Llamas use social information from conspecifics and humans to solve a spatial detour task.

Learning by observing others (i.e. social learning) is an important mechanism to reduce the costs of individual learning. Social learning can occur between conspecifics but also heterospecifics. Domestication processes might have changed the animals' sensitivity to human social cues and recent research indicates that domesticated species are particularly good in learning socially from humans. Llamas (Lama glama) are an interesting model species for that purpose. Llamas were bred as pack animals, which requires close contact and cooperative behaviour towards humans. We investigated whether llamas learn socially from trained conspecifics and humans in a spatial detour task. Subjects were required to detour metal hurdles arranged in a V-shape to reach a food reward. Llamas were more successful in solving the task after both a human and a conspecific demonstrated the task compared to a control condition with no demonstrator. Individual differences in behaviour (i.e. food motivation and distraction) further affected the success rate. Animals did not necessarily use the same route as the demonstrators, thus, indicating that they adopted a more general detour behaviour. These results suggest that llamas can extract information from conspecific and heterospecific demonstrations; hence, broadening our knowledge of domesticated species that are sensitive to human social behaviour.

Horses wait for more and better rewards in a delay of gratification paradigm.

Self-control, defined as the ability to forgo immediate satisfaction in favor of better pay-offs in the future, has been extensively studied, revealing enormous variation between and within species. Horses are interesting in this regard because as a grazing species they are expected to show low self-control whereas its social complexity might be linked to high self-control abilities. Additionally, self-control may be a key factor in training and/or coping with potentially stressful husbandry conditions. We assessed horses' self-control abilities in a simplified delay of gratification test that can be easily implemented in a farm setting. In Experiment 1, we gave horses (N = 52) the choice between an immediately available low-quality reward and a delayed high-quality reward that could only be obtained if the horse refrained from consuming the immediate reward. Different experimenters (N = 30) that underwent prior training in the procedures, tested horses in two test phases either with their eyes visible or invisible (sunglasses). Twenty horses waited up to the maximum delay stage of 60 s while all horses performed worse in the second test phase. In Experiment 2, we improved the test procedure (i.e., one experimenter, refined criterion for success), and tested 30 additional horses in a quality and quantity condition (one reward vs. delayed bigger reward). Two horses successfully waited for 60 s (quality: N = 1, quantity: N = 1). Horses tolerated higher delays, if they were first tested in the quantity condition. Furthermore, horses that were fed hay ad libitum, instead of in a restricted manner, reached higher delays. Coping behaviors (e.g., looking away, head movements, pawing, and increasing distance to reward) facilitated waiting success and horses were able to anticipate the upcoming delay duration as indicated by non-random distributions of giving-up times. We found no correlations between owner-assessed traits (e.g., trainability and patience) and individual performance in the test. These results suggest that horses are able to exert self-control in a delay of gratification paradigm similar to other domesticated species. Our simplified paradigm could be used to gather large scale data, e.g., to investigate the role of self-control in trainability or success in equestrian sports.

Intra- and interspecific variation in self-control capacities of parrots in a delay of gratification task.

Forgoing immediate satisfaction for higher pay-offs in the future (delayed gratification) could be adaptive in situations that wild animals may encounter. To explain species-differences in self-control, hypotheses based on social complexity, feeding ecology, brain size and metabolic rate have been proposed. To explore these hypotheses in a comparative setting, we tested three macaw species (neotropical parrots)-great green macaws (N = 8), blue-throated macaws (N = 6), blue-headed macaws (N = 6)-and the distantly related African grey parrots (afrotropical parrots; N = 8) in a modified rotating tray task, in which subjects are required to inhibit consuming a constantly available low-quality reward in favour of a high-quality reward that becomes available only after an increasing delay (min. 5 s, max. 60 s). All four species successfully waited for a minimum of 8.3 s ± 11.7 s (group level mean ± SD) with African greys reaching a delay of 29.4 ± 15.2 s, and great green macaws-as best performing macaw species-tolerating delays of 20 s ± 8 s. The best performing African grey individual reached a maximum delay of 50 s, whereas, a great green and a blue-throated macaw tolerated a delay of 30 s max. Females tolerated higher maximum delays than males. Engaging in distraction behaviours enhanced waiting performance across species and all birds were able to anticipate the waiting duration. Our results suggest that both feeding and socio-ecological complexity may be a factor in self-control, but further systematically collected comparative data on self-control of different (parrot) species are required to test the evolutionary hypotheses rigorously.

No evidence for a relationship between breed cooperativeness and inequity aversion in dogs.

Inequity aversion, the resistance to inequitable outcomes, has been demonstrated in a wide variety of animal species. Inequity aversion was hypothesised to have co-evolved with cooperation but only limited evidence supports this. Dogs provide a suitable model species to test this hypothesis as dogs were previously shown to be inequity averse and dog breeds vary in the extent to which they were selected for cooperativeness. Here, we compared the response of 12 individuals of "cooperative worker" breeds with that of 12 individuals of "independent worker" breeds in the "paw task" previously used to demonstrate inequity aversion in dogs. We also compared the two breed groups' subsequent social behaviours in a food tolerance test and free interaction session. Although subjects in both breed groups were inequity averse, we found no considerable difference between the groups in the extent of the negative response to inequity or in the impact of the inequity on subsequent social behaviours. However, we found differences between the breed groups in the response to reward omission with cooperative breeds tending to work for longer than independent breeds. Additionally, in the free interaction session, individuals of cooperative breeds spent more time in proximity to their partner in the baseline condition than individuals of independent breeds. Overall, our results do not provide support for the hypothesis that inequity aversion and cooperation co-evolved. However, they illuminate potential differences in selection pressures experienced by cooperative worker and independent worker dog breeds throughout their evolutionary history.

Dogs wait longer for better rewards than wolves in a delay of gratification task: but why?

Self-control has been shown to be linked with being cooperative and successful in humans and with the g-factor in chimpanzees. As such, it is likely to play an important role in all forms of problem-solving. Self-control, however, does not just vary across individuals but seems also to be dependent on the ecological niche of the respective species. With dogs having been selected to live in the human environment, several domestication hypotheses have predicted that dogs are better at self-control and thus more tolerant of longer delays than wolves. Here we set out to test this prediction by comparing dogs' and wolves' self-control abilities using a delay of gratification task where the animals had to wait for a predefined delay duration to exchange a low-quality reward for a high-quality reward. We found that in our task, dogs outperformed the wolves waiting an average of 66 s vs. 24 s in the wolves. Food quality did not influence how long the animals waited for the better reward. However, dogs performed overall better in motivation trials than the wolves, although the dogs' performance in those trials was dependent on the duration of the delays in the test trials, whereas this was not the case for the wolves. Overall, the data suggest that selection by humans for traits influencing self-control rather than ecological factors might drive self-control abilities in wolves and dogs. However, several other factors might contribute or explain the observed differences including the presence of the humans, which might have inhibited the dogs more than the wolves, lower motivation of the wolves compared to the dogs to participate in the task and/or wolves having a better understanding of the task contingencies. These possible explanations need further exploration.

Parrots Voluntarily Help Each Other to Obtain Food Rewards.

Helping others to obtain benefits, even at a cost to oneself, poses an evolutionary puzzle [1]. While kin selection explains such "selfless" acts among relatives, only reciprocity (paying back received favors) entails fitness benefits for unrelated individuals [2]. So far, experimental evidence for both prosocial helping (providing voluntary assistance for achieving an action-based goal) and reciprocity has been reported in a few mammals but no avian species [3]. In order to gain insights into the evolutionary origins of these behaviors, the capacity of non-mammalian species for prosociality and for reciprocity needs to be investigated. We tested two parrot species in an instrumental-helping paradigm involving "token transfer." Here, actors could provide tokens to their neighbor, who could exchange them with an experimenter for food. To verify whether the parrots understood the task's contingencies, we systematically varied the presence of a partner and the possibility for exchange. We found that African grey parrots voluntarily and spontaneously transferred tokens to conspecific partners, whereas significantly fewer transfers occurred in the control conditions. Transfers were affected by the strength of the dyads' affiliation and partially by the receivers' attention-getting behaviors. Furthermore, the birds reciprocated the help once the roles were reversed. Blue-headed macaws, in contrast, transferred hardly any tokens. Species differences in social tolerance might explain this discrepancy. These findings show that instrumental helping based on a prosocial attitude, accompanied but potentially not sustained by reciprocity, is present in parrots, suggesting that this capacity evolved convergently in this avian group and mammals.

Parrots do not show inequity aversion.

Inequity aversion, the negative reaction to unequal treatment, is considered a mechanism for stabilizing cooperative interactions between non-kin group members. However, this might only be adaptive for species that switch cooperative partners. Utilizing a comparative approach, inequity aversion has been assessed in many mammalian species and recently also in corvids and one parrot species, kea, revealing mixed results. To broaden our knowledge about the phylogenetic distribution of inequity aversion, we tested four parrot species in the token exchange paradigm. We varied the quality of rewards delivered to dyads of birds, as well as the effort required to obtain a reward. Blue-headed macaws and African grey parrots showed no reaction to being rewarded unequally. The bigger macaws were less willing to exchange tokens in the "unequal" condition compared to the "equal high" condition in which both birds obtained high quality rewards, but a closer examination of the results and the findings from the control conditions reveal that inequity aversion does not account for it. None of the species responded to inequity in terms of effort. Parrots may not exhibit inequity aversion due to interdependence on their life-long partner and the high costs associated with finding a new partner.

Dogs and wolves do not differ in their inhibitory control abilities in a non-social test battery.

Being able to inhibit certain behaviours is of clear advantage in various situations. In particular, it has been suggested that inhibitory control plays a role in problem-solving and cooperation. Interspecific differences in the capacity for inhibitory control have been attributed to social and ecological factors, while one additional factor, namely domestication, has received only little attention so far. Dogs are an interesting species to test the effects of socio-ecological factors and also the influence of domestication on inhibitory control abilities. While dogs might have been selected for enhanced inhibition skills during domestication, the predictions derived from their socio-ecological background are reversed. Wolves are cooperative hunters and breeders, while dogs predominately scavenge and raise their young alone, accordingly, it would be predicted that dogs show impaired inhibitory control abilities since they no longer rely on these coordinated actions. To test these hypotheses, we assessed inhibitory control abilities in dogs and wolves raised and kept under similar conditions. Moreover, considering the problem of context-specificity in inhibitory control measures, we employed a multiple-test-approach. In line with previous studies, we found that the single inhibition tests did not correlate with each other. Using an exploratory approach, we found three components that explained the variation of behaviours across tests: motivation, flexibility, and perseveration. Interestingly, these inhibition components did not differ between dogs and wolves, which contradicts the predictions based on their socio-ecological backgrounds but also suggests that at least in tasks with minimal human influence, domestication did not affect dogs' inhibitory control abilities, thus raising questions in regard to the selection processes that might have affected inhibitory control abilities during the course of domestication.

Assessing African grey parrots' prosocial tendencies in a token choice paradigm.

Prosociality is defined as a voluntary, typically low-cost behaviour that benefits another individual. Social tolerance has been proposed as a potential driver for its evolution, both on the proximate and on the ultimate level. Parrots are an interesting species to study such other-regarding behaviours, given that they are highly social and stand out in terms of relative brain size and cognitive capacity. We tested eight African grey parrots in a dyadic prosocial choice test. They faced a choice between two different tokens, a prosocial (actor and partner rewarded) and a selfish (only actor rewarded) one. We found that the birds did not behave prosocially when one subject remained in the actor role; however, when roles were alternated, the birds' prosocial choices increased. The birds also seemed to reciprocate their partner's choices, given that a contingency between choices was observed. If the food provisioned to the partner was of higher quality than that the actor obtained, actors increased their willingness to provide food to their partner. Nonetheless, the control conditions suggest that the parrots did not fully understand the task's contingencies. In sum, African grey parrots show the potential for prosociality and reciprocity; however, considering their lack of understanding of the contingencies of the particular tasks used in this study, the underlying motivation for the observed behaviour remains to be addressed by future studies, in order to elucidate the phylogenetic distribution of prosociality further.

Dogs' reaction to inequity is affected by inhibitory control.

Inequity aversion is thought to act as a mechanism to ensure cooperation and has been studied in many different species, consistently revealing inter-individual variation. Inhibitory control has been proposed to act as one factor responsible for this variation since individuals need to inhibit performing the required action and/or refuse rewards in order to exhibit inequity aversion. Here, we investigated if dogs' sensitivity to inequity is affected by their capacity for inhibitory control, assessed in a test battery and questionnaire. Overall, dogs showing high compulsivity scores (i.e. repetitive behaviours independent of feedback) were more motivated to participate in the inequity task independent of the rewarding scheme. Dogs were more sensitive to inequity and individual contrast if they exhibited a slower decision speed in the inhibition tasks. Furthermore, less persistent and more impulsive dogs were more sensitive to reward inequity, potentially due to having a lower tolerance level for frustration. Results indicate that aspects of inhibitory control can explain the variation in dogs' inequity response, highlighting one of the mechanisms underlying responses to inequity. Emphasising the importance to design paradigms, which allow us to disentangle capacities to recognise inequity from the inability to react to it due to poor inhibitory control abilities.

Measures of Dogs' Inhibitory Control Abilities Do Not Correlate across Tasks.

Inhibitory control, the ability to overcome prepotent but ineffective behaviors, has been studied extensively across species, revealing the involvement of this ability in many different aspects of life. While various different paradigms have been created in order to measure inhibitory control, only a limited number of studies have investigated whether such measurements indeed evaluate the same underlying mechanism, especially in non-human animals. In humans, inhibitory control is a complex construct composed of distinct behavioral processes rather than of a single unified measure. In the current study, we aimed to investigate the validity of inhibitory control paradigms in dogs. Sixty-seven dogs were tested in a battery consisting of frequently used inhibitory control tests. Additionally, dog owners were asked to complete an impulsivity questionnaire about their dog. No correlation of dogs' performance across tasks was found. In order to understand whether there are some underlying behavioral aspects explaining dogs' performance across tests, we performed principle component analyses. Results revealed that three components (persistency, compulsivity and decision speed) explained the variation across tasks. The questionnaire and dogs' individual characteristics (i.e., age and sex) provided only limited information for the derived components. Overall, results suggest that no unique measurement for inhibitory control exists in dogs, but tests rather measure different aspects of this ability. Considering the context-specificity of inhibitory control in dogs and most probably also in other non-human animals, extreme caution is needed when making conclusions about inhibitory control abilities based on a single test.

What Are the Ingredients for an Inequity Paradigm? Manipulating the Experimenter's Involvement in an Inequity Task with Dogs.

Cooperation is only beneficial if the outcome is equally shared between individuals involved in the cooperative interaction. A mechanism to limit the development of unequal cooperation is inequity aversion, the negative reaction to unequal treatment. While inequity aversion has been studied extensively across many animal species, it remains unclear whether inequity aversion elicited in experimental settings is directed to the cooperative partner animal or rather to the experimenter distributing the rewards unequally. In the current study we aimed to further investigate whether the presence of an experimenter distributing rewards is essential in order to elicit inequity aversion in dogs. We tested 22 dog dyads in an inequity task, requiring dyads to alternately press a buzzer in order to receive rewards of equal or unequal value. We manipulated the extent of the experimenter's involvement in the task: in the experimenter-present version an experimenter gave a command to the dogs to press the buzzer and delivered the rewards by pushing the bowls into the dogs' enclosure. In contrast, in the experimenter-absent version, no experimenter was visible and the buzzer and bowls were moved from behind a curtain. We found that dogs did not respond to the unequal treatment regardless of the experimenter's involvement in the task. Nonetheless, we found that dogs based their behavior on frustration and social facilitation in the experimenter-absent version of the task, suggesting that a social interaction with an experimenter may be one aspect necessary to elicit inequity aversion. One potential explanation for the absence of inequity aversion in the experimenter-present version of the task might be the reward delivery method. Using separate sets of reward bowls for each dog instead of a shared bowl could have removed a potentially important competitive aspect (i.e., shared resource) from the inequity paradigm. In addition, delivering the rewards via bowls, rather than directly handing the rewards to the dogs, might have caused dogs to perceive the task as less cooperative. These results suggest that both the presence of an experimenter causing inequity and the inclusion of a competitive or cooperative element in the task may be basic requirements for eliciting inequity aversion.

Reward type and behavioural patterns predict dogs' success in a delay of gratification paradigm.

Inhibiting an immediate behaviour in favour of an alternative but more advantageous behaviour has been linked to individual success in life, especially in humans. Dogs, which have been living in the human environment for thousands of years, are exposed to daily situations that require inhibition different in context from other non-domesticated species. One task regularly used to study inhibitory control is the delay of gratification task, which requires individuals to choose between an immediate option of lower value and a delayed option of higher value. We tested sixteen dogs in a non-social delay of gratification task, conducting two different conditions: a quality and a quantity condition. While the majority of dogs failed to wait for more than 10 s, some dogs tolerated delays of up to 140 s, while one dog waited for 15 minutes. Moreover, dogs had more difficulties to wait if the reward increased in terms of quantity than quality. Interestingly, dogs were able to anticipate the delay duration and some dogs developed behavioural patterns that predicted waiting, which seems similar in some respects to 'coping-strategies' found in children, chimpanzees and parrots. Our results indicate that strategies to cope with impulsivity seem to be consistent and present across animal taxa.

Inequity Aversion Negatively Affects Tolerance and Contact-Seeking Behaviours towards Partner and Experimenter.

Inequity aversion has been proposed to act as a limiting factor for cooperation, thus preventing subjects from disadvantageous cooperative interactions. While a recent study revealed that also dogs show some sensitivity to inequity, the underlying mechanisms of this behaviour are still unclear. The aim of the current study was threefold: 1) to replicate the study by Range et al. (2009, PNAS, 106, 340-345); 2) to investigate the emotional mechanisms involved in the inequity response by measuring the heart rate and 3) to explore the link between inequity aversion and cooperation in terms of behaviours shown towards the partner dog and towards the experimenter who caused the inequity. Dog tested in dyads were alternately asked to give their paw and were either equally or unequally rewarded by the experimenter. After each social test condition, we conducted food tolerance tests and free interaction tests in which the subjects' social behaviour towards the partner and the experimenter were observed. As in the previous study, subjects refused to continue giving their paw when only the partner was rewarded, but not when both dogs were rewarded with rewards of different quality. Although subjects did not react to this quality inequity during the test, we did find reduced durations of food sharing in the subsequent tolerance test, indicating that dogs perceived the inequity but were not able to react to it in the test context. Moreover, subjects avoided their partner and the experimenter more during the free interaction time following unequal compared to equal treatment. Despite the clear behavioural reactions to inequity, we could not detect any changes in heart rate. Results suggest that inequity aversion might in fact be mediated by simple emotional mechanisms: sharing a negative experience, like inequity, might reduce future cooperation by decreasing the likelihood of proximity being maintained between partners.