Life History Adaptations to Seasonality.

Seasonality creates a template for many natural processes and evolutionary adaptations. Organisms are often faced with an annual cycle consisting of a productive (favorable) and unproductive period. This yearly cycle along with other seasonal variations in abiotic factors and associated biotic interactions form strong selection pressures shaping the scheduling of annual activities and the developmental stages and modes of life through the year. Annual decisions impact trade-offs that involve both current and future reproductive value (RV), and life history theory provides the foundation to understand these linkages between phenology and an organism's full life. Annual routine models further allow for multiple annual decisions to be optimized and predicted with respect to lifetime consequences. Studies of life history adaptations to seasonality are concerned with questions such as: within the productive season, should growth come first, followed by reproduction, or the other way around? What is the best time to diapause or migrate, and how will this timing impact other life history traits? Should energy reserves be built, to transfer resources from 1 year to the next, and allow for the spatial and temporal freedom of capital breeding? If offspring value is low during parts of the productive season, what is then the best alternative to reproduction: accumulate stores, grow, or wait in safety? To help answer these and other questions, I provide an overview of key theoretical concepts and some of the main life schedules, annual routines, and trade-offs involved. Adaptations to the unproductive period include diapause (dormancy), embryonic resting stages (eggs, seeds), energy reserves, and seasonal migrations. Adaptations to the productive window include rapid growth, high reproductive effort, capital breeding, and reproduction entrained to the annual cycle and with precise timing. Distinct annual routines, large body size, energy storage capacities, and parental care are also adaptations to seasonality. Phenotypic plasticity and state-dependence are important parts of these traits and are adaptations in their own. I give particular attention to timing of breeding and the associated birth-time dependent contributions to fitness. Seasonality in offspring value impacts the scheduling of growth, storage, and reproduction and may create parent-offspring conflicts over breeding timing. A combined offspring and parent value perspective should be adopted more broadly, also because of the management implications. I further argue for strategic but careful use of latitudinal (and altitudinal) gradients, and more attention to the role of seasonally varying predation risk as a selective force.