The year-class phenomenon and the storage effect in marine fishes

Abstract Factors contributing to population growth through strong year-class formation have driven a century of directed research in fisheries science. A central discovery of Hjort's paradigm was that multiple generations overlap and longevity is matched with frequency of strong recruitments. Here, I elaborate on this tenet by examining how intra-population modalities in spawning and early habitat use favour population resiliency. A modern theory that has application is the storage effect [Warner, R.R., Chesson, P.L., 1985. Coexistence mediated by recruitment fluctuations – a field guide to the storage effect. Am. Nat. 125, 769–787], whereby spawning stock biomass accumulates each year so that when early survival conditions are favourable, stored egg production can result in explosive population growth. I review two early life history behaviours that contribute to the storage effect: split cohorts (i.e., seasonal pulses of eggs and larvae) and contingent behaviour (i.e., dispersive and retentive patterns in early dispersal). Episodic and pulsed production of larvae is a common feature for marine fishes, well documented through otolith microstructure and hatch-date analyses. In temperate and boreal fishes, early and late spawned cohorts of larvae and juveniles may have differing fates dependent upon seasonal and inter-annual fluctuations in weather and climate. Often, a coastal fish may spawn for a protracted period, yet only a few days' egg production will result in successful recruitment. In these and other instances, it is clear that diversity in spawning behaviour can confer resilience against temporal variations in early survival conditions. Although many factors contribute to intra-population spawning modalities, size and age structure of adults play an important role. Contingent structure, an idea dating to Hjort (herring contingents) and Gilbert (salmon contingents), has been resurrected to describe the diversity of intra-population modalities observed through otolith microchemical and electronic tagging approaches. Retentive and dispersive behaviours confer resiliency against early survival conditions that vary spatially. Examples of contingent structure are increasingly numerous for diadromous fishes. Here, a nursery habitat associated with a contingent behaviour may make a small contribution in a given year, but over a decade contribute significantly to spawning stock biomass. For flatfish and other marine fishes, contingent structure is probable but not well documented. Proximate factors leading to contingent structure are poorly known, but for diadromous fishes, time of spawning and early life history energetic thresholds is hypothesized to lead to alternative life cycles. Here again time of spawning may lead to the storage effect by hedging against spatial variance in early vital rates. Managing for the storage effect will be promoted by conservation of adult age structure and early habitats upon which both strong and weak year-classes rely.