The herring gull complex is not a ring species

Under what circumstances speciation in sexually reproducing animals can occur without geographical disjunction is still controversial. According to the ring-species model, a reproductive barrier may arise through ‘isolation by distance ’ when peripheral populations of a species meet after expanding around some uninhabitable barrier. The classical example of this kind of speciation is the herring gull (Larus argentatus) complex, with a circumpolar distribution in the Northern Hemisphere. Based on mitochondrial DNA variation among 21 gull taxa, we show that members of this complex differentiated largely in allopa-try following multiple vicariance and long-distance-colonization events, not primarily through isolation by distance. Reproductive isolation evolved more rapidly between some lineages than between others, irres-pective of their genetic distance. Extant taxa are the result of divergent as well as reticulate evolution between two ancestral lineages originally separated in a North Atlantic refugium and a continental Eura-sian refugium, respectively. Continental birds expanded along the entire north Eurasian coast and via Beringia into North America. Contrary to the ring-species model, we find no genetic evidence for a closure of the circumpolar ring through colonization of Europe by North American herring gulls. However, closure of the ring in the opposite direction may be imminent, with lesser black-backed gulls about to coloniz