Sunday, July 16, 2023

[Herpetology • 2023] Cordylus momboloensis • A New Species of Green-eyed Cordylus Laurenti, 1768 (Squamata: Cordylidae) from the west-central Highlands of Angola, and the Rediscovery of Cordylus angolensis (Bocage, 1895)

 

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


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

Keywords: Africa, Cordyliformes, distribution, Sauria, taxonomy

Cordylus angolensis: live neotype (A) and habitat at the collecting site of Taqueta Mountain, Angola (B);
and Cordylus 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).

Holotype (PEM R25217) 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.

Cordylus momboloensis sp. nov.
 Mombolo girdled lizard; Lagarto-espinhoso do Mombolo

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).

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);
Cordylus momboloensis sp. nov. —F MNCN 50760 (paratype), G FKH 0125 (paratype), H FKH 0127 (paratype), I FKH 0128 (paratype), J FKH 0129, K PEM R25217 (holotype), L MNCN 50761 (allotype);
C. namakuiyus—M PEM R25218, 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 R19784.
Photos: Pedro Vaz Pinto (A–N, P–V), Timóteo Júlio (O, W–X), and William R. Branch (Y).

   

Live specimens of various species of Cordylus:
 A C. machadoi (PEM R25218 – 6 km north of Humpata, Huíla Province, Angola; photo: William R. Branch); B C. namakuiyus (PEM R18005 – between Namibe and Omahua lodge, Namibe Province, Angola; photo: William R. Branch); C C. phonolithos (CAS 263581, holotype – vicinity of N’Dolondolo, Serra da Neve, Namibe Province, Angola; photo: Luis M. P. Ceríaco); D C. rhodesianus (near Chimoio, Mozambique; photo: David Maguire);
E C. tropidosternum (Kayalekera, Malawi; photo: Luke Verburgt); F C. jonesii (Umbabat Private Nature Reserve, Mpumalanga Province, South Africa; photo: Darren Pietersen); G C. vittifer (NMB R8523 – Farm: Dipka 220, Vrede district, Free State Province, South Africa; photo: Michael F. Bates); H C. cordylus (NMB R8540 – Ha Sehlaba, Lesotho; photo: Michael F. Bates). Representatives of all four Cordylus clades recovered by the current genetic analysis are included.
 

Michael F. Bates, Javier Lobón-Rovira, Edward L. Stanley, William R. Branch and Pedro Vaz Pinto. 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: 599-646. DOI: 10.3897/vz.73.e95639