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Reputation is a key component in social interactions of group-living animals and appears to play a role in the establishment of cooperation. Animals can form reputations of individuals by directly interacting with them or by observing them interact with a third party, i.e., eavesdropping. Previous research has focused on whether dogs (Canis lupus familiaris) can eavesdrop on humans because of their ability to cooperate with humans, however the results are mixed and if they can eavesdrop, it is unknown whether this ability evolved during the domestication process or whether it was inherited from their ancestor, wolves (Canis lupus). Our study investigated whether equally hand-raised, pack-living dogs and wolves can form reputations of humans in a food-giving situation through indirect and/or direct experience. The experimental procedure comprised three parts: baseline (to test whether the subject preferred a person prior to the experiment), observation and testing. In the observation phase, the subject observed two humans interact with a dog demonstrator-one acted generously and fed the dog, and the other acted selfishly and refused to feed the dog. The subject could then choose which person to approach in the test phase. In the following experience phase, the animals interacted directly with the same two humans who behaved either in a generous or selfish manner. Then, they were again given a choice whom to approach. We found that dogs and wolves, at the group level, did not differentiate between a generous or selfish partner after indirect or direct experience, but wolves were more attentive towards the generous person during the observation phase and some dogs and wolves did prefer the generous partner, at least after indirect and direct experience was combined. Our study suggests that reputation formation may be more difficult than expected for animals and we emphasise the importance of context when studying reputation formation in animals.

Eavesdropping is the acquisition of information by observing third-party interactions. Considering dogs' (Canis lupus familiaris) dependence on humans, it would be beneficial for them to eavesdrop on human interactions to choose an appropriate partner to associate with. Previous studies have found that dogs preferred a human who acted generously or cooperatively towards another human over one who acted selfishly or non-cooperatively, however they often did not control for potential location biases. This study controlled for local enhancement and investigated whether dogs derive and act on information about unfamiliar humans through reputation-like inferences by observing third-party interactions. 42 dogs participated in the experiment, which consisted of an observation phase and a test phase. In the observation phase, the animals observed a human with a box of food ask for help to open it from two people-one was helpful and the other was not. The test phase consisted of the impossible task and a choice test. Half of the sample was tested in the experimental condition and the other half was tested in the side control condition, where the two people swapped positions before the test phase. The results of the impossible task showed that dogs only looked at the helpful person first when the people stayed on the same side as they did in the observation phase. In the choice test, dogs chose at random, regardless of whether the people stayed on the same side or swapped positions. Our findings provide tentative support for a local enhancement interpretation of eavesdropping.

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.

Persistence in object manipulation has been consistently associated with problem-solving success and it is known to be affected, at the individual level, by life experience. Differences in life experiences are particularly poorly studied in the problem-solving context and mainly refer to the comparison between wild and captive animals. Dogs represent interesting study subjects, since dog populations differ widely in their life experiences. In this comparative study we investigated subjects' persistence when presenting a novel object containing food that could not be accessed (impossible task) to three dog populations with very diverse life experiences: free-ranging village dogs (in Morocco), pet dogs (in Vienna) and captive pack living dogs (Wolf Science Center-WSC). We found that pet dogs and captive dogs (WSC) were more manipulative and persistent than free-ranging dogs. The low persistence of free ranging-dogs is unlikely the effect of a lack of exposure to objects, since they are confronted with many human' artefacts in their environment daily. Instead, we suggest that the higher persistence of captive dogs and pet dogs in comparison to free-ranging dogs might be due to their increased experience of human-mediated object interaction. This provides subjects with a socially guided experience in manipulating and interacting with objects increasing their motivation to engage in such tasks.

Prosociality is important for initiating cooperation. Interestingly, while wolves rely heavily on cooperation, dogs' do so substantially less thus leading to the prediction that wolves are more prosocial than dogs. However, domestication hypotheses suggest dogs have been selected for higher cooperation, leading to the opposing prediction- increased prosocial tendencies in dogs. To tease apart these hypotheses we adapted a paradigm previously used with pet dogs to directly compare dogs and wolves. In a prosocial choice task, wolves acted prosocially to in-group partners; providing significantly more food to a pack-member compared to a control where the partner had no access to the food. Dogs did not. Additionally, wolves did not show a prosocial response to non-pack members, in line with previous research that social relationships are important for prosociality. In sum, when kept in the same conditions, wolves are more prosocial than their domestic counterpart, further supporting suggestions that reliance on cooperation is a driving force for prosocial attitudes.

Food preferences may be driven by a species' ecology. Closely related species such as dogs and wolves may have evolved preferences for different foods owing to their differing foraging styles. Wolves have been shown to be more persistent in problem-solving experiments and more risk-prone in a foraging task. A possible element affecting these (and other) results is a potential wolf-dog difference in food preferences. To address this possibility, we tested similarly raised and kept dogs and wolves in two different food choice tasks, a classic two-choice task and a multiple-choice paradigm. We predicted that if dogs have adapted to a more opportunistic, scavenging foraging style, they would show a weaker preference for meat over starch rich foods (such as kibble) and be less affected by hunger than wolves. Alternatively, given the recentness of the new niche dogs have created, we predicted no substantial differences between dogs' and wolves' food preferences. We found that our subjects did not differ in their preference for meat over kibble in either paradigm. However, wolves' (but not dogs') choice patterns were affected by satiation, with wolves being less "selective" when hungry. Furthermore, when fed before testing, wolves were more selective than dogs. These differences were more noticeable in the multiple-choice paradigm than the two-choice task, suggesting that the former, novel paradigm may be more sensitive and better capable of evaluating food preferences in a diverse range of species. Overall, we found that the distinct differences in wolves' and dogs' ecology and foraging styles do not appear to have affected their food preferences and thus, differences in food preferences are unlikely to have influenced results of previous experiments demonstrating wolf-dog differences in cognitive skills.

Post-conflict affiliative interactions have been widely investigated in primates but not extensively in other species. Using the Post Conflict-Matched Control (PC-MC) comparison method, this study investigated the patterns of post-conflict opponent affiliation (POA) of a captive family group of 19 arctic wolves (Canis lupus arctos), investigating the correlation with various factors. We found that POAs occurred mainly in the non-feeding context and more often when the victim was dominant and the aggressor subordinate. Furthermore, POAs were more likely to have been initiated by the victim than the aggressor. Victims' stress related behaviours occurred more in PC than MC periods, and more after high vs. low intensity aggressions but they were not more likely to occur after conflicts between wolves with a stronger social bond and POAs did not reduce their rate of occurrence. Our results showed that re-aggression was twice less frequent when a friendly interaction occurred between the aggressor and the victim, and consistent with this, victims engaged in POAs more often than the aggressor. Overall, our results support the hypothesis that POAs in wolves may have been selected for as a mechanism to avoid conflict escalation, which could lead to social disruption and hence jeopardize cooperative activities. The high relatedness among individuals in the pack and the greater dependence of all members on cooperation in breeding and hunting may reduce the importance of 'relationship quality' as a mediating factor of POAs, although dominance relationships, which are directly linked to the risks of further conflicts, do play an important role.

Animals utilize behavioral signals across a range of different contexts in order to communicate with others and produce probable behavioral outcomes. During play animals frequently adopt action patterns used in other contexts. Researchers have therefore hypothesized that play signals have evolved to clarify communicative intent. One highly stereotyped play signal is the canid play bow, but its function remains contested. In order to clarify how canid puppies use play bows, we used data on play bows in immature wolves (ages 2.7-7.8 months) and dogs (ages 2 to 5 months) to test hypotheses evaluated in a previous study of adult dogs. We found that young dogs used play bows similarly to adult dogs; play bows most often occurred after a brief pause in play followed by complementary highly active play states. However, while the relative number of play bows and total observation time was similar between dog and wolf puppies, wolves did not follow this behavioral pattern, as play bows were unsuccessful in eliciting further play activity by the partner. While some similarities for the function of play bows in dog and wolf puppies were documented, it appears that play bows may function differently in wolf puppies in regards to re-initiating play.

Social play is known as a cooperative interaction between individuals involving multiple mechanisms. However, the extent to which the equality of individuals' play styles affects the interaction has not been studied in many species. Dyadic play between wolf puppies, as well as between puppies and adults, was studied to investigate both self-handicapping and offensive behaviors to determine the extent to which wolves engage in play styles where one individual does not dominate the play. Our results did not support the hypothesized '50:50' rule, which suggests that more advantaged individuals should show higher rates of self-handicapping behaviors in order to facilitate play with others. Adult wolves performed significantly less self-handicapping behaviors than their puppy partners, and they performed significantly more offensive behaviors than their puppy partners. While the '50:50' rule was not supported at any time during our study period, dyads consisting of two puppies had significantly more equal play than dyads consisting of one puppy and one adult. These results suggest that wolf puppies are more likely to play on equal terms with similarly-aged play partners, while the dominance status of the partners dictates offensive and self-handicapping behaviors between animals of different ages.

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.

Empathy covers a wide range of phenomena varying according to the degree of cognitive complexity involved; ranging from emotional contagion, defined as the sharing of others' emotional states, to sympathetic concern requiring animals to have an appraisal of the others' situation and showing concern-like behaviors. While most studies have investigated how animals reacted in response to conspecifics' distress, dogs so far have mainly been targeted to examine cross-species empathic responses. To investigate whether dogs would respond with empathy-like behavior also to conspecifics, we adopted a playback method using conspecifics' vocalizations (whines) recorded during a distressful event as well as control sounds. Our subjects were first exposed to a playback phase where they were subjected either to a control sound, a familiar whine (from their familiar partner) or a stranger whine stimulus (from a stranger dog), and then a reunion phase where the familiar partner entered the room. When exposed to whines, dogs showed a higher behavioral alertness and exhibited more stress-related behaviors compared to when exposed to acoustically similar control sounds. Moreover, they demonstrated more comfort-offering behaviors toward their familiar partners following whine playbacks than after control stimuli. Furthermore, when looking at the first session, this comfort offering was biased towards the familiar partner when subjects were previously exposed to the familiar compared to the stranger whines. Finally, familiar whine stimuli tended to maintain higher cortisol levels while stranger whines did not. To our knowledge, these results are the first to suggest that dogs can experience and demonstrate "empathic-like" responses to conspecifics' distress-calls.

A number of studies have recently investigated personality traits in non-human species, with the dog gaining popularity as a subject species for research in this area. Recent research has shown the consistency of personality traits across both context and time for adult dogs, both when using questionnaire based methods of investigation and behavioural analyses of the dogs' behaviour. However, only a few studies have assessed the correspondence between these two methods, with results varying considerably across studies. Furthermore, most studies have focused on adult dogs, despite the fact that an understanding of personality traits in young puppies may be important for research focusing on the genetic basis of personality traits. In the current study, we sought to evaluate the correspondence between a questionnaire based method and the in depth analyses of the behaviour of 2-month old puppies in an open-field test in which a number of both social and non-social stimuli were presented to the subjects. We further evaluated consistency of traits over time by re-testing a subset of puppies. The correspondence between methods was high and test- retest consistency (for the main trait) was also good using both evaluation methods. Results showed clear factors referring to the two main personality traits 'extroversion,' (i.e. the enthusiastic, exuberant approach to the stimuli) and 'neuroticism,' (i.e. the more cautious and fearful approach to the stimuli), potentially similar to the shyness-boldness dimension found in previous studies. Furthermore, both methods identified an 'amicability' dimension, expressing the positive interactions the pups directed at the humans stranger, and a 'reservedness' dimension which identified pups who largely chose not to interact with the stimuli, and were defined as quiet and not nosey in the questionnaire.

Prosociality has received increasing interest by non-human animal researchers since the initial discoveries that suggested it is not a uniquely human trait. However, thus far studies, even within the same species, have not garnered conclusive results. A prominent suggestion for this disparity is the effect methodology can have on prosocial responses in animals. We recently found evidence of prosociality in domestic dogs towards familiar conspecifics using a bar-pulling paradigm, in which a subject could pull a rope to deliver food to its partner. Therefore, the current study aimed to assess whether dogs would show a similar response in a different paradigm, based on the token exchange task paradigm frequently used with primates. In this task, dogs had the option to touch a token with their nose that delivered a reward to an adjacent receiver enclosure, which contained a familiar conspecific, a stranger or no dog at all. Crucially, we also included a social facilitation control condition, whereby the partner (stranger/familiar) was present but unable to access the food. We found that the familiarity effect remained consistent across tasks, with dogs of both the bar-pulling and token choice experiments providing more food to familiar partners than in a non-social control and providing less food to stranger partners than this same control. However, in contrast to our previous bar-pulling experiment, we could not exclude social facilitation as an underlying motive in the current task. We propose this difference in results between tasks may be related to increased task complexity in the token choice paradigm, making it harder for dogs to discriminate between the test and social facilitation conditions. Overall our findings suggest that subtle methodological changes can have an impact on prosocial behaviours in dogs and highlights the importance of controlling for social facilitation effects in such experiments.

Inhibitory control i.e. blocking an impulsive or prepotent response in favour of a more appropriate alternative, has been suggested to play an important role in cooperative behaviour. Interestingly, while dogs and wolves show a similar social organization, they differ in their intraspecific cooperation tendencies in that wolves rely more heavily on group coordination in regard to hunting and pup-rearing compared to dogs. Hence, based on the 'canine cooperation' hypothesis wolves should show better inhibitory control than dogs. On the other hand, through the domestication process, dogs may have been selected for cooperative tendencies towards humans and/or a less reactive temperament, which may in turn have affected their inhibitory control abilities. Hence, based on the latter hypothesis, we would expect dogs to show a higher performance in tasks requiring inhibitory control. To test the predictive value of these alternative hypotheses, in the current study two tasks; the 'cylinder task' and the 'detour task', which are designed to assess inhibitory control, were used to evaluate the performance of identically raised pack dogs and wolves. Results from the cylinder task showed a significantly poorer performance in wolves than identically-raised pack dogs (and showed that pack-dogs performed similarly to pet dogs with different training experiences), however contrary results emerged in the detour task, with wolves showing a shorter latency to success and less perseverative behaviour at the fence. Results are discussed in relation to previous studies using these paradigms and in terms of the validity of these two methods in assessing inhibitory control.

Understanding of other's actions as goal-directed is considered a fundamental ability underlying cognitive and social development in human infants. A number of studies using the habituation-dishabituation paradigm have shown that the ability to discern intentional relations, in terms of goal-directedness of an action towards an object, appears around 5 months of age. The question of whether non-human species can perceive other's actions as goal-directed has been more controversial, however there is mounting evidence that at least some primates species do. Recently domestic dogs have been shown to be particularly sensitive to human communicative cues and more so in cooperative and intentional contexts. Furthermore, they have been shown to imitate selectively. Taken together these results suggest that dogs may perceive others' actions as goal-directed, however no study has investigated this issue directly. In the current study, adopting an infant habituation-dishabituation paradigm, we investigated whether dogs attribute intentions to an animate (a human) but not an inanimate (a black box) agent interacting with an object. Following an habituation phase in which the agent interacted always with one of two objects, two sets of 3 trials were presented: new side trials (in which the agent interacted with the same object as in the habituation trial but placed in a novel location) and new goal trials (in which the agent interacted with the other object placed in the old location). Dogs showed a similar pattern of response to that shown in infants, looking longer in the new goal than new side trials when they saw the human agent interact with the object. No such difference emerging with the inanimate agent (the black box). Results provide the first evidence that a non-primate species can perceive another individual's actions as goal-directed. We discuss results in terms of the prevailing mentalisitic and non-mentalistic hypotheses regarding goal-attribution.

Social referencing is a process whereby an individual uses the emotional information provided by an informant about a novel object/stimulus to guide his/her own future behaviour towards it. In this study adult dogs were tested in a social referencing paradigm involving a potentially scary object with either their owner or a stranger acting as the informant and delivering either a positive or negative emotional message. The aim was to evaluate the influence of the informant's identity on the dogs' referential looking behaviour and behavioural regulation when the message was delivered using only vocal and facial emotional expressions. Results show that most dogs looked referentially at the informant, regardless of his/her identity. Furthermore, when the owner acted as the informant dogs that received a positive emotional message changed their behaviour, looking at him/her more often and spending more time approaching the object and close to it; conversely, dogs that were given a negative message took longer to approach the object and to interact with it. Fewer differences in the dog's behaviour emerged when the informant was the stranger, suggesting that the dog-informant relationship may influence the dog's behavioural regulation. Results are discussed in relation to studies on human-dog communication, attachment, mood modification and joint attention.

Dogs appear to be sensitive to human ostensive communicative cues in a variety of situations, however there is still a measure of controversy as to the way in which these cues influence human-dog interactions. There is evidence for instance that dogs can be led into making evaluation errors in a quantity discrimination task, for example losing their preference for a larger food quantity if a human shows a preference for a smaller one, yet there is, so far, no explanation for this phenomenon. Using a modified version of this task, in the current study we investigated whether non-social, social or communicative cues (alone or in combination) cause dogs to go against their preference for the larger food quantity. Results show that dogs' evaluation errors are indeed caused by a social bias, but, somewhat contrary to previous studies, they highlight the potent effect of stimulus enhancement (handling the target) in influencing the dogs' response. A mild influence on the dog's behaviour was found only when different ostensive cues (and no handling of the target) were used in combination, suggesting their cumulative effect. The discussion addresses possible motives for discrepancies with previous studies suggesting that both the intentionality and the directionality of the action may be important in causing dogs' social biases.