Migration patterns and recorded emigration of the Great Cormorant phalacrocorax carbo sinensis in the largest French colony of Lac de Grand-Lieu: density-dependent factors operating at different time and geographical scales

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anglais. The number of wintering Great Cormorants Phalacrocorax carbo sinensis in France has strongly increased since the 1970s, mainly due to the protection of the 'continental' sinensis subspecies in countries north of France. This increase has led to the establishment of a pioneering inland breeding colony in western France, while previously only the largely marine 'Atlantic' P. c. carbo subspecies occurred on the Channel coast. The marine subspecies was attracted and bred in this new inland settlement of sinensis, which rapidly became the largest colony in France. This paper analyses the migration pattern of birds from this colony by analysis of the dispersal of colour-ringed birds between 1989 and 2008. Interestingly, besides a classic south-west migration pattern, birds from this colony also displayed a pattern heading north-east, up to countries such as The Netherlands, from where the founders (sinensis) of this colony probably originated. Sightings and recoveries revealed that about 25% of the adults and 19% of the first-year birds headed north-east. Due to this north-east migration direction, the overall annual dispersal point was located only 50 km south-west of the colony, although in December and January this midpoint was located about 320 km south-west of the colony. The birds largely avoided Brittany, presumably to avoid competition with individuals of the carbo subspecies, and the main wintering areas of sinensis from other colonies, both in France (east, centre and south) and in Spain. Over the years 1989-2008, in the breeding period, the mean dispersal distance was shorter for adults than for young birds (54 km vs. 144 km, respectively) but in winter adult birds migrated further than young ones (305 km vs. 221 km, respectively). The mean annual dispersal distance in winter varied from 106 km to 527 km (all age-classes taken together). Migratory distance was not related to mean winter temperature. For adults, dispersal distance correlated with the annual number of breeding pairs in the Grand-Lieu colony between 1990 and 2003, but not between 2004-2008. Emigration (breeding in another colony) was recorded up to 2011 to 11 inland colonies and one coastal colony (founded more recently than Grand-Lieu), nine of them in France, two in Spain and one in The Netherlands. Annual emigration rate was negatively related to colony size in Grand-Lieu. The study points to the existence of density-dependent effects on distribution patterns of Cormorants outside the breeding season but also suggests connectivity and interaction among colonies that are hundreds of kilometres apart.