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Heliobolus crawfordi MARQUES, CERÍACO, HEINICKE, CHEBOURI, CONRADIE, TOLLEY & BAUER, 2022

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Higher TaxaLacertidae, Eremiadinae, Sauria, Lacertoidea, Squamata (lizards)
Subspecies 
Common NamesE: Crawford-Cabral’s bushveld lizard
Portuguese: lagartixa de Crawford-Cabral 
SynonymHeliobolus crawfordi MARQUES, CERÍACO, HEINICKE, CHEBOURI, CONRADIE, TOLLEY & BAUER 2022
Eremias lugubris — BOCAGE 1867, 1895 [part]
Eremias lugubris — BOULENGER 1921 [part]
Eremias lugubris — MONARD 1937 [part]
Eremias lugubris — MERTENS 1938 [part]
Lamperemias lugubris — SZCZERBAK 1975 [part]
Heliobolus lugubris — BRANCH 1998 [part]
Heliobolus lugubris — MARQUES et al. 2018 [part]
Heliobolus lugubris — BRANCH et al. 2019 [part] 
DistributionAngola (central coastal regions).

Type locality: N’Dolondolo, Namibe Province, Angola,
(-13.8133°S, 13.1362°E), 681 m elevation.  
Reproduction 
TypesHolotype: CAS 266269, adult male, collected by Luis M.P. Ceríaco, Suzana A. Bandeira and Ishan Agarwal, on 21 November 2016.
Paratypes (n = 16): AMNH-R 41567, 41570, 41587; CAS 266267, 266268, 266271, 266273, 266275; PEM R21625, 22058, 22059, 24004, 24019, 24024, 24025, 24168. 
DiagnosisDiagnosis: Medium-sized lizard, identified to genus by the following combination of characters: well-developed limbs, slender body, elongated snout, long tail, and a distinct collar on ventral region of neck. Heliobolus crawfordi sp. nov. can be distinguished from other members of its genus by the following combination of characteristics: (1) slender body of medium-size, mean SVL 52.2 mm; (2) long-tailed (mean 121.9 mm), tail more than twice body length; (3) midbody scale rows 70–90 (mean: 78); (4) low number of subdigital lamellae under the fourth toe (mean: 22); (5) parietals usually separated; (6) cranial shields not ornamented and temporal shield smooth; (7) lateral dark marking extending through the ear to the posterior margin of the eye; (8) presence of bright yellow dots ventrolaterally.

Comparisons: “Heliobolus crawfordi is distinguished from H. neumanni by possessing a higher number of midbody scale rows (70–90 vs. 40–42). Heliobolus crawfordi sp. nov. is distinguished from H. nitidus by possessing a higher number of midbody scale rows (70–90 vs. 52–64) and by color pattern (background light brown to orange-brown above versus background greenish, especially on the flanks). Heliobolus crawfordi sp. nov. is distinguished from H. spekii by having the cranial shields not ornamented and temporal shield smooth (versus cranial shields ornamented and temporal shield keeled). The morphological differences between H. crawfordi sp. nov. and H. lugubris are much more subtle, indicative of the close phylogenetic relationship between the two species. Molecular data and distribution (H. crawfordi sp. nov. in western Angola versus H. lugubris in south-eastern Angola, Namibia, Botswana, Zimbabwe, South Africa, and Mozambique) are the best proxies for its identification, as is the presence of bright yellow spots ventrolaterally (absent in H. lugubris). Heliobolus crawfordi sp. nov. can potentially also be distinguished from H. lugubris by the presence of 1–3 rows of granules between the supraoculars and supraciliaries (versus only one row), by having a lower average number of collar plates (7.1 [6–14] vs. 8.5 [6–11]), a lower number of subdigital lamellae under the fourth toe (22 [16–26] vs. 25.3 [23–28]) and having the parietal scales usually separated (versus usually in contact). Juveniles of H. crawfordican be distinguished from H. lugubris by the presence of a continuous white-yellow vertebral stripe (vs. a discontinuous vertebral stripe). Heliobolus crawfordi sp. nov. can be distinguished from H. bivari by having, on average, a lower number of subdigital lamellae under the fourth toe (22 [16–26] vs. 26.6 [21–34]), parietal scales usually separated (vs. usually in contact), a lateral dark marking through the ear to the posterior margin of the eye (vs. faded or totally absent), and presence of bright yellow spots ventrolaterally (vs. absent).”

Variation: Presence of the ventrolateral yellow spots is variable from being obvious to faded in some individuals.

Coloration: Juveniles black above and below with a continuous yellow vertebral stripe on the dorsum and two broken stripes above the arms; tail sandy-orange; symmetrically arranged yellow to orange spots and markings on the top of the head and snout, labials present some white markings.

Background coloration is brown to orange-brown with three continuous beige dorsal stripes, with a series of fairly broad transverse dark brown markings between stripes. Dorsolateral dark brown markings continue through the ear to the posterior margin of the eye. Bright yellow spots present ventrolaterally. Vertebral stripe dividing on the neck (in a Y-shape), extending anteriorly to the posterior margins of the parietals, and extending posteriorly to the lower back, where it fades out. Lower back uniform light orange; tail orange-grey dorsally. A pale ventrolateral band arises on the posterior margin of the subocular, extending posteriorly through the ear to the hindlimb insertion; pale yellow anteriorly, becoming beige posteriorly. Head uniformly light brown. Supralabials yellow with some dark infusions; dark infusions becoming more prominent on subocular and posterior supralabials, forming a narrow dark brown stripe that continues posteriorly to the ear opening. Limbs light brown to orange-brown, speckled with large pale spots dorsally. The venter is homogeneous dirty white, except for the orangish palmar regions of the hands and feet.  
CommentNatural history: Juveniles seem to imitate Anthia (Carabidae) ground beetles through their iconic white and yellow markings on a black body, their adopting an “arched” position when walking, and imitating the general shape of a beetle.

Diet: Little is known about the natural history of this species, but its ecological habits and behavior are expected to be similar to other Heliobolus species, being oviparous and insectivorous. 
EtymologySpecies named after the Portuguese mammalogist João Crawford-Cabral (Funchal, 1929–2020), a researcher at the former Instituto de Investigação Científica de Angola (IICA), Sá da Bandeira [currently Lubango], Huíla Province, Angola, and the recently defunct Instituto de Investigação Científica Tropical (IICT), Lisbon, Portugal. Crawford-Cabral played a pivotal role in the establishment and development of the zoological collections of IICA, currently housed at the Instituto Superior de Ciências de Educação (ISCED) in Lubango, Huíla Province, Angola, as well as in the publication of several important syntheses of biogeographic analyses on Angolan vertebrates. The specific epithet is a patronym in the masculine genitive singular. 
References
  • Marques MP, Ceríaco LMP, Heinicke MP, Chehouri RM, Conradie W, Tolley KA, Bauer AM 2022. The Angolan bushveld lizards, genus Heliobolus Fitzinger, 1843 (Squamata: Lacertidae): Integrative taxonomy and the description of two new species. Vertebrate Zoology 72: 745–769 - get paper here
 
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