Secondary contact zones have been formed between several pairs of avian species and subspecies in northern and north-eastern Iran during the post-Pleistocene and Holocene periods. Three subspecies groups out of the four in the great tit (Parus major), major, bokharensis and cinereus, are believed to have come into local or regional secondary contact in north-eastern Iran. Parus major intermedius is also known from this region and has long been suggested to have a hybrid origin from hybridization between the yellow western (major) subspecies group and the grey-coloured eastern (bokharensis or cinereus) subspecies group based on its intermediate plumage coloration. Here, we investigated the evidence of intergradation between subspecies groups and the evolutionary origin of P. m. intermedius using mitochondrial DNA sequences and microsatellites, and approximate Bayesian computation to test competing scenarios for the demographic history of the populations. Our analyses indicate a divergence origin for intermedius that resulted from expansion of the major subspecies group. Low mitochondrial diversity and high genetic differentiation in comparison with central populations suggest that intermedius is a peripheral population. Microsatellite data show no signs of nuclear admixture between the bokharensis and major subspecies groups. However, one phenotypically intermedius specimen had bokharensis mtDNA and major nuclear DNA in the assumed hybrid zone (Lotf Abad), supporting past introgression.

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