Integrative taxonomy of Afrotropical Ornithodoros (Ornithodoros) (Acari: Ixodida: Argasidae)

Afrotropical Ornithodoros (Ornithodoros) ticks are revised based on qualitative morphology of females and nymphs, as well as tarsus I shape outlines of females measured in a geometric morphometric framework. These lines of evidence corroborate lineages based on 16S rRNA nucleotide sequence data. Four previously unrecognized species are described, along with a revived nomen nudum that was previously considered a synonym. Afrotropical Ornithodoros (Ornithodoros) now comprise ten species. Ornithodoros moubata and Ornithodoros porcinus are separated from three other species in southern Africa (Ornithodoros compactus, Ornithodoros phacochoerus n. sp., Ornithodoros waterbergensis n. sp.), with O. porcinus restricted to central east Africa. Known species boundaries for Ornithodoros apertus and O. compactus are supported. Ornithodoros savignyi are separated from three other species in South Africa and Namibia, with O. savignyi restricted to north Africa. Neumann's Ornithodoros pavimentosus nom. rev. are resurrected from synonymy as a species that occur in Bushmanland, Namaqualand and Namibia, while Ornithodoros kalahariensis n. sp. occur in Kalahari thornveld, and Ornithodoros noorsveldensis n. sp. occur in Noorsveld thicket of South Africa. Detailed descriptions are given for each species along with high resolution images and point map distributions. Support is provided for speciation driven by riverine barriers, Pliocene uplift and differential arid tolerance. Exaggerated tarsus I shape in the O. savignyi group suggests adaptation to fossorial habits and soil type. Conversely, reduced tarsus I shape in the O. moubata group is suggested as an evolutionary consequence of the life history change from soil to warthog burrows. This study represents an integrative (iterative) approach to delimiting Afrotropical Ornithodoros (Ornithodoros) species, and provides the first application of tarsus I shape outlines in a geometric morphometric framework for testing species boundaries.