Ichnotropis capensis (Smith, 1838) sensu lato

Figures 20, 21, 22, 23; Table 6

Taxonomic note.

Ichnotropis capensis was originally described from the “ sandy deserts around Latakoo ”, which corresponds to the present-day Kuruman area in the Northern Cape province, South Africa. Since its original description, no additional specimens have been collected from the type locality or proximate areas. The nearest known record today is from Giya Camp in southern Botswana, approximately 340 km north of the type locality. This gap in distribution may be due to limited sampling effort in the region (Tolley et al. 2023), or alternatively, it may reflect uncertainty or inaccuracy in the locality information provided by Smith (1838), who was traveling extensively across the northern provinces of South Africa at the time. Due to the uncertainty surrounding the type locality and the fact that the type specimen remains unaccounted for in the BMNH (P. Campbell, pers. comm. 23 January 2024), the designation of a neotype is recommended to further stabilise the taxonomic status of Ichnotropis capensis. However, we refrain from taking this action at present, because we lack comparative material from the southern and eastern parts of the species’ distribution, including areas near the type locality of Ichnotropis longipes (Mazoë and the region between Umtali and Marandellas). Although we have examined photographs of the I. longipes type specimens (all subadults), we choose not to assign this name to any of the remaining I. capensis clades until additional material becomes available and a neotype for I. capensis can be designated.

Jacobsen et al. (2010) noted the presence of disjunct populations of Ichnotropis capensis between the east coast of southern Africa and the interior. Populations from the eastern coastal regions of South Africa and adjacent Mozambique form a distinct monophyletic clade that differs genetically by 4.1–6.4 % 16 S uncorrected p distance from other I. capensis populations. Although this coastal material could be referred to Ichnotropis macrolepidota Peters, 1854 — originally described from Lourenço Marques (now Maputo), Mozambique — our analysis indicates that the observed genetic differences fall within the expected range of intraspecific variation and likely reflect geographic separation between populations. This was further supported by the lack of clear morphological or geographical separation. As such, we consider Ichnotropis macrolepidota a junior synonym of I. capensis. Nevertheless, if future studies demonstrate that the eastern coastal populations represent a distinct operational taxonomic unit (OTU), the name I. macrolepidota should be resurrected for it.

Synonymy.

Algyra capensis Smith, 1838: 94; Tropidosaura Dumerelii Smith, 1849: appendix 7; Ichnotropis macrolepidota Peters, 1854: 617; Ichnotropis longipes Boulenger, 1902: 17.

Type.

BMNH 1865.5.4.56, collected from the ‘Sandy deserts around Latakoo’ [= Kuruman], Northern Cape, South Africa by Andrew Smith.

General description.

A medium-sized lacertid with a narrow and depressed snout. Head scalation strongly striated. Nostril pierced between three nasals; the supranasals are in broad contact behind the rostral; single frontonasal, as broad as long; paired prefrontal scales in broad contact medially; prefrontal separated from the anterior supraocular (only in contact in 15 out of 245 specimens examined) and separated from supraciliaries by a smaller scale; two large supraoculars, which are separated from the supraciliaries by one (very rarely two) row of small scales (4–9) and preceded by a cluster of 3–10 smaller scales; two loreal scales present, which are separated from the anterior supraocular by 2–3 scales; 1–2 post-supraoculars; subocular in contact with the lip; 3–6 (mostly four) supralabials in front of the subocular; 5–8 (mostly six) infralabials; five chin shields, with the anterior three in broad contact; 3–5 (mostly four) supraciliaries; 25–42 (average: 36.7) midbody scale rows; 8–10 (average: 8.8) longitudinal rows of enlarged ventral plates; 20–31 (average: 25.8) transverse ventral scale rows; 16–26 (average: 21.6) subdigital lamellae under the 4 th toe; 6–15 femoral pores per thigh. Size: Adult specimens varied from 40.0– 67.8 mm (mean: 54.6 mm) SVL and 69.5–149.0 mm (mean: 110.4 mm) TAIL. Largest female: 65 mm SVL (NMZB-UM 9228 – Umtali, Zimbabwe); largest male: 67.8 mm SVL (BE_ RMCA _Vert.R.7785 – Dilolo, DRC). Colouration (Fig. 20): In males, the flanks feature a striking, broad black longitudinal band that originates at the tip of the snout, passes through the eye, and extends posteriorly well beyond the hind limbs, gradually fading towards the tip of the tail. This black band is bordered by two distinct white stripes: The upper stripe begins just behind the eye, while the lower stripe originates at the rostral plate, crosses the tympanum, and runs parallel to the black band along the length of the body. Below the lower white stripe, a vivid reddish-orange stripe is especially prominent on the anterior flanks. A secondary short black line also originates at the snout, runs across the supralabials along the side of the head, and terminates anterior to the insertion of the forelimbs. The main black band on the flank is often scattered with small white spots, particularly towards the posterior end of the body. In breeding males, the white stripes on the head and neck, as well as the gular region, become infused with a bright yellow hue. The dorsal surface is a rich reddish-brown, adorned with scattered dark brown speckling. Females exhibit a more subdued colouration, with an overall grey-brown tone that is lighter on the ventral side. A single, less pronounced dark black stripe originates at the snout, passes through the eye, and continues along the flanks, gradually fading towards the tail. Juveniles and subadults are often grey in colouration with a white dorsolateral stripe. The venter is mostly white, but some specimens exhibit grey colouration with scattered black specks.

Distribution.

Widespread, occurring across several countries in southern Africa, including Angola, Namibia, Botswana, Zambia, Zimbabwe, Mozambique, and parts of South Africa and Malawi (Fig. 19). Historical records from north-eastern Angola and DRC assigned to I. capensis or I. overlaeti need to be re-evaluated in light of this study and might be assignable to either I. tanganicana, I. bivittata or I. longicorpa sp. nov. (see new species description below).

Habitat and Natural History.

This species prefers arid to mesic savanna habitats. It is a diurnal lizard, actively foraging for small invertebrates such as termites, spiders, beetles, and grasshoppers. Females lay up to nine eggs per clutch, typically during the summer months from October to November. The eggs measure approximately 5.5–7.0 mm by 8.5–9.5 mm. The incubation period ranges from 56 to 77 days, with hatchlings emerging between January and March. Females may produce up to two clutches within a single breeding season.