No full-text available
To read the full-text of this research,
you can request a copy directly from the authors.
Down but not out: Supine postures as facilitators of play in domestic dogs
Abstract
We used two sets of videotaped data of playing domestic dog dyads to determine whether rolling over during play served as a signal of submission or whether it was a combat manouevre adopted as part of an ongoing play sequence. Our results provide strong support for the latter. In the absence of any overt indication of agonism, the frequency with which rollovers occurred was determined primarily by play bout length. The discrepancy in partner size had no effect on the probability that rollovers would occur and there was no evidence that smaller dogs were more likely to rollover or to sustain a supine posture for longer, if they did. The supine phase of rollovers was significantly skewed to short durations. Most rollovers were either defensive (evading a nape bite) or offensive (launching an attack). None could be categorized as submissive. We conclude that asymmetries in the performance of rollovers cannot be assumed to point to asymmetries in the relationships between play partners.
... Using the supine posture as a measure of symmetry in the play fighting among dogs revealed marked asymmetry leading to the conclusion that the role of reciprocity in play fighting has been exaggerated [71]. However, an analysis of when rolling over to supine occurs during play fighting that took into account the partner's movements, revealed that nearly 90% of all instances were used as a combat tactic, mostly to avoid being bitten on the nape [72]. Moreover, analysis of the duration of remaining supine, the presence of associated gestures (e.g., ear and tail positions) and whether supine defense led to immobility or to counterattacks, all supported the conclusion that rolling over to supine serves a combat function, with no aspects of the action showing signs of ritualization that may indicate submission. ...
... Moreover, analysis of the duration of remaining supine, the presence of associated gestures (e.g., ear and tail positions) and whether supine defense led to immobility or to counterattacks, all supported the conclusion that rolling over to supine serves a combat function, with no aspects of the action showing signs of ritualization that may indicate submission. Of the small percentage of supine maneuvers that did exhibit a pattern of ritualization indicative of them being used for communication, they did so in a context suitable for soliciting play, not for submission [72]. Viewed as a combat tactic, the differences in the frequency of use of the supine maneuver between partners were best accounted for by differences in body size and so the accessibility of the body targets competed over (i.e., nape, throat). ...
... However, instead of remaining inert as in the previous context, it immediately lunges up and bites at its partner's throat. That is, from the outset, the act of rolling over is executed as a defensive maneuver that is then coupled with a counterattack [72]. The common action in these two scenarios is rolling over onto the back, but whereas in the former the predominant functional value of the action is one of communication, in the other, it is as a combat tactic. ...
During competitive interactions, such as fighting and predation, animals perform various actions, some of which are easy to characterize and label, some of which are reliably repeated. Such 'behavior patterns' are often the measures of choice when comparing across species and experimental contexts. However, as Bob Blanchard and others have pointed out, such measurements can be misleading as in competitive interactions in which the animals compete for some advantage, often the biting or otherwise contacting a particular target on the opponent's body. In this context, the animals' behavior is better analyzed in terms of the tactics of attack and defense deployed by the combatants to gain or avoid contact with those targets. Several examples are shown to reveal that this is an important distinction as simply scoring predefined behavior patterns can obscure the dynamic context in which the actions are performed. This can lead to confounding species and experimental differences and the mislabeling of combat actions as communicatory signals.
Copyright © 2015. Published by Elsevier Inc.
... Likewise, behaviors traditionally thought of as "submissive" signals (eg, rolling over onto one's back during interactions) instead appear to act as maneuvers to continue a play sequence. 42 Rolling over is a determinant of bout length of play that, in the absence of frank agonistic behavior, serves to avoid nape bites. Rolling over can also be a tactical move to retrieve control of the play bout and determine its subsequent direction. ...
... Asymmetries in rollovers depended wholly on the specific play context, and rollovers were associated with shortened bout intervals and pauses (eg, the dog that rolls over determines when play pauses and that dog's next behavior determines whether play resumes), not with commonly used measures of submission or asymmetry in relationships. 42 Detailed, frame-by-frame analysis of behaviors that dogs exhibit when approached by another dog indicates that the specific signals sent and the familiarity of the dogs determine the behavioral response. 37 When meeting an unfamiliar dog, dogs significantly more often exhibited nose licking, freezing, paw lifting, and head turning and made themselves smaller than they did when meeting a familiar dog. ...
There is now a large body of research in veterinary behavioral medicine that is clinically relevant and could enrich patients' and practitioners' lives. Too often, however, this research is published in journals that may not be readily available to veterinarians in private practice. Four important topics in the area of veterinary behavioral medicine for which belief has not kept pace with the published data are the unmet need for behavioral medicine in veterinary practice, the veterinary experience as a contributor to fear and distress in dogs and cats, social signaling in dogs and the ongoing "dominance" debate, and punishment as an intervention to change behavior. The present article seeks to provide a critical overview of recent research that is shifting existing paradigms on these topics and should alter the way veterinarians observe and care for patients.
... Because many dog social behaviours have not yet been studied adequately in their own right, it is likely that the functions of those behaviours are not fully understood. Recent controversies over the roles of play bows (Byosiere et al., 2016) and roll overs (Norman et al., 2015;Smuts et al., 2015) illustrate this point. Nor is labelling behaviours a priori as dominant or submissive entirely appropriate, since if and how dominance applies to companion dogs (e.g., van Kerkhove, 2004;Bradshaw et al., 2009), particularly those in transient social groups with few resources to defend, is debatable and has only recently received significant empirical attention (e.g., Schilder et al., 2014;Smuts, 2014;Trisko and Smuts, 2015;Trisko et al., 2016). ...
... From standing, rolls onto back or side with forelegs pointing in air or pulled in close to the chest; genitals exposed (see Bradshaw and Lea, 1992;Norman et al., 2015;Scott, 1950). Run/leap self-present (I, R) ...
This study examines the activity budgets and social behaviours initiated and received by 69 focal dogs in an off-leash dog park for 400 seconds after entry, a time of high activity about which little is known. Using motivationally-neutral labels for social behaviour categories, we describe the frequency of behaviours, and correlations among them. We then examine these relationships in the context of proposed functions for some behaviours in dogs, in terms of information gathering and communication, including visual and tactile signalling. Time spent with other dogs decreased rapidly over the visit, and much of this early interaction involved greeting the park newcomer. Snout-muzzle contact behaviours were ubiquitous, while other behaviours were rarely observed, including aggressive behaviours. Correlations among certain non-contact behaviours initiated and received by focal dogs are consistent with their function as visual signals that may influence the continuation and form of social interactions, and their possible role in social mimicry (i.e., play bow and pull-rear away). Age, sex, and number of dogs present in the park influenced specific aspects of dogs' activity budgets, and a few behaviours. This ethological study provides fundamental data on dog social behaviour in dog parks, about which surprisingly little has been published.
... For example, although rolling over to supine during play in dogs has been subsumed as part of the behavioural measures thought to signify dominance asymmetries (Bauer & Smuts, 2007), detailed analysis of the correlated actions by both participants suggests that rolling over is mostly used as a defensive tactic. However, rolling over on the back can also be used as an invitation to play in dogs (Norman et al., 2015) and a number of other species, such as in black bear cubs (Burghardt & Burghardt, 1972) and juvenile vervet monkeys (Pellis et al., 2014b). In this way, such an action can have multiple functions during play (Smuts et al., 2015), suggesting that commonly recognized 'signals', such as the play bow in dogs, need to be evaluated empirically (Byosiere et al., 2016) to ensure that the different functional uses can be discerned. ...
... To calculate the play asymmetry index (PAI), we clustered the play patterns as offensive, self-handicapping and neutral (see Appendix, Table A1). Although during play fighting some patterns can be categorized as both offensive and defensive according to the different contexts in which they are performed (Norman et al., 2015;Pellis & Pellis, 1997), here we wanted to evaluate the direction of each 'offensive' pattern (Appendix , Table A1). So, if a dog tried to bite a playmate and the playmate tried to counterattack with a bite, the outcome of the interaction was balanced, which 2 ðnumber of action units agreed by both codersÞ ðnumber of action units coded by V:M:Þ þ ðnumber of action units coded by F:M:Þ means that the two players have a similar opportunity to get the upper-hand during their playful encounters and this ensures reciprocity. ...
Play fighting, the most iconic form of social play, is often punctuated by specific signals, such as the relaxed open mouth (ROM) display, limiting the risk of misunderstanding between playmates. Although there is general consensus that the ROM of dogs is a ritualized version of play biting, the empirical demonstration of the actual ritualization of ROM has been lacking. We videorecorded and analysed 118 playful sessions involving 24 Czechoslovakian wolfdogs (12 females; 12 males), which is a breed of domestic dog, Canis lupus familiaris, showing wolf-like behavioural traits. By using an integrated approach of different techniques (dog facial action coding system, an unsupervised cluster analysis and the Levenshtein distance), we empirically demonstrate that the ROM is intrinsically different from the play biting action in this breed of dog. Contrary to the play bite, during ROM, the recruitment of muscular action units for each facial display was more consistent, conspicuous and intra- and interindividually stereotyped. Moreover, a sequential analysis revealed that the ROM usually preceded playful offensive patterns, thus underlining the real metacommunicative function of the signal. Finally, by running a linear mixed model, we found that the most balanced sessions were punctuated by the most prolonged performance of ROM, thus revealing the efficiency of the facial signal in maintaining a balanced session. In conclusion, through the processes of formalization, simplification and emphasis, an ordinary precursor behaviour (i.e. play biting) has been taken out of context and transformed into an extraordinary, derived behaviour (i.e. ROM) specifically designed to attract receivers' attention and modulate playful social interactions in dogs.
... This allowed to calculate the Play Asymmetry Index (PAI). Although play fighting patterns can be classified as both offensive and defensive according to the contexts in which they occur (Pellis & Pellis 1997;Norman et al. 2015), for the PAI calculation we needed to evaluate the direction of each 'offensive' pattern. So, if a dog tries to push a playmate (offensive) and the playmate tries to defend itself by counterattacking with a bite (offensive), the outcome of the interaction is considered balanced, which means that the two subjects have a similar opportunity to gain advantage during their playful encounters thus producing reciprocity. ...
Dogs engage in play behavior at every age and the play bow is their most iconic playful posture. However, the function of this posture is still under debate. Here, we selected the Czechoslovakian Wolfdog as a model breed to clarify the function of the play bow. We analyzed frame-by-frame 118 sessions of 24 subjects and recorded 76 play bow events. We found that all the play bows were performed in the visual field of the playmate suggesting that the sender takes into account the attentional state of the receiver when releasing the signal. By drawing survival curves and using log-rank test we found that play bow was mainly performed during a short pause in an ongoing session and that its performance triggered the playmate's reaction again. These findings show that play bow functions in restoring the partner motivation to play. Finally, by using a sequential analysis and a generalized mixed model, we found no evidence
... This allowed to calculate the Play Asymmetry Index (PAI). Although play fighting patterns can be classified as both offensive and defensive according to the contexts in which they occur (Pellis and Pellis 1997;Norman et al. 2015), for the PAI calculation we needed to evaluate the direction of each "offensive" pattern. So, if a dog tries to push a playmate (offensive) and the playmate tries to defend itself by counterattacking with a bite (offensive), the outcome of the interaction is considered balanced, which means that the 2 subjects have a similar opportunity to gain advantage during their playful encounters thus producing reciprocity. ...
Dogs engage in play behavior at every age and the play bow is their most iconic playful posture. However, the function of this posture is still under debate. Here, we selected the Czechoslovakian Wolfdog as a model breed to clarify the function of the play bow. We analyzed frame-by-frame 118 sessions of 24 subjects and recorded 76 play bow events. We found that all the play bows were performed in the visual field of the playmate suggesting that the sender takes into account the attentional state of the receiver when releasing the signal. By drawing survival curves and using log-rank test we found that play bow was mainly performed during a short pause in an ongoing session and that its performance triggered the playmate’s reaction again. These findings show that play bow functions in restoring the partner motivation to play. Finally, by using a sequential analysis and a generalized mixed model, we found no evidence supporting the metacommunicative function of the play bow. The signal did not necessarily precede a contact offensive behavior (e.g., play biting, play pushing) and it was not affected by the level of asymmetry of the play session. In conclusion, in Czechoslovakian Wolfdogs play bow can be considered a visual signal useful to maintain the motivation to play in the receiver. Therefore, we suggest that the mismatched number of play bows emitted by the two players in a given session can be predictive of their different motivation to play.
... However, given the variety of ways in which different species incorporate cooperation in their play fighting (Pellis & Pellis, 2017), a necessary first step for crossspecies comparisons is to characterize the species-typical targets and the associated combat tactics. Doing so provides a framework for identifying actions that are not compatible with combat and so are likely present to facilitate cooperation (e.g., Norman, Pellis, Barrett, & Henzi, 2015;Pellis & Pellis, 2016;Pellis, Pellis, Barrett, & Henzi, 2014). ...
Play fighting in many species involves partners competing to bite one another while avoiding being bitten. Species can differ in the body targets that are bitten and the tactics used to attack and defend those targets. However, even closely related species that attack and defend the same body target using the same tactics can differ markedly in how much the competitiveness of such interactions is mitigated by cooperation. A degree of cooperation is necessary to ensure that some turn-taking between the roles of attacker and defender occurs, as this is critical in preventing play fighting from escalating into serious fighting. In the present study, the dyadic play fighting of captive troops of 4 closely related species of Old World monkeys, 2 each from 2 genera of Papio and Mandrillus, was analyzed. All 4 species have a comparable social organization, are large bodied with considerable sexual dimorphism, and are mostly terrestrial. In all species, the target of biting is the same – the area encompassing the upper arm, shoulder, and side of the neck – and they have the same tactics of attack and defense. However, the Papio species exhibit more cooperation in their play than do the Mandrillus species, with the former using tactics that make biting easier to attain and that facilitate close bodily contact. It is possible that species differences in how rigidly dominance relationships are maintained are expressed in the play of juveniles by altering the balance between competition and cooperation.
... For example, rolling over on to the back and assuming a supine posture, which is a frequent manoeuvre during play fights (Bauer and Smuts, 2007;Fox, 1969), also has an apparent submissive function in dogs and wolves (Lorenz, 1943;Schenkel, 1967). However, in both dogs (Norman et al., 2015) and wolves (Cordoni, 2009), this behaviour has been found to be more consistently used as a combat tactic than as a submission signal. Similarly, jacky dragons (Amphibolurus muricatus) were found to strategically use the same signals to escalate or de-escalate a conflict, depending on the context of the signals produced by their opponent (Van Dyk and Evans, 2008). ...
Dominance hierarchies can reduce conflict within social groups and agonistic signals can help to establish and maintain these hierarchies. Behaviours produced by subordinates in response to aggression are often assumed to function as signals of submission, however, these behaviours may serve other purposes, for example, defence or escape. For a behaviour to act as a submission signal, the receiver must respond by reducing their likelihood of further aggression towards the signaller. In the current study, we examine the receiver response to a putative signal of submission, the head up display, within established social groups of the cooperatively breeding fish, the daffodil cichlid (Neolamprologus pulcher). We found that when subordinate signallers produce the head up display in response to aggression from the breeder male, he exhibited a longer latency to behave aggressively towards that individual again. We also report that head up displays are rarely produced without being elicited by aggression, and the number of head up displays correlates with the amount of aggression received. Our results demonstrate that the head up display is used as a signal of submission in the daffodil cichlid and provide insight into intragroup communication in an emerging model system for the study of social behaviour.
... The study of emotion is not my expertise, so I leave the nuances of this argument to others. However, recent work on two overt behaviours in dogs -the "roll-over" (Norman et al., 2015) and the "play bow" (Byosiere et al., 2016) -may be instructive. In both cases, these common dog behaviours have been assigned a priori to functional categories (i.e., as signals of submission and meta-communication, respectively). ...
... Also, as we have already noted, within a species variation in reciprocation is likely present, based on age, sex, and dominance relationships. For example, animals not familiar with one another may have different rates of reciprocation than those that are familiar with one another (e.g., Bauer & Smuts, 2007;Norman, Pellis, Barrett, & Henzi, 2015). Nonetheless, it should be noted that the examples cited above for spider monkeys, macaques, rats, degus, and the various species of pigs all involved animals that were familiar with one another. ...
Play fighting is a common form of play reported among species of mammals, birds, and some other taxa. The competition present in play fighting revolves around gaining some advantage, such as biting a partner without being bitten. The behavior simulated during play fighting need not be restricted to that present in adult serious fighting, but can involve competitive interactions derived from amicable behavior, such as sex and social grooming, or from nonsocial competition, such as predation. What unifies play fighting, irrespective of the functional behavior being simulated, is that it involves some degree of reciprocity, or turn taking, that requires that the competition be attenuated by cooperation. However, there are several different ways in which cooperation can be inserted into playful interactions, and these vary in use across different species. The moderation of competition with cooperation forces animals to monitor their own actions and those of their partners, and this common feature appears to be one vehicle through which the experience of play fighting in the juvenile period can train animals for greater psychological resilience. The monitoring and contextual adjustment of actions influences the development of executive functions of the brain, which, in turn, leads to the development of more adaptable adults.
... For example, the Eshkol-Wachman Movement Notation method (Eshkol & Wachman, 1958) is a system in which body position in space and time is coded, providing a record of the temporal sequence of movements and their organization. These records can be performed separately on interacting organisms, and utilized to examine the relative spatiotemporal structure of behavior between individuals (Carrier, Leca, Pellis, & Vasey, 2015;Norman, Pellis, Barrett, & Henzi, 2015;Pasztor, Smith, MacDonald, Michener, & Pellis, 2001;Pellis, 1982). A major advantage of this system is that the topography of specific behaviors can be characterized in great detail. ...
Social interactions form the basis of a broad range of functions related to survival and mating. The complexity of social behaviors and the flexibility required for normal social interactions make social behavior particularly susceptible to disruption. The consequences of developmental insults in the social domain and the associated neurobiological factors are commonly studied in rodents. Though methods for investigating social interactions in the laboratory are diverse, animals are typically placed together in an apparatus for a brief period (under 30 min) and allowed to interact freely while behavior is recorded for subsequent analysis. A standard approach to the analysis of social behavior involves quantification of the frequency and duration of individual social behaviors. This approach provides information about the allocation of time to particular behaviors within a session, which is typically sufficient for detection of robust alterations in behavior. Virtually all social species, however, display complex sequences of social behavior that are not captured in the quantification of individual behaviors. Sequences of behavior may provide more sensitive indicators of disruptions in social behavior. Sophisticated analysis systems for quantification of behavior sequences have been available for many years; however, the required training and time to complete these analyses represent significant barriers to high-throughput assessments. We present a simple approach to the quantification of behavioral sequences that requires minimal additional analytical steps after individual behaviors are coded. We implement this approach to identify altered social behavior in rats exposed to alcohol during prenatal development, and show that the frequency of several pairwise sequences of behavior discriminate controls from ethanol-exposed rats when the frequency of individual behaviors involved in those sequences does not. Thus, the approach described here may be useful in detecting subtle deficits in the social domain and identifying neural circuits involved in the organization of social behavior.
... However, the 50-50 rule does not fit for all species in which it has been tested. For example, many researchers demonstrated that, contrary to the expectations, dogs engage in unbalanced playful sessions (Bauer & Smuts 2007;Ward et al. 2008;Norman et al. 2015;. Obviously, the degree of asymmetry that characterizes the playful session can be influenced by factors such as species, context and the relationship quality shared by the players, so such asymmetry will vary according to the different social functions of play. ...
Although play fighting has been studied for over a century in both human and non-human animals, quantitative data on marine mammals are still scarce. Here, we investigated play fighting in South American sea lions (Otaria flavescens), one of the most sexually dimorphic species with an extreme polygynous mating system, high levels of both intra- and intersexual competition. All these features make South American sea lions a
good model species to test some predictions on play fighting. Our results indicate play is restricted to juveniles, being inhibited among adults, and as to be expected in a species that shows a high degree of sexual dimorphism, it is mainly expressed in males. Even though playful interactions were punctuated by competitive behaviours, animals played in a highly symmetric way and were able to adjust their competitive playful interactions in a flexible manner and so reduce the risk of escalation to a minimum level. They were highly selective in their choice of playmates by limiting the number of players per session and playing more with agematched companions and friends. All these factors taken together are probably at the basis of the low risk of escalation recorded during the study. This result is predictive of a high ability and motivation of these
animals to engage in play behaviour which can have a possible role not only in the acquisition of dominance status, but also in establishing and maintaining social relationships, an unexpected role in a so highly competitive species.
... However, after this attack, the disadvantaged partners were able to launch a counterattack approximately 30% of the time. One of several dog studies paired a medium-sized female dog with multiple partners to look at whether or not roll-over behavior within play is a submissive signal [21]. Smaller dogs did not roll onto their backs more, and rolling over behavior was typically categorized as 'defensive' to avoid attacks or 'offensive' to launch attacks, rather than submissive. ...
... However, after this attack, the disadvantaged partners were able to launch a counterattack approximately 30% of the time. One of several dog studies paired a medium-sized female dog with multiple partners to look at whether or not roll-over behavior within play is a submissive signal [21]. Smaller dogs did not roll onto their backs more, and rolling over behavior was typically categorized as 'defensive' to avoid attacks or 'offensive' to launch attacks, rather than submissive. ...
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.
... This signal redundancy is less necessary in wolves who play less competitively and in which the standing-over posture used in the playful context seems not to serve to assert dominance (Bekoff, 1975) as it occurs also in other carnivore species (e.g., bears, Burghardt & Burghardt, 1972). Similarly, in dogs, it has been recently demonstrated that the roll-over posture (an animal rolls over onto its back by exposing its ventrum) during play is not predictive of the real dominance relationship characterizing the dyad (Norman et al., 2015). One of the potential factors explaining why wolves do not use as sophisticated a set of signals (e.g., the play bow) to maintain a playful mood could be due to the fact that play activity is less frequent between individuals who greatly differ in social rank (Cordoni, 2009). ...
The concept of peace, with its corollary of behaviours, strategies and social implications, is commonly believed as a uniquely human feature. Through a comparative approach, we show how social play in animals may have paved the way for the emergence of peace. By playing fairly, human and nonhuman animals learn to manage their social dynamics in a more relaxed and tolerant way that results in a more effective management of conflicts. We show that play promotes tolerance, cooperation , fairness and reciprocity, which are essential elements of the so-called positive peace. This kind of peace is reached through an evolving process in which individuals continually modify social relationships to attain peaceful coexistence. In conclusion, we assume that the concept of peace has deep biological roots that constitute the basis for more sophisticated cultural constructions.
... In this commentary, we compare and contrast Norman et al.s' findings on rollovers during dog play (Norman et al., 2015; the "target article") with our work on dog play fighting (Bauer and Smuts, 2007;Ward et al., 2008). We first review our major findings and then correct some errors in the target article's descriptions of our work. ...
Cambridge Core - Educational Psychology - The Cambridge Handbook of Play - edited by Peter K. Smith
An animal's welfare state is intrinsically linked to its affective state. Evidence suggests that sentient, conscious animals can experience a range of affective states, such as pain, fear or boredom as well as positive affects like joy, curiosity, satiation or lust. In the behavioural assessment of animal welfare, there is increasing recognition that it is not simply which behaviours an animal engages in but also the quality of its movement. Kinematics is an approach which is being more widely applied to the behavioural assessment of animal welfare. Kinematics is a field of mechanics that describes the movement of points on a body by defining these points in a coordinate system and precisely tracking how they change in terms of space and time. A major opportunity exists for using kinematic technology to inform our understanding of the emotional state of animals. This review argues that kinematics is a useful methodology for identifying and characterising movement indicative of an animal's affective state. It demonstrates that kinematics: i) appears useful in detecting subtleties in the expression of affective states; ii) could be used in conjunction with, and add extra information to, affective tests (for example, an approach/avoidance paradigm); and iii) could potentially, eventually, be developed into an automated affective state detection system for improving the welfare of animals used in research or production. Furthering our knowledge of animal affective states using kinematics requires engagement from many areas of science outside of animal welfare, such as sports science, computer science, engineering and psychology.
Due to their playful propensity, dogs are a good model to test some hypotheses about play dynamics (length, asymmetry, features of players) and communication (play bow [PBOW]; relaxed open-mouth [ROM] display). We video-recorded 203 play sessions between dogs in an off-leash dog park in Palermo, Italy. Contrary to the expectation, play asymmetry (particularly high in this species) did not differ between stranger and familiar dogs, thus suggesting the limited role of play in forming dominance relationships. Asymmetry negatively affected the duration of the session, whereas the increasing number of players was positively linked to the duration of playful interactions. The number of PBOWs exchanged by players may exert a certain influence on the session length as well. PBOWs were performed independently from the kind of play (locomotor vs. contact) the dogs were engaging in. Conversely, ROMs were preferentially emitted during contact play when "face-to-face" interactions were more likely. Body closeness is also required in case opening the mouth has not a signal function but only preludes a bite. However, in the 82% of cases play bites did not follow a ROM, thus suggesting that dogs place ROMs in the appropriate context to optimize signal detectability. In conclusion, 2 tactics may concur in coping with the asymmetry and unpredictability of play sessions in dogs. First, whenever the asymmetry increases dogs shorten the duration of their sessions thus limiting the risk of possible escalation. Second, dogs make use of a good communicative system based on the reciprocal exchange of playful signals. (PsycINFO Database Record
During play fighting, animals make a variety of movements, some of which seem unrelated to the ongoing actions of the partner. Among such movements are the jumps and rotations reported in many species of Old World monkeys. In the present study, videotaped sequences of jumps and rotations performed by juvenile vervet monkeys were analyzed. Using the Eshkol-Wachman Movement Notation (EWMN) the sequences were described frame-by-frame revealing that about 82% of the jumps and rotations were correlated with the movements performed by the partner, consistent with the view that these movements are used as tactics of attack and defense. The majority of the remaining 18% occurred in contexts in which the performer solicited playful attacks from the partner. A small number of the rest were performed in a manner consistent with the movements being imposed by the performer to increase the difficulty in contacting the partner. The same distribution was present in both captive and free-living monkeys. Thus the analyses show that while most jumps were combat-related, these movements can occur in a variety of functional contexts.
Macaques are the most geographically widespread and behaviorally diverse primate genus, and
although macaque species share the same, basic social structure, they display broad interspecific
variation in patterns of adult social behavior. Based on these patterns, macaque species have been
arranged along a 4-grade scale for social style. At one end of the scale, there are grade 1 species (e.g.,
Japanese macaques) that have highly hierarchical and despotic social systems, and at the other end,
grade 4 species (e.g., Tonkean macaques) that have more relaxed and egalitarian social systems. We
predicted that a species from the more despotic end of the spectrum should have more competitive
play fights and that a species from the egalitarian end, more cooperative ones. A detailed analysis of
videotaped sequences of juvenile play fighting in Tonkean and Japanese macaques was used to
characterize the targets and tactics of attack and defense. Even though the two species have a similar
behavioral repertoire, there are significant differences in how that repertoire is used and these
differences are consistent with one species having more competitive interactions than the other.
Contrasting multi-animal play fights versus pairs showed that the more cooperative style of the
Tonkean macaques is further exaggerated. The results suggest that differences in styles of attack and
defense in play fighting may be influenced by differences in the species’ social systems.
Abstract - During play fighting, animals make a variety of movements, some of which seem unrelated to the ongoing actions of the partner. Among such movements are the jumps and rotations reported in many species of Old World monkeys. In the present study, videotaped sequences of jumps and rotations performed by juvenile vervet monkeys were analyzed. Using the Eshkol-Wachman Movement Notation (EWMN) the sequences were described frame-by-frame revealing that about 82% of the jumps and rotations were correlated with the movements performed by the partner, consistent with the view that these movements are used as tactics of attack and defense. The majority of the remaining 18% occurred in contexts in which the performer solicited playful attacks from the partner. A small number of the rest were performed in a manner consistent with the movements being imposed by the performer to increase the difficulty in contacting the partner. The same distribution was present in both captive and free-living monkeys. Thus the analyses show that while most jumps were combat-related, these movements can occur in a variety of functional contexts.
To identify genetic changes underlying dog domestication and reconstruct their early evolutionary history, we generated high-quality genome sequences from three gray wolves, one from each of the three putative centers of dog domestication, two basal dog lineages (Basenji and Dingo) and a golden jackal as an outgroup. Analysis of these sequences supports a demographic model in which dogs and wolves diverged through a dynamic process involving population bottlenecks in both lineages and post-divergence gene flow. In dogs, the domestication bottleneck involved at least a 16-fold reduction in population size, a much more severe bottleneck than estimated previously. A sharp bottleneck in wolves occurred soon after their divergence from dogs, implying that the pool of diversity from which dogs arose was substantially larger than represented by modern wolf populations. We narrow the plausible range for the date of initial dog domestication to an interval spanning 11-16 thousand years ago, predating the rise of agriculture. In light of this finding, we expand upon previous work regarding the increase in copy number of the amylase gene (AMY2B) in dogs, which is believed to have aided digestion of starch in agricultural refuse. We find standing variation for amylase copy number variation in wolves and little or no copy number increase in the Dingo and Husky lineages. In conjunction with the estimated timing of dog origins, these results provide additional support to archaeological finds, suggesting the earliest dogs arose alongside hunter-gathers rather than agriculturists. Regarding the geographic origin of dogs, we find that, surprisingly, none of the extant wolf lineages from putative domestication centers is more closely related to dogs, and, instead, the sampled wolves form a sister monophyletic clade. This result, in combination with dog-wolf admixture during the process of domestication, suggests that a re-evaluation of past hypotheses regarding dog origins is necessary.
The Eshkol-Wachmann movement notation is used to describe motor sequences in the interactions of both golden jackals (Canis aureus) and Tasmanian devils (Sarcophilus harrisii). Motor behavior is rigorously described in terms of the elementary movements of limb segments. The same movements are described in four coordinate systems: in relation to the animal’s own body, in relation to the environment, in relation to a partner, and in relation to the topography of the contact point with the partner on the animal’s own body.
We videotaped behaviour in four litters of domestic dogs to explore social play and the development of relationships within litters. We collected data when the puppies were between 3 and 40 weeks of age, but collection times varied by litter. We divided data analysis into three time periods to coincide approximately with critical periods in the early social development of dogs. Early play-partner preferences were associated with preferences in later time periods, and the tendency for puppies to prefer specific partners increased over time. Play did not conform to 50–50 symmetry of roles between partners, which some researchers claim is necessary to sustain play. In the later juvenile period (time 3), dogs who engaged in high rates of offense behaviours (e.g. chasing, forcing partners down) also initiated play at higher rates, implying that winning during play may become more important as puppies mature. Self-handicapping behaviours were positively associated with play signalling, suggesting that, like play signals, self-handicapping may function to indicate playful intent. In mixed-sex dyads, males initiated play, engaged in offense behaviours, and self-handicapped more than females. Females were more likely to initiate with females across all time periods, but males were more likely to initiate with males only in time 3. We discuss results from mixed- and same-sex interactions with reference to inter- and intrasexual competition. The types of offense and self-handicapping behaviours displayed were similar across litters, suggesting that the expression of these behaviours may follow a similar ontogeny in puppies in general.
Social play behavior is receiving increasing attention from ethologists and other scientists who are interested in social development and social organization. To date, however, few quantitative data have been amassed concerning play, and it is not uncommon to read about unsupported contentions being accepted as fact. The purpose of this chapter is to summarize some aspects of play behavior, and to show that more rigorous quantitative analyses are needed in order for interested workers to gain a fuller understanding of this category of behavior.
In the diagnosis and treatment of behavioural disorders in multi-cat households, it is often assumed that a dominance hierarchy exists between the cats ( e.g. Crowell-Davis, 2002). While such hierarchies are probably commonplace among dogs, what evidence there is to support the existence of social hierarchies in groups of domestic cats has mainly been gathered from reproductively entire animals, such as single sex laboratory colonies, and free-ranging aggregations of ferals. For example, Natoli et al. (2001) used receipt of “submissive” (defensive) behaviour to construct a weakly linear hierarchy in a group of 14 farm cats, but this did not correspond to the hierarchy derived from receipt of affiliative behaviour. We have investigated the alternative hypotheses that apparent dominance hierarchies in multi-cat households may actually be based upon territorial behaviour, or some other undetermined social system.
Lekking male greater sage-grouse (Centrocercus urophasianus) compete with neighbours not only by strutting to attract females but also by directly challenging other males. These challenges include approaching another male and adopting an anti-parallel orientation at close quarters ('facing past encounter') and fighting, in which the birds strike one another with their wings. Facing past encounters and facing past encounters that led to fights in free-living sage-grouse were videotaped and analysed to test predictions arising from two sets of hypotheses to account for the features of such encounters. They could be used to assess or threaten opponents (index signal or threat signal hypotheses) or they may be the result of a stalemate in which one bird's attempts to gain an vantage point for attack are neutralised by counter moves by the other bird (combat hypothesis). Frame-by-frame analyses of both facing past encounters and fights were used to extract data to test specific predictions arising from the three hypotheses. The results, overall, support the hypothesis that the facing past orientation arises from combat. However, the results also suggest that, once in the anti-parallel orientation, opportunities emerge for communication to take place.
The postnatal development of agonistic behavior patterns in the wolf, coyote and grey fox are described, and in the red and Arctic fox which were first observed at a later age. Piloerection and back arching were present in all species, the latter behavior being especially well developed in the grey fox. The orientation of attack was almost exclusively directed at the cheek in the grey fox, but in other canids the shoulder hackle area, throat and muzzle area were also attacked; prolonged bouts of jaw-muzzle wrestling occurred in the wolf, scruff-wrestling in wolf and coyote and dog, and cheek-wrestling in the grey fox during agonistic play (play-fighting). Orientation of attack was correlated with distinctive body markings in various species. The contribution of social experience in the ontogeny of scruff-oriented attack was demonstrated in hand-raised isolated dogs and visually deprived group-raised dogs. The significance of head-turning, looking away and avoidance of eye contact in the wolf and dog is discussed.
Play behavior was observed in a captive group of four juvenile Hamadryas baboons at the Brookfield Zoo, Brookfield, Illinois, U.S.A.. The group consisted of a male, aged 37 months, a female, aged 28 months and two young males aged 16 and 11 months at the beginning of the 3 month study. Tape recorded commentary of 1006 play bouts was analyzed, and this data was supplemented with material gathered by other methods. Three play behaviors - chase, face-off, and wrestling - and two
The term "dominance" is widely used in the academic and popular literature on the behavior of domestic dogs, especially in the context of aggression. Although dominance is correctly a property of relationships, it has been erroneously used to describe a supposed trait of individual dogs, even though there is little evidence that such a trait exists. When used correctly to describe a relationship between 2 individuals, it tends to be misapplied as a motivation for social interactions, rather than simply a quality of that relationship. Hence, it is commonly suggested that a desire 'to be dominant' actually drives behavior, especially aggression, in the domestic dog. By contrast, many recent studies of wolf packs have questioned whether there is any direct correspondence between dominance within a relationship and agonistic behavior, and in contrast to wolves, hierarchical social structures have little relationship with reproductive behavior in feral dog packs. Nor do the exchanges of aggressive and submissive behavior in feral dogs, originally published by S. K. Pal and coworkers, fit the pattern predicted from wolf behavior, especially the submissive behavior observed between members of different packs. In the present study of a freely interacting group of neutered male domestic dogs, pairwise relationships were evident, but no overall hierarchy could be detected. Since there seems to be little empirical basis for wolf-type dominance hierarchies in dogs, the authors have examined alternative constructs. Parker's Resource Holding Potential (RHP) appears to be less useful when applied to domestic dogs than to other species, although it has the advantage of incorporating the concept of subjective resource value (V) as a factor influencing whether or not conflicts are escalated. The authors propose that associative learning, combined with V, can provide more parsimonious explanations for agonistic behavior in dogs than can the traditional concept of dominance.
In this paper, the development of social play behavior in coyotes, wolves, and beagles is discussed, as are the ways in which "play intention" is communicated and a "play mood" maintained. Coyotes were observed to fight more and play less than wolves or beagles of the same age, and the differential development of social behavior in these canids may play some role in determining the later species-typical social organization observed in adults.
Analysis of the dynamics of the ontogeny of social interaction is of critical importance in order that behavioral development may be comprehended in its own right, and the relationship between infant and adult behavior understood. In this review, general concepts of behavioral development in mammals are discussed and analyzed, and the many variables that are involved are considered. When it is impossible to control or observe the social interaction of the developing organism in its natural environment, captive subjects should be used. There is increasing evidence that results obtained with the latter are related to social organization observed in the wild. Play behavior is operationally defined on the basis of a comprehensive review of the literature and of personal observations of the social development of canids. The essential 'need' for social interaction during infant life is discussed, as is the phenomenon of behavioral neoteny. The concept of metacommunication, and its relationship to social development are analyzed, and the role of ritualization in the evolution of metacommunicative signals is considered.
Attack by dominant male colony mice on intruders included chasing and lateral attack behaviors, while the corresponding intruder behaviors were flight, boxing, and checking. Both of these are similar to the attack and defensive behaviors of colony rats and intruders. However, mice did not show a significant constraint on bites to ventral areas, and the rat defensive behavior of lying on the back, which is effective because of this constraint, was rare; the corresponding “on-top” behavior of attackers was almost absent in mice. These findings strongly support the view that intraspecific attack and defensive behaviors, and target sites for bites, are interrelated factors facilitating effective but nonlethal agonistic interactions in muroid rodents.
This paper describes elements in the social behaviour of the laboratory rat, mouse, hamster and Guinea-pig. These elements are divided into postures, which are static, and acts, which involve movement. A total of 45 of these elements are mentioned, most of which are common, with only slight modification, to all four species. Apart from these the guinea pig differs in not having a true Upright Posture and also in showing a male sexul display "Rumba". The postures are classified under broad motivational headings. A number of general concepts are discussed, for example the relation of convulsions to flight behaviour, the reduction of incoming aggressive stimuli in submissive postures, "Cut-Off", and the inhibition of biting in the more social species.
Submission in the wolf and dog is defined on the basis ot its motivation: submission is the effort of the inferior to attain friendly or harmonic social integration.
Submission functions as an appeal or a contribution to social integration, but only if it meets a corresponding attitude in the superior. The form of submissive behavior in wolf and dog is ritualized and symbolized cub-behavior. Two main forms of submissive behavior occur in wolf and dog: active submission, derived from begging for milk or food, and passive submission, derived from the posture which the cub adopts when cleaned by its mother.
The definition of submission is generally applicable to vertebrates living in groups based on intimacy and a social hierarchical order. The concept of submission as the role of the defeated in the terminal phase of fight with the function to inhibit automatically aggression in the superior should be dismissed. In vertebrates at least three types of conflict with different terminal phases occur: (1). Severe fight based on intolerance; ends with flight by the inferior or with his death. (2). Ritualized fight over a privilege; ends with the “giving-up-the-claim ritual” of the inferior, which automatically blocks the aggression of the superior. (3). Minor conflict in closed groups; settled by submissive behavior of the inferior.
In closed vertebrate groups, intermediate forms between (1) and (3) occur, depending on the proportion between activated intimacy and intolerance.
Animal social play represents an important tool for self- and social-assessment purposes during the juvenile phase. Nevertheless, this activity may continue into adulthood as well providing immediate benefits to the playmates. In this study, I investigated the dynamics of adult play in a wolf colony hosted at the Pistoia Zoo (Italy). The study wolves performed social play to a greater extent compared to solitary play. Play distribution was not affected by relationship quality (measured by body contact and agonistic support frequencies) and aggression levels. Probably, in wolves other behavioural strategies are employed for strengthening inter-individual relationships and reducing conflicts among fellows. Play was distributed throughout the entire group independently of the sex of playmates. The absence of sexual-dimorphism in play may be linked to the fact that in the wolf pack males and females share the same roles and behavioural repertoire. Rank distance between conspecifics negatively correlated with play distribution: by playing wolves with closest ranking positions tested each other for acquiring information on skills of possible competitor and gaining hierarchical advantage over it. Finally, in agreement with previous studies, my findings showed that wolves significantly reduced their playful activity during contests of high conflict of interests such as mating period and feeding time.
An analysis is made of the social behaviour of the male laboratory rat using the following methods. One rat is introduced into the home cage of another. One observer records the series of elements shown by each rat. These results are tabulated in sequence tables of elements. The tables are analysed by calculating an "expected" value for each cell and comparing this with the observed value. An ethogram is built up showing the relationships of the elements seen when the rats are close together and indicating the possible motivation of these elements in terms of the interaction of Aggression and Flight. It is shown that the Flight motivated elements fall into two groups, one leading to Crouch and the other leading to Submissive Posture. The occurrence of grooming and digging as displacement activities is shown and is contrasted to the occurrence of mounting which appears to be separately aroused in male-male situations. A group of elements occurring when the rats are at a distance from each other, and showing conflict between approach and avoidance, is described. It is suggested that there are two main Flight pathways, one leading to a Submissive Posture and the other to Crouch or Retreat, the occurrence of these is related to two types of behaviour seen in the wild, intra-colonial and territorial. Finally, the possible occurrence of an Approach component other than Aggression or Mating, which might be called a social drive, is suggested.
The writer first discusses the innate and endogenous factors in instinctive responses of animals, pointing out that the various external signals which release the responses summate in their effect in a non-Gestalt manner. In both domesticated animals and civilized man, the complex of stimuli capable of releasing instinctive behavior expands, decreasing the precision of the instinct and diminishing the ability to survive in the natural environment. And so the development of reason (contrasted with instinct) is both an advance and a decadence, an advantage and a danger. Only by purposeful exercise of this reason itself, and by voluntarily limiting his liberty, can man make up for his loss of guiding instincts and become master of his destiny. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
House mice have been reported rarely to perform the supine behavior pattern as a defensive tactic during intraspecific fighting. However, in this study of intraspecific fighting by male mice, it is shown that mice do indeed rotate to supine. This maneuver is used to evade or extricate themselves from bites to the lower dorsum by the attacking opponent. Once free from the bite the defender does not remain supine, but will immediately turn to prone and flee. Remaining motionless in the supine position may serve a submissive function in other species, but this does not seem to be the case for mice. The present findings illustrate that the supine tactic is a dynamic maneuver for defense of body areas targeted by the opponent. © 1992 Wiley-Liss, Inc.
This study concerns the form of combative interactions between wolf pups and how this form relates to strategy. Filmed records were analysed for dyadic interactions among three pups between the ages of two weeks and five months. By isolating and recombining the variables of body pitch, snout contact placement, and mutual orientation, combative strategies leading to unbalancing (falling) of one partner were evaluated. It was found that consistent dyadic configurations, bite locations and patterns of leg sweeps could be related to three distinct styles of falling (side, back and forward falls). Forward falls developed last, and represent a controlled strategy of active defense. “Top” and “bottom” pups can switch roles but show position–dependent regularities in behavior. These biomechanically defined regularities clarify some problems associated with the constructs of play and dominance–submission.
Play-fighting appears to involve the behavior patterns of attack and defense otherwise seen in serious fighting. The degree of similarity, however, depends on the body targets attacked and defended during these forms of fighting. For many taxa, including diverse mammalian families and some birds, the same targets are attacked and defended during both play-fighting and serious fighting. However, for several species of muroid rodents, the targets of play-fighting are not the same as those of serious fighting. In these cases, the tactics of attack and defense are also different. It is argued that for these muroid species the playful targets have arisen from amicable behavior (e.g., social investigation, greeting, allogrooming) rather than, as appears to be the case in so many other taxa, from agonistic behavior. These data strongly suggest that “play-fighting” has evolved from different precursors in different taxa and thus has multiple origins. Furthermore, these data have an important bearing on the universal applicability of many of the suggested functions of play-fighting and also on how such behavior is to be described and classified.
Play-fighting is often difficult to differentiate from inhibited or immature serious fighting because both may utilize many of the same behavior patterns. In the rat the two behaviors involve different targets of attack. During play-fighting, snout or oral contact is directed at the opponent's nape of the neck, whereas during serious fighting, male residents mostly direct their bites at the intruder's rump. Although similar to those used in serious fighting, the behavior patterns used during play-fighting are modified to achieve the different targets of attack. Even though the tactics of attack and defense appear more adult-like with increasing age, the playful targets persist well into adulthood.
Adult male rats living together form dominance relationships, with one dominant and the remainder adopting subordinate roles. In previous studies, it was shown that in adult male pairs, the subordinate rat initiates more playful contacts and retains a more juvenile response to the playful contacts by the dominant. In this experiment, triads were used to examine the play between subordinate males. The subordinates directed fewer playful contacts to each other than to the dominant rat, and there was a symmetrical play relationship between the subordinates. After the dominant was removed from the colony, one subordinate became the dominant. Playful interactions amongst these pairs increased, with the subordiante intiating more playful contacts than the dominant. Furthermore, from a similarly low frequency of juvenile-type response to playful contact to each other when in triads, the subordinate in the dyads increased its frequency of juvenile responses to the dominant partner. This supports the hypothesis that the playful behavior of subordinate male rats towards the dominant is an adaptive response, playful behavior of subordinate male rats towards the dominant is an adaptive response, serving a “friendship maintenance” function. Finally, when in triads, one subordinate was more playful with the dominant than the other subordinate. It was the least playful subordinate that was the most likely to become the dominant. This sugests that within a colony, not all subordinates are the same. © 1993 Wiley-Liss, Inc.
Male and female laboratory rats invariably investigate a novel conspecific placed in their home cages. In Experiment 1, mature
male rats were exposed in their home cages to active and inactive juvenile males. Inactive juveniles were pretreated with
haloperidol to induce behavioral stasis in a normally upright, quadrupedal stance. In repeated daily observations, males exposed
to active juveniles displayed significantly longer intervals of investigation than did males exposed to inactive juveniles.
In Experiment 2, mature males and females were repeatedly exposed to active and inactive castrate females. Males investigated
significantly longer than did females, active female castrates were investigated significantly longer than were inactive female
castrates, and sex of subject interacted significantly with activity-nonactivity of the social stimulus animal. In Experiment
3, mature males and females were repeatedly exposed to active and inactive castrate males. Males investigated significantly
longer than did females, active male castrates were investigated significantly longer than were inactive male castrates, and
sex of subject interacted significantly with activity-nonactivity of the social stimulus animal. The results demonstrate that
sexual dimorphism in persistence of social investigation may be interpreted as a sex difference in response to normal movement
cues of a stimulus complex characterizing a conspecific.
Play fighting in many species contains behavioral elements of direct aggression; that is, those behavior patterns that are used to threaten and contact opponents. Although many investigators have alleged that there are play signals that can be used to unambiguously distinguish playful from nonplayful aggression, the existence of such signals is mainly anecdotal and correlational. For most species, such signals are either not present, or are not present with sufficient regularity to function as unambiguous markers of play. In most playful encounters, participants seem able to use contextual cues to assertain whether the actions of the partner are affiliative or not. Where play promoting signals do appear to exist, they are often general purpose affiliative signals rather than ones specific to play. Such signals are most often associated with situations where the playful intentions of the performer may be ambiguous to the recipient. In these situations, such signals appear to be crucial in avoiding escalation to serious aggression. The danger of escalating from play to aggression is most often reported as animals become sexually mature and stronger. Also, it is in early adulthood that members of a social group compete for dominance. In these situations, a signal “this is play” may not be sufficient. Instead, signals that can retroactively appease a play partner with “I was only kidding, ” may be very important so as to manipulate, and thus obfuscate, the true intent of the action. When present, the signals occurring in play are highly variable in form and context, which is what would be expected for such subtle social manipulation. We suggest that species with more complex social relationships, where individuals are likely to probe and test their relationships with play fighting, are most likely to be the ones that make the most sophisticated use of such signals.
The paper provides a quantitative assessment for free-living baboons of: (1) The age changes and sex differences in the amount of social play and in the relative frequency of social play movement patterns; and (2) The relationships between individuals in play. Males tended to play more, were rougher, and more often took the active part in play than did females. Animals of both sexes increasingly took the active part in play with age. Some categories of movement patterns were characteristic of male play only. Factors which affected choice of play partner included age, sex, ‘maternal club’ and sibling relations. Which partner initiated and terminated play, or vocalized, was largely determined by the sex and relative age of the partners. The results are discussed in relation to the possible functions of social play in behavioural development.
Gestures and facial displays are involved in regulating many aspects of mammal social life such as aggression, dominance–subordinate relationships, appeasement and play. Playful activity is an interesting behaviour for examining the role of signals as intentional communication systems. When animals play they perform patterns that are used in other serious contexts. To avoid miscommunication, many species have evolved signals to maintain a playful mood. Bonobos, Pan paniscus, with their flexible social relationships and playful propensity, may represent a good model species to test some hypotheses on adult play signalling. I analysed the potential roles of facial play expressions and solitary play in soliciting and regulating social play and found that adult bonobos used the play face (relaxed open-mouth display) in a selective manner. Play faces were more frequent during social than solitary play and, within social play, polyadic sessions (even though less frequent than dyadic sessions) were characterized by a higher frequency of signals. Following the rule of play intensity matching, play faces were more frequent when the two players matched in age and size (sessions among adults). Moreover, among dyads there was a positive correlation between the frequency of aggressive interactions performed and the frequency of play signals used, thus suggesting that signals are crucial in play negotiations among individuals showing high baseline levels of aggression. Finally, solitary play, especially when it involved pirouettes and somersaults, had an important role in triggering social play.
Social play involves a dynamic combination of competition and cooperation, yet few studies have systematically evaluated the cooperative side of play. We studied dyadic play in domestic dogs to investigate factors influencing variation in cooperative play strategies like self-handicapping and role reversal. Dyadic play bouts were videotaped and coded for asymmetric behaviours. We predicted that variation in play style would reflect salient aspects of the canine social system, including dominance relationships and age and size differences, but not sex differences. Our results refute the 50:50 rule proposed by some researchers, which asserts that participants must equalize their behaviour to maintain a playful atmosphere. We observed divergence from 50:50 symmetry to varying degrees across dyads. This variability was especially linked to dominance and age advantages, such that higher-ranking and/or older dogs generally showed higher proportions of attacks and pursuits and lower proportions of self-handicapping than their disadvantaged play partners. These results contradict the notion that more advantaged individuals consistently relinquish their advantage to facilitate play. Role reversals did occur, but certain social conventions apparently dictated which behaviours could be used during role reversals. For example, role reversals occurred during chases and tackles, but never during mounts, muzzle bites or muzzle licks, suggesting that these latter behaviours may be invariant indicators of formal dominance during play in domestic dogs. Play signalling was linked to self-handicapping behaviour but not to attack/pursuit behaviour, indicating that perhaps self-handicapping and play signalling work together to communicate playful intent and reinforce existing roles.
Observations on the agonistic behaviour of 12 free-ranging dogs from two neighbouring groups were recorded in Katwa town, India. Both intra- and inter-group agonistic encounters were recorded on a seasonal basis. Mean (±S.E.) seasonal number of intra-group agonistic encounters of individual dog was greatest in winter (13.33±1.89) and then in late monsoon (12.33±1.99), when the females were lactating and in oestrus, respectively. Similarly, the mean (±S.E.) seasonal number of inter-group agonistic encounters of individual dog was greatest in winter (32.25±4.43) and then in late monsoon (27.75±2.01). There was a significant difference between the intra- and inter-group agonistic encounters. Dominance hierarchies were established among the adult dogs of either sex based on aggressive encounters. Although individual differences in agonism were observed, overall levels of aggression were higher among the adult females than for other groups. In contrast, overall levels of submission were higher among the juvenile males than for other groups. The results from this study suggest that reproductive season, sex and age have a significant effect on the agonistic behaviour of free-ranging dogs.
Schenkel (1947, 1967) was the first to describe ‘ritualized fighting’ in wolves. The current study identifies a set of movement patterns employed during such interactions. The relations between the interactants' movements and the contribution of each individual to these relations are emphasized. Three relational variables are employed to describe interactions: relative distance, relative orientation, and the points of opposition between the interactants. These variables in combination form a three-dimensional interaction space in which a single point describes the momentary state of the configuration of the wolves. The maintenance of four relative configurations plus five transitions between such configurations comprised a consistent set of behavioural patterns. These regular patterns of relational movements indicate that each interactant's movements are constrained in part by a set of rules related to the simultaneous movements of the social partner. In addition, the description of the actual movements in the environment by the individual interactants revealed role-dependent individual contributions by the two interactants to the coordination and management of the relational variables.
Biologists, breeders and trainers, and champion sled dog racers, Raymond and Lorna Coppinger have more than four decades of experience with literally thousands of dogs. Offering a scientifically informed perspective on canines and their relations with humans, the Coppingers take a close look at eight different types of dogsâhousehold, village, livestock guarding, herding, sled-pulling, pointing, retrieving, and hound. They argue that dogs did not evolve directly from wolves, nor were they trained by early humans; instead they domesticated themselves to exploit a new ecological niche: Mesolithic village dumps. Tracing the evolution of today's breeds from these village dogs, the Coppingers show how characteristic shapes and behaviorsâfrom pointing and baying to the sleek shapes of running dogsâarise from both genetic heritage and the environments in which pups are raised. For both dogs and humans to get the most out of each other, we need to understand and adapt to the biological needs and dispositions of our canine companions, just as they have to ours.
Actions called play signals have evolved in many species in which social play has been observed. Despite there being only few empirical demonstrations, it generally is accepted that play signals are important in the initiation ("I want to play") and maintenance ("I still want to play") of ongoing social play. In this study I consider whether a specific and highly stereotyped signal, the bow, is used to maintain social play in adult and infant domestic dogs, infant wolves, and infant coyotes. To answer this question the temporal placement of bows relative to actions that are also used in other contexts (dominance or predatory encounters) such as biting accompanied by rapid side-to-side shaking of the head was analyzed to determine if bows performed during ongoing social play are used to communicate the message "I want to play despite what I am going to do or just did -- I still want to play." The nonrandom occurrence of bows supports the hypothesis that bows are used to maintain social play in these canids when actions borrowed from other contexts, especially bites accompanied by rapid side-to-side shaking of the head, are likely to be misinterpreted.
The social cognitive capacities of dogs, including their communication skills and use of visual attention cues, have recently been investigated in numerous experimental studies. This paper reports on research of domestic dog behavior in a natural setting, which shows sensitivity to the visual attention of their partners when engaged in dyadic rough-and-tumble play. The sequential behaviors and head-direction of both dogs were noted throughout the bouts. The behaviors were differentially used according to the partner's posture. Play signals were sent nearly exclusively to forward-facing conspecifics; attention-getting behaviors were used most often when a playmate was facing away, and before signaling an interest to play. In addition, the mode of attention-getter matched the degree of inattentiveness of the playmate: stronger attention-getters were used when a playmate was looking away or distracted, less forceful ones when the partner was facing forward or laterally. In other words, these dogs showed attention to, and acted to manipulate, a feature of other dogs that mediates their ability to respond: which feature in human interaction is called "attention".
Attack of dominant colony males of an albino rat (Rattus norvegicus) strain, on introduced strangers, produced a non-random distribution of bites, with ventral trunk virtually never bitten. Also, vibrissae-contact of attacker and defender interfered with bites to the defender's head and upper back. The specific agonistic reactions of attacking and defending rats appeared to involve strategies based on these limitation on attack: defenders utilized 'boxing' and lying 'on-the-back' behaviour, which interposed ventral trunk and vibrissae between attacker and defender. In turn, the 'lateral display' permitted attackers to circumvent the defender's behaviour. Limitations on attack therefore appeared to underlie the specific agonistic behaviour of both attacking and defending rats.
A quantitative comparison is given of aggressive play and aggression in terms of the duration and type of movement patterns employed, and the relationships between individuals. The mean length of contact in play was greater than in aggression, whereas chases tended to be longer in adult male aggression than in play, Linear hierarchies describing the overall direction of interactions between individuals were divisible into two sorts, according to the frequency with which roles were reversed. The regions of the body bitten and the proportions in which six types of movement pattern were used were related to the type of hierarchy involved. The results are discussed with reference to the possible functions of social play in the development of movement patterns and individual relationships.
Play fighting is a frequent activity of juvenile rats and appears to show marked variability amongst individuals in that some rats play a great deal and others very little. This study attempted to identify some of the factors involved in producing this individual variability. The major influence over an individual's frequency of play as a juvenile was found to be the frequency of play by the partner. That is, play appears to be contagious, in that a high playing animal stimulates its partner to play frequently as well. In male juveniles, but seemingly not in female juveniles, the subsequent adult status of one partner as dominant influences the subordinate-to-be to initiate more playful contacts. In addition to these extrinsic influences, however, there appear to be intrinsic factors that influence whether an individual is a high or low playing animal. One intrinsic factor appears to be 'boldness', so that bolder animals tend to initiate more playful contacts. Higher players tend to be more susceptible to the stereotypy-inducing effects of the dopamine agonist, apomorphine, and tend to be more dependent upon the playful activity of the partner to maintain their own high levels of play. Both of these characteristics are consistent with other studies comparing bold and timid rats. Boldness, however, only seems to influence how much play a rat will exhibit, not how much play it is capable of exhibiting. Neonatal testosterone augmentation increases juvenile play fighting but not apomorphine susceptibility, suggesting that a high player need not be a bold animal. The total frequency of play an individual is capable of initiating appears to depend upon perinatal exposure to androgens. Boldness and the playfulness of the partner appear to modulate the expression of this hormonally set value.
Comparative analysis of mammalian genomes provides important insight into the structure and function of genes. However, the comparative analysis of gene sequences from individuals of the same and different species also provides insight into the evolution of genes, populations, and species. We exemplify these two uses of genomic information. First, we document the evolutionary relationships of the domestic dog to other carnivores by using a variety of DNA-based information. A phylogenetic comparison of mitochondrial DNA sequences in dogs and gray wolves shows that dogs may have originated from multiple wolf populations at a time much earlier than suggested by the archaeologic record. We discuss previous theories about dog development and evolution in light of the new genetic data. Second, we review recent progress in dog genetic mapping due to the development of hypervariable markers and specific chromosome paints. Extensive genetic homology in gene order and function between humans and dogs has been discovered. The dog promises to be a valuable model for identifying genes that control morphologic differences between mammals as well as understanding genetically based disease.
Bekoff [J. Consci. Stud. 8 (2001) 81] argued that mammalian social play is a useful behavioral phenotype on which to concentrate in order to learn more about the evolution of fairness. Here, we build a game theoretical model designed to formalize some of the ideas laid out by Bekoff, and to examine whether 'fair' strategies can in fact be evolutionarily stable. The models we present examine fairness at two different developmental stages during an individual's ontogeny, and hence we create four strategies-fair at time 1/fair at time 2, not fair at time 1/not fair at time 2, fair at time 1/not fair at time 2, not fair at time 1/fair at time 2. Our results suggest that when considering species where fairness can be expressed during two different developmental stages, acting fairly should be more common than never acting fairly. In addition, when no one strategy was evolutionarily stable, we found that all four strategies we model can coexist at evolutionary equilibrium. Even in the absence of an overwhelming database from which to test our model, the general predictions we make have significant implications for the evolution of fairness.
Evolution of Canine Social Behavior Play Fighting
- R Abrantes
Abrantes, R., 2005. Evolution of Canine Social Behavior. Dogwise Publishing, Wenatchee, WA. Aldis, O., 1975. Play Fighting. Academic Press, NY.
Genome sequencing highlights the dynamic early history of dogs Homeostatic motor processes in mammalian interactions: a chore-ography of display
- A H Freedman
- I Gronau
- R M Schweizer
- D Vecchyo
- E Han
- P M Silva
- M Galav-Erni
- Z Fan
- P Marx
- B Lorente-Galdos
- H Beale
- O Ramirez
- F Hormozdiari
- C Alkan
- C Vila
- K Squire
- E Geffen
- J Kusak
- A R Boyko
- H G Parker
- C Lee
- V Tadigotla
- A Siepel
- C D Bustamante
- T T Harkins
- S F Nelson
- E A Ostrander
- T Bonet
- R K Wayne
- J Novembre
Freedman, A.H., Gronau, I., Schweizer, R.M., Vecchyo, D., Han, E., Silva, P.M., Galav-erni, M., Fan, Z., Marx, P., Lorente-Galdos, B., Beale, H., Ramirez, O., Hormozdiari, F., Alkan, C., Vila', C., Squire, K., Geffen, E., Kusak, J., Boyko, A.R., Parker, H.G., Lee, C., Tadigotla, V., Siepel, A., Bustamante, C.D., Harkins, T.T., Nelson, S.F., Ostrander, E.A., Marques-Bonet, T., Wayne, R.K., Novembre, J., 2014. Genome sequencing highlights the dynamic early history of dogs. PLoS Genet. 10, e1004016. Golani, I., 1976. Homeostatic motor processes in mammalian interactions: a chore-ography of display. In: Bateson, P.P.G., Klopfer, P.H. (Eds.), Perspectives in Ethology, vol. 2. Plenum, New York, pp. 69–134.