In this paper, we describe a technique to evaluate the evolutionary dynamics of the timing of spawning for iteroparous species. The life cycle of the species consists of three life stages, embryonic, juvenile and adult whereby the transitions of life stages (gametogenesis, birth and maturation) occur at species-specific sizes. The dynamics of the population is studied in a semi-chemostat environment where the inflowing food concentration is periodic (annual). A dynamic energy budget-based continuous-time model is used to describe the uptake of the food, storage in reserves and allocation of the energy to growth, maintenance, development (embryos, juveniles) and reproduction (adults). A discrete-event process is used for modelling reproduction. At a fixed spawning date of the year, the reproduction buffer is emptied and a new cohort is formed by eggs with a fixed size and energy content. The population consists of cohorts: for each year one consisting of individuals with the same age which die after their last reproduction event. The resulting mathematical model is a finite-dimensional set of ordinary differential equations with fixed 1-year periodic boundary conditions yielding a stroboscopic map. We will study the evolutionary development of the population using the adaptive dynamics approach. The trait is the timing of spawning. Pairwise and mutual invasibility plots are calculated using bifurcation analysis of the stroboscopic map. The evolutionary singular strategy value belonging to the evolutionary endpoint for the trait allows for an interpretation of the reproduction strategy of the population. In a case study, parameter values from the literature for the bivalve Macoma balthica are used.Keywords: adaptive dynamics; bifurcation analysis; bivalve Macoma balthica; dynamic energy budget-structured model; iteroparous species
INTRODUCTIONIn temperate regions, many species show a specific timing of reproduction within the seasonal cycle. In this paper, we develop a technique to evaluate the evolutionary dynamics of the timing of reproduction of iteroparous species. We assume that the annual reproduction occurs at a fixed date in the year, and the evolution of this life-history trait is studied. Our approach combines physiologically structured population modelling to describe the dynamics at the ecological time scale with the adaptive dynamics (AD) approach which occurs at an evolutionary time scale. The developed technique can be used in a bifurcation analysis to explore how the reproduction strategy of the population depends on individual and environmental properties. The life cycle of our model individual consists of three life stages: embryos (no feeding, no reproduction),