Abstract: Imitation can be defined as the copying of behavior. To a biologist, interest in imitation is focused on its adaptive value for the survival of the organism, but to a psychologist, the mechanisms responsible for imitation are the most interesting. For psychologists, the most important cases of imitation are those that involve demonstrated behavior that the imitator cannot see when it performs the behavior (e.g., scratching one's head). Such examples of imitation are sometimes referred to as opaque imitation because they are difficult to account for without positing cognitive mechanisms, such as perspective taking, that most animals have not been acknowledged to have. The present review first identifies various forms of social influence and social learning that do not qualify as opaque imitation, including species-typical mechanisms (e.g., mimicry and contagion), motivational mechanisms (e.g., social facilitation, incentive motivation, transfer of fear), attentional mechanisms (e.g., local enhancement, stimulus enhancement), imprinting, following, observational conditioning, and learning how the environment works (affordance learning). It then presents evidence for different forms of opaque imitation in animals, and identifies characteristics of human imitation that have been proposed to distinguish it from animal imitation. Finally, it examines the role played in opaque imitation by demonstrator reinforcement and observer motivation. Although accounts of imitation have been proposed that vary in their level of analysis from neural to cognitive, at present no theory of imitation appears to be adequate to account for the varied results that have been found.

Abstract: Considered an indicator of self-awareness, mirror self-recognition (MSR) has long seemed limited to humans and apes. In both phylogeny and human ontogeny, MSR is thought to correlate with higher forms of empathy and altruistic behavior. Apart from humans and apes, dolphins and elephants are also known for such capacities. After the recent discovery of MSR in dolphins (Tursiops truncatus), elephants thus were the next logical candidate species. We exposed three Asian elephants (Elephas maximus) to a large mirror to investigate their responses. Animals that possess MSR typically progress through four stages of behavior when facing a mirror: (i) social responses, (ii) physical inspection (e.g., looking behind the mirror), (iii) repetitive mirror-testing behavior, and (iv) realization of seeing themselves. Visible marks and invisible sham-marks were applied to the elephants' heads to test whether they would pass the litmus “mark test” for MSR in which an individual spontaneously uses a mirror to touch an otherwise imperceptible mark on its own body. Here, we report a successful MSR elephant study and report striking parallels in the progression of responses to mirrors among apes, dolphins, and elephants. These parallels suggest convergent cognitive evolution most likely related to complex sociality and cooperation.

Abstract: Whereas the way animals perceive human contact has been particularly examined in pet animals, a small amount of investigations has been done in domestic ungulates. It was nevertheless assumed that, as pet animals, non-aggressive forms of tactile contact were as well rewarding or positive for these species, even though the features of intraspecific relationships in pet animals and domestic ungulates may be to some extent different.We test here the hypothesis that horses may not consider physical handling by humans as a positive event. When comparing different early human-foal interactions, we found that early exposure to a motionless human enhanced slightly foals reactions to humans whereas forced stroking or handling in early life did not improve later human-foal relation. Foals that were assisted during their first suckling (e.g., brought to the dam's teat) even tended to avoid human approach at 2 weeks, and physical contact at 1 month of age.We argue that interspecies differences may exist in how tactile stimulation is perceived. It may be important for the establishment of a bond that a young animal is active in the process and able, through its behavioral responses, to help define what is positive for it. This way of investigation may have important general implications in how we consider the development of social relations, both within and between species.

Abstract: Reconciliation is a conflict resolution mechanism that is common to many gregarious species with individualized societies. Reconciliation repairs the damaged relationship between the opponents and decreases postconflict (PC) anxiety. The “integrated hypothesis” links the quality of the opponents' relationship to PC anxiety, since it proposes that conflicts among partners with high relationship quality will yield high levels of PC anxiety, which in turn will lead to an increased likelihood of reconciliation. We tested the integrated hypothesis in captive chimpanzees (Pan troglodytes) in the Arnhem Zoo, The Netherlands. We applied the standard PC/matched control (MC) method. Our results mostly support the integrated hypothesis, in that more valuable and compatible partners (i.e., males and frequent groomers) reconciled more often than less valuable and weakly compatible partners (i.e., females and infrequent groomers). In addition, PC anxiety was higher after conflicts among males than among females. Emotional arousal thus appears to be a mediator facilitating reconciliation. However, in contrast to the predictions derived from the integrated hypothesis, PC anxiety appeared only in aggressees, and not in aggressors, of conflicts. This suggests that while relationship quality determines PC anxiety, it is dependent on the role of the participants in the conflict.

Abstract: The trap-tube problem is used to assess whether an individual is able to foresee the outcome of its actions. To solve the task, an animal must use a tool to push a piece of food out of a tube, which has a trap along its length. An animal may learn to avoid the trap through a rule based on associative processes, e.g. using the distance of trap or food as a cue, or by understanding relations between cause and effect. This task has been used to test physical cognition in a number of tool-using species, but never a non-tool-user. We developed an experimental design that enabled us to test non-tool-using rooks, Corvus frugilegus. Our modification of the task removed the cognitive requirements of active tool use but still allowed us to test whether rooks can solve the trap-tube problem, and if so how. Additionally, we developed two new control tasks to determine whether rooks were able to transfer knowledge to similar, but novel problems, thus revealing more about the mechanisms involved in solving the task. We found that three out of seven rooks solved the modified trap-tube problem task, showing that the ability to solve the trap-tube problem is not restricted to tool-using animals. We found no evidence that the birds solved the task using an understanding of its causal properties, given that none of the birds passed the novel transfer tasks.