No abstract

Social foragers can use either a 'producer' strategy, which involves searching for food, or a 'scrounger' strategy, which involves joining others' food discoveries. While producers rely on personal information and past experience, we may ask whether the tendency to forage as a producer is related to being a better learner. To answer this question, we hand-raised house sparrow (Passer domesticus) nestlings that upon independence were given an individual-learning task that required them to associate colour signal and food presence. Following the testing phase, all fledglings were released into a shared aviary, and their social-foraging tendencies were measured. We found a significant positive correlation between individual's performance in the individual-learning task and subsequent tendency to use searching (producing) behaviour. Individual-learning score was negatively correlated with initial fear of the test apparatus and with body weight. However, the correlation between individual learning and searching remained significant after controlling for these variables. Since it was measured before the birds entered a social group, individual-learning ability could not be the outcome of being a producer. However, the two traits may be initially associated, or individual learning could facilitate producing behaviour. To our knowledge, this is the first evidence that associates individual-learning abilities with social-foraging strategies in animal groups.

In frequency-dependent games, strategy choice may be innate or learned. While experimental evidence in the producer -scrounger game suggests that learned strategy choice may be common, a recent theoretical analysis demonstrated that learning by only some individuals prevents learning from evolving in others. Here, however, we model learning explicitly, and demonstrate that learning can easily evolve in the whole population. We used an agent-based evolutionary simulation of the producer -scrounger game to test the success of two general learning rules for strategy choice. We found that learning was eventually acquired by all individuals under a sufficient degree of environmental fluctuation, and when players were phenotypically asymmetric. In the absence of sufficient environmental change or phenotypic asymmetries, the correct target for learning seems to be confounded by game dynamics, and innate strategy choice is likely to be fixed in the population. The results demonstrate that under biologically plausible conditions, learning can easily evolve in the whole population and that phenotypic asymmetry is important for the evolution of learned strategy choice, especially in a stable or mildly changing environment.

Although there has been extensive research on the evolution of individual decision making under risk (when facing variable outcomes), little is known on how the evolution of such decision-making mechanisms has been shaped by social learning and exploitation. We presented socially foraging house sparrows with a choice between scattered feeding wells in which millet seeds were hidden under 2 types of colored sand: green sand offering ~80 seeds with a probability of 0.1 (high risk-high reward) and yellow sand offering 1 seed with certainty (low risk-low reward). Although the expected benefit of choosing variable wells was 8 times higher than that of choosing constant wells, only some sparrows developed a preference for variable wells, whereas others developed a significant preference for constant wells. We found that this dichotomy could be explained by stochastic individual differences in sampling success during foraging, rather than by social foraging strategies (active searching vs. joining others). Moreover, preference for variable or constant wells was related to the sparrows' success during searching, rather than during joining others or when picking exposed seeds (i.e., they learn when actively searching in the sand). Finally, although for many sparrows learning resulted in an apparently maladaptive risk aversion, group living still allowed them to enjoy profitable variable wells by occasionally joining variable-preferring sparrows.

Choosing from whom to learn is an important element of social learning. It affects learner success and the profile of behaviors in the population. Because individuals often differ in their traits and capabilities, their benefits from different behaviors may also vary. Homophily, or assortment, the tendency of individuals to interact with other individuals with similar traits, is known to affect the spread of behaviors in humans. We introduce models to study the evolution of assortative social learning (ASL), where assorting on a trait acts as an individual-specific mechanism for filtering relevant models from which to learn when that trait varies. We show that when the trait is polymorphic, ASL may maintain a stable behavioral polymorphism within a population (independently of coexistence with individual learning in a population). We explore the evolution of ASL when assortment is based on a nonheritable or partially heritable trait, and when ASL competes with different non-ASL strategies: oblique (learning from the parental generation) and vertical (learning from the parent). We suggest that the tendency to assort may be advantageous in the context of social learning, and that ASL might be an important concept for the evolutionary theory of social learning. K E Y W O R D S :Assortment, homophily, model choice strategies, phenotypic asymmetry, social learning strategies.