Parapatric diversification after post-glacial range expansion in the gall fly Urophora cardui (Tephritidae)

Aim Primary and secondary genetic clines in post-glacial colonized regions have different implications for biogeographic distributions and the origin of species. Primary clines arise in situ after colonization as adaptive responses to environmental gradients, while secondary clines are caused by contact between vicariant lineages. Here we analyse primary versus secondary origin of a genetic cline in the tephritid fly Urophora cardui in Jutland, Denmark, in a post-glacial landscape. Location Western Palaearctic. Methods Phylogeographic and demographic analyses of U. cardui based on mitochondrial DNA (mtDNA) genealogies, hierarchical genetic variance tests based on allozymes and distribution analysis of a rare allele from the Jutland cline. Results There was no phylogeographic divergence between the Jutland population of U. cardui north of the cline and neighbouring western European regional populations, which all shared the common western European mtDNA haplotype H1. At nuclear loci, by contrast, the North Jutland population was diverged above the mean level of divergence among regional populations and had no loss of genetic variation. A rare allozyme allele that was frequent in the cline area (up to 45%) and was missing north of the cline also occurred at low frequency (0-14%) elsewhere in the sampling range. Shallow phylogeographic divergence was observed between Russian and western European populations and between English and continental populations. Main conclusions The genetic variation patterns support primary cline evolution and parapatric divergence in Jutland following a demographic expansion of a western European ancestral source population of U. cardui, and suggest cryptic refugia and/or selection in other European population assemblages. The patterns of intra-specific regional divergence are discussed with respect to the interpretation of cryptic refugia in Europe after the most recent ice age.