Learning to be different: acquired skills, social learning, frequency dependence, and environmental variation can cause behaviourally mediated foraging specializations

Question: How does the ability to improve foraging skills by learning, and to transfer that learned knowledge, affect the development of intra-population foraging specializations? Features of the model: We use both a state-dependent life-history model implemented by stochastic dynamic programming (SDPM) and an individual-based model (IBM) to capture the dynamic nature of behavioural preferences in feeding. Variables in the SDPM include energy reserves, skill levels, energy and handling time per single prey item, metabolic rate, the rates at which skills are learned and forgotten, the effect of skills on handling time, and the relationship between energy reserves and fitness. Additional variables in the IBM include the probability of successful weaning, the logistic dynamics of the prey species with stochastic recruitment, the intensity of top-down control of prey by predators, the mean and variance in skill levels of new recruits, and the extent to which learned information can be transmitted via matrilineal social learning. Key range of variables: We explore the effects of approaching the time horizon in the SDPM, changing the extent to which skills can improve with experience, increasing the rates of learning or forgetting of skills, changing whether the learning curve is constant, accelerating (‘J’-shaped) or decelerating (‘r’-shaped), changing both mean and maximum possible energy reserves, changing metabolic costs of foraging, and changing the rate of encounter with prey. Conclusions: The model results show that the following factors increase the degree of prey specialization observed in a predator population: (1) Experience handling a prey type can substantially improve foraging skills for that prey. (2) There is limited ability to retain complex learned skills for multiple prey types. (3) The learning curve for acquiring new foraging skills is accelerating, or J-shaped. (4) The metabolic costs of foraging are high relative to available energy reserves. (5) Offspring can learn foraging skills from their mothers (matrilineal social Correspondence: M.T. Tinker, US Geological Survey, Western Ecological Research Center, Long Marine Laboratory, 100 Shaffter Road, Santa Cruz, CA 95060, USA. e-mail: [email protected] Consult the copyright statement on the inside front cover for non-commercial copying policies. Evolutionary Ecology Research, 2009, 11: 841–869 © 2009 M. Tim Tinker learning). (6) Food abundance is limited, such that average individual energy reserves are low Additionally, the following factors increase the likelihood of alternative specializations co-occurring in a predator population: (1) The predator exerts effective top-down control of prey abundance, resulting in frequency-dependent dynamics. (2) There is stochastic variation in prey population dynamics, but this variation is neither too extreme in magnitude nor too ‘slow’ with respect to the time required for an individual forager to learn new foraging skills. For a given predator population, we deduce that the degree of specialization will be highest for those prey types requiring complex capture or handling skills, while prey species that are both profitable and easy to capture and handle will be included in the diet of all individuals. Frequency-dependent benefits of selecting alternative prey types, combined with the ability of foragers to improve their foraging skills by learning, and transmit learned skills to offspring, can result in behaviourally mediated foraging specialization, and also lead to the co-existence of alternative specializations. The extent of such specialization is predicted to be a variable trait, increasing in locations or years when intra-specific competition is high relative to inter-specific