Abstract: Comparative experimental studies of imitative learning have focused mainly on primates and birds. However, cetaceans are promising candidates to display imitative learning as they have evolved in socioecological settings that have selected for large brains, complex sociality, and coordinated predatory tactics. Here we tested imitative learning in killer whales, Orcinus orca. We used a ‘do-as-other-does’ paradigm in which 3 subjects witnessed a conspecific demonstrator’s performance that included 15 familiar and 4 novel behaviours. The three subjects (1) learned the copy command signal ‘Do that’ very quickly, that is, 20 trials on average; (2) copied 100 % of the demonstrator’s familiar and novel actions; (3) achieved full matches in the first attempt for 8–13 familiar behaviours (out of 15) and for the 2 novel behaviours (out of 2) in one subject; and (4) took no longer than 8 trials to accurately copy any familiar behaviour, and no longer than 16 trials to copy any novel behaviour. This study provides experimental evidence for body imitation, including production imitation, in killer whales that is comparable to that observed in dolphins tested under similar conditions. These findings suggest that imitative learning may underpin some of the group-specific traditions reported in killer whales in the field.

Abstract: Equine welfare can be enhanced by applying species specific training. This may incorporate social learning, as horses are highly social and social stimuli are of primary importance. Social learning is comparable to individual learning in its learning mechanisms, differing primarily in the way it is stimulated. Our initial study showed that horses of different breeds (N = 38) follow humans after observing other horses doing so, but only if the observed horse was familiar to and higher ranking than the observer (Fisher's exact test: N = 12, P = 0.003). A second study showed that horses and ponies (N = 25) learned to pull a rope to open a feeding apparatus after observing demonstrations by conspecifics, again, only if the demonstrating horse was older and higher ranking than the observer (Fisher's combination test, N = 3, v2 = 27.71, p = 0.006). Our third approach showed that horses and ponies (N = 24) learned to press a switch to open a feeding apparatus after observing a familiar person (GzLM: N = 24, z = 2.33, P = 0.02). Most recently, we confronted horses and ponies (N = 50) with persons demonstrating different techniques for opening a feeding apparatus. In this study we investigated whether the horses would copy the demonstrators' techniques or apply their own. Here only some horses copied the technique, and most of the successful learners used their mouths irrespective of the demonstrators' postures (Chi Square Test: N = 40, df = 2, &#967;2 = 31.4, p < 0.001). In all the approaches social stimuli elicited learning processes in the test horses, while only a few individuals in the control groups mastered the tasks by individual learning. The following behaviour observed in the initial study may have been facilitated by a social stimuli (social facilitation), and the opening of the feed boxes in the subsequent studies appear to be mostly the result of enhancement (social enhancement). Some horses may have used the social stimuli at first and continued their learning process by individual trial and error. However, the horses were also selective in whom and some in how to copy. This may have been conditioned (socially conditioned) or the result of simple forms of reasoning on the reliability of the particular information provided by demonstrators of certain social ranks or social positions, as high ranking and familiar horses and familiar persons were copied and some imitated exactly.
Lay person message: Traditional riding instructions suggest that horses learn by observing other horses. For example, older, more experienced driving horses are used for initial training of young driving horses. We have shown that horses indeed use learning stimuli provided by other horse, as well as by humans. Horses readily accept stimuli observed in high ranking and familiar horses, and familiar persons. Such stimuli elicit learning processes which are comparable to individual learning. We suggest applying social learning whenever possible, as it is much faster and less stressful than individual learning, where learners experience negative outcomes in trial and error learning.

Abstract: There are numerous reports of novel learned behaviour patterns in animal populations, yet the factors influencing the invention and spread of these innovations remain poorly understood. Here we investigated to what extent the pattern of spread of innovations in captive groups of starlings, Sturnus vulgaris, could be predicted by knowledge of individual and social group variables, including association patterns, social rank orders, measures of neophobia and asocial learning performance. We presented small groups of starlings with a series of novel extractive foraging tasks and recorded the latency for each bird to contact and solve each task, as well as the orders of contacting and solving. We then explored which variables best predicted the observed diffusion patterns. Object neophobia and social rank measures characterized who was the first of the group to contact the novel foraging tasks, and the subsequent spread of contacting tasks was associated with latency to feed in a novel environment. Asocial learning performance, measured in isolation, predicted who was the first solver of the novel foraging tasks in each group. Association patterns did not predict the spread of solving. Contact latency and solving duration were negatively correlated, consistent with social learning underlying the spread of solving. Our findings indicate that we can improve our understanding of the diffusion dynamics of innovations in animal groups by investigating group-dependent and individual variables in combination. We introduce novel methods for exploring predictors of the origin and spread of behavioural innovations that could be widely applied.

Abstract: Our understanding of how, why, and the circumstances under which animals use social information has been facilitated by three principal areas of research, social learning, public information use and social eavesdropping. With few exceptions, these related concepts have remained remarkably distinct within the literature, with little discussion or integration among them. Are these distinctions warranted? We tackle the issue by exploring similarities and differences between the concepts with respect to how animals gather and use social information, the type of information gathered, how information is packaged, and the relative payoffs to individuals involved. We contend that none of the currently dominant paradigms, social learning, public information use, or social eavesdropping, provide a unifying theme for studying social information use. Instead, we favour the central characteristic of the three concepts, social information use, as the overarching umbrella, and advocate a broader conceptual framework for understanding more comprehensively how animals behave with their social environments. Our intention is not to revolutionize the fields of social learning, public information use or social eavesdropping, but rather to stimulate discussion among researchers investigating the abilities of animals to extract information from the social environment.

Abstract: The question whether a certain species is or is not able to imitate has received much recent attention. However, the ability to copy a demonstrated action might depend not only on the imitative ability of the observer but also on its attention, a factor widely neglected in discussions and experiments. Since attention differs between species as well as between individuals, it is likely to influence the amount and type of information different species and/or observers may extract from a given demonstration. We studied attention in common marmosets, Callithrix jacchus. In a series of experimental sessions, individual marmosets watched different conspecific models that were searching, manipulating an object and feeding. The observers could see the demonstration through two observation holes, which allowed us to measure exactly how often and for how long they watched the model. Marmosets were more attentive towards conspecifics engaged in a problem-solving task than an exploring model. Individual variation in attention was large, ranging from less than 10% to over 49%. Attention also depended on the identity of the model with an increase in attention if it was of the opposite sex. Overall, attention of marmosets was short with a median duration of 6 s while the model manipulated an object. This study measured the real interest of the observer towards a model, which could be an important variable in social-learning experiments.