Frédérique Dubois

Problem-solving performance is correlated with reproductive success in a wild bird population

Although interindividual variation in problem-solving ability is well documented, its relation to variation in fitness in the wild remains unclear. We investigated the relationship between performance on a problem-solving task and measures of reproductive success in a wild population of great tits, Parus major. We presented breeding pairs during the nestling provisioning period with a novel string-pulling task requiring the parents to remove an obstacle with their leg that temporarily blocked access to their nestbox. We found that nests where at least one parent solved the task had higher nestling survival until fledging than nests where both parents were nonsolvers. Furthermore, clutch size, hatching success and fledgling number were positively correlated with speed in solving the task. Our study suggests that natural selection may directly act on interindividual variation in problem-solving performance. In light of these results, the mechanisms maintaining between-individual variation in problem-solving performance in natural populations need further investigation.

Chapter 2 Social Foraging and the Study of Exploitative Behavior

Publisher Summary A large number of social interactions whether cooperative or exploitative revolve around foraging, whether social insects provisioning the communal nest or vampire bats regurgitating blood meals to a companion when needed. Research on animal cooperation, for instance, has often resorted to food rewards as a means of studying the behavioral rules that promote cooperation or defection. Many ecological problems at the population level involve foraging. The link between foraging and population phenomena is direct. Optimal foraging theory relies heavily on simple optimality as a means to formalize foraging problems. The field of social foraging theory is subdivided into four main categories: (1) group membership, (2) decisions within patches, (3) descriptive models of phenotypic diversity, and (4) producer–scrounger (PS) decisions. The PS approach is useful in exploring questions of resource defense. In particular, it provides a convenient means of investigating how the characteristics of a resource affect the extent to which it should be appropriated by others. Because there can be no defense without having others attempting to appropriate, the PS game provides a means to investigate the whole process of resource defense.

Learning in a game context: strategy choice by some keeps learning from evolving in others

Behavioural decisions in a social context commonly have frequency-dependent outcomes and so require analysis using evolutionary game theory. Learning provides a mechanism for tracking changing conditions and it has frequently been predicted to supplant fixed behaviour in shifting environments; yet few studies have examined the evolution of learning specifically in a game-theoretic context. We present a model that examines the evolution of learning in a frequency-dependent context created by a producer–scrounger game, where producers search for their own resources and scroungers usurp the discoveries of producers. We ask whether a learning mutant that can optimize its use of producer and scrounger to local conditions can invade a population of non-learning individuals that play producer and scrounger with fixed probabilities. We find that learning provides an initial advantage but never evolves to fixation. Once a stable equilibrium is attained, the population is always made up of a majority of fixed players and a minority of learning individuals. This result is robust to variation in the initial proportion of fixed individuals, the rate of within- and between-generation environmental change, and population size. Such learning polymorphisms will manifest themselves in a wide range of contexts, providing an important element leading to behavioural syndromes.

Long-term social bonds promote cooperation in the iterated Prisoner's Dilemma

Reciprocal altruism, one of the most probable explanations for cooperation among non-kin, has been modelled as a Prisoners Dilemma. According to this game, cooperation could evolve when individuals, who expect to play again, use conditional strategies like tit-for-tat or Pavlov. There is evidence that humans use such strategies to achieve mutual cooperation, but most controlled experiments with non-human animals have failed to find cooperation. One reason for this could be that subjects fail to cooperate because they behave as if they were to play only once. To assess this hypothesis, we conducted an experiment with monogamous zebra finches (Taeniopygia guttata) that were tested in a two-choice apparatus, with either their social partner or an experimental opponent of the opposite sex. We found that zebra finches maintained high levels of cooperation in an iterated Prisoners Dilemma game only when interacting with their social partner. Although other mechanisms may have contributed to the observed difference between the two treatments, our results support the hypothesis that animals do not systematically give in to the short-term temptation of cheating when long-term benefits exist. Thus, our findings contradict the commonly accepted idea that reciprocal altruism will be rare in non-human animals.

Mate-choice copying by female zebra finches, Taeniopygia guttata: what happens when model females provide inconsistent information?

For a long time, mate-choice copying was thought to be restricted to lekking and polygynous species. Yet, recent experimental studies revealed that social information can play a role in the evolution of mate preferences in monogamous species with biparental care. However, this phenomenon has been demonstrated only under particular conditions and the prevalence and importance of this phenomenon therefore remains to be evaluated. In particular, previous laboratory experiments have consisted in exposing test females to only one paired male at a time, while under natural conditions monogamous females are likely to observe the choice of several females before making a decision. Thus, in the present study, female zebra finches (Taeniopygia guttata) could observe two model females simultaneously, that provided either inconsistent or consistent information, depending on whether they were interacting with different types of males or with males of a same phenotype. We found that the relative importance given to private and social information on females’ preferences varied with the consistency of social information and females significantly changed their preference only when social information was consistent. There was, nevertheless, a large variation in their responses. We suggest that such variations could be due to the fact that the benefits of mate-choice copying are frequency-dependent, and that this constrain would further contribute to limit the use of social information in monogamous species.

Individual differences in sampling behaviour predict social information use in zebra finches

When animals have to decide where to forage, what to eat or with whom to mate, they can base their decisions on either socially or personally acquired information. In accordance with theoretical predictions, there is experimental evidence that animals adjust the weight they give to both sources of information depending on circumstances. Notably, several studies have demonstrated that individuals rely more on social information when personal information is difficult to acquire or unreliable, because these conditions leave them uncertain as how to behave. Yet, even when individuals are exposed to the same conditions, they generally differ widely in the value they attribute to social and personal information. These differences suggest that the tendency to rely on social information would also depend on intrinsic characteristics that affect, for instance, individual efficiency in collecting personal information. To address this issue, we conducted laboratory experiments using female zebra finches (Taeniopygia guttata) and we tested them under three consecutive conditions. First, we evaluated their reliance on social information in a mating context and in a foraging context. Then, we measured their efficiency in acquiring personal information by recording their sampling behaviour when searching for hidden food. We found that females that sampled their environment less actively consistently relied on social information to a greater extent compared with those that invested more in sampling. Contrary to what is generally assumed, then, our study demonstrates that social information use is not entirely flexible and context dependent.

Food sharing among retaliators: sequential arrivals and information asymmetries

Many animals share food, that is, to tolerate competitors at a defensible clump. Most accounts of resource sharing invoke special evolutionary processes or ecological circumstances that reduce their generality. Surprisingly, the Hawk–Dove game has been unable to address in a simple and general way why so many group foraging animals share food. We modify the Hawk–Dove game by allowing a finder the opportunity of retaliating if joiners escalate and by considering the consequences of information asymmetries concerning resource value among players. Introducing the first change, the retaliator strategy was sufficient to predict widespread sharing in habitats where food clumps are of intermediate richness. When information asymmetry between finder and joiner is created by allowing the quality of clumps to vary, we show that the conditions for sharing are even more easily met and apply to a wider range of resource qualities. Our model therefore offers one of the most parsimonious and potentially general evolutionary accounts of the origin of non-aggressive resource sharing.

Mate choice copying in monogamous species: should females use public information to choose extrapair mates?

Empirical evidence that females can copy each others mating preferences comes predominantly from research on lekking species, but recent laboratory studies revealed that public information can also play a role in the evolution of mate preferences in monogamous species with biparental care. Although the question of why monogamous females copy each others mating preferences is still debated, it has been suggested that public information could be used by females to assess and choose extrapair mates. Since there is only indirect empirical support for this hypothesis, I developed a game-theoretic model to explore the possible roles that mate choice copying may play in mediating extrapair behaviours in monogamous species. Like previous game-theoretic models on mate choice copying, the model predicts that only females that have a high probability of obtaining a better partner should copy the mating decisions of others. On the other hand, unlike previous games that applied to lekking species, the present model predicts that mate choice copying can be advantageous even when there is no variation among females in their discrimination ability and they all can assess the quality of potential partners without error. Moreover, as mate choice copying essentially benefits high-quality males, males of lower quality would be interested in developing defensive tactics to discourage females from seeking extrapair copulations, thereby contributing to intraspecific and interspecific variations in the rate of extrapair paternity.

Frequency-dependent payoffs and sequential decision-making favour consistent tactic use.

Although natural selection should have favoured individuals capable of adjusting the weight they give to personal and social information according to circumstances, individuals generally differ consistently in their individual weighting of both types of information. Such individual differences are correlated with personality traits, suggesting that personality could directly affect individuals’ ability to collect personal or social information. Alternatively, the link between personality and information use could simply emerge as a by-product of the sequential decision-making process in a frequency-dependent context. Indeed, when the gains associated with behavioural options depend on the choices of others, an individuals sequence of arrival could constrain its choice of options leading to the emergence of correlated behaviours. Any factor such as personality that affects decision order could thus be correlated with information use. To test this new explanation, we developed an individual-based model that simulates a group of animals engaged in a game of sequential frequency-dependent decision: a producer–scrounger game. Our results confirm that the sequence of decision, in this case enforced by the order in which animals enter a foraging area, consistently influences their mean tactic use and their individual plasticity, an outcome reminiscent of the correlation reported between personality and social information use.

Audience Effect Alters Male Mating Preferences in Zebra Finches (Taeniopygia guttata)

The social environment of animals strongly influences the mating preferences of both the choosing and the observing individuals. Notably, there is recent evidence that polygamous males decrease their selectivity when being observed by competitors in order to direct their rivals’ attention away from their true interest and, consequently, reduce sperm competition risk. Yet, other mechanisms, whose importance remains unexplored, could induce similar effects. In monogamous species with mutual choice, particularly, if males adjust their selectivity according to the risk of being rejected by their preferred mate, they should as well become less selective when potential rivals are present. Here, we investigated whether the presence of bystanders modifies male mating preferences when the risk of sperm competition is low, by carrying out mate-choice experiments with male zebra finches (Taeniopygia guttata) whose preferences for two females were measured twice: with and without an audience. We found that the presence of potential rivals had no effect on the males’ choosiness. However, with an audience, they spent more time with the female that was considered as the less attractive one in the control condition. These findings support the hypothesis that monogamous males alter their mate choice decisions in the presence of a male audience to reduce the risk of remaining unpaired. Thus, our results indicate that several explanations can account for the changes in male preferences due to the presence of competitors and highlight the importance of assessing the relative role of each mechanism potentially involved, to be able to make conclusions about the effect of an audience on signal evolution.