Changing food conditions and size declines in a North Sea forage fish

Declining body sizes are prevalent in marine fish. While these declines have been suggested to be a response to increasing temperatures, the evidence is mixed and the underlying causes of observed declines often unknown. Here, we explore drivers of spatio-temporal patterns in size in lesser sandeel (Ammodytes marinus), an important prey for seabirds and marine mammals, focusing on ongoing size declines in the North Sea. We combine experimental and field data with ecological theory to develop a biologically realistic dynamic energy budget model that explicitly models feeding, metabolism and energy allocation to produce daily predictions of size during the juvenile growth season from 1979 to 2016. When forced with daily temperature estimates and zooplankton data from the Continuous Plankton Recorder, model predictions reproduce observed spatio-temporal patterns in size well. Our results suggest that the most plausible driver of observed size declines in the western North Sea is declining prey densities. In contrast, the direct effect of temperature on sandeel size is small, but interacts with local prey availability so that the effect varies in both size and direction over space. Our results thus suggest that to understand effects of climate change on fish size we need to account for both direct physiological effects and changes in resource availability. Finally, we use the model to show that early-life phenology and turbidity (via its impact on intake rates in the visually foraging sandeel) may also impact sandeel size, highlighting the importance of broadening our view of potential drivers of size declines.