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Fraser, O. N., & Bugnyar, T. (2010). Do Ravens Show Consolation? Responses to Distressed Others. PLoS ONE, 5(5), e10605.
Abstract: <sec> <title>Background</title> <p>Bystander affiliation (post-conflict affiliation from an uninvolved bystander to the conflict victim) may represent an expression of empathy in which the bystander consoles the victim to alleviate the victim's distress (“consolation”). However, alternative hypotheses for the function of bystander affiliation also exist. Determining whether ravens spontaneously offer consolation to distressed partners may not only help us to understand how animals deal with the costs of aggressive conflict, but may also play an important role in the empathy debate.</p> </sec><sec> <title>Methodology/Principal findings</title> <p>This study investigates the post-conflict behavior of ravens, applying the predictive framework for the function of bystander affiliation for the first time in a non-ape species. We found weak evidence for reconciliation (post-conflict affiliation between former opponents), but strong evidence for both bystander affiliation and solicited bystander affiliation (post-conflict affiliation from the victim to a bystander). Bystanders involved in both interactions were likely to share a valuable relationship with the victim. Bystander affiliation offered to the victim was more likely to occur after intense conflicts. Renewed aggression was less likely to occur after the victim solicited affiliation from a bystander.</p> </sec><sec> <title>Conclusions/Significance</title> <p>Our findings suggest that in ravens, bystanders may console victims with whom they share a valuable relationship, thus alleviating the victims' post-conflict distress. Conversely victims may affiliate with bystanders after a conflict in order to reduce the likelihood of renewed aggression. These results stress the importance of relationship quality in determining the occurrence and function of post-conflict interactions, and show that ravens may be sensitive to the emotions of others.</p> </sec>
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Bugnyar, T., & Heinrich, B. (2006). Pilfering ravens, Corvus corax, adjust their behaviour to social context and identity of competitors. Anim. Cogn., 9(4), 369–376.
Abstract: Like other corvids, food-storing ravens protect their caches from being pilfered by conspecifics by means of aggression and by re-caching. In the wild and in captivity, potential pilferers rarely approach caches until the storers have left the cache vicinity. When storers are experimentally prevented from leaving, pilferers first search at places other than the cache sites. These behaviours raise the possibility that ravens are capable of withholding intentions and providing false information to avoid provoking the storers' aggression for cache protection. Alternatively, birds may refrain from pilfering to avoid conflicts with dominants. Here we examined whether ravens adjust their pilfer tactics according to social context and type of competitors. We allowed birds that had witnessed a conspecific making caches to pilfer those caches either in private, together with the storer, or together with a conspecific bystander that had not created the caches (non-storer) but had seen them being made. Compared to in-private trials, ravens delayed approaching the caches only in the presence of storers. Furthermore, they quickly engaged in searching away from the caches when together with dominant storers but directly approached the caches when together with dominant non-storers. These findings demonstrate that ravens selectively alter their pilfer behaviour with those individuals that are likely to defend the caches (storers) and support the interpretation that they are deceptively manipulating the others' behaviour.
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Heinrich, B., & Bugnyar, T. (2007). Just how smart are ravens? Sci Am, 296(4), 64–71.
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Schloegl, C., Kotrschal, K., & Bugnyar, T. (2007). Gaze following in common ravens, Corvus corax: ontogeny and habituation. Anim. Behav., 74(4), 769–778.
Abstract: Co-orientation with others by using their gaze direction is considered to be adaptive for detecting food or predators or monitoring social interactions. Like the great apes, common ravens are capable of following human experimenters' gaze direction not only into distant space but also behind visual barriers. We investigated the ontogenetic development of these abilities by confronting birds with a human foster parent looking up (experiment 1) and behind visual barriers (experiment 3) and their modification by habituation (experiments 2 and 4). We tested a group of 12 hand-reared ravens during their first 10 months of life. Ravens responded to others' look-ups soon after fledging but could track their gaze behind a visual barrier only 4 months later, at the age they usually become independent from their parents. Furthermore, ravens quickly ceased responding to repeated look-ups by the model, but did not habituate to repeated gaze cues directed behind a barrier. Our findings support the idea that the two modes of gaze following reflect different cognitive levels in ravens and, possibly, have different functions.
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Bugnyar, T., & Kotrschal, K. (2004). Leading a conspecific away from food in ravens ( Corvus corax)? Anim. Cogn., 7(2), 69–76.
Abstract: Active misleading of conspecifics has been described as a social strategy mainly for primates. Here we report a raven leading a competitor away from food in a social foraging task. Four individuals had to search and compete for hidden food at color-marked clusters of artificial food caches. At the beginning of the experiment, a subordinate male found and exploited the majority of the food. As a result, the dominant male displaced him from the already opened boxes. The subordinate male then developed a pattern, when the loss of reward to the dominant got high, of moving to unrewarded clusters and opening boxes there. This diversion often led the dominant to approach those unrewarded clusters and the subordinate then had a head start for exploiting the rewarded boxes. Subsequently, however, the dominant male learned not to follow the subordinate to unrewarded clusters and eventually started searching for the reward himself. These interactions between the two males illustrate the ravens' potential for deceptively manipulating conspecifics. We discuss under which circumstances ravens might use this capacity.
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Bugnyar, T., & Kotrschal, K. (2001). Movement coordination and signalling in ravens (Corvus corax): an experimental field study. Acta. Ethol., 3(2), 101–109.
Abstract: Vagrant non-breeding ravens frequently attract conspecifics to rich ephemeral food sources. There, grouping may allow them to overcome the defence of territorial breeders. Here, we focus on ravens making use of regular food supplies in a game park, where they divert food from the provision of park animals. We investigated if ravens foraging in the Cumberland game park (Grünau, Austria) are attentive towards one another when they experience some unpredictability in food provisioning. We confronted a group of 30-50 ravens with two different treatments. Ten minutes ahead of the feeding of either wolves or wild boars we showed buckets containing pieces of meat to the ravens flying overhead. In the reliable cue treatment (RCT), the meat was placed next to one of the two enclosures, whereas in the unreliable cue treatment (UCT), the buckets were placed simultaneously in front of both enclosures though only in one of the enclosures were the animals fed 10 min later. Thus, during RCT but not during UCT, ravens could predict where food would become available. Only during UCT, ravens moved in large groups between the two feeding sites. Many ravens moving at the same time in the same direction may indicate some co-ordination in space and time, which is most likely achieved by social attraction among individuals. Furthermore, the number of ravens approaching and leaving, respectively, a feeding site cross-correlated with a temporary increase in the rate of a food-associated call, the yell. This suggests that in addition to watching each other, calling may have contributed to group formation. Possible benefits of group formation during food inspection are discussed.
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Bugnyar, T., & Kotrschal, K. (2002). Observational learning and the raiding of food caches in ravens, Corvus corax: is it `tactical' deception? Anim. Behav., 64(2), 185–195.
Abstract: Group-foraging ravens scatter-hoard when they are competing for food and, to some extent, also raid the caches made by others. We investigated the effects of observational spatial memory on individual caching and raiding tactics. With captive ravens, we found visual observation was essential for locating and raiding the caches of conspecifics. Both captive and free-ranging ravens, food cachers as well as potential cache raiders, responded to each other's presence. Cachers withdrew from conspecifics and most often placed their caches behind structures, obstructing the view of potential observers. Raiders watched inconspicuously and kept at a distance to cachers close to their cache sites. In response to the presence of potential raiders or because of their initial movements towards caches, the cachers frequently interrupted caching, changed cache sites, or recovered their food items. These results suggest that ravens, regardless of whether they act as cachers or raiders, are capable of withholding information about their intentions and, hence, manipulate the other bird's attention either to prevent or to achieve social-learning opportunities. Such interactions may qualify as `tactical' deception and may have created a considerable pressure selecting for social cognition in ravens. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.
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Schwab, C., Bugnyar, T., Schloegl, C., & Kotrschal, K. (2008). Enhanced social learning between siblings in common ravens, Corvus corax. Anim. Behav., 75(2), 501–508.
Abstract: It has been suggested that social dynamics affect social learning but empirical support for this idea is scarce. Here we show that affiliate relationships among kin indeed enhance the performance of common ravens, Corvus corax, in a social learning task. Via daily behavioural protocols we first monitored social dynamics in our group of captive young ravens. Siblings spent significantly more time in close proximity to each other than did nonsiblings. We subsequently tested birds on a stimulus enhancement task in model-observer dyads composed of both siblings and nonsiblings. During demonstration the observer could watch the model manipulating one particular object (target object) in an adjacent room. After removing the model, the observer was confronted with five different objects including the former target object. Observers from sibling dyads handled the target object for significantly longer periods of time as compared with the other four available objects, whereas observers from nonsibling dyads did not show a preference for the target object. Also, siblings matched the model's decision to cache or not to cache objects significantly more often than did nonsiblings. Hence, siblings were likely to attend to both, the behaviour of the model (caching or noncaching) and object-specific details. Our results support the hypothesis that affiliate relations between individuals affect the transmission of information and may lead to directed social learning even when spatial proximity has been experimentally controlled for.
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Schmidt, J., Scheid, C., Kotrschal, K., Bugnyar, T., & Schloegl, C. (2011). Gaze direction – A cue for hidden food in rooks (Corvus frugilegus)? Behavioural Processes, 88(2), 88–93.
Abstract: Other individual's head- and eye-directions can be used as social cues indicating the presence of important events. Among birds, ravens and rooks have been shown to co-orient with conspecifics and with humans by following their gaze direction into distant space and behind visual screens. Both species use screens to cache food in private; also, it had been suggested that they may rely on gaze cues to detect hidden food. However, in an object-choice task, ravens failed to do so, and their competitive lifestyle may have prevented them from relying on these cues. Here we tested closely related and cooperative rooks. Food was hidden in one of two cups and the experimenter gazed at the baited cup. In a second experiment, we aimed to increase the birds’ motivation to choose correctly by increasing the investment needed to obtain the reward. To do so, the birds had to pull on a string to obtain the cup. Here, the birds as a group tended to rely on gaze cues. In addition, individual birds quickly learned to use the cue in both experiments. Although rooks may not use gaze cues to find hidden food spontaneously, they may quickly learn to do so.
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Bugnyar, T. (2011). Knower–guesser differentiation in ravens: others' viewpoints matter. Proc. Roy. Soc. Lond. B Biol. Sci., 278(1705), 634–640.
Abstract: Differentiating between individuals with different knowledge states is an important step in child development and has been considered as a hallmark in human evolution. Recently, primates and corvids have been reported to pass knower–guesser tasks, raising the possibility of mental attribution skills in non-human animals. Yet, it has been difficult to distinguish ‘mind-reading’ from behaviour-reading alternatives, specifically the use of behavioural cues and/or the application of associatively learned rules. Here, I show that ravens (Corvus corax) observing an experimenter hiding food are capable of predicting the behaviour of bystanders that had been visible at both, none or just one of two caching events. Manipulating the competitors' visual field independently of the view of the test-subject resulted in an instant drop in performance, whereas controls for behavioural cues had no such effect. These findings indicate that ravens not only remember whom they have seen at caching but also take into account that the other's view was blocked. Notably, it does not suffice for the birds to associate specific competitors with specific caches. These results support the idea that certain socio-ecological conditions may select for similar cognitive abilities in distantly related species and that some birds have evolved analogous precursors to a human theory-of-mind.
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