Claudia A. F. Wascher

Corvids can decide if a future exchange is worth waiting for

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.

Waiting for better, not for more: corvids respond to quality in two delay maintenance tasks

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.

Heart Rate Modulation by Social Contexts in Greylag Geese (Anser anser)

Physical activity is generally considered as most relevant for modulating heart rate (HR). The authors show here that HR is not only modulated by physical activity but even more by social contexts. HR modulation in three free-ranging, socially embedded, male greylag geese fitted with implanted radiotransmitters was investigated. Measured HR ranged from 40 beats per minute (bpm) during rest to a maximum over 400 bpm during takeoff. Almost the same maximum HRs (400 bpm) were reached during social interactions, which however, generally require less bodily action. Mean HR during social interactions (agonistic interactions, vocalizations) was significantly higher than during behaviors with a less obvious social context (e.g., resting, comfort or feeding behavior), but with comparable physical activity involved. The authors also found significant and consistent differences in HR between the three focal individuals, probably because of individual behavioral phenotype. Our results show that social context has a strong modulatory effect on the sympathico-adrenergic activity in a social bird and conclude that particularly the latter may pose considerable energetic costs. (PsycINFO Database Record (c) 2008 APA, all rights reserved).

You sound familiar: carrion crows can differentiate between the calls of known and unknown heterospecifics.

In group-living animals, it is adaptive to recognize conspecifics on the basis of familiarity or group membership as it allows association with preferred social partners and avoidance of competitors. However, animals do not only associate with conspecifics but also with heterospecifics, for example in mixed-species flocks. Consequently, between-species recognition, based either on familiarity or even individual recognition, is likely to be beneficial. The extent to which animals can distinguish between familiar and unfamiliar heterospecifics is currently unclear. In the present study, we investigated the ability of eight carrion crows to differentiate between the voices and calls of familiar and unfamiliar humans and jackdaws. The crows responded significantly more often to unfamiliar than familiar human playbacks and, conversely, responded more to familiar than unfamiliar jackdaw calls. Our results provide the first evidence that birds can discriminate between familiar and unfamiliar heterospecific individuals using auditory stimuli.

Did tool-use evolve with enhanced physical cognitive abilities?

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.

Heart rate responses to agonistic encounters in greylag geese, Anser anser

As in most social groups, agonistic interactions of various intensities are common in a goose flock. This may cause social stress, modulating heart rate (HR), which may serve as a measure of energetic investment and also of individual emotional involvement. We investigated HR responses to social encounters in 24 free-living greylag geese in an intact social environment. We recorded 1602 social interactions of various intensities in which the focal individual either attacked another member of the flock or was attacked itself. We analysed five HR parameters (mean HR, maximum HR, HR increase, duration until maximum, time until the baseline value was reached again). Generally, HR scaled positively with increasing intensity of agonistic interactions as well as with increasing duration. Individuals showed higher HRs when attacking than when being attacked. In addition, focal individuals responded with a greater HR increase when confronted with an opponent winning a higher percentage of interactions than itself. Repeated agonistic interactions against a specific opponent were related to greater HR responses than single events and focal individuals responded more strongly to male opponents than to females. Our results indicate a differential HR response depending on the intensity and duration of an interaction as well as the identity of the opponent. This differential physiological investment may reflect differences in emotional involvement depending on the social context of a particular agonistic interaction.

Heart rate responses to induced challenge situations in greylag geese (Anser anser)

Adequate short-term responses to stressors are of great importance for the health and well-being of individuals and factors modulating the physiological stress response (e.g., controllability, suddenness, familiarity) of a stimulus are well described under laboratory conditions. In the present study we aimed at investigating the stress response in greylag geese (Anser anser) in the field, confronting individuals with naturally occurring stressors. We measured beat-to-beat heart rate (HR) via fully implanted transmitters during three different experimental challenges: (1) catching and holding, (2) confrontation with a model predator, and (3) approach by different humans. We compared this to a control period and HR during agonistic encounters, a naturally occurring stressor. All three experimental situations evoked a HR increase. Highest HR responses were elicited by catching and holding the animals. In the third experiment, HR responses were greatest when the geese were approached by a human stranger (i.e., somebody the geese have never seen before). Hence, geese discriminated between different kinds of stressors and adjusted their physiological response depending on the type of stressor. Our results show that geese were able to discriminate between individual humans. In line with a number of lab studies, we suggest that particularly the controllability of certain situation determines the intensity of the HR response, also in a natural setting in the field.

Heart Rate during Conflicts Predicts Post-Conflict Stress-Related Behavior in Greylag Geese

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 individuals 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 individuals 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.

With whom to dine? Ravens' responses to food-associated calls depend on individual characteristics of the caller

Upon discovering food, common ravens, Corvus corax, produce far-reaching ‘haa’ calls or yells, which are individually distinct and signal food availability to conspecifics. Here, we investigated whether ravens respond differently to ‘haa’ calls of known and unknown individuals. In a paired playback design, we tested responses to ‘haa’ call sequences in a group containing individually marked free-ranging ravens. We simultaneously played call sequences of a male and a female raven in two different locations and varied familiarity (known or unknown to the local group). Ravens responded strongest to dyads containing familiar females, performing more scan flights above and by perching in trees near the respective speaker. Acoustic analysis of the calls used as stimuli showed no sex-, age- or familiarity-specific acoustic cues, but highly significant classification results at the individual level. Taken together, our findings indicate that ravens respond to individual characteristics in ‘haa’ calls, and choose whom to approach for feeding, i.e. join social allies and avoid dominant conspecifics. This is the first study to investigate responses to ‘haa’ calls under natural conditions in a wild population containing individually marked ravens.

Familiarity with the experimenter influences the performance of Common ravens (Corvus corax) and Carrion crows (Corvus corone corone) in cognitive tasks.

Highlights • We compared the results of corvids in experiments with familiar/unfamiliar humans.• We investigated behavioural reactions towards familiar and unfamiliar humans.• Corvids performed significantly better in experiments with familiar humans.• Corvids did not show more neophobia towards unfamiliar humans.• Hence, familiarity positively affected the experimental performance of corvids.