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Scheiber, I. B. R., WeiB, B. M., Hirschenhauser, K., Wascher, C. A. F., Nedelcu, J. T., & Kotrschal, K. (2008). Does 'Relationship Intelligence' Make Big Brains in Birds? Open Biol J, 1, 6–8 (3).
Abstract: Lately, Emery et al. developed a bird-specific modification of the *social brain hypothesis*, termed *relationship intelligence hypothesis*. Although the idea may be valuable, we doubt that it is supported by sufficient evidence and critically discuss some of the arguments raised by the authors in favour of their new idea.
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Range, F., Bugnyar, T., Schlogl, C., & Kotrschal, K. (2006). Individual and sex differences in learning abilities of ravens. Behav. Process., 73(1), 100–106.
Abstract: Behavioral and physiological characteristics of individuals within the same species have been found to be stable across time and contexts. In this study, we investigated individual differences in learning abilities and object and social manipulation to test for consistency within individuals across different tasks. Individual ravens (Corvus corax) were tested in simple color and position discrimination tasks to establish their learning abilities. We found that males were significantly better in the acquisition of the first discrimination task and the object manipulation task, but not in any of the other tasks. Furthermore, faster learners engaged less often in manipulations of conspecifics and exploration of objects to get access to food. No relationship between object and social manipulation and reversal training were found. Our results suggest that individual differences in regard to the acquisition of new tasks may be related to personalities or at least object manipulation in ravens.
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Schloegl, C., Kotrschal, K., & Bugnyar, T. (2008). Modifying the object-choice task: Is the way you look important for ravens? Behav. Process., 77(1), 61–65.
Abstract: Most animals seem to have difficulties in using gaze cues to find hidden food in object-choice tasks. For instance, chimpanzees usually fail in these tests, even though they are capable of following other's gaze geometrically behind barriers. Similar to chimpanzees, common ravens are skilled in tracking other's gaze but fail in object-choice tasks. We here explored whether procedural modifications, which had been used successfully in chimpanzees, would also yield positive results in ravens. In our modifications (a) the experimenter approached the cup while gazing at it, (b) the gaze cue was accompanied by a sound and (c) the experimenter could actually see the food while giving the gaze cue. Two out of seven birds performed above chance level in some of these conditions. However, we ascribe this improvement to the individuals' learning ability rather than to an understanding of the communicative nature of the task. This interpretation is further supported by results of a follow-up experiment suggesting that ravens may not rely on conspecifics' gaze cues for finding food caches in a natural foraging context. In sum, our results suggest that ravens may not transfer their gaze follow abilities to foraging situations involving hidden food.
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Wascher, C. A. F., Fraser, O. N., & Kotrschal, K. (2010). Heart Rate during Conflicts Predicts Post-Conflict Stress-Related Behavior in Greylag Geese. PLoS ONE, 5(12), e15751.
Abstract: Background
Social stressors are known to be among the most potent stressors in group-living animals. This is not only manifested in individual physiology (heart rate, glucocorticoids), but also in how individuals behave directly after a conflict. Certain ‘stress-related behaviors’ such as autopreening, body shaking, scratching and vigilance have been suggested to indicate an individual's emotional state. Such behaviors may also alleviate stress, but the behavioral context and physiological basis of those behaviors is still poorly understood.
Methodology/Principal Findings
We recorded beat-to-beat heart rates (HR) of 22 greylag geese in response to agonistic encounters using fully implanted sensor-transmitter packages. Additionally, for 143 major events we analyzed the behavior shown by our focal animals in the first two minutes after an interaction. Our results show that the HR during encounters and characteristics of the interaction predicted the frequency and duration of behaviors shown after a conflict.
Conclusions/Significance
To our knowledge this is the first study to quantify the physiological and behavioral responses to single agonistic encounters and to link this to post conflict behavior. Our results demonstrate that ‘stress-related behaviors’ are flexibly modulated by the characteristics of the preceding aggressive interaction and reflect the individual's emotional strain, which is linked to autonomic arousal. We found no support for the stress-alleviating hypothesis, but we propose that stress-related behaviors may play a role in communication with other group members, particularly with pair-partners.
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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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Kralj-Fiser, S., Scheiber, I. B. R., Blejec, A., Moestl, E., & Kotrschal, K. (2007). Individualities in a flock of free-roaming greylag geese: behavioral and physiological consistency over time and across situations. Horm Behav, 51(2), 239–248.
Abstract: The concept of personality implies individual differences in behavior and physiology that show some degree of repeatability/consistency over time and across contexts. Most studies of animal personality, particularly studies of individuals' variation in physiological mechanisms, have been conducted on selected individuals in controlled conditions. We attempted to detect consistent behaviors as well as physiological patterns in greylag ganders (Anser anser) from a free-roaming flock living in semi-natural conditions. We tested 10 individuals repeatedly, in a handling trial, resembling tests for characterization of “temperaments” in captive animals. We recorded the behavior of the same 10 individuals during four situations in the socially intact flock: (1) a “low density feeding condition”, (2) a “high density feeding condition”, (3) a “low density post-feeding situation” and (4) while the geese rested. We collected fecal samples for determination of excreted immuno-reactive corticosterone (BM) and testosterone metabolites (TM) after handling trials, as well as the “low density feeding” and the “high density feeding” conditions. BM levels were very highly consistent over the repeats of handling trials, and the “low density feeding condition” and tended to be consistent over the first two repeats of the “high density feeding condition”. Also, BM responses tended to be consistent across contexts. Despite seasonal variation, there tended to be inter-test consistency of TM, which pointed to some individual differences in TM as well. Aggressiveness turned out to be a highly repeatable trait, which was consistent across social situations, and tended to correlate with an individual's resistance during handling trials. Also, “proximity to the female partner” and “sociability” – the average number of neighboring geese in a close distance while resting – were consistent. We conclude that aggressiveness, “affiliative tendencies” and levels of excreted corticosterone and testosterone metabolites may be crucial factors of personality in geese.
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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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