North by north-west: climate change and directions of density shifts in birds.

There is increasing evidence that climate change shifts species distributions towards poles and mountain tops. However, most studies are based on presence-absence data, and either abundance or the observation effort has rarely been measured. In addition, hardly any studies have investigated the direction of shifts and factors affecting them. Here, we show using count data on a 1000 km south-north gradient in Finland, that between 1970-1989 and 2000-2012, 128 bird species shifted their densities, on average, 37 km towards the north north-east. The species-specific directions of the shifts in density were significantly explained by migration behaviour and habitat type. Although the temperatures have also moved on average towards the north north-east (186 km), the species-specific directions of the shifts in density and temperature did not correlate due to high variation in density shifts. Findings highlight that climate change is unlikely the only driver of the direction of species density shifts, but species-specific characteristics and human land-use practices are also influencing the direction. Furthermore, the alarming results show that former climatic conditions in the north-west corner of Finland have already moved out of the country. This highlights the need for an international approach in research and conservation actions to mitigate the impacts of climate change.