depressed. Head 1.3 times longer (21.6 mm) than broad (16.5 mm). HH 8.7 mm. SEL 8.2 mm. Nasals in median broad contact; entire frontonasal lozenge-shaped, broader than long, separated from frontal by enlarged prefrontals (in median contact, forming a suture), separated from rostral by nasals, separated from loreal by prefrontals; frontal in contact with first and second supraoculars, followed by a pair of frontoparietals in broad, median contact; a distinctive keyhole-shaped interparietal in broad contact with the frontoparietals, separating anterior parietals; intrusion of a scale in broad contact with interparietal and occipitals, thereby separating posterior parietals; right posterior parietal scale is fragmented; parietal window visible; 10 rugose occipital scales; 10 elongated nuchal scales. Four supraoculars and three supraciliaries. Nasals large, with nostril pierced centrally on upper margin. Loreal in contact with preocular, nasal and first two supralabials; three suboculars, well separated from the lip by the third, fourth and fifth supralabials. Rostral twice as broad as deep; supralabials 6; infralabials 6; chin shields 5. Mental twice as broad as long; gulars smooth, enlarged and forming transverse rows posteriorly, with 17 gulars between the posterior extent of the jaws. Dorsal scales rectangular, rugose, strongly and obtusely keeled; dorsals and laterals in 25 transverse and 23 longitudinal rows; ventrals squarish, smooth, in 23 transverse and 17 longitudinal rows. Scales on dorsum of limbs large, strongly keeled and spinose with thin, non-imbricated osteoderms; subdigital lamellae under fourth toe 15; SAL 28.9 mm. AGD 32.4 mm. HML 9.3 mm, RUL 7.9 mm; FL 12.9 mm; TFL 10.6 mm; LTL 9.8 mm; femoral pores seven; generation glands 16���17. Tail with whorls of large, elongate, strongly keeled, spinose and acuminate scales, spines directed posteriorly; largest spines in dorsolateral position. Cranial osteology. The parietal is pentagonal with two short, well-spaced posteromedial processes bracing a very small, laterally flattened posteriomedial supraoccipital process (Fig. 7). The premaxilla is unpaired and bears seven pleurodont teeth and five foramina, with a dorsal process that extends posteriorly to be clasped by the nasals, which themselves insert into an unpaired frontal posteriorly and prefrontals posterolaterally. The maxilla is typically scinciform, with a deeply grooved crista dentalis and 20 pleurodont teeth. A laminar lacrimal lies medial to the facial process of the maxilla, extending from the posterioventral process of the prefrontal to the anterior process of the jugal. No palpebral is present, though the prefrontal has a small, flattened, laterally projecting tubercle that supports the anteriormost superorbital osteoderm in much the same way. The jugal is triangular in cross-section and asymmetrically T-shaped, with a tapering anterior process and a broad, truncated posterior process that extends along and past the posterior edge of the maxilla. Edentate pterygoids extend back to connect with the quadrates, becoming C-shaped in cross-section posterior to the epipterygoid condyle. The squamosal is curved and blade-like, circular in cross-section anteriorly, becoming flattened posteriorly, where it articulates with the cephalic condyle of the quadrate and the supratemporals. Supratemporals are flattened, sickle-shaped and unfused with the paroccipital processes. The bones of the braincase are unfused, suggesting that this individual may be a subadult. The prootic bears an extended alar process, a well-developed, anteriorly expanded crista prootica, and a very weak supratrigeminal process. Basipterygoid processes are well developed and flattened. The lower jaw has a large adductor fossa, a flattened and medially curved retroarticular process, a medially open Meckelian canal and a dentary with a strong subdental shelf, 22 mandibular teeth, and 11 dentary foramina. Postcranial osteology. The holotype has 25 presacral, two sacral and 16 postsacral vertebrae (Fig. 7). There are five cervical ribs, three sternal ribs and two xiphisternal ribs. The asternal ribs are asymmetric, with four long ribs with ossified costal cartilage and six short asternal ribs on the left side and five long and seven short ribs on the right. The first three cervical ribs are distally flattened with bifid cartilaginous projections. The pelvic girdle is well developed and flattened. No iliac tubercle is present. There is a well-developed, ventrally angled pubic tubercle directly anterior to the obturator foramen. Both hypoischium and hyperischium are well developed. Pubic bones are well separated by a bifurcate prepubic cartilage. The sternal plate is broad and lacks a fontanelle. Interclavicle cruciform, clavicles rod-like and flattened dorsally. The epicoracoid is narrow and curved, connecting the scapular ray to the primary and secondary coracoid rays, but not to the anterior process of the scapular. The condyles of long bones are unfused and the metatarsals and metacarpels are not fully developed, suggesting that this individual is not fully adult. Digits display the typical squamate phalangeal arrangement of 2-3-4-5-3 for the manus and 2-3-4-5-4 for the pes. Osteoderms. Scales of the dorsal and temporal regions of the skull and the ventrolateral aspects of the jaws are underlain with rugose osteoderms (Fig. 8). These osteoderms are fused to the proximal parietal, frontal and postorbital bones, although the mesokinetic and metakinetic joints appear unobstructed and flexible. The tail, legs and dorsal and lateral aspects of the body are covered in osteoderms (Fig. 9). The dorsal and lateral trunk are protected by noncontiguous, rectangular, 100-200 ��m thick osteoderms that become increasingly keeled and mucronate laterally. The caudal osteoderms are significantly thicker (up to 500��m), sharply spined and arranged in imbricate transverse whorls. The whole limbs are covered by imbricate circular/rhomboid osteoderms, keeled and mucronate dorsally, plate-like ventrally. The gular and ventral regions lack osteoderms. Coloration. Dorsum orange-brown, fading to dirty yellow laterally. Head orange-brown; supralabials and infralabials yellowish; a dark-brown bar extends from the posterior aspect of the eye to the temporals. The base of the tail is brown, with an orange coloration similar to that the laterally that extends towards the tip; dorsum of limbs dark brown. Laterally, a dark-brown line extends from the neck towards the insertion of the forelimbs. The body venter is cream and subcaudal surface is faded orange. TABLE 2. Variation in meristic counts for specimens of Cordylus phonolithos sp. nov., C. namakuiyus, C. machadoi and C. angolensis. All measurements are presented in millimeters (mm). Data of C. namakuiyus retrieved from Stanley et al. (2016); data for C. machadoi retrieved from Stanley et al. (2016) and two additional specimens from this study; data for C. phonolithos sp. nov. and C. angolensis collected in this study. Variation. Variation in meristic counts of the type series is reported in Table 2. The single juvenile paratype agrees almost entirely with the holotype, although, the loreal and the preocular scales are fused and the right posterior parietal is entire. Measurements of the paratype are the following: SVL 49.9 mm; HL 15.2 mm; HW 10.7 mm; HH 6.4 mm; SEL 5.8 mm; SAL 20.2 mm; AGD 23.1 mm; HML 6 mm; RUL 5.4 mm; FL 7.9 mm; TFL 7.8 mm; LTL 8 mm. Distribution. The new species is known only from N���Dolondolo, at the base of the Serra de Neve Inselberg in Namibe Province, Angola. The observational record cited by Stanley et al. (2016), originally identified as C. namakuyius from ���Sera [sic] de Neve��� (= Serra da Neve) is likely to correspond to C. phonolithos sp. nov. Habitat and Natural history notes. This species was found in granite outcrops in ���sparse Miombo��� forest (Fig. 10), dominated by trees of the genera Brachystegia and Jubernardia (Grandvaux-Barbosa 1970). The juvenile paratype was found under a small granite rock on a larger rock, while the holotype was collected from within a crevice in a granite boulder. The latter was exposed outside of the crevice, quickly sheltering in response to our presence. CT scanning of the holotype revealed a myriapod carapace in its digestive tract. Etymology. The specific epithet ��� phonolithos ��� is a noun in apposition from the Greek ��� phono��� = sound + ��� lithos��� = rock, which means ���sound stone���. In the local Mucobal language the type locality name, ���N���Dolondolo��� means literally ���rock that sounds like a bell��� or ���bell��� and stems from the presence of a large and famous phonolite stone at the locality. Phonolites are rare igneous volcanic stones of intermediate composition between felsic and maphic, with aphanitic to porphyritic texture that produce a very distinctive metallic sound when hit, similar to the ringing of a metallic bell. We suggest ���N���Dolondolo Girdled Lizard��� and ���Lagarto Espinhoso de N���Dolondolo��� as the English and Portuguese common names, respectively, for this species., Published as part of Marques, Mariana P., Cer��aco, Luis M. P., Stanley, Edward L., Bandeira, Suzana A., Agarwal, Ishan & Bauer, Aaron M., 2019, A new species of Girdled Lizard (Squamata: Cordylidae) from the Serra da Neve Inselberg, Namibe Province, southwestern Angola, pp. 503-524 in Zootaxa 4668 (4) on pages 509-518, DOI: 10.11646/zootaxa.4668.4.4, http://zenodo.org/record/3449886, {"references":["Stanley, E. L., Ceriaco, L. M. P., Bandeira, S., Valerio, H., Bates, M. F. & Branch, W. R. (2016) A review of Cordylus machadoi (Squamata: Cordylidae) in southwestern Angola with the description of a new species from the Pro-Namib desert. Zootaxa, 4061 (3), 201 - 226. https: // doi. org / 10.11646 / zootaxa. 4061.3.1","Branch, W. R. (1998) Field Guide to Snakes and Other Reptiles of Southern Africa, 3 rd edition. Ralph Curtis Books, Sanibel Island, Florida, 399 pp.","Broadley, D. G. & Branch, W. R. (2002) A review of the small east African Cordylus (Sauria: Cordylidae), with the description of a new species. African Journal of Herpetology, 51, 9 - 34. https: // doi. org / 10.1080 / 21564574.2002.9635459","Stanley, E. L., Bauer, A. M., Jackman, T. R., Branch, W. R. & Mouton, P. le F. N. (2011) Between a rock and a hard polytomy: Rapid radiation in the rupicolous girdled lizards (Squamata: Cordylidae). Molecular Phylogenetics and Evolution, 58, 53 - 70. https: // doi. org / 10.1016 / j. ympev. 2010.08.024","Grandvaux-Barbosa L. A. (1970) Carta fitogeorafica de Angola. Instituto de Investigacao Cientifica de Angola, Luanda, 323 pp."]}

Trachylepis raymondlaurenti Marques, Cer��aco, Bandeira, Pauwels & Bauer sp. nov. (Figs. 5, 6) Mabuya megalura (de Witte 1953: 107) Mabuya megalura subsp. (Laurent 1964: 74) Trachylepis megalura (Broadley & Cotterill 2004: 42 [partim]) Trachylepis cf. megalura (Cer��aco et al. 2016b: 71; 2018b: 423; Marques et al. 2018: 264) Holotype. CAS 258401 (Figs. 5, 6), adult female collected in the Giant Sable Sanctuary of Cangandala National Park (- 9.84606�� N, 16.7223�� E, WGS-84; elevation 1101 m above sea level) (Fig. 4), Malanje Province, Angola, by Mariana Marques, Luis Cer��aco, Suzana Bandeira, Edward Stanley and Jens Vindum on 17 September 2015. Paratypes. MD 5309, adult female collected in Alto Cu��lo (- 10.01667�� N, 19.55�� E, WGS-84; elevation 1101 m), Lunda Sul Province, Angola, by a local in June 1954; MCZ R-67627, adult female with the same collection data as the former; RBINS 2691 (formerly RBINS 6998), adult female collected in Lusinga (- 8.933333�� N, 27.2�� E, WGS-84; elevation 1810 m), Upemba National Park, DRC, during the Mission G. - F. de Witte on 16 March 1947; RBINS 2692 (formerly RBINS 6999), adult male with locality data as for the former but collected on 9���17 April 1947; RBINS 2693 (formerly RBINS 7000), adult male with locality data as for RBINS 2691 but collected on 21 June���21 July 1947; RBINS 2694 (formerly RBINS 7001), adult male with the same locality data as for RBINS 2691 but collected in December 1947; RBINS 2695 (formerly RBINS 7002), adult female with the same locality data as for RBINS 2691 but collected in January 1948. Diagnosis. Trachylepis raymondlaurenti sp. nov. is readily distinguished from all other congeners by the following combination of characters: (1) slender body of medium size, SVL up to 79.3 mm; (2) long-tailed, up to 215.8 mm, greater than twice body length; (3) well-defined neck constriction; (4) color homogeneously greyishbrown dorsally, with some dark speckles on sides of tail; ventral surface with some scattered dark speckles near the flanks and under the tail; supralabials and infralabials dotted with white; no distinct longitudinal or transverse dorsal bands; (5) MSR 24���28, SAD 48���53, SAV 50���58; (6) KDS 0���3; (7) scales on palms of hands and soles of feet smooth; (8) supranasals separated; and (9) prefrontals separated. Holotype description. Individual in good condition. Gravid female, with 10 fetuses in advanced stage of development. Habitus and relative sizes of head, body and tail comparable to Trachylepis megalura. Body cylindrical and slender with a well-defined neck and thin limbs; tail very long, its length greater than twice the SVL, smoothly tapering. Fore- and hind limbs overlap when depressed against the body. SVL 79.3 mm, TL 152.3 mm. HL 13.1 mm, with relatively long snout (HL/HW 65.1%). Additional measurements are presented in Table 2. Ear opening small, without anterior lobes. Rostral visible from above. Small nostrils set posteriorly so that postnasal effectively borders nostril. Supranasals without contact. Frontonasal wider than long, in contact with loreal scale. Prefrontals pentagonal, separated from one another, each in contact with the following head shields: frontonasal, loreal, first and second supraocular, second supraciliary and frontal. Two loreals. Frontal longer than the distance between anterior tip of frontal and tip of snout. Frontal in contact with three supraoculars on each side. Two frontoparietals, in contact with each other and the frontal, third and fourth supraoculars, parietal and interparietal. Frontoparietal plus interparietal length greater than frontal length. Interparietal twice as long as wide, with a visible parietal foramen. Parietals of same length as frontoparietals. Parietals not in contact with one another. Five supraciliaries, second largest. Seven supralabials, fifth being subocular. Six infralabials. Postmental bordering seven scales. Transparent scale present in lower eyelid, as is usual for Trachylepis. Tympanum visible, at same level as mouth. Dorsal scales each with three smooth keels, sometimes very indistinct. Ventral scales smooth. MSR 28, SAD 52, SAV 58. Limbs with five digits; scales on soles of hands and feet smooth. Relative length of fingers IV>III>II>V>I, relative length of toes IV>III>V>II>I. Finger-IV lamellae 13, Toe-IV lamellae 17. Color in life is homogenous light greenish-brown on the entire dorsum; head, neck, legs and tail uniformly light grey-brown. No visible spots, markings or dorsolateral stripes. Gular region white, remaining ventral areas yellowish, except the brown palmar regions of hand and feet. In preservative, background color of flanks and upper side of head, neck, dorsum, legs and tail homogenously brown, with some white speckles running from the labials through the anterior half of the tail. Above the labials the head is uniformly brown. Eyelids dark-brown. Ventral surfaces uniformly whitish with some scattered dark speckles near the flanks and under the tail. Variation. Variation in measurements and scalation of the paratypes of Trachylepis raymondlaurenti sp. nov. are reported in Table 2. Head scalation similar to holotype in all paratypes except for prefrontals, which vary considerably; in RBINS 2 691 and RBINS 2695 the prefrontals are separated by an additional median scale, and in RBINS 2693 they are separated. Coloration of paratypes agrees entirely with that of holotype. Hatchlings have a reddish tail and hind legs (Fig. 7). Comparison with other species. Trachylepis raymondlaurenti sp. nov. differs from all congeners, with the exception of T. megalura and T. boulengeri, in being a slender skink with a well-marked neck, a very long tail (TL generally more than twice the SVL), and a low number of midbody scale rows (24���28) in comparison with the majority of Trachylepis species (Broadley 2000). Trachylepis raymondlaurenti is readily distinguishable from T. megalura by its coloration and scalation. Trachylepis megalura is characterized by a striped pattern with a number of fine dorsal longitudinal black or white stripes (grey-brown, light orange-brown or golden or red-brown) including a very characteristic prominent white stripe on the flanks, bordered above by a black stripe running from the upper lip along the entire body onto the tail, while the new species is characterized by the lack of the lateral stripe on the flanks, being homogeneously greyish-brown dorsally with scattered dark speckles. In terms of scalation, the supranasals and prefrontals are never in contact in T. raymondlaurenti sp. nov., while in T. megalura they are both in contact. Trachylepis boulengeri shares more morphological similarities with the new species than with T. megalura. Both Trachylepis boulengeri and T. raymondlaurenti have a uniform grey-brown background, sometimes with a few scattered dark flecks, but T. boulengeri also exhibits a black streak from below the eye to the ear opening, which is not visible in T. raymondlaurenti. In both taxa the supranasals and prefrontals are without contact, although T. boulengeri frequently has only one pretemporal (Broadley 2000), whereas T. raymondlaurenti has two. Trachylepis boulengeri has five to 11 keels per dorsal scale, whereas T. raymondlaurenti and T. megalura have only three weak keels, sometimes nearly indistinct. Distribution. Trachylepis raymondlaurenti sp. nov. appears to be restricted to the southeastern DRC in the Katanga Region, and to northern Angola, with records for Malanje and Lunda-Norte provinces. The species potentially occurs in the neighboring provinces of Bi��, Lunda-Sul and Moxico in Angola, and in northern parts of Zambia. Habitat and Natural History notes. The holotype was found while it was basking under a shrub of Cryptosepalum maraviense Oliv. (Fabaceae) in a Miombo woodland habitat (Fig. 8). Dissection revealed ten welldeveloped fetuses in the holotype (Fig. 7), indicating that the young of this diurnal, viviparous species are likely born in late September or October. The fetuses, and presumably neonates, have bright reddish-orange tails that are proportionally shorter than in the adults. The relative length changes in tail length during ontogeny (and presumably also does so in T. megalura), explain why the holotype of Eumeces massaianus, a probable subadult, has a proportionally shorter tail than the types of Euprepes megalurus, a fact which contributed to its early recognition as a distinct taxon. Etymology. The species is named after Raymond F. Laurent (1917���2005), Belgian herpetologist who specialized in African amphibians and reptiles and contributed significantly to current knowledge of the Angolan and Congolese herpetofaunas. We propose the English name "Laurent���s Long Tailed Skink", the Portuguese name "Lagartixa de Cauda Longa de Laurent", and the French name "Scinque �� longue queue de Laurent"., Published as part of Marques, Mariana P., Cer��aco, Luis M. P., Bandeira, Suzana, Pauwels, Olivier S. G. & Bauer, Aaron M., 2019, Description of a new long-tailed skink (Scincidae: Trachylepis) from Angola and the Democratic Republic of the Congo, pp. 51-68 in Zootaxa 4568 (1) on pages 59-62, DOI: 10.11646/zootaxa.4568.1.3, http://zenodo.org/record/2599228, {"references":["de Witte, G. - F. (1953) Exploration du Parc National de l'Upemba. Fol. 6. Reptiles. Institut des Parcs Nationaux du Congo Belge, Brussels, 322 pp.","Laurent, R. (1964) Reptiles et amphibiens de l'Angola (troisieme contribution). Publicacoes Culturais. Companhia de Diamantes de Angola, 67, 1 - 165.","Broadley, D. G. & Cotterill, F. P. D. (2004) The reptiles of southeast Katanga, an overlooked ' hot spot'. African Journal of Herpetology, 53 (1), 35 - 61. https: // doi. org / 10.1080 / 21564574.2004.9635497","Ceriaco, L. M. P., Marques, M. P. & Bandeira, S. A. (2016 b) Anfibios e Repteis do Parque Nacional da Cangandala. Instituto Nacional da Biodiversidade e Areas de Conservacao, Angola & Museu Nacional de Historia Nacional e da Ciencia da Universidade de Lisboa, Portugal, 97 pp.","Marques, M. P., Ceriaco, L. M. P., Blackburn, D. C. & Bauer, A. M. (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 (2), 1 - 501.","Broadley, D. G. (2000) A review of the genus Mabuya in southeastern Africa (Sauria: Scincidae). African Journal of Herpetology, 49 (2), 87 - 110. https: // doi. org / 10.1080 / 21564574.2000.9635437"]}

African pygmy toads of the genusPoyntonophrynusare some of the least known species of African toads. The genus comprises ten recognized species endemic to sub-Saharan Africa, five of which are restricted to southwestern Africa. Recent field research in Angola provided new material for three species ofPoyntonophrynus, including a morphologically distinctive population from the Serra da Neve Inselberg. Based on a combination of external morphology, high-resolution computed tomography scanning, and molecular phylogenetic analysis, the Serra da Neve population is described as new species that is nested within the genus. The most striking character that differentiates the newly described species from its congeners is the lack of a tympanic middle ear, a condition common in the family Bufonidae, but so far not known forPoyntonophrynus. The description of this new species from southwestern Angola reinforces the biogeographic importance of the region and further suggests that southwestern Africa is the cradle of diversity for this genus.

Stanley, Edward L., Ceríaco, Luis M. P., Bandeira, Suzana, Valerio, Hilaria, Bates, Michael F., Branch, William R. (2016): A review of Cordylus machadoi (Squamata: Cordylidae) in southwestern Angola, with the description of a new species from the Pro-Namib desert. Zootaxa 4061 (3): 201-226, DOI: http://doi.org/10.11646/zootaxa.4061.3.1

Cordylus namakuiyus sp. nov. (Figs. 1–6, Table 2) Holotype. CAS 254912, an adult female collected from a large rock outcrop near Caraculo, on the road from Lubango and Namibe, Namibe province, Angola [15 °0' 59.40 "S, 12 ° 38 ' 31.30 "E; 503 m elevation], collected by E. L. Stanley, S. de Sá, S. Bandeira, H. Valerio, A. L. Kuhn, J. V. Vindum and L. Ceríaco, on 6 December 2013. Paratypes. Eight specimens: CAS 254913 – 14, two fetuses taken from the holotype, with the same data; CAS 256529, one juvenile collected at the same locality as the holotype [15 °0' 57.3 "S, 12 ° 38 ' 32.6 "E; 509 m elevation] by the same collectors; CAS 254754, adult female, CAS 254755, adult male, CAS 256530, juvenile, CAS 256531, adult female, all collected on the outskirts of Pico Azevedo [15 ° 28 ' 33.2 "S, 12 ° 27 ' 45.7 "E; 421 m elevation] by the same collectors of the holotype on 7 December 2013; PEM R 18005, adult female, collected in a low rock outcrop bordering the road between Namibe and Omahua lodge [15 ° 59 ' 48.5 "S, 12 ° 24 ' 24.6 "E; 308 m elevation], Namibe Province, Angola, by W. R. Branch, K. A. Tolley and G. J. Measey on 20 January 2009. Additional material. Forty-nine specimens. One specimen: TM 40430, adult female from 49 km ESE of Tombua, Namibe Province, Angola [15 ° 53 'S, 12 ° 16 'E], collected by W. D. Haacke on 31 March 1971; and a series of 48 specimens collected by A. S. Vernay, H. Lang and R. Boulton between 25 April and 4 August 1925, locality “ Angola ”: Adult females: AMNH R47274, 47276, 47278, 47279, 47282 –84, 47286, 47287, 47289-95, 47297, 47298, 47299, 47304, 47305, and 47315. Adult males: AMNH R47275, 47277, 47280, 47281, 47285, 47288, 47296, 47300 –03, 47306 –09, and 47311. Juveniles: AMNH R47310, 47312– 14, and 47316 – 23. Diagnosis. A relatively large species of Cordylus, identified to genus by the following combination of characters: four-limbed, rupicolous, viviparous, strongly depressed triangular head and body, osteoderms present, rhomboidal, imbricate and keeled dorsal scales present, occipitals non-spinose, and spinose caudal and limb scales enlarged (Branch 1998; Broadley and Branch 2002; Stanley et al., 2011). Cordylus namakuiyus sp. nov. differs from all other species in the genus except for C. vittifer and C. machadoi, by the presence of a transverse row of elongated dorsal scales immediately posterior to occipitals (vs. absence); from C. vittifer by possessing an incomplete row of pre-occipital scales between posterior parietal and occipital scales (absent in C. vittifer), and by having infralabials that are moderately-deeply corrugated (vs. usually smooth); from C. machadoi by having a large, keyhole-shaped interparietal in broad contact with frontoparietals, thereby separating anterior parietals (vs. small, diamond-shaped interparietal not in contact with frontoparietal, thereby never completely separating anterior parietals), temporal scales that are strongly keeled (vs. weakly keeled), fewer transverse gular rows (14–17 vs. 17– 24), light brown dorsal body coloration (vs. darker brown), and by the absence of dark speckles on throat and ventral body surfaces (vs. presence), presence of osteoderms on throat and ventral surfaces (vs. absence); from C. angolensis (based on original description; Bocage 1895) by having fewer ventral transverse scale rows (21–24 vs. 27), light brown dorsal body coloration (vs. brown with blackish speckles over paler dorsal ground coloration), and by the absence of longitudinal series of whitish speckles along dorsal surface (vs. presence of two longitudinal series of small whitish speckles along dorsum), and presence of a loreal (vs. absence). Description of holotype. SVL 101 mm. Head and body depressed. Head 1.2 × as long as broad. Nasals in median broad contact; frontonasal lozenge-shaped, as broad as long, separated from frontal by enlarged prefrontals (in median contact, forming a suture), separated from rostral by nasals, separated from loreal by prefrontals; frontal in contact with first and second supraoculars, followed by a pair of large frontoparietals in broad, median contact; a distinctive keyhole-shaped interparietal in broad contact with the frontoparietals, separating anterior parietals; parietal eye visible; a row of 10 rugose occipitals. A row of 11 elongated nuchal scales. Four supraoculars and three supraciliaries. Nasals large, with nostril pierced centrally on upper margin. Loreal in contact with preocular, nasal and first two supralabials; three suboculars, well separated from the lip by the fourth and fifth supralabials. Rostral twice as broad as deep; supralabials five; infralabials five; chin shields five. Mental twice as broad as long; gulars smooth, enlarged and forming transverse rows posteriorly, with 16 gulars between the posterior extent of the jaws. Dorsal scales rectangular, rugose, obtusely keeled, mucronate and serrate posteriorly; dorsals and laterals in 25 transverse and 24 longitudinal rows; ventrals squarish, smooth, in 21 transverse and 15 longitudinal rows. Scales on limbs above large, strongly keeled and spinose; subdigital lamellae under fourth toe 14; femoral pores five; generation glands absent. Tail with whorls of large, elongate, strongly keeled, spinose and serrated scales, spines directed posteriorly and largest superolaterally. Cranial osteology. The parietal is pentagonal with a well-developed, unbifurcated medio-posterior process that extends past the postero-lateral processes. The premaxilla is unpaired and contains seven pleurodont teeth and five foramina, with a dorsal process that extends posteriorly to be clasped by the nasals, which themselves insert into an unpaired frontal. The maxilla is typically scinciform, with a deeply grooved crista dentalis and 16 pleurodont teeth. No palpebral is present, though the prefrontal has a small, flattened, laterally projecting tubercle that supports the anterior-most superorbital osteoderm in much the same way. The jugal is triangular in crosssection and asymmetrically T-shaped, with a tapering anterior process and a broad, truncated posterior process that extends along and past the posterior edge of the maxilla. The lacrimal bone is small, flattened and extends along the postero-ventral boundary of the maxilla and prefrontal. Edentate pterygoids extend back to connect with the quadrates, becoming C-shaped in cross-section posterior to the epipterygoid condyle. The squamosal is curved and blade-like, circular in cross-section anteriorly, becoming flattened posteriorly, where it articulates with the cephalic condyle of the quadrate and the braincase. Supratemporals are flattened, reniform and fused to the paroccipital processes. The supraoccipital has a strong sagittal crest that extends posteriorly to contact the ventral surface of the medio-posterior process of the parietal. The prootic bears an extended alar process and a well-developed, rhomboid christa prootica, and a very weak supratrigeminal process. Basipterygoid processes are well developed and flattened. The lower jaw possesses a large adductor fossa, a highly flattened and medially extended retroarticular process, a medially open meckelian canal and a dentary with a strong subdental shelf, 21 mandibular teeth, and nine dentary foramina. Postcranial osteology. The holotype possesses 25 presacral and 17 postsacral vertebrae with the last 21 mm (roughly 20 %) of the tail made up of regenerated cartilage. There are five cervical ribs, three sternal ribs, two xiphisternal ribs, five long asternal ribs with ossified costal cartilage, and eight short asternal ribs. The first three cervical ribs are distally flattened with bifid cartilaginous projections. Sacral vertebrae are fused asymmetrically, with the left diapophysis formed from the fused sacral ribs, while the right diapophysis is formed by the fusing of the second sacral and first caudal ribs. The next two pairs of caudal ribs are well developed and angled anteriorly, with subsequent ribs becoming increasingly smaller and posteriorly angled. Pelvic girdle is well developed and flattened. No iliac tubercle is present. There is a well-developed, ventrally angled pubic tubercle directly anterior to the obturator foramen. Both hypoischium and hyperischium are well developed. Pubic bones are well separated by triangular prepubic cartilage. The sternal plate is broad and lacks a fontanelle. Interclavicle cruciform, clavicles rod-like and flattened dorsally. The epicoracoid is narrow and curved, connecting the scapular ray to the primary and secondary coracoid rays, but not to the anterior process of the scapular. Limb bones well developed. Digits display the typical squamate phalangeal arrangement of 3–5 – 4 – 3 – 2. A large sesamoid is found in the palm of the hands. The fifth metatarsal possesses an elongated medial process at midbody. Osteoderms. Scales of the dorsal and temporal regions of the skull and the ventrolateral aspects of the jaws are underlain with rugose osteoderms. These osteoderms fuse to the proximal parietal, frontal and postorbital bones, although the mesokinetic and metakinetic joints appear unobstructed and flexible. The entire body is covered in robust osteoderms, with only the femoral pores and the axillary, inguinal and cloacal regions unarmoured. Large, rectangular, imbricate osteoderms protect the dorsal and lateral aspect of the trunk, becoming more keeled and mucronate laterally. Unkeeled, imbricate osteoderms cover the entire venter, from the gulars to the cloaca, rectangular on the throat and abdomen, becoming rhomboid around the sternum and cloaca. Several abdominal osteoderms appear to be fractured. Caudal osteoderms are thick, sharply spined and arranged in imbricate transverse whorls. Limbs covered by imbricate circular/rhomboid osteoderms, keeled and mucronate dorsally, plate-like ventrally. Colouration of holotype. Dorsal coloration is medially brown, blending to a straw-yellow laterally. Head yellow-brown; supra and infralabials yellowish; a dark-brown bar extends from the posterior aspect of the eye to the temporals. The base of the tail is medially brown, surrounded by a straw-yellow coloration that extends towards the tip and eventually dominates the tail; the dorsal aspect of the limbs is brown. Laterally a dark-brown line, similar to the eye-mask coloration, extends from the neck towards the insertion of the forelimbs. The venter is yellowish-white, cream on the tail; the underside of the hind limbs, cloacal region and tail are cream, grading into yellowish on the anterior half of the tail. Variation. Variation in scalation and body measurements of C. namakuiyus sp. nov. are reported in Tables 2 and 3. The sole male paratype CAS 254755 has 12 generation glands on the ventral surface of each thigh. One female paratype (CAS 254754) differs from the holotype in having the loreal and preocular fused. The medioposterior parietal process is slightly bifurcated distally in CAS 254754, CAS 265529 and PEM R 18005, and does not extend back beyond the posterolateral processes in CAS 25630. Five premaxillary foramina are present in all type specimens except CAS 254754 (two) and CAS 256531 (four). Specimens CAS 254755 and PEM R 18005 have 15 maxillary teeth, and CAS 256531 has 17. All paratypes have 21 mandibular teeth, except CAS 254754 (20), CAS 256530 (18), and CAS 254755 and CAS 256529 (16). The supratemporals are not fused to the paroccipital processes in CAS 254754, CAS 256529, and CAS 256530. Five of the paratypes have full, unautotomised tails: CAS 254755 has 29 postsacral vertebrae, CAS 25474, CAS 256529 each have 27, and the two fetuses, CAS 254913 and CAS 254914, have 25 and 26 respectively. One specimen, CAS 256531, has 26 presacral vertebrae. CAS 254755, CAS 256529 and CAS 256530 each possess four pairs of long asternal ribs, while CAS 256529 and PEM R 18005 each have seven short asternal ribs. None of the paratypes exhibit the asymmetrical sacral fusion of the holotype, but rather display the normal condition of having the sacral diapophyses formed from the sacral ribs. The hyperischium of CAS 254755 extends anteriorly to connect with the prepubic cartilage. Etymology. The specific epithet “ namakuiyus ” is the masculine latinised form of namakuiya, which means “thorny” in the Herero language, referring to the sharp spines on the limbs and tail of this species. Suggested common name: Kaokoveld Girdled Lizard. Distribution. Currently, the new species is known only from the south and central parts of Namibe Province, Angola, southwest of the Leba escarpment, at elevations of 215–509 m a.s.l. Localities. Angola: AMNH R 47274 –47321. Namibe Province, outskirts of Caraculo: CAS 254912 – 14 [15 °0' 59.4 "S, 12 ° 38 ' 31.3 "E; 503 m elevation], CAS 256529 [15 °0' 57.3 "S, 12 ° 38 ' 32.6 "E; 509 m elevation]; outskirts of Pico Azevedo: CAS 254754 –55, 256530, 256531) [15 ° 28 ' 33.2 "S, 12 ° 27 ' 45.7 "E; 421 m elevation]; Road from Namibe to Omahua lodge PEM R 18005 [15 ° 59 ' 48.5 "S, 12 ° 24 ' 24.6 "E; 308 m elevation]; 49 km ESE of Tombua: TM 40430 (15 ° 53 'S, 12 ° 16 'E; 215 m elevation). Habitat and natural history notes. This species is found in gently sloping crevices of granite outcrops in the arid Kaokoveld. When approached, specimens retreat into their fissure as far as possible, wedging themselves head-first and protecting their head and flanks with their spiny tails. As with all other cordylines, C. namakuiyus sp. nov. is viviparous; the holotype contained two large fetuses (SVL 49.5 % of the holotype). The fetuses appear to be approaching full-term, suggesting a spring/early summer parturition period. Although these fetuses lack any body osteoderms, the slightly larger neonates (SVL 55.6–56.4 % of the holotype) possess significant dermal ossification, suggesting that the young rapidly accumulate osteodermal armour following parturition. One of the Pico Azevedo specimens was found occupying the same rock crack as an adult Chondrodactylus pulitzerae (Schmidt), and PEM R 18005 from Iona National Park inhabited the same crack as an adult Chondrodactylus fitzsimonsi (Loveridge).