Costs and benefits of flocking in foraging white-fronted geese (Anser albifrons): effects of resource depletion

This study investigated the costs and benefits of flocking in white-fronted geese Anser albifrons foraging on rice grains in Japan. The time budgets of focal geese were recorded, and the effects of flock size on the proportions of time spent in vigilant and agonistic behaviour were tested. The results showed that the decline in vigilance level and consequent increase in foraging time were beneficial results of flocking whereas agonistic interactions, a potential cost of flocking, did not increase with increasing flock size. However, seasonal variation in flock size suggested that exploitative competition could be a cost of flocking; the sizes of flocks in spring, when resource depletion had progressed, were significantly reduced compared with those in autumn. An experimental increase in rice density resulted in a significant increase in flock size. We conclude that the flock size of foraging white-fronted geese is a result of compromise between a constant benefit of flocking (i.e. decline in vigilance level) and a cost of flocking varying with food abundance (i.e. exploitative competition).     

Ontogenie und Funktion der Rangordnung innerhalb einer Geschwisterschar von Graugänsen (Anser anser L.)

Between the siblings of a family of greylag geese (Anser anser L.) a rank order is established by fighting during the first days after hatching, and also by other, more complex interactions. The position of each goose in the rank order can be defined by three groups of behaviour patterns (aggressive behaviour, “greetings”, escape responses). A clear-cut ranking relation between two individuals is a prerequisite for their close cohesion. The survival function of rank order lies in diminishing the intensity and frequency of agonistic interactions, in diminishing social stress situations and in guaranteeing the close family cohesion.     

Social behaviour and heart rate in tammar wallabies (Macropodidae: Macropus eugenii)

In an enclosed group of tammar wallabies the behaviour of all individuals and the heart rate (HR) of three males were recorded simultaneously. The social structure was characterized by the dominance relationships between the males and the sexual preference of all males for the oldest female. During non-oestrus, dominance was obvious when a male intervened terminating the sexual behaviour of a subdominant one. The relatively low amount of agonistic behaviour between the males is assumed to be due to the clear dominance relationships. Locomotion as well as sexual and agonistic behaviour were accompanied by an acceleration of HR. The HR pattern depended on the intensity of the behaviour, its duration and the animal's identity. Commonly performed sexual behaviour and approach/retreat encounters gave rise to HR values not different from HR values during locomotor activity without interactive components. Social events that occurred infrequently—like fights, and the mating of an oestrous female—were accompanied by strong accelerations of HR indicating high cardiac effort. The mild cardiac activity during common social encounters is consistent with a strategy of maximizing energy conservation. HR patterns during specific behaviour could not entirely be accounted for by the energetic costs of activity. Spreading the forelimbs in response to specific stimuli—like the fly-over of raptors—was indicative of a strong HR response even if the animal was motionless. During specific social encounters like the sexual behaviour of the alpha-male following an intervention—HR responses revealed that arousal might exceed motor activity in affecting HR. Individual differences of these HR responses are attributed to the age, experience and social status of a male.     

Inter-individual relationships in proboscis monkeys: a preliminary comparison with other non-human primates

This is the first report on inter-individual relationships within a one-male group of proboscis monkeys (Nasalis larvatus) based on detailed identification of individuals. From May 2005 to 2006, focal and ad libitum data of agonistic and grooming behaviour were collected in a forest along the Menanggul River, Sabah, Malaysia. During the study period, we collected over 1,968 h of focal data on the adult male and 1,539 h of focal data on the six females. Their social interactions, including agonistic and grooming behaviour, appeared to follow typical patterns reported for other colobines: the incidence of social interaction within groups is low. Of 39 agonistic events, 26 were displacement from sleeping places along the river, 6 were the α male threatening other monkeys to mediate quarrels between females and between females and juveniles, and 7 were displacement from feeding places. Although the agonistic behaviour matrix based on the 33 intra-group agonistic events (excluding events between adults and juveniles and between adults and infants) was indicative of non-significant linearity, there were some specific dominated individuals within the group of proboscis monkeys. Nonetheless, grooming behaviour among adult females within a group were not affected by the dominance hierarchy. This study also conducted initial comparisons of grooming patterns among proboscis monkeys and other primate species. On the basis of comparison of their grooming networks, similar grooming patterns among both-sex-disperse and male-philopatric/female-disperse species were detected. Because adult females in these species migrate to groups repeatedly, it may be difficult to establish the firm grooming exchange relationship for particular individuals within groups, unlike in female-philopatric/male-disperse species. However, grooming distribution patterns within groups among primate species were difficult to explain solely on the basis of their dispersal patterns. Newly immigrated females in some species including proboscis monkeys are eager to have social interactions with senior group members to improve their social position.     

Physiological implications of pair-bond status in greylag geese

In group-living vertebrates, reliable social allies play a decisive role in dealing with stressors. For example, support by social allies is known to dampen glucocorticoid responses. It remains unknown, however, how social embedding affects the sympatho-adrenergic axis as indicated by heart rate (HR) in non-human animals. We studied the relationships between HR, pair-bond status and distance from the pair-partner in twenty-five free-ranging greylag geese (Anser anser) in a natural social environment. In three individuals, we investigated HR responses following partner loss. Overall, we found a context- and sex-dependent difference in HR between paired and unpaired individuals, paired males having a lower HR during agonistic encounters, and unpaired females having a lower HR during resting. Also, in paired females HR increased with increasing distance from the partner. Our data suggest that HR is modulated by pair-bond status in greylag geese in a context- and sex-dependent manner, which may be representative for social vertebrates in general. Despite the low sample size, the present study indicates that proper social embedding may optimize an individual's physiological investment in the social domain. This reduces individual energy expenditure and may benefit health and reproductive success.     

Heart rate during conflicts predicts post-conflict stress-related behavior in greylag geese

Background

Social stressors are known to be among the most potent stressors in group-living animals. This is not only manifested in individual physiology (heart rate, glucocorticoids), but also in how individuals behave directly after a conflict. Certain ‘stress-related behaviors’ such as autopreening, body shaking, scratching and vigilance have been suggested to indicate an individual''s emotional state. Such behaviors may also alleviate stress, but the behavioral context and physiological basis of those behaviors is still poorly understood.

Methodology/Principal Findings

We recorded beat-to-beat heart rates (HR) of 22 greylag geese in response to agonistic encounters using fully implanted sensor-transmitter packages. Additionally, for 143 major events we analyzed the behavior shown by our focal animals in the first two minutes after an interaction. Our results show that the HR during encounters and characteristics of the interaction predicted the frequency and duration of behaviors shown after a conflict.

Conclusions/Significance

To our knowledge this is the first study to quantify the physiological and behavioral responses to single agonistic encounters and to link this to post conflict behavior. Our results demonstrate that ‘stress-related behaviors’ are flexibly modulated by the characteristics of the preceding aggressive interaction and reflect the individual''s emotional strain, which is linked to autonomic arousal. We found no support for the stress-alleviating hypothesis, but we propose that stress-related behaviors may play a role in communication with other group members, particularly with pair-partners.     

An ethological analysis of types of agonistic interaction in a captive group of Java-monkeys (Macaca fascicularis)

The primate literature provides many indications not only that the nature of dyadic interactions is to a large extent determined by the relations of the interacting animals with others and between these others, but also of the existence of polyadic interactions in which more than two individuals are simultaneously involved. The objectives of the present study are to obtain a quantitative categorization of the agonistic interaction types of captive Java-monkeys and an analysis of their dynamics. After having described the agonistic behaviour patterns of Java-monkeys we shall discuss the categorization of agonistic interaction types (depending on the number of involvees: “dyads”, “triads” and “polyads”), the way in which these types can be further differentiated on the basis of the nature and the direction of the behaviours shown (e.g., different types of alliances), and the existence of so-called “sub-directed” behaviours (i.e., non-agonistic behaviours which are shown towards a dominant third animal more or less simultaneously with aggressive behaviour directed towards an opponent). The analysis indicates that agonistic behaviour is different both in its form and its regulation in interactions of different complexity. This research was supported in part by a government grant (i.e.: Beleidsruimte project: 16-21-06, “Brain and Behaviour”) to the first author. The investigation was supported by a grant from the Beleidsruimtemiddelen Hersenen en Gedrag to the first author.     

Social Use of Facial Expressions in Hylobatids

Non-human primates use various communicative means in interactions with others. While primate gestures are commonly considered to be intentionally and flexibly used signals, facial expressions are often referred to as inflexible, automatic expressions of affective internal states. To explore whether and how non-human primates use facial expressions in specific communicative interactions, we studied five species of small apes (gibbons) by employing a newly established Facial Action Coding System for hylobatid species (GibbonFACS). We found that, despite individuals often being in close proximity to each other, in social (as opposed to non-social contexts) the duration of facial expressions was significantly longer when gibbons were facing another individual compared to non-facing situations. Social contexts included grooming, agonistic interactions and play, whereas non-social contexts included resting and self-grooming. Additionally, gibbons used facial expressions while facing another individual more often in social contexts than non-social contexts where facial expressions were produced regardless of the attentional state of the partner. Also, facial expressions were more likely ‘responded to’ by the partner’s facial expressions when facing another individual than non-facing. Taken together, our results indicate that gibbons use their facial expressions differentially depending on the social context and are able to use them in a directed way in communicative interactions with other conspecifics.     

Integrating personality research and animal contest theory: aggressiveness in the green swordtail Xiphophorus helleri

Aggression occurs when individuals compete over limiting resources. While theoretical studies have long placed a strong emphasis on context-specificity of aggression, there is increasing recognition that consistent behavioural differences exist among individuals, and that aggressiveness may be an important component of individual personality. Though empirical studies tend to focus on one aspect or the other, we suggest there is merit in modelling both within- and among-individual variation in agonistic behaviour simultaneously. Here, we demonstrate how this can be achieved using multivariate linear mixed effect models. Using data from repeated mirror trials and dyadic interactions of male green swordtails, Xiphophorus helleri, we show repeatable components of (co)variation in a suite of agonistic behaviour that is broadly consistent with a major axis of variation in aggressiveness. We also show that observed focal behaviour is dependent on opponent effects, which can themselves be repeatable but were more generally found to be context specific. In particular, our models show that within-individual variation in agonistic behaviour is explained, at least in part, by the relative size of a live opponent as predicted by contest theory. Finally, we suggest several additional applications of the multivariate models demonstrated here. These include testing the recently queried functional equivalence of alternative experimental approaches, (e.g., mirror trials, dyadic interaction tests) for assaying individual aggressiveness.     

THE SHORT-TERM BEHAVIORAL REACTIONS OF BOTTLENOSE DOLPHINS TO INTERACTIONS WITH BOATS IN DOUBTFUL SOUND, NEW ZEALAND

Doubtful Sound is home to one of the southernmost resident populations of bottlenose dolphins ( Tursiops sp.). This population regularly interacts with scenic cruises. During these interactions, dolphins tend to horizontally and vertically avoid vessels, especially when the behavior of these vessels is intrusive. This study aimed at understanding the behavioral reactions of individuals to these interactions that lead to the disruption of the school's behavioral state. Observing the behavioral events performed by individuals during an interaction can help define the short-term reactions elicited by the boat presence. I recorded the behavioral events performed by all individuals of focal schools. The frequency of occurrence of all events was compared depending on the presence of vessels, their behavior, and the behavioral state of the focal school. Dolphins tended to perform more side flops while interacting with powerboats, a behavior which may be used as a non-vocal communication tool. Moreover, the movement of dolphins became more erratic during interactions with all types of vessels. These effects increased when the boats were more intrusive while interacting. This study shows that the impact of interaction with boats can be minimized if the vessels respect the guidelines in place.     

Lateralization in monogamous pairs:wild geese prefer to keep their partner in the left hemifield except when disturbed

Behavioural lateralization, which reflects the functional specializations of the two brain hemispheres, is assumed to play an important role in cooperative intraspecific interactions. However, there are few studies focused on the lateralization in cooperative behaviours of individuals, especially in a natural setting. In the present study, we investigated lateralized spatial interactions between the partners in life-long monogamous pairs. The male-female pairs of two geese species (barnacle, Branta leucopsis, and white-fronted, Anser albifrons geese), were observed during different stages of the annual cycle in a variety of conditions. In geese flocks, we recorded which visual hemifield (left/right) the following partner used to monitor the leading partner relevant to the type of behaviour and the disturbance factors. In a significant majority of pairs, the following bird viewed the leading partner with the left eye during routine behaviours such as resting and feeding in undisturbed conditions. This behavioural lateralization, implicating the right hemisphere processing, was consistent across the different aggregation sites and years of the study. In contrast, no significant bias was found in a variety of geese behaviours associated with enhanced disturbance (when alert on water, flying or fleeing away when disturbed, feeding during the hunting period, in urban area feeding and during moulting). We hypothesize that the increased demands for right hemisphere processing to deal with stressful and emergency situations may interfere with the manifestation of lateralization in social interactions.     

Effects of Passive Social Support in Juvenile Greylag Geese (Anser anser): A Study from Fledging to Adulthood

In general, support by social allies may reduce stress, increase an individual's status and facilitate access to resources. In Greylag geese (Anser anser), offspring stay with their parents for an entire year or even longer and thereby enjoy social support in encounters with other flock members. We investigated the influence of spatial distance to one's allies on the outcome of agonistic encounters in a natural flock situation for a total of 12 sibling groups after fledging. In addition, we tested two groups of hand‐raised juvenile geese over a time span of 11/2 yr. Passive (i.e. not interfering) human supporters of different familiarity were placed at a standard distance during food provisionings, which produced a tight flock situation. Success in agonistic interactions increased with decreasing distance to members of their social unit. The hand‐raised juveniles were more successful in agonistic interactions and showed increased feeding rates when accompanied by a familiar human than when alone or with a non‐familiar human. The effect of the presence of familiar humans on success in agonistic encounters significantly decreased with increasing age, while feeding rates remained elevated. The positive effects of social support were particularly evident in females. We conclude that social support has similar effects and functions in the highly social greylag geese as reported for social mammals.     

The behaviour of the common hamster (Cricetus cricetus) under zoo conditions

Studies on the behaviour of the common hamster are very rare and fragmentary. From August till the middle of September 2005 we observed behaviour of six individuals of the common hamster (one female and five juveniles: three males and two females) and from the second part of September till October 2005 we observed behaviour of two juvenile males after they were separated from the group. The “Focal animal sampling” was used as an observation method, and the total time of observation amounted to 75 hours. During our observation 7707 bouts of behaviour were recorded. Both social and non-social behaviour were categorised. The number of the non-social behaviour prevailed significantly over the number of the social behaviour (n=496; 6.4%). The most frequent observed non-social behaviour was: head rearing (n=2019; 28%). Among interindividual interactions “escape” (45.4%; n=225) and “vocalisation” (15.7%; n=78) were noticed most frequently. Play-fighting, direct identification, following and hugging were the only aspects of the social behaviour characteristic of the young.The behaviour repertoire of the common hamster seems to be similar to the behaviour of the golden hamster (Mesocricetus auratus).     

Physiological correlates of social behaviour in Bighorn sheep: a field study using electrocardiogram telemetry

Telemetered heart rates (HR) were correlated with social interactions of adult free-ranging Bighorn sheep (0vi.s canadensis canadensis). Minimal cardiac responses suggest that economy of expression is a pervasive feature of social behaviour in this species. Commonly performed courtship, dominance and agonistic displays elicited few HR responses in either sender or receiver; those that did occur, were transitory and usually accompanied motor reactions by the subject. Intense behavioural acts that occurred infrequently, such as mount attempts, rush-charges and clashes, evoked maximal HR increases in both performer and receiver. These findings are consistent with the complex social behaviour of Bighorn sheep and a life-strategy demanding frugal investment of resources in maintenance, in order to maximize growth and reproduction.     

'Better off alone than in bad company': agonistic colour display in mimetic juveniles of two ephippid species

Comparative field observations of agonistic interactions in juvenile leaf-mimicking Platax orbicularis and Chaetodipterus faber (Ephippidae) were conducted in coastal waters of the Pacific and Atlantic Oceans. Similar agonistic behaviour was observed in the two species, in which individuals stopped displaying their mimetic colouration during encounters with conspecifics, to display conspicuous colours, such as transverse stripes along the body. These events were observed occasionally, almost invariably in individuals of smaller body size. Larger-bodied individuals of both species spent less time in agonistic displays. The absolute size of the fish, however, did not appear to affect the outcome of the encounter, suggesting that dominance is a temporary condition, based on the relative size of the opponents during encounters.     

The ontogeny of agonistic behaviour in the blue gourami, Trichogaster trichopterus (Pisces,Anabantoidei)

The development of agonistic behaviour in the blue gourami, Trichogaster trichopterus, was studied from hatching to four months of age. Larval motor patterns appear first as simple, uncoordinated, and irregular movements and gradually become regular and well coordinated, forming integrated complex patterns, such as swimming, surfacing, and feeding. Most motor patterns used in agonistic interactions are first exhibited in non-social contexts. Initial interactions between young fish consist of Approach and investigation in a feeding context, in some cases followed by nibbling, Biting, and Chasing by one fish. Later, initial Approaches are separated from Biting and Chasing by Lateral Displays, circling, Tail Beating, and Fin Tugging, and by Appeasement behaviour that inhibits Chasing and Biting by the opponent.Agonistic behaviour of socially isolated fish, when paired in dyadic encounters as adults, indicates that the role of social experience as an integrator and regulator of social behaviour may change progressively during development and become more important as ontogeny proceeds.     

Coping with Conflict during Initial Encounters in Chimpanzees

When strangers meet, conflict is likely to arise from incompatibilities in motivations and expectations, as well as from the absence of predictability in interactions. This study explores the mechanisms that mitigate aggression, permit mutual evaluation, establish tolerance, and facilitate the development of social bonds between unfamiliar chimpanzees ( Pan troglodytes ). Data collected during the initial half-hour of the introduction of 33 unfamiliar pairs (16 female-male and 17 female-female) at the Yerkes Primate Center and Detroit Zoo were used to assess the sequence of events during initial encounters and the function of different behaviour patterns in the formation of relationships. Initial encounters followed a regular sequence of interactions (i.e. a 'species-typical etiquette'), commencing with agonistic behaviour, followed by brief friendly touches, and finally allogrooming. Brief friendly touches, which are associated with conciliatory and tension-reducing functions in established relationships, appeared to serve an evaluative function related to status differentiation during initial interactions. In fact, individuals more closely matched in competitive ability (inferred from their eventual dominance rank distance) exchanged friendly touches at higher rates than those more obviously mismatched. These evaluative touches may represent a safer alternative to more assertive interactions when assessments of competitive ability are required in situations likely to escalate. Allogrooming promoted the development of tolerance and relaxed proximity; it was, in fact, effective in reducing agonism and negative-outcome approaches. Allogrooming may take on a particularly important role in reducing agonism because it begins the commerce of benefits that balance the dispersive competitive forces and enable cohesive social groupings.     

Fights at the Dinner Table: Agonistic Behavior in Pachydiplax longipennis (Odonata: Libellulidae) at Feeding Sites

Aggressive behavior of Pachydiplax longipennis during foraging was quantified by observing focal individuals on arrays of artificial perches. Pachydiplax apparently aggressively defend, for up to several hours at a time, one or a few feeding perches. Seventeen percent of all behaviors included agonistic actions, e.g., chasing or physical contact. The frequency of interactions was correlated positively with ambient temperature, solar radiation, prey density and density of other dragonflies. Both sexes initiated and responded to intra- and interspecific aggression; intraspecific interactions were more intense, however. Males had significantly higher interaction rates and fighting success than females, and intraspecific male–male contests were particularly intense. When prey were visibly localized, contest winners commonly gained perches closer to the prey swarm, and aggressive behavior was apparently correlated with feeding opportunity. Despite the frequency of aggression, these dragonflies allocated only about 19 s, on average, to agonistic behavior during 30-min observation periods. This and other costs appear small compared to foraging benefits of occupying a favorable perch, although at a very high interaction intensity high energy costs and lower intake reduce the net energy gain.