Evidence for time-dependent calculations about future rewards is scarce in non-human animals. In non-human primates, only great apes are comparable with humans. Still, some species wait for several minutes to obtain a better reward in delayed exchange tasks. Corvids have been shown to match with non-human primates in some time-related tasks. Here, we investigate a delay of gratification in two corvid species, the carrion crow (Corvus corone) and the common raven (Corvus corax), in an exchange task. Results show that corvids success decreases quickly as delay increases, with a maximal delay of up to 320 s (more than 5 min). The decision to wait rests both on the quality of the prospective reward and the time required to obtain it. Corvids also apply tactics (placing the reward on the ground or caching it) that probably alleviate costs of waiting and distract their attention during waiting. These findings contrast previous results on delayed gratification in birds and indicate that some species may perform comparably to primates.

Self-control, that is, overcoming impulsivity towards immediate gratification in favour of a greater but delayed reward, is seen as a valuable skill when making future-oriented decisions. Experimental studies in nonhuman primates revealed that individuals of some species are willing to tolerate delays of up to several minutes in order to gain food of a higher quantity or quality. Recently, birds (carrion crows, Corvus corone, common ravens, Corvus corax, Goffin cockatoos, Cacatua goffiniana) performed comparably to primates in an exchange task, contradicting previous notions that birds may lack any impulse control. However, performance differed strikingly with the currency of exchange: individuals of all three species performed better when asked to wait for a higher food quality, rather than quantity. Here, we built on this work and tested whether the apparent difference in levels of self-control expressed in quality versus quantity tasks reflects cognitive constraints or is merely due to methodological limitations. In addition to the exchange paradigm, we applied another established delay maintenance methodology: the accumulation task. In this latter task, food items accumulated to a maximum of four pieces, whereas in the exchange task, an initial item could be exchanged for a reward item after a certain time delay elapsed. In both tasks, birds (seven crows, five ravens) were asked to wait in order to optimize either the quality or the quantity of food. We found that corvids were willing to delay gratification when it led to a food reward of higher quality, but not when waiting was rewarded with a higher quantity, independent of the experimental paradigm. This study is the first to test crows and ravens with two different paradigms, the accumulation and the exchange of food, within the same experiment, allowing for fair comparisons between methods and species. (Stevens & Stephens, 2010), ranging from foraging decisions (Kacelnik, 2003; reviewed in Stephens & Anderson, 2001) to social interactions, for example mate choice (Sozou & Seymour, 2003) or reciprocity in cooperative events (Stevens & Hauser, 2004). Going for the immediately available but less preferred option instead of postponing action in favour of an overall better but delayed reward is defined as impulsivity, whereas self-control refers to the opposite strategy (Ainslie, 1974;Kalenscher, Ohmann, & Güntürkün, 2006;Logue, Chavarro, Rachlin, & Reeder, 1988). Europe PMC Funders GroupFrom an economical point of view, the preference for a maximum payoff should be selected for (Noë, Hooff, & Hammerstein, 2001) 2005). It has been commonly suggested that temporal discounting is a critical factor in intertemporal decisions (Kacelnik & Bateson, 1996;Kalenscher & Pennartz, 2008;Stevens & Stephens, 2010). Accordingly, future rewards are subjectively rated less valuable the longer the delay until they are received, because delay is associated with uncertainty for realization of the benefits and probability of loss. Alternatively, it has been argued th...

Sensitivity to inequity is considered to be a crucial cognitive tool in the evolution of human cooperation. The ability has recently been shown also in primates and dogs, raising the question of an evolutionary basis of inequity aversion. We present first evidence that two bird species are sensitive to other individuals' efforts and payoffs. In a token exchange task we tested both behavioral responses to inequity in the quality of reward (preferred versus non-preferred food) and to the absence of reward in the presence of a rewarded partner, in 5 pairs of corvids (6 crows, 4 ravens). Birds decreased their exchange performance when the experimental partner received the reward as a gift, which indicates that they are sensitive to other individuals' working effort. They also decreased their exchange performance in the inequity compared with the equity condition. Notably, corvids refused to take the reward after a successful exchange more often in the inequity compared with the other conditions. Our findings indicate that awareness to other individuals' efforts and payoffs may evolve independently of phylogeny in systems with a given degree of social complexity.

The use and manufacture of tools have been considered to be cognitively demanding and thus a possible driving factor in the evolution of intelligence. In this study, we tested the hypothesis that enhanced physical cognitive abilities evolved in conjunction with the use of tools, by comparing the performance of naturally tool-using and non-tool-using species in a suite of physical and general learning tasks. We predicted that the habitually tool-using species, New Caledonian crows and Galápagos woodpecker finches, should outperform their non-tool-using relatives, the small tree finches and the carrion crows in a physical problem but not in general learning tasks. We only found a divergence in the predicted direction for corvids. That only one of our comparisons supports the predictions under this hypothesis might be attributable to different complexities of tool-use in the two tool-using species. A critical evaluation is offered of the conceptual and methodological problems inherent in comparative studies on tool-related cognitive abilities.

Self-control is critical for both humans and nonhuman animals because it underlies complex cognitive abilities, such as decision-making and future planning, enabling goal-directed behavior. For instance, it is positively associated with social competence and life success measures in humans. We present the first review of delay of gratification as a measure of self-control in nonhuman primates, corvids (crow family) and psittacines (parrot order): disparate groups that show comparable advanced cognitive abilities and similar socio-ecological factors. We compare delay of gratification performance and identify key issues and outstanding areas for future research, including finding the best measures and drivers of delayed gratification.Our review therefore contributes to our understanding of both delayed gratification as a measure of self-control and of complex cognition in animals.

Animals are expected to be judicious in the use of the energy they gain due to the costs and limits associated with its intake. The management of energy expenditure (EE) exhibited by animals has previously been considered in terms of three patterns: the constrained, independent and performance patterns of energy management. These patterns can be interpreted by regressing daily EE against maintenance EE measured over extended periods. From the multiple studies on this topic, there is equivocal evidence about the existence of universal patterns in certain aspects of energy management. The implicit assumption that animals exhibit specifically one of three discrete energy management patterns, and without variation, seems simplistic. We suggest that animals can exhibit gradations of different energy management patterns and that the exact pattern will fluctuate as their environmental context changes. To investigate these ideas, and for possible large‐scale patterns in energy management, we analysed long‐term heart rate data—a strong proxy for EE—across and within individuals in 16 species of birds, mammals and fish. Our analyses of 292 individuals representing 46,539 observation‐days suggest that vertebrates typically exhibit predominantly the independent or performance energy patterns at the across‐individual level, and that the pattern does not associate with taxonomic group. Within individuals, however, animals generally exhibit some degree of energy constraint. Together, these findings indicate that across diverse species, some individuals supply more energy to all aspects of their life than do others, however all individuals must trade‐off deployment of their available energy between competing functions. This demonstrates that within‐individual analyses are essential for the interpretation of energy management patterns. We also found that species do not necessarily exhibit a fixed energy management pattern but rather temporal variation in their energy management over the year. Animals’ energy management exhibited stronger energy constraint during periods of higher EE, which typically coincided with clear and key life cycle events such as reproduction, suggesting an adaptive plasticity to respond to fluctuating energy demands. A http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.13264/suppinfo is available for this article.

Simply observing other individuals interacting has been shown to affect subsequent behaviour and also hormones in 'bystander' individuals. However, immediate physiological responses of an observer have been hardly investigated. Here we present results on individuals' heart rate (HR) responses during various situations, which occur regularly in a flock of greylag geese (Anser anser, e.g. agonistic encounters, vehicles passing by). We recorded simultaneously HR and behaviour of 21 semi-tame free-roaming geese, equipped with fully implanted transmitters. We considered 304 social and 81 non-social events during which the focal individuals did not respond behaviourally. Independent of the spatial distance to the event, these HR responses were significantly greater in social contexts (e.g. departing or landing geese, agonistic interactions) than in non-social situations (e.g. vehicles passing by, thunder). Focal individuals showed a significantly higher maximum HR as well as a greater HR increase in response to agonistic interactions, in which the pair partner or a family member was involved, as compared with a non-affiliated goose. Also, HR was significantly higher when the bystander watched non-affiliated geese interacting, which were higher ranking than the focal. We conclude that these differences are due to different relevance of the recorded events for the focal individual, depending on the individuals involved in the observed interaction.

The requirements of living in social groups, and forming and maintaining social relationships are hypothesized to be one of the major drivers behind the evolution of cognitive abilities. Most empirical studies investigating the relationships between sociality and cognition compare cognitive performance between species living in systems that differ in social complexity. In this review, we ask whether and how individuals benefit from cognitive skills in their social interactions. Cognitive abilities, such as perception, attention, learning, memory, and inhibitory control, aid in forming and maintaining social relationships. We investigate whether there is evidence that individual variation in these abilities influences individual variation in social relationships. We then consider the evolutionary consequences of the interaction between sociality and cognitive ability to address whether bi-directional relationships exist between the two, such that cognition can both shape and be shaped by social interactions and the social environment. In doing so, we suggest that social network analysis is emerging as a powerful tool that can be used to test for directional causal relationships between sociality and cognition. Overall, our review highlights the importance of investigating individual variation in cognition to understand how it shapes the patterns of social relationships.This article is part of the theme issue 'Causes and consequences of individual differences in cognitive abilities'.