Can collective memories shape fish distributions? A test, linking space-time occurrence models and population demographics

Social learning can be fundamental to cohesive group living, and schooling fishes have proven ideal test subjects for recent work in this field. For many species, both demographic factors, and inter- (and intra-) generational information exchange are considered vital ingredients in how movement decisions are reached. Yet key information is often missing on the spatial outcomes of such decisions, and questions concerning how migratory traditions are influenced by collective memory, density-dependent and density-independent processes remain open. To explore these issues, we focused on Atlantic herring Clupea harengus, a long-lived, dense-schooling species of high commercial importance, noted for its unpredictable shifts in winter distribution, and developed a series of Bayesian space-time occurrence models to investigate wintering dynamics over 23 yr, using point-referenced fishery and survey records from Icelandic waters. We included covariates reflecting local-scale environmental factors, temporally-lagged prey biomass and recent fishing activity, and through an index capturing distributional persistence over time, derived two proxies for spatial memory of past wintering sites. The previous winter's occurrence pattern was a strong predictor of the present pattern, its influence increasing with adult population size. Although the mechanistic underpinnings of this result remain uncertain, we suggest that a 'wisdom of the crowd' dynamic may be at play, by which navigational accuracy towards traditional wintering sites improves in larger and/or denser, better synchronized schools. Wintering herring also preferred warmer, fresher, moderately stratified waters of lower velocity, close to hotspots of summer zooplankton biomass, our results indicative of heightened environmental sensitivity in younger cohorts. Incorporating spatiotemporal correlation structure and time-varying regression coefficients improved model performance, and validation tests on independent observations one-year ahead illustrate the potential of uniting demographic information and non-stationary models to quantify both the strength of collective memory in animal groups and its relevance for the spatial management of populations.