Welwitschia: Phylogeography of a living fossil, diversified within a desert refuge.

Welwitschia mirabilis is one of the most extraordinary plant species on earth. With a fossil record of 112 My and phylogenetically isolated within the order Gnetales, the monotypic genus Welwitschia has survived only in the northern Namib Desert in Angola and Namibia. Despite its iconic role, the biogeography, ecological niche, and evolutionary history of the species remain poorly understood. Here we present the first comprehensive map of the strongly disjunct species range, and we explore the genetic relationships among all range fragments based on six SSR markers. We also assess the variation of the environmental niche and habitat preference. Our results confirm genetic divergence, which is consistent with the hypothetical existence of two subspecies within Welwitschia. We identify an efficient geographical barrier separating two gene pools at 18.7°S in northern Namibia. We also identify further diversification within each of the two subspecies, with several different gene pools in ten isolated range fragments. Given the presence of well-isolated populations with unique gene pools and the association with different bioclimatic variables, rock types, and habitats within arid river catchments, we can hypothesize that the present intraspecific diversity may have evolved at least in part within the present refuge of the northern Namib Desert.