The Angolan bushveld lizards, genus Heliobolus Fitzinger, 1843 (Squamata: Lacertidae): Integrative taxonomy and the description of two new species

The genus Heliobolus comprises four recognized species, all endemic to sub-Saharan Africa. Of these, only Heliobolus lugubris occurs in southern Africa, its distribution extending from Angola in the west to Mozambique in the east and reaching as far south as parts of northern South Africa. Like many of the reptile species that occur in southern Africa, Heliobolus lugubris is poorly studied, and preliminary investigation suggested that it may contain cryptic diversity. The present work focusses on the Angolan population of H. lugubris and uses an integrative taxonomic approach based on morphological, coloration and DNA sequence data. The results indicate that some of the current and historical specimens of H. lugubris from Angola do not correspond to the nominotypical form, and that differences between specimens suggest the presence of two additional species, described here as Heliobolus bivari sp. nov. from the southernmost xeric/desertic regions and plateau of Namibe Province, southwestern Angola and H. crawfordi sp. nov. from the Serra da Neve inselberg north through the sub-desert coastal regions of northern Namibe, Benguela, and Kwanza Sul provinces. Nominotypical Heliobolus lugubris is confirmed to occur in Cuando Cubango Province, southeastern Angola.