An arms race between producers and scroungers can drive the evolution of social cognition.

The "social intelligence hypothesis" states that the need to cope with complexities of social life has driven the evolution of advanced cognitive abilities. It is usually invoked in the context of challenges arising from complex intragroup structures, hierarchies, and alliances. However, a fundamental aspect of group living remains largely unexplored as a driving force in cognitive evolution: the competition between individuals searching for resources (producers) and conspecifics that parasitize their findings (scroungers). In populations of social foragers, abilities that enable scroungers to steal by outsmarting producers, and those allowing producers to prevent theft by outsmarting scroungers, are likely to be beneficial and may fuel a cognitive arms race. Using analytical theory and agent-based simulations, we present a general model for such a race that is driven by the producer-scrounger game and show that the race's plausibility is dramatically affected by the nature of the evolving abilities. If scrounging and scrounging avoidance rely on separate, strategy-specific cognitive abilities, arms races are short-lived and have a limited effect on cognition. However, general cognitive abilities that facilitate both scrounging and scrounging avoidance undergo stable, long-lasting arms races. Thus, ubiquitous foraging interactions may lead to the evolution of general cognitive abilities in social animals, without the requirement of complex intragroup structures.

Western scrub-jays allocate longer observation time to more valuable information.

When humans mentally reconstruct past events and imagine future scenarios, their subjective experience of mentally time travelling is accompanied by the awareness of doing so. Despite recent popularity of studying episodic memory in animals, such phenomenological consciousness has been extremely difficult to demonstrate without agreed behavioural markers of consciousness in non-linguistic subjects. We presented western scrub-jays (Aphelocoma californica) with a task requiring them to allocate observing time between two peepholes to see food being hidden in either of two compartments, one where observing the hiding location was necessary to later relocate the food, and another where food could easily be found without watching. Jays first separately experienced these consequences of possessing information in each compartment and subsequently, once given a choice, made more looks and spent more time looking into the compartment where information was necessary than into the compartment where it was unnecessary. Thus, the jays can collect information to solve a future problem. Moreover, they can differentiate sources of information according to their potential value and modify behaviour to efficiently collect important, usable information. This is the first evidence of metacognition in a species that passes the behavioural criteria for both retrospective and prospective mental time travel.

Parents and offspring in an evolutionary game: the effect of supply on demand when costs of care vary.

Current models of parent-offspring communication do not explicitly predict the effect of parental food supply on offspring demand (ESD). However, existing theory is frequently interpreted as predicting a negative ESD, such that offspring beg less when parental supply is high. While empirical evidence largely supports this interpretation, several studies have identified the opposite case, with well-fed offspring begging more than those in poorer condition. Here, we show that signalling theory can give rise to either a negative or a positive ESD depending on the precise form of costs and benefits. Introducing variation among parents in the cost of care, we show that the ESD may change sign depending upon the quantitative relation between two effects: (i) decreased supply leads to increased begging because of an increase in marginal fitness benefit of additional resources to offspring, (ii) decreased supply leads to reduced begging because it is associated with a decrease in parental responsiveness, rendering begging less effective. To illustrate the interplay between these two effects, we show that Godfray's seminal model of begging yields a negative ESD when care is generally cheap, because the impact of supply on the marginal benefits of additional resources then outweighs the associated changes in parental responsiveness to begging. By contrast, the same model predicts a positive ESD when care is generally costly, because the impact of care costs on parental responsiveness then outweighs the change in marginal benefits.

Problems faced by food-caching corvids and the evolution of cognitive solutions.

The scatter hoarding of food, or caching, is a widespread and well-studied behaviour. Recent experiments with caching corvids have provided evidence for episodic-like memory, future planning and possibly mental attribution, all cognitive abilities that were thought to be unique to humans. In addition to the complexity of making flexible, informed decisions about caching and recovering, this behaviour is underpinned by a motivationally controlled compulsion to cache. In this review, we shall first discuss the compulsive side of caching both during ontogeny and in the caching behaviour of adult corvids. We then consider some of the problems that these birds face and review the evidence for the cognitive abilities they use to solve them. Thus, the emergence of episodic-like memory is viewed as a solution for coping with food perishability, while the various cache-protection and pilfering strategies may be sophisticated tools to deprive competitors of information, either by reducing the quality of information they can gather, or invalidating the information they already have. Finally, we shall examine whether such future-oriented behaviour involves future planning and ask why this and other cognitive abilities might have evolved in corvids.

Context-dependent flight speed: evidence for energetically optimal flight speed in the bat Pipistrellus kuhlii?

1. Understanding the causes and consequences of animal flight speed has long been a challenge in biology. Aerodynamic theory is used to predict the most economical flight speeds, minimizing energy expenditure either per distance (maximal range speed, Vmr) or per time (minimal power speed, Vmp). When foraging in flight, flight speed also affects prey encounter and energy intake rates. According to optimal flight speed theory, such effects may shift the energetically optimal foraging speed to above Vmp. 2. Therefore, we predicted that if energetic considerations indeed have a substantial effect on flight speed of aerial-hawking bats, they will use high speed (close to Vmr) to commute from their daily roost to the foraging sites, while a slower speed (but still above Vmp) will be preferred during foraging. To test these predictions, echolocation calls of commuting and foraging Pipistrellus kuhlii were recorded and their flight tracks were reconstructed using an acoustic flight path tracking system. 3. Confirming our qualitative prediction, commuting flight was found to be significantly faster than foraging flight (9.3 vs. 6.7 m s(-1)), even when controlling for its lower tortuosity. 4. In order to examine our quantitative prediction, we compared observed flight speeds with Vmp and Vmr values generated for the study population using two alternative aerodynamic models, based on mass and wing morphology variables measured from bats we captured while commuting. The Vmp and Vmr values generated by one of the models were much lower than our measured flight speed. According to the other model used, however, measured foraging flight was faster than Vmp and commuting flight slightly slower than Vmr, which is in agreement with the predictions of optimal flight speed theory. 5. Thus, the second aerodynamic model we used seems to be a reasonable predictor of the different flight speeds used by the bats while foraging and while commuting. This supports the hypothesis that bats fly at a context-dependent, energetically optimal flight speed.

The adaptive value of parental responsiveness to nestling begging.

Despite extensive theoretical and empirical research into offspring food solicitation behaviour as a model for parent-offspring conflict and communication, the adaptive value of parental responsiveness to begging has never been tested experimentally. Game theory models, as well as empirical studies, suggest that begging conveys information on offspring state, which implies that parental investment can be better translated to fitness by responding to begging when allocating resources rather than by ignoring it. However, this assumption and its underlying mechanisms have received little or no attention. Here we show by experiments with hand-raised house sparrow (Passer domesticus) nestlings that a 'responsive parent' will do better than a hypothetical 'non-responsive' mutant (that provides similar food amounts, but irrespective of begging). This is neither because food-deprived nestlings convert food to mass more efficiently, however, nor because responsiveness reduces costly begging. Rather, responsiveness to begging is adaptive because it reduces two opposing risks: one is wasting time when returning too soon to feed already satiated nestlings and the other is repeatedly overlooking some nestlings as a result of the stochastic nature of a random, non-responsive strategy. This study provides the first experimental evidence for the adaptive value of parental responsiveness to offspring begging.