Protected areas are one of the major tools used in the conservation of biodiversity, but animals are unlikely always to remain within these human‐made boundaries. Understanding when and why species choose to leave protected areas can help us to improve the effectiveness of these management tools. Here, we investigate the use of protected and non‐protected areas by two migratory species undergoing rapid wintering population changes in northwest Europe: Whooper Swans Cygnus cygnus and Bewick's Swans Cygnus columbianus bewickii. Global positioning system tags were fitted to 15 Whooper Swans in winter 2008/09 and to 18 Bewick's Swans from winter 2013/14 to 2014/15 at the Ouse Washes Special Protection Area (an internationally important roost for wintering waterbirds) and on adjacent fields in southeast England. Here, swans feed on farmland during the day but return to designated reserves to roost at night, where they receive protection from predators and disturbance within managed roost habitats. When swans roost elsewhere at alternative sites, they may face more adverse conditions, and so understanding the extent and causes of the use of alternative roosts is important for swan conservation efforts. The alternative roosting proportion, defined as the proportion of nights spent outside protected reserves, was 0.237 for Bewick's Swans and challenging to quantify accurately for Whooper Swans. A generalized additive mixed model to model repeated measurements on individuals showed that the proportion of time that Bewick's Swans spent at alternative roosts correlated positively with river level and negatively with temperature. Competition and foraging flight distances are thought to drive these relationships, as swans seek access both to roost space and to nearby feeding habitats. Our findings improve our understanding of the environmental conditions under which migratory waterbirds may choose to roost outside protected areas.
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