Nucras broadleyi, Branch & Conradie & Pinto & Tolley, 2019

Branch, William R., Conradie, Werner, Pinto, Pedro Vaz & Tolley, Krystal A., 2019, Another Angolan Namib endemic species: a new Nucras Gray, 1838 (Squamata: Lacertidae) from south-western Angola, Amphibian & Reptile Conservation 13 (2), pp. 82-95 : 86-91

Nucras broadleyi sp. nov.

Angolan Sandveld Lizard

urn:lsid:zoobank.org:act:C82E3A75-96FF-4D2A-9B52-3ABF4B58BC2B

( Figures 4–6 )

Chersonymy. Nucras tessellata var. taeniolata ( Bocage 1895: 30) , Nucras tessellata var. taeniolata ( Boulenger 1910: 474) , Nucras tessellata var. holubi ( Boulenger 1917: 210) , Nucras intertexta var. holubi ( Boulenger 1920: 20) , Nucras tessellata ( Monard 1937: 73; Laurent 1964: 56), Nucras ornata ( Broadley 1965: 23) , Nucras tessellata ( Broadley 1972: 30; Ceríaco et al. 2016: 56; Burger 2014: 171), Nucras aff. tessellata ( Marques et al. 2018: 221; Branch et al. 2019: 317).

Type material. The type series is comprised of the three most recently collected specimens, which are housed in PEM and TM.

Holotype. A subadult male ( PEM R 24005, AG 018 ), 10 km west of Lola , edge of Bentiaba River valley, Namibe Province, Angola (-14.29028, 13.53056, WGS 84, 802 m asl). Collected by W. R. Branch, P. Vaz Pinto, and J.S. de Almeida on 2 November 2015.

Paratypes (2). a) A subadult female ( PEM R 24157, AG 166 ), 8.8 km southwest of Farm Mucungo , Namibe Province, Angola (-14.80167, 12.41917, WGS 84, 385 m asl). Collected by W. R. Branch, P. Vaz Pinto, and J. S. de Almeida on 8 November 2015 . b) An unsexed adult ( TM 40392 ), “ 34 km S of Moçâmedes to Porto Alexandre , Angola, 1512 Ca ” (= 34 km S Namibe to Tômbwa), Namibe Province, Angola (approx. -15.48220, 12.18289). Collected by W.D. Haacke on 30 March 1971 .

Additional referred material: The following additional material was used to expand the description of variation within the species: a) an adult male (MD 1967, Laurent 1964), “km 34 de la route de Moçâmedes à Sa da Bandeira” (= 34 km from Namibe on Lubango road , -15.03333, 12.41667), collected 24 October 1949 , b) MBL 646 , 647 a, 647b ( Bocage 1895: 30) from Maconjo (approx. -15.01667, 13.20000), c ) BM 1970.6.29.10–11 ( Boulenger 1910: 474) from Ponang Kuma (= Donguena, approx. -17.01667, 14.71667), and d ) MHNC 91.0524 ( Monard 1937) from Capelongo (approx. -14.88333, 15.083333), collected April 1933 .

Etymology. The specific epithet is a patronym in honor of Donald G. Broadley for his numerous contributions to the herpetofauna of Africa. Don (as most of us knew him) was the first to recognize the Angolan population as a separate species ( Broadley 1972). The name is constructed in the masculine genitive.

Diagnosis. Assigned to Nucras due to a well-defined collar (absent in Ichnotropis ), toes not serrated or fringed (versus serrated or fringed in Meroles ), subdigital lamellae smooth (versus keeled in Pedioplanis and Heliobolus ), subocular bordering lip, the nostril is pierced between two nasals, nasal well separated from upper labial, and dorsal scales small, smooth, and juxtaposed.

The new species can be diagnosed from other Nucras species based on a combination of the following characters: series of transversely enlarged plates present under forearm (versus absent or only feebly enlarged in Nucras lalandii ), a small series (0–6) of small granules present between supraciliaries and supraoculars (versus mostly absent in N. boulengeri and N. lalandii ), 23–29 lamellae under 4 th toe (versus less than 22 in N. lalandii ), dorsum with a series of longitudinal pale stripes (versus dark cross bands present in N. lalandii and N. scalaris or a series of pale vertebral spots, sometimes forming irregular transverse bands in N. intertexta or lack of any dorsal patterns in N. aurantiaca ), four pale stripes on nape with outer stripes forming a.

Bloemfontein

Museum Province Province Province Province Province Province Province Province Province National Cape Cape Cape Cape Cape Cape Cape

NMB

–

,

, Northern Western, Northern,, Northern Northern, Western,, Western Kamanjab,

KwaZulu-Natal Cape

, Western

Museum Locality South Africa Africa South South Africa South Africa South Africa GenBank Africa South South Africa Namibia, South Africa Africa South Elizabeth Port

–

1 005257 951543 951540 951542 951541 632229 871208 547691 871206 871207 547718

RAG PEM HG HF HF HF HF EF DQ HF DQ DQ HF,

Science 4 005232 951535 951536 751398 547725 547732 547733 547759 of ND HG — HF — HF — FR HF HF HF HF Academy 005211 951559 951556 951558 951557 080358 871152 547772 871148 871149 547800 California S 16 HG HF HF HF HF AF DQ HF DQ DQ HF

CAS

–

: number 206725 R 18745 R 16873 R 11574 R 16872 209602 R 18376

abbreviations Museum accession CAS PEM PEM NMB PEM NA NA NMNW CAS PEM Museum. accession 5584 08 069 - 20650 21061 20687 0531

08

6001 6067 049

analysis Field ID

AMB KTH MB MB MB MH GW AMB AMB SVN genetic

in

Samples

used

Species tessellata tessellata tessellata tessellata tessellata longicaudata australis australis capensis capensis suborbitalis.

)

continued

2

Table

(Genus Nucras Nucras Nucras Nucras Nucras Outgroup Latastia Australolacerta Australolacerta Ichnotropis Ichnotropis Meroles

continuous light stripe with the outer edges of the parietals (similar to Broadley’s (1972) N. tessellata tessellata var. “T;” differs from N. livida and N. tessellata where the outer stripes often do not form a continuous light stripe with the outer edges of the parietals; differs from N. caeiscaudata and N. ornata where there are only three longitudinal stripes present on nape and sometimes the vertebral ones are absent), well defined occipital scale separating parietals (versus reduced or absent in northern Namibia N. holubi , which is referred to as N. intertexta damarana Parker ; as well as absent in N. caesicaudata ), parietal foramen absent (often present in all other species except N. taeniolata ), and postnasals separated (usually fused in N. taeniolata ).

In the phylogenetic analysis, the uncorrected p -distances show that this clade differs by>8% for 16S,>14% for ND4, and>1% for RAG1 sequence divergence from other members of the N. tessellata clade.

Description of Holotype ( Fig. 4 ). Body relatively slender (SVL approx. 4.5 times the head length, tail truncated), with hindlimbs larger than forelimbs (femur of hind limb equal to length of tibia); head narrow and elongated (56% longer than wide) with narrow pointed but blunt snout, that is slightly longer than distance from back of eye to rear of ear opening. Rostrum protruding and visible from below. Nasals paired and in contact (0.2 mm suture length), not swollen, nostril directed backwards separating postnasals. Frontonasal single, wider than long (1.1 × 1.8 mm). Prefrontals paired and in broad median contact with one another (0.6 mm suture length), wider than long (1.1 × 1.2 mm). Frontal entire, longer than wide (2.7 × 1.9 mm). Two large rounded supraoculars, both in contact with the frontal, with anterior supraocular preceded by a single large scale in contact with prefrontal, frontonasal, and posterior loreal, with posterior supraocular bordered by a single large scale in contact with parietal and frontoparietal. Paired frontoparietal in broad contact (1.3 mm suture length), nearly as wide as long (1.7 × 1.5 mm). Parietals twice as long as wide (3.1 × 1.8 mm), fully separate by a large, pentagonal interparietal (2.5 × 1.2 mm) that is twice as long as wide, slightly shorter than frontoparietals and nearly equal to length of frontonasal and prefrontal combined. Small subtriangular occipital (0.5 × 0.7 mm). Two loreals, second much larger than first. Six supraciliaries on each side, 1 st is the longest. A single minute granule scale between supraocular and supracilliares on right side, none on left side. Four supralabials anterior to subocular and three supralabials posterior to subocular, on both sides. Subocular slightly elevated medial and bordering the lip, its lower border being shorter than the upper. Three temporal scales, first longer than others, smooth. Tympanic shield as wide as long, border of ear opening. No ear lobes. Lower eyelid with transparent brille formed by five larger scales, surrounded by numerous smaller scales. Lower eyelid separated from subocular and enlarged temporal scales by a series of 10 smaller scales. Small scale above 3 rd supralabial separating the posterior loreal and subocular.

Enlarged scale bordering 1 st post subocular, supralabial, and the subocular. Six infralabials on both sides, with 3 rd being longest; four enlarged pairs of chin shields, last largest and first three in broad contact. Twenty-four gular scales in a straight line between symphysis of chin shields and median collar plate, equal in size except last 4–5 larger. Collar free, comprising seven enlarged plates (median subtriangular) and extending slightly onto side of neck as a crease, bordered by 2–3 smaller scales. Dorsal scales small, juxtaposed, granular, smooth, larger on sides toward ventrals. Midbody scales 42. Ventral plates eight longitudinal and 28 transverse rows (from collar to groin), plates of the innermost rows longer than broad, with outer row notably smaller than other rows, transverse row of ventrals across chest just behind collar longer than broad; preanal scales irregular, median ones larger. Scales on upper surface of forearm large, smooth or slightly keeled. Scales on lower surface of forearm with eight enlarged plates, at least twice the width of scales on upper forearm. Scales on upper surface of tibia rhombic, subimbricate, smooth, and much larger than dorsal scales. Tibia below with a series of large plates. Subdigital lamellae under fourth toe 23R/ 25L. Femoral pores 13R/ 15L. Dorsal scales on tail oblique, strongly keeled diagonally, and truncate behind, ventral scales on tail obtusely keeled.

Coloration. Dorsum with eight pale cream to white dorsolateral longitudinal stripes, separated by dark brown to black stripes. These stripes are more boldly patterned anteriorly, fading posteriorly. No light vertebral stripe. The two pale paravertebral stripes are separated by a very narrow strip of darker scales that starts on the interparietal through the occipital scale and fades posteriorly onto body and tail. The dorsolateral stripe extending along outer borders of parietals continues onto the tail. It is followed by the upper lateral stripe extending from posterior of the eye onto the head through the mid-temporal with a brief break above the ear opening, and continues onto the tail. The lower lateral stripe starts at the subocular, through the

ear opening, broken briefly above the arm, after which it continues all the way onto the tail. Ventrum white and lower limbs oblique white. Fore limbs upper surface black with scattered pale blotches. Hind limbs light brown with pale blotches. Upper surface of tail red-brown, similar to hind limbs. Scales bordering the orbit are black edged.

Variation ( Figs. 5–6 ). Meristic and escalation data are summarized in Table 1. The largest specimen examined is (BM 1907.6.29.10) 74 + 144 mm (tail regenerated). Regarding coloration, there seem to be three main variations among material examined: 1) 8–9 longitudinal stripes as in holotype (in PEM R24005, MBL 647a, 647b, MHNC 91.0524–5), 2) 4–5 pale longitudinal stripes broken up posteriorly with flanks spotted (in BM 1970.6.29.10–11, TM 40392, MD 1967), and 3) broken paravertebral stripes, continuous dorsolateral line and barred flanks (in PEM R24157), similar to N. intertexta .

Distribution. Found only in semi-arid south-western Angola, throughout much of Namibe Province and extending onto the escarpment of southern Hu í la and Cunene Provinces ( Fig. 1 ). Known localities include: Maconjo ( Bocage 1895: 30), Ponang Kuma (=Donguena) ( Boulenger 1910: 472), 34 km from Namibe on Lubango road ( Laurent 1964: 56), 34 km south of Tombwa (TM 40397), 8.8 km southwest of Farm Mucungo (this study), 10 km west of Lola (this study), and Capelongo ( Monard 1937: 73). The locality of Caconda ( Bocage 1895) extends the species distribution further north into Huíla Province, but the specimens could not be critically evaluated by Broadley (1972) and are now presumably lost.

Habitat. The species appears to be associated with mopane woodlands, dry savannas, and semi-desert shrublands ( Barbosa 1970). The new material was found in sandy plains with scattered low granite outcrops, with varying degrees of short grass cover and scattered bushes. Vegetation included Colophospermum mopane , Ficus sp., Senegalia (=Acacia) mellifera, Commiphora sp., Boscia foetida , and Salvadora persica . The confirmed historical records were also obtained within the dry woodland zone, even though the possible occurrence of the species in Caconda would place the species above 1,500 m asl and well into the mesic conditions of Brachystegia habitats ( Barbosa 1970).

Figures

Fig. 4. Nucras broadleyi sp. nov. A – holotype, adult male, PEM R24005 (AG 18) in life; B – general habitat photo of type locality, 10 km west of Lola, edge of Bentiaba River valley, Namibe Province, Angola; C – lateral close-up of head of holotype; D – dorsal close-up of head of holotype; E – ventral close-up of head of holotype (Photos: Bill Branch).

Fig. 5. Nucras broadleyi sp. nov. A – paratype, adult female, PEM R24157 (AG 166) dorsal view; B – ventral view; C – dorsal close-up of head of paratype; D – ventral close-up of head of paratype; E – lateral close-up of head of paratype; F – general habitat photo of type locality, 8.8 km southwest of Farm Mucongo, Namibe Province, Angola (Photos: Bill Branch).

Fig. 6. Variation of Nucras broadleyi nov. sp. dorsal color pattern.A – TM 40392 from “34 km S of Moçâmedes to Porto Alexandre;” B – BM 1970.6.29.10 from Ponang Kuma (=Donquena); C – MHNC 91.0524 from Capelongo; D – MD 1967 from “km 34 de la route de Moçâmedes à Sa da Bandeira” (Photos: A,B – Bill Branch, C, D – Luis Ceríaco).

Fig.1. Map showing distribution of Nucras broadleyi sp. nov. in Angola.