Ecological segregation and population structuring of the cormorant Phalacrocorax carbo in Europe, in relation to the recent introgression of continental and marine subspecies

Populations of the continental Great Cormorant P. c. sinensis have expanded from north-eastern Europe towards the western part of the range of the marine P. c. carbo breeding in the United Kingdom and France. The aim of the study was to test the hypothesis of ecological segregation between subspecies by analysing the structuring of the European populations. Sequencing the mtDNA of 231 birds belonging to 20 colonies revealed 38 haplotypes based on 25 polymorphic sites (5.76% sequence divergence). P. c. sinensis (S) was well confirmed, but usual P. c. carbo formed two coastal populations, the real P. c. carbo C mainly in the western part of the range (United Kingdom, coastal France), and also in Norway and Sardinia, and N, branched to the Japanese Cormorant P. capillatus and probably isolated by glaciations, mainly present in the Nordic range (Norway, but also on the coasts from Sweden to Brittany), we named P. c. norvegicus. In a variable position in the trees but close to C is a group of undetermined origin haplotypes, named U, also present in both traditional ranges. The new tree-nesting colonies in Brittany are clearly a mixture of S and the two clades C and N previously described as P. c. carbo, with a decreasing proportion of C + N between 1993 (67%), 1996 (60%) and 2002 (33%) for the pioneering Grand-Lieu colony. These results confirmed the current introgression of continental populations in the western range, with probable hybridization. Although the subspecies can switch habitats locally due to social behaviour and migrations, the ecological segregation between the two usual subspecies appears to be largely confirmed in Europe.