Phenological mismatch, carryover effects, and marine survival in a wild steelhead trout Oncorhynchus mykiss population

Climate-driven changes in the oceans, such as shifts in prey timing and abundance, could influence variability in population productivity of marine fishes. For example, according to the match/mismatch hypothesis, the temporal matching of the young salmon outmigration from freshwater to the ocean relative to the timing of availability of their prey could influence their marine survival. Indeed, understanding patterns and processes of marine survival is particularly pressing in many salmon and steelhead trout populations due to recent declines. To determine whether phenological mismatches between juvenile salmonids and their prey could contribute to low ocean survival, we analyzed the migration timing and ocean survival of 22,116 tagged juvenile steelhead trout Oncorhynchus mykiss over 12 years from the Wind River, Washington State, USA. We used a Bayesian multilevel modelling approach with variable selection to assess how survival was associated with body size, river exit date, the biological spring transition date (the day when northern zooplankton first appeared in the coastal region near the Columbia River estuary), and the degree of mismatch (the effect of the interaction between individual outmigration timing and biological spring transition date). The variables with the highest probability of contributing to individual survival were fish size (100%), river exit date (99%), the interaction between year and river exit date (91%), and the biological spring transition date (64%). Fish that were larger than average at outmigration had higher ocean survival, providing further evidence that freshwater growing conditions have carryover effects on marine survival. Years with greater annual phenological mismatches such as those years with late biological spring transition dates (i.e., occurring after June 1st), or warm sea surface temperatures, had sufficiently low marine survival to compromise recovery goals. Substantial intra-annual variation in outmigration timing buffered the population from inter-annual variation in optimal outmigration timing. Collectively these findings indicate that freshwater growing conditions, migration timing, and the timing of highquality food availability in the nearshore coastal environment work in concert to influence individual survival and annual smolt-to-adult returns.