Cordylus momboloensis, Bates & Lobón-Rovira & Stanley & Branch & Vaz Pinto, 2023

Bates, Michael F., Lobon-Rovira, Javier, Stanley, Edward L., Branch, William R. & Vaz Pinto, Pedro, 2023, A new species of green-eyed Cordylus Laurenti, 1768 from the west-central highlands of Angola, and the rediscovery of Cordylus angolensis (Bocage, 1895) (Squamata: Cordylidae), Vertebrate Zoology 73, pp. 599-646 : 599

Cordylus momboloensis sp. nov.

Figs 4 , 5 , 6 , 7 , 8 , 9 , 11 , 13 , 14 , 15

Chresonymy.

Cordylus angolensis not Bocage, 1895: Broadley and Branch (2002: 10) partim; Branch et al. (2005: 137) partim; Broadley (2006: 1) partim; Stanley et al. (2011: 67) partim; Greenbaum et al. (2012: 32) partim: Mombolo; Reissig (2014: 33 + map) partim; Stanley et al. (2016: 202 + map) partim.

Holotype.

PEM R25217, an adult male from Sandula (Mom-bolo), Cuanza Sul Province, Angola (-12.1789°, 15.0292°; 1975 m a.s.l.) collected on 6 November 2016 by William R. Branch, Ninda Baptista and Pedro Vaz Pinto.

Paratypes.

Allotype: MNCN 50761, an adult gravid female from Sandula (Mombolo), Cuanza Sul Province, Angola (-12.1811°, 15.0316°; 2095 m a.s.l.) collected 29 May 2019 by Pedro Vaz Pinto. Other paratypes: AMNH R47333 (adult male), AMNH R47331 (adult female), AMNH R47332 (adult female), AMNH R47334 (adult female) and AMNH R47335 (subadult female), all from Mombolo, Angola (exact site unknown, but likely at the Chapman’s farm of Sandula, same co-ordinates as for the holotype) collected on 11 September 1925 by 'H. Chapman and A. Chapman’ during the Vernay Angola Expedition. MNCN 50760 (male), FKH 0127 (male), FKH 0128 (male), FKH 0125 (female), from Vondo (Uassamba Mountain), Cuanza Sul Province, Angola (-11.2415°, 14.6288°; 1989 m a.s.l) collected 3 February 2019 by Pedro Vaz Pinto.

Additional referred material.

TM 46476, an adult female from 3 km W of Condé, Cuanza Sul Province, Angola (about -10.8516°, 14.6387°; 1236 m a.s.l.) collected on 25 May 1974 by Wulf D. Haacke. FKH 0129, a juvenile from Vondo ( Uassamba Mountain ), Cuanza Sul Province, Angola (-11.2415°, 14.6288°; 1989 m a.s.l) collected 3 February 2019 by Pedro Vaz Pinto .

Diagnosis.

(Where variation in additional material falls outside the range of type material, this is indicated in parentheses.) A medium to large rupicolous Cordylus with a moderately depressed head and body. Referred to Cordylus (rather than any other species of Cordylidae ) by the following combination of characters: head distinct from body, two pairs of large and well developed limbs (body serpentiform, head indistinct from body, and limbs rudimentary in Chamaesaura ), scales on back large and keeled (granular in Platysaurus , partly granular in Pseudocordylus and Hemicordylus ), non-spinose occipitals [and post-occipitals] (occipitals spinose in Smaug ), 23-24 transverse dorsal scale rows (40-43 in Ninurta ; 31-46 in Karusasaurus ; 15-16 in Ouroborus ); loreal absent (present in Smaug , Ninurta , Ouroborus , Karusasaurus , and Namazonurus ).

Cordylus momboloensis sp. nov. is distinguishable from other members of its genus by the following combination of characters: (1) back dark brown with a paravertebral series of pale markings; (2) top of head with pale blotches; (3) iris of the eye mostly pale green (blue-green to some eyes), with a brown ring around the pupil; (4) scales of the first transverse row of dorsals similar in appearance to those of the row behind; (5) loreal shield absent; (6) nostril pierced in the posterior part of a large nasal, situated behind the suture of rostral and first supralabial, always well separated from the first supralabial, and usually separated from the preocular; (7) an interrupted row of non-spinose occipitals consisting of 2-3 scales on either side of head; (8) a row of 5-6 non-spinose post-occipitals, the median scales of which are in contact with the posterior parietals and separate the occipitals; (9) Frontonasal separated from the frontal by a pair of prefrontals, each of which exceeds it in size; (10) Anterior pair of parietals usually in contact anteriorly; (11) dorsolateral and lateral scales weakly to moderately spinose; (12) tail spinose, more weakly so distally; (13) dorsal scale rows transversely 22-24; (14) dorsal scale rows longitudinally 18-23; (15) ventral scale rows transversely 22-27; (16) ventral scale rows longitudinally 12-13 [rarely 11 or 14]; (17) subdigital lamellae on 4th toe 11-15; (18) femoral pores per thigh 5-8 in males, 4-6 in females [rarely 0; 7 on one side in a non-type specimen, TM 46476]; (19) differentiated femoral scales [generation glands] per thigh in males 25-37; (20) premaxillary teeth 9.

Its status as a new species is supported by monophyly with high levels of support from a suite of three mitochondrial and six nuclear markers (see above); and it differs from C. angolensis , the most similar species genetically and morphologically, by an uncorrelated ND2 p-distance of 9.22% (Table 3 ).

Comparisons with other Cordylus species.

The new species differs from all other Cordylus in having nine (rather than seven, rarely six) premaxillary teeth, and from most other congeners (except C. angolensis , C. ukingensis , C. macro-pholis , and C. vittifer [occasionally has a loreal]) in lacking a loreal. It differs from C. ukingensis and C. macropholis by virtue of its weakly keeled or smooth versus strongly keeled (even spinose in C. macropholis ) gulars, as well as by having its nostril pierced near the middle of the posterior part of the nasal scale (versus infero-posteriorly). Differs from C. tropidosternum by having smooth or weakly keeled versus keeled gulars, and having the nostril well separated from the first supralabial (not in contact or near-contact); from C. rhodesianus by having distinctly rugose versus finely rugose to smooth upper head shields, 22-24 versus 25-29 transverse rows of dorsals, and a straight versus curved sulcus dividing the posterior part of the nasal; from C. jonesii by having its nostril pierced halfway up the posterior edge of the nasal rather than towards the centre; and a straight versus curved sulcus dividing the posterior part of the nasal; and from C. marunguensis which has the nostril pierced centrally on the lower margin of the nasal. Distinguished from C. vittifer , C. machadoi , C. namakuiyus and C. phonolithos by always lacking a loreal scale, having most or all scales of the first transverse row of dorsals of similar length (rather than longer) than those of the next row, and by having a pair of paravertebral rows of pale greenish-cream spots or blotches versus a lack of these.

The new species is most similar to C. angolensis , but distinguished from it as follows: in live specimens the iris of the eye is largely pale green to blueish-green versus brown; top of head distinctly marked by pale blotches versus mostly plain brown; throat usually dirty white versus cream; a medially-interrupted row of occipital scales (2-3 on either side of head) is present, never forming a continuous row, each scale usually smaller than a post-occipital versus a continuous row of 6 occipitals across the back of the head; lower numbers of ventral scale rows longitudinally (usually 12-13 vs 14); relatively longer forearms (forearm/SVL 0.20-0.25 versus 0.15-0.19); higher numbers of generation glands in males (25-37 per thigh versus 19-25); and higher numbers of premaxillary teeth (nine versus seven).

Description of holotype.

PEM R25217 (Figs 4 - 6 , 11 , 13 , Tables 5 , 6 ). External morphology: Head and body moderately depressed. SVL 85.1 mm. Tail 77.5 mm (tip truncated and healed), 91.1% SVL. Head length 25.3 mm, 1.19 times as long as wide (21.4 mm), head depth 9.2 mm, 43.2% head width. Upper head shields coarsely rugose, not striated. Nasals in moderate contact, their posterior halves separated by the frontonasal, which is quadrangular and about half the size of a prefrontal; prefrontals in moderate contact with one another, and contacting the supranasals, preoculars and anterior supraoculars on either side; frontal in contact with first and second supraoculars, followed by a pair of frontoparietals in median contact; interparietal quadrangular, separating posterior one-third of anterior parietals, and anterior one-third of posterior parietals, the latter being distinctly larger than the anterior ones; occipitals two on one side of head, three on the other, interrupted by a slightly curved row of six post-occipitals; occipitals and post-occipitals rugose, occasionally striated but essentially non-spinose. Four supraoculars and three supraciliaries (postocular also in contact with posterior supraocular). Nasals large, with nostril pierced more-or-less centrally in the posterior part of the scale; nostril separated from preocular by a distance of about half its diameter, distinctly separated from first supralabial by a distance of more than half the height of the latter scale (i.e., at least half the greatest extent of the nostril); nasal divided posteriorly at the level of the middle of the nostril, and also divided below the nostril, forming a small postnasal. Loreal absent. Lower eyelid with 6-7 vertically enlarged septa; suboculars 4 left and 3 right, well separated from the lip. Rostral 2.3 times as broad as deep; supralabials on left side six (four anterior to median subocular), right side five (three anterior to median subocular); infralabials 6; sublabials 5. Mental 1.9 times as broad as long; gulars vary considerably in size and shape, from elongate, sometimes rectangular, on the sides of the throat, to squarish, oval and irregular, sometimes pentagonal or hexagonal, smooth and at most subimbricate, forming about 12 transverse rows (from first row in line with angle of jaws to last distinct row posterior to pair of sublabials), and 26 longitudinal rows between posterior angles of jaws; 5 chin shields (including a tiny elongate scale medially) in contact with 1st pair of sublabials.

Dorsal scales closely-set but often juxtaposed, rectangular, rugose, moderately keeled (less so on middle of back), some are also weakly spinose dorso-laterally, occasionally weakly serrated at their posterior edges; laterals occasionally subimbricate, often oval, rugose, sharply keeled and moderately spinose, no additional spines or serrations on the free ends of scales; dorsals (including laterals) in 22 transverse rows (excluding one half-row) and 23 longitudinal rows (including row of small vertebral scales which is interrupted at parts); on the central part of the belly the paired rows of mesial ventrals are rectangular (transversely), the next row on either side with only slightly transversely rectangular scales, the others being squarish, with two rows of lateral ventrals on either side consisting of longitudinally-rectangular scales; ventrals mostly smooth, but the outermost lateral row on either side, and an additional row of ‘pseudo-ventrals’ on either side, with some obtusely keeled scales; ventrals in 22 transverse and 12 longitudinal rows (excluding a row of oval, keeled scales on either side best considered ‘pseudo-ventrals’); a pair of enlarged pre-cloacal plates is followed anteriorly (before the ventrals) by 2-3 transverse rows of much smaller plates.

Scales on upper parts of hindlimbs are large, strongly keeled and strongly spinose; scales of forelimbs large, moderately keeled and moderately spinose; scales under fourth (longest) toe 13 on left foot, 14 on right; scales under fourth (longest) finger 10 on left hand, 11 on right; femoral pores 7 on each thigh, of moderate size with distinct plugs of yellowish secretion; differentiated glandular femoral scales on thighs 26 on left, 27 right. Tail whorled, dorsally with large, elongate, rugose, strongly keeled, spinose, weakly serrated scales, with spines directed backwards and longest superolaterally; supracaudals strongly keeled throughout most of their length, subcaudals basally distinctly keeled mainly on the distal half of tail. Scales on palms of hands obtusely keeled, on soles of feet moderately keeled; supradigital scales of hands weakly keeled, of feet moderately keeled; subdigital scales of hands and feet weakly keeled or smooth.

Colour of holotype (in life): The upper parts of the head, back, flanks, limbs and tail were mostly dark brown, with irregular pale or light greenish to blueish-green markings on either side of the midline (about six pairs) and on top of the head; venter whitish to cream; throat dirty white (Fig. 13 ). After preservation: As above, but back with indications of dark bands, and pale dorsal markings and venter have faded to a dirty white colour.

Variation in paratypes.

(n = 10). (Figs 4 - 6 , 8 - 9 , 14 ; Tables 5 , 6 ). Note: In AMNH R47333 the following characters could not be examined as the snout had been crushed/damaged: loreal, septa in eyelid, suboculars, infralabials and mental.

External morphology: All paratypes are similar to the holotype, but differ as follows: SVL 76.1-90.5 mm (males), 60.7-89.5 mm (females). Original tail shorter than SVL (87.5-97.6%, n = 9). Head 1.05-1.21 times as long as wide, head height 0.416-0.582 times head width. Nasals in narrow contact in FKH 0125, MNCN 50760 and MNCN 50761 (allotype; posterior three-quarters separated by frontonasal). Frontonasal tiny in AMNH R47333, slightly smaller than a prefrontal in FKH 0127, MNCN 50760 and MNCN 50761, about equal in size to a prefrontal in FKH 0125 and FKH 0129; prefrontals in broad contact in MNCN 50760 and FKH 0128, in narrow contact in MNCN 50761; interparietal irregular and separating anterior parietals in FKH 0125, kite-shaped (with most acute angle anteriorly) in MNCN 50760, completely separating posterior parietals in AMNH R47331, completely separating anterior parietals in MNCN 50760, and separating most of the anterior parietals in AMNH R47332 (the anterior part of the interparietal is followed, after a short gap, by an elongate lozenge-shaped scale which separates the anterior extremity of the anterior parietals and contacts the suture between frontoparietals); anterior and posterior parietals similar in size in AMNH R47331, 47333-4. Occipitals 2-3 on either side of head, interrupted medially by the row of post-occipitals; a slightly (AMNH R47335) to distinctly (AMNH R47332-4, MNCN 50760) curved row of six (five in FKH 0128) post-occipitals; nasal not divided below the nostril; a tiny elongate scale is present between the nasal, preocular and first supralabial in AMNH R47332 (nostrils damaged/reamed in AMNH R47334). A few occipitals with weak keels in FKH 0125. Nostril separated from preocular by more than half its diameter on left side of head and less than half on right (FKH 0125, MNCN 50760), about half on left and less than half on the right in FKH 0127-8, and less than half on both sides in FKH 0129; and nostril separated from first supralabial by a distance greater than half its height on left and equal on right (MNCN 50760), equal to half the distance on the left in FKH 0125, and exceeding the height of the supralabial on the left in FKH 0129. Lower half of preocular with a vertical suture medially in MNCN 50761 (allotype). Lower eyelid with 5-7 vertical septa; suboculars 3-4 (two on left side of head in FKH 0125, usual two posterior scales apparently fused). Rostral 2.21-2.60 times as broad as deep [AMNH only]; supralabials 5-6; infralabials 5-6, but 4 on right side of head of AMNH R47331. Anterior pair of sublabials of FKH 0125 completely separated by a moderate-sized elongate scale that contacts the mental; 4-5 chin shields (including small median scale) in contact with 1st pair of sublabials. Mental 1.52-1.75 times as broad as long [AMNH only]; gulars subimbricate to juxtaposed, irregular throughout in FKH 0125, FKH 0127, MNCN 50760 and MNCN 50761 (allotype), a few lateral gulars with obtuse and feeble median keels, but FKH 0125, FKH 0128 and FKH 0129 also have feebly keeled gulars centrally, in 16-22 longitudinal rows between posterior angles of jaws.

Dorsals plus laterals in 22-24 transverse rows and 18-22 longitudinal rows; ventrals in 22-27 transverse and 12-13 [11 in FKH 0128, 14 in AMNH R47333] longitudinal rows (an additional row of ‘pseudo-ventrals’ on either side between ventrals and laterals in AMNH R47331 and 47332 only).

Scales under fourth (longest) toe 11-15, under fourth (longest) finger 10-12 [AMNH only]; femoral pores per thigh 5-8 in males, 4-6 (0 in AMNH R47334) in females; differentiated glandular femoral scales in males only, 25-37 per thigh. In AMNH R47332 the basal subcaudals are obtusely keeled or smooth. Supradigital scales of hands smooth to moderately keeled, of feet smooth to strongly keeled; in AMNH R47331 subdigital scales of feet strongly keeled and occasionally spinose, especially posteriorly.

Colour of paratypes (in life): Colouration of the Sandula and Vondo paratypes was similar to that of the holotype. The belly had a slightly blueish tinge.

Colour of paratypes (in preservative): AMNH R47331 is the most distinctly marked of the old ‘Mombolo’ series (Fig. 14 ): The upper parts of the head, back, flanks, limbs and tail are medium to dark brown; there are five dark transverse bands across the back from nape to tail, separated by narrower bands of paler colour, including three pairs of large and distinct cream-coloured paravertebral blotches, together with a series of less distinct but similarly pale blotches dorso-laterally; and a few cream spots are present on the top of the head. In the other AMNH specimens the upper parts of the head, back, flanks, limbs and tail are medium to dark brown, but without distinct dark markings or pale spots (pattern may have faded). Venters of all AMNH specimens are cream to pale tan.

Cranial skeleton : (Fig. 8 ; segmented mesh files of paratype AMNH R47334 and TM 46476 can be found here (tinyurl.com/CangolensisAMNH) and here (tinyurl.com/CordylusAngola2). Morphosource links to tomogram stacks (https://doi.org/10.17602/M2/M530010 and https://doi.org/10.17602/M2/M530028). Consistent characters between both specimens are reported here, with any differences noted. The scales of the dorsal and temporal regions of the skull and the ventrolateral aspects of the jaws overlie rugose osteoderms. These osteoderms fuse to the proximal parietal, frontal and postorbital bones, although the mesokinetic and metakinetic joints appear unobstructed and flexible. Lateral maxilla and anterior aspect of the premaxilla lack osteoderms. The parietal is pentagonal, with five osteoderms that underlie the parietal shields fused to its dorsal surface, and tapers to a bifid medioposterior process, which articulates with the sagittal crest of the supraocciptial. Three large osteoderms are fused to the frontal, which is unpaired and clasped by the parietal at its posterolateral edge. The upper temporal fenestra is obscured anteriorly by a large osteoderm fused to the dorsal surface of the postorbital bone, posteriorly by two unfused rectagonal osteoderms that overlie the squamosal. Premaxilla is unpaired and contains nine pleurodont teeth, with a dorsal process that extends posteriorly to intersect the nasals (absent due to damage in AMNH R47334), which themselves overlie the frontal. The maxillae are typically scinciform, with a deeply grooved crista dentalis, a deep fossa that accommodates the lacrimal sac, and 20-21 (TM 46476) to 18-19 (AMNH R47334) teeth. Teeth display pleurodont attachment and are unicuspid, with a slight concave surface where they connect with the mandibular teeth. No palpebral is present but the prefrontals possess a small protuberance, forming a shelf that directly underlies the anteriormost superorbital osteoderm. 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. The lacrimal bone is small, flattened and oval, slightly bicuspid anteriorly, the two processes meeting prefrontal cusps to bracket the lacrimal duct. Pterygoids are edentate and 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 laterally flattened, ovoid and not fused with the elongate paraoccipital processes. The posterior aspect of the prootic in the smaller AMNH R47334 is not fully fused with the otooccipital, resulting in a deep groove along the dorsal aspect of the paraocciptal processes. Quadrates very broad with a pronounced ridge and concave region at the entire lateral edge in TM 46476 (right quadrate of AMNH R47334 partially damaged). The supraoccipital has a strong sagittal crest that extends posteriorly to contact the ventral surface of the medioposterior 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 that is closed posteriorly by a large splenial, and a dentary with a strong subdental shelf; 22 (AMNH R47334) and 25-26 (TM 46476) mandibular teeth, and 6-7 dentary foramina.

Postcranial skeleton : Segmented mesh files of AMNH R47333 and TM 46476 can be found here: tinyurl.com/CangAMNHSkel and here tinyurl.com/Cang2Skel. Mor-phosource links to tomogram stacks (https://doi.org/10.17602/M2/M15862 and https://doi.org/10.17602/M2/M529986). The axial Cordylus skeleton is comprised of 25 presacral vertebrae: eight cervical, three sternal, two xiphisternal, five long asternal ribs with ossified costal cartilage, then seven short asternal ribs (the ribs immediately anterior to the sacral vertebrae in AMNH R47333 are highly reduced), two sacral and 24 caudal vertebrae (incomplete, regenerated tail in AMNH R47333). Cervical ribs 4-6 are distally flattened and biphid, with the ventral processes more elongated. Pubis flattened and curved with a large, ventrally-angled pectineal tubercle. Pubic symphysis flattened and triangular, separating the pubes entirely. Hyperischiam and hypoischium well developed. Sternal plate broad, lacking a fontanelle. Interclavicle cruciform. Digits display a typical phalangeal pattern of 2-3-4-5-3 for the manus and 2-3-4-5-4 for the pes.

Osteoderms (Fig. 9 ): The dorsal trunk is covered in rectangular, dorsomedially keeled, imbricate osteoderms, each roughly three times longer than its width. The dorsal osteoderms are arranged in whorls, becoming more oval and better separated laterally. The nuchal osteoderms are spined posterior to the tympanic opening. Ventral osteoderms are absent in AMNH R47333, and delicate and plate-like in TM 46476, grading from rhomboid in the gular region to mildly imbricated and cycloid in the pectoral region, square and non-imbricate in the abdominal region, to pentagonal in the cloacal region. The forelimbs are covered in keeled, imbricate, rhomboid osteoderms, except for the axillary, antecubical and palmar regions, which are unarmoured. The hindlimbs are covered in rhomboid osteoderms, except for the ventral femoral, popliteal and plantar regions. Hindlimb osteoderms are unkeeled on the anterior thigh, become more spinose posteriorly and distally. The caudal osteoderms are large, robust and arranged in imbricated whorls, feebly keeled and mucronate along the dorsal and ventral aspects, becoming more heavily spined laterally.

Variation in additional material (n = 2).

(Figs 4 - 6 , 8 - 9 , 15 ; Tables 5 , 6 )

External morphology: TM 46476 (female). SVL 106.0 mm. Original tail 106.7% SVL. Head 1.25 times as long as wide, head height 0.469 times head width. Nasals in broad contact. Interparietal bulb-shaped with a long neck, completely separating the anterior parietals. Anterior and posterior parietals similar in size and shape. Nostril in contact with preocular, nasal scale apparently not divided, and no postnasal present. Lower half of preocular with a vertical suture medially, suggesting partial fusion with loreal. Lower eyelid with five vertically enlarged septa. Rostral 2.8 times as broad as deep. Supralabials 5 left, 7 right. Mental 1.49 times as long as broad. Gulars mostly rectangular and obtusely and feebly keeled, imbricate to subimbricate, forming nine transverse rows (from first row in line with angle of jaws to last distinct row posterior to pair of sublabials), and 23 longitudinal rows between posterior angles of jaws. Four chin shields in contact with first pair of sublabials. Dorsals in 24 transverse and 20 longitudinal rows. Ventrals in 22 transverse and 12 (plus one row of ‘pseudo-ventrals’ on either side) longitudinal rows. Pair of cloacal plates preceded by two transverse rows of much smaller plates. Scales under fourth toe 13 (both sides), under fourth finger 12 (both sides). Femoral pores 7 left, 5 right; no differentiated femoral scales. Subdigital scales of feet strongly keeled and spinose, especially posteriorly.

FKH 0129 (juvenile). Data for head only, see Figs 4 - 6 and Table 5 . Supra-labials 6. Infralabials 7 left, 6 right. Suboculars 3. Occipi-tals 2 on either side of head.

Colour of additional material (in life). TM 46476: Upperparts medium brown with a paired series of irregular cream paravertebral blotches from nape to base of tail; top of head with a few small, scattered, cream markings. FKH 0129 was similar to the type series.

Size.

Largest male (MNCN 50760, Vondo): SVL 90.5 mm, tail length 88.4 mm (original), total length 178.9 mm, head length 23.1 mm, head width 21.3 mm, head height 11.9 mm. Largest female (MNCN 50761 allotype, Vondo): SVL 89.5 mm, tail length 85.4 mm (original), total length 174.9 mm, HL 22.3 mm, HW 20.5 mm, HD 11.8 mm. A non-type female (TM 46476, 3 km W of Condé), referred to this species, measures SVL 106.0, mm, tail length 113 mm (original), total length 219 mm, HL 26.8 mm, HW 22.2 mm, HD 12.7.

Etymology.

The name ‘momboloensis’ derives from the local name ( “m’bolo”) for Mombolo which in Umbundo - main language in central Angola - is applied to bread or staple food. It is a reference to the fact that the region in question consists of a fertile and relatively large highland plateau (surrounded by mountains, in Luanza Sul Province). Most of the available specimens, including the holotype as well as the first series of specimens of this species collected during the Vernay Angola Expedion in 1925, were collected in Mombolo.

Habitat.

Although no specific habitat information is available for the material housed at the AMNH, it is likely that it may have been obtained at the same site as the recent material from Sandula. At both the Sandula and Vondo localities, the species proved to be rupiculous and specimens were found sheltering inside cracks and crevices in rocky outcrops (igneous rocks in Sandula and granite in Vondo) present in fairly steep slopes above 1900 m a.s.l. (Fig. 11D ). The surrounding habitat at Sandula and Vondo is montane grassland. The specimen from Condé was reportedly found in a rock outcrop surrounded by grassland and sparse miombo savannah (see Fig. 15B ).

Distribution.

This species was collected at Sandula (Monte Verde) and Vondo (Mount Uassamba), both in Cuanza Sul Province on high mountain peaks at 1975-2095 m a.s.l., and an individual was briefly observed at Morro do Pundo (-12.4464°, 13.9171°) in Benguela Province, at approximately 1000 m elevation near the town of Bocoio (Fig. 12 ). The Bocoio specimen quickly disappeared into a deep crevice and only a tail tip was secured. The latter was analysed for 16S and the specimen was referred to this species. The two peaks at Sandula and Vondo are situated about 110 km apart in highlands that include several mountain chains that have never been surveyed before, suggesting that the species is widespread on the west-central Angolan plateau in suitable mountain habitat, while the specimen from 3 km W of Condé places the species further north at elevations of about 1236 m a.s.l. in miombo woodlands (Fig. 12 ).

The AMNH material was collected by 'H. Chapman and A. Chapman’ in 1925 and simply labeled as ‘Mombolo’. This location, however, is rather vague as the region of Mombolo covers about 1000 km2, and included three farms belonging to the Chapman family. A very detailed discussion on the problems in relation to another rupiculous lizard, Afroedura bogerti , collected under identical circumstances ( Loveridge 1944b), is provided by Branch et al. (2017). It can be noted that Hill and Carter (1941) stated that "Boulton and Charles Chapman went to Namba in the Mombolo region." Two of Chapman’s farms were located on the separate mountain chains of Namba and Monte Verde (Sandula), suggesting that the geographic location could be confidently narrowed down to those two sites. However, several recent surveys conducted on both mountains have so far resulted in Cordylus being found only at Sandula, suggesting that this was the original locality where the Chapman specimens were collected.

Conservation.

Little is known about the biology of this species, but habitat at the four sites where it has been found appears to be relatively undisturbed. Given the current knowledge, if an assessment was to be done, it is likely to result in a classification of Least Concern based on IUCN (2022a) Standards and Petitions Committee criteria.

Figures

Figure 4. Dorsal views of the head in Angolan Cordylus. Cordylus angolensis - A CHL 611, B CHL 615, C MNCN 50649, D CHL 613, E MNCN 50648 (neotype); C. momboloensis sp. nov. - F MNCN 50760 (paratype), G FKH 0125 (paratype), H FKH 0127 (paratype), I FKH 0128 (paratype), J FKH 0129, K PEM R 25217 (holotype), L MNCN 50761 (allotype); C. namakuiyus - M PEM R 25218, N CHL 616, O CHL 609; C. phonolithos - P CHL 852, Q CHL 1056, R MNCN 50651, S CHL 851, T CHL 1057, U MNCN 50652, V CHL 848; and C. machadoi - W CHL 701, X CHL 700, Y PEM R 19784. Photos: Pedro Vaz Pinto (A-N, P-V), Timoteo Julio (O, W-X), and William R. Branch (Y).

Figure 5. Lateral views of the head in Angolan Cordylus. Cordylus angolensis - A CHL 611, B CHL 615, C MNCN 50649, D CHL 613, E MNCN 50648 (neotype); C. momboloensis sp. nov. - F MNCN 50760 (paratype), G FKH 0125 (paratype), H FKH 0127 (paratype), I FKH 0128 (paratype), J FKH 0129, K PEM R 25217 (holotype), L MNCN 50761 (allotype); C. namakuiyus - M PEM R 25218, N CHL 616, O CHL 609; C. phonolithos - P CHL 852, Q CHL 1056, R MNCN 50651, S CHL 851, T CHL 1057, U MNCN 50652, V CHL 848; and C. machadoi - W CHL 701, X CHL 700, Y PEM R 19784. Photos: Pedro Vaz Pinto (A-N, P-V), Timoteo Julio (O, W-X), and William R. Branch (Y).

Figure 6. Ventral views of the head in Angolan Cordylus. Cordylus angolensis - A CHL 611, B CHL 615, C MNCN 50649, D CHL 613, E MNCN 50648 (neotype); C. momboloensis sp. nov. - F MNCN 50760 (paratype), G FKH 0125 (paratype), H FKH 0127 (paratype), I FKH 0128 (paratype), J FKH 0129, K PEM R 25217 (holotype), L MNCN 50761 (allotype); C. namakuiyus - M PEM R 25218, N CHL 616, O CHL 609; C. phonolithos - P CHL 852, Q CHL 1056, R MNCN 50651, S CHL 851, T CHL 1057, U MNCN 50652, V CHL 848; and C. machadoi - W CHL 701, X CHL 700, Y PEM R 19784. Photos: Pedro Vaz Pinto (A-N, P-V), Timoteo Julio (O, W-X), and William R. Branch (Y).

Figure 7. Arrangement of the nasal shield, nostril and scales of the snout in Cordylus: A C. angolensis (CHL 615 - Taqueta Mountain, Angola); B C. momboloensis sp. nov. (AMNH R 47333 - ' Mombolo', Angola); C C. machadoi (TM 40133 - 1.4 km NE of Humbia, Angola); D C. namakuiyus (PEM R 25218 - 10.4 km S of Rio Mucungo on tar road to Bentiaba, Namibe Province, Angola); E C. phonolithos (CAS 263581, holotype - vicinity of N'Dolondolo, Serra da Neve, Angola); F C. vittifer (NMB R 8721 - Farm Goede Hoop 182, Mpumalanga Province, South Africa); G C. tropidosternum (PEM R 13288 - Moebase village, Zambesiaca Province, Mozambique); H C. jonesii (NMB R 8656 - Farm Doorenwaard 313, Limpopo Province, South Africa); I C. rhodesianus (NMB R 10034 - York Forest Land, Zimbabwe); J C. cordylus (NMB R 9525 - De Hoop Nature Reserve, Western Cape Province, South Africa). Photos: Javier Lobon-Rovira (A), Michael F. Bates (B, C, F, H, I, J), William R. Branch (D, G), and Luis M. P. Ceriaco (E).

Figure 8. Computed tomography images depicting the cranial osteology of the five species of Cordylus from Angola.

Figure 9. Computed tomography generated images showing the osteology of the five Angolan Cordylus species. Osteoderms are highlighted in blue-green.

Figure 11. Cordylus angolensis: live neotype (A) and habitat at the collecting site of Taqueta Mountain, Angola (B); and C. momboloensis sp. nov.: live holotype (C) and habitat at the type locality of Sandula, Angola (D). Photos: Pedro Vaz Pinto (A, B), and William R. Branch (C, D).

Figure 12. Geographical distribution of Cordylus angolensis (dark blue) and C. momboloensis sp. nov. (green) in Angola. The distribution of the three species of the C. machadoi group (C. machadoi [red], C. namakuiyus [pale blue], C. phonolithos [yellow]) is also indicated. Type localities are indicated by stars; for C. angolensis, the original type locality of Caconda is indicated by a black star, while the neotype locality is indicated by a dark blue star. Locality records are from the present study (see species accounts and Appendix 1) as well as Bocage (1895), Laurent (1964), Stanley et al. (2016), Marques et al. (2019), and Lobon-Rovira et al. (2022 b).

Figure 13. Holotype (PEM R 25217) of Cordylus momboloensis sp. nov. from Sandula (Mombolo), Cuanza Sul Province, Angola. A dorsal and ventral aspects, B lateral (right), dorsal and ventral aspects of the head, C underside of the foot, D femoral region of the legs. Although not evident because of the way the holotype was photographed, its tail is intact. Photos: Pedro Vaz Pinto.

Figure 14. Paratypes of Cordylus momboloensis sp. nov. from ' Mombolo', Angola (from left: AMNH R 47333, R 47332, R 47331). Photo: Wulf D. Haacke.

Figure 15. A, C Adult female Cordylus momboloensis sp. nov. (TM 46476) from 3 km W of Conde, Cuanza Sul Province, Angola. B Habitat shown is in the vicinity of Conde. Photos: Wulf D. Haacke (A, C), and William R. Branch (B).