Long-term shifts in phenology, thermal niche, population size, and their interactions in marine pelagic copepods

Under climatic warming many species shift their seasonal timing of life cycle events (phenology) and seasonal abundance distribution, but whether they maintain the same thermal niche is still poorly understood. Here, we studied multidecadal trends in abundance and phenology of seven major copepod species across three stations (Stonehaven (SH), Helgoland Roads (HR), and Plymouth L4) on the North-West European shelf, spanning similar to 6.5 degrees of latitude. All seven species consistently occupied colder temperatures at the northern station compared to the southerly station, but they maintained the same realized thermal niche over years. Expected phenological shifts (i.e., earlier when warmer) in some stations were obscured possibly by the long-term drop of copepod density in spring-summer, which may be due to a variation in the food/predators abundance. The ongoing spring-summer declines in abundance (similar to 50%) of many North Atlantic pelagic species over the last five decades, as found in recent studies, may have also influenced the metrics of seasonal timing. To separate the seasonal timing of life events from that of seasonal abundance distribution, we used a time series of egg production rate (EPR) of Calanus helgolandicus at L4, and found that this shifted later into the summer-autumn over the last 30 yr of warming, coincident with declining spring-summer food and increasing predator abundance. Overall, direct temperature effects do appear to influence the seasonal timing of the copepods, but to explain impacts at individual stations or long-term trends in population size or phenology, understanding the changing balance of food and predators appears to be critical.