Interacting effects of unobserved heterogeneity and individual stochasticity in the life-history of the Southern fulmar

Abstract

Individuals are heterogeneous in many ways. Some of these differences are incorporated as individual states (e.g., age, size, breeding status) in population models. However, substantial amounts of heterogeneity may remain unaccounted for, due to genetic, maternal, or environmental factors. Such unobserved heterogeneity (UH) affects the behavior of heterogeneous cohorts via intra-cohort selection and contributes to inter-individual variance in demographic outcomes such as longevity and lifetime reproduction. Variance is also produced by individual stochasticity, due to random events in the life cycle of wild organisms, yet no study thus far has attempted to decompose the variance in demographic outcomes into contributions from unobserved heterogeneity and individual stochasticity for an animal population in the wild. We developed a stage-classified matrix population model for the Southern fulmar breeding on Ile des Pétrels, Antarctica. We applied multi-event, multi-state markrecapture methods to estimate a finite mixture model accounting for UH in all vital rates and Markov chain methods to calculate demographic outcomes. Finally, we partitioned the variance in demographic outcomes into contributions from unobserved heterogeneity and individual stochasticity. We identify three UH groups, differing substantially in longevity, lifetime reproductive output, age at first reproduction, and in the proportion of the life spent in each reproductive state.

14% of individuals at fledging have a delayed but high probability of recruitment and extended reproductive lifespan.

67% of individuals are less likely to reach adulthood, recruit late and skip breeding often but have the highest adult survival rate.

19% of individuals recruit early and attempt to breed often. They are likely to raise their offspring successfully, but experience a relatively short lifespan.

Unobserved heterogeneity only explains a small fraction of the variances in longevity (5.9%), age at first reproduction (3.7%) and lifetime reproduction (22%). UH can affect the entire life cycle, including survival, development, and reproductive rates, with consequences over the lifetime of individuals and impacts on cohort dynamics. The respective role of unobserved heterogeneity versus individual stochasticity varies greatly among demographic outcomes. We discuss the implication of our finding for the gradient of life-history strategies observed among species and argue that individual differences should always be accounted for in demographic studies of wild populations.