Paraphyly of the widespread generalist red fox (Vulpes vulpes): introgression rather than recent divergence of the arid-adapted Ruppell's fox (Vulpes rueppellii)?

Understanding of the evolutionary history of two closely related canid sister taxa, the geographically restricted, arid-adapted Ruppell's fox (Vulpes rueppellii) and the widespread generalist red fox (Vulpes vulpes), has been hampered by limited sampling in the biogeographically complex region of North Africa and the Middle East. We sequenced mitochondrial DNA (mtDNA) cytochrome b and D-loop fragments from 116 samples for both species and combined these data with previously published sequences, resulting in 459 haplotypes. Obtained phylogenies showed high support for most branches, including for a newly described 'Palearctic clade' that includes North African and Asian individuals from both species. All V. rueppellii individuals fell within the Palearctic clade, forming two previously undescribed subclades that were intermingled with, but not shared with V. vulpes. Our robust placement of V. rueppellii within V. vulpes renders the latter paraphyletic. We propose three scenarios that could explain these observations: (1) rapid, recent speciation of V. rueppellii from V. vulpes, (2) incomplete lineage sorting, or (3) ancient divergence followed by introgression and secondary mtDNA similarity. The third scenario is in best agreement with evidence from the fossil record, and morphometric and ecological distinctiveness between the two taxa, and therefore seems most likely.