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de Waal, F. B., & Luttrell, L. M. (1986). The similarity principle underlying social bonding among female rhesus monkeys. Folia Primatol (Basel), 46(4), 215–234.
Abstract: Twenty adult female rhesus monkeys (Macaca mulatta) were observed over a three-year period. They lived in a mixed captive group with kinship relations known for three generations. The study's aim was to test Seyfarth's [J. theor. Biol. 65: 671-698, 1977] model of rank-related grooming and to investigate two other possible determinants of social bonding, i.e. relative age and the group's stratification into two social classes. Data on affiliation, coalitions, and social competition were collected by means of both focal observation and instantaneous time sampling. Whereas certain elements of the existing model were confirmed, its explanatory principles were not. Social competition did not result in more contact among close-ranking females (the opposite effect was found), and the relation between affiliative behavior and coalitions was more complex than predicted. Based on multivariate analyses and a comparison of theoretical models, we propose a simpler, more encompassing principle underlying interfemale attraction. According to this 'similarity principle', rhesus females establish bonds with females whom they most resemble. The similarity may concern genetical and social background, age, hierarchical position and social class. Effects of these four factors were independently demonstrated. The most successful model assumed that similarity factors influence female bonding in a cumulative fashion.
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de Waal, F. B. (1977). The organization of agonistic relations within two captive groups of Java-monkeys (Macaca fascicularis). Z. Tierpsychol., 44(3), 225–282.
Abstract: The paper offers a detailed quantitative descripition of the distribution of agonistic activities over the members of two groups of Java-monkeys (Macaca fascicularis). These groups lived in captivity and were well-established: i.e. they had an extensive network of genealogical relationships. The study pays special attention to agonistic interactions with three or more participants. Its main purpose is an analysis of the way dyadic agonistic relations (e.g. dominance relations) are affected by third group members and the relations among these. The paper presents data on the ontogeny of 'dependent dominance', the 'control role' of the alpha-male, and the functions of different types of alliances.
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Heitor, F., do Mar Oom, M., & Vicente, L. (2006). Social relationships in a herd of Sorraia horses Part I. Correlates of social dominance and contexts of aggression. Behav. Process., 73(2), 170–177.
Abstract: Factors related to dominance rank and the functions of aggression were studied in a herd of Sorraia horses, Equus caballus, under extensive management. Subjects were 10 adult mares 5-18 years old and a stallion introduced into the group for breeding. Dominance relationships among mares were clear, irrespective of rank difference, and remained stable after introduction of the stallion. The dominance hierarchy was significantly linear and rank was positively correlated with age and total aggressiveness. Higher-ranking mares received lower frequency and intensity of agonistic interactions. Nevertheless, higher-ranking dominants were not more likely to elicit submission from their subordinates than lower-ranking dominants. Neither close-ranking mares nor mares with less clear dominance relationships were more aggressive towards each other. Agonistic interactions seemed to be used more importantly in regulation of space than to obtain access to food or to reassert dominance relationships. Contexts of aggression were related to mare rank. The results suggest that dominance relationships based on age as a conventional criterion were established to reduce aggressiveness in a herd where the costs of aggression are likely to outweigh the benefits.
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Hemelrijk, C. K., & Wantia, J. (2005). Individual variation by self-organisation. Neurosci Biobehav Rev, 29(1), 125–136.
Abstract: In this paper, we show that differences in dominance and spatial centrality of individuals in a group may arise through self-organisation. Our instrument is a model, called DomWorld, that represents two traits that are often found in animals, namely grouping and competing. In this model individual differences grow under the following conditions: (1) when the intensity of aggression increases and grouping becomes denser, (2) when the degree of sexual dimorphism in fighting power increases. In this case the differences among females compared to males grow too, (3) when, upon encountering another individual, the tendency to attack is 'obligate' and not conditional, namely 'sensitive to risks'. Results resemble phenomena described for societies of primates, mice, birds and pigs.
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Johnstone, R. A. (2001). Eavesdropping and animal conflict. Proc. Natl. Acad. Sci. U.S.A., 98(16), 9177–9180.
Abstract: Fights between pairs of animals frequently take place within a wider social context. The displays exchanged during conflict, and the outcome of an encounter, are often detectable by individuals who are not immediately involved. In at least some species, such bystanders are known to eavesdrop on contests between others, and to modify their behavior toward the contestants in response to the observed interaction. Here, I extend Maynard Smith's well known model of animal aggression, the Hawk-Dove game, to incorporate the possibility of eavesdroppers. I show that some eavesdropping is favored whenever the cost of losing an escalated fight exceeds the value of the contested resource, and that its equilibrium frequency is greatest when costs are relatively high. Eavesdropping reduces the risk of escalated conflict relative to that expected by chance, given the level of aggression in the population. However, it also promotes increased aggression, because it enhances the value of victory. The net result is that escalated conflicts are predicted to occur more frequently when eavesdropping is possible.
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Earley, R. L., & Dugatkin, L. A. (2002). Eavesdropping on visual cues in green swordtail (Xiphophorus helleri) fights: a case for networking. Proc Biol Sci, 269(1494), 943–952.
Abstract: Aggressive contests probably occur in networking environments where information about fighting ability is conveyed both to an opponent and to individuals peripheral to the fight itself, the bystanders. Our primary aim was to investigate the relative influences of eavesdropping and prior social experience on the dynamics of aggressive contests in Xiphophorus helleri. A bystander's ability to witness an encounter was manipulated using clear, one-way mirror, and opaque partitions. After watching (or not watching) the initial contest, the bystander encountered either the winner or loser of the bout. Treatment comparisons of bystander-winner or bystander-loser contest dynamics indicated the presence or absence of winner, loser, or eavesdropping effects. Winner and loser effects had negligible influences on bystander contest dynamics. Eavesdropping significantly reduced the bystander's propensity to initiate aggression, escalate, and win against seen winners regardless of whether the watched bout had escalated or not. Though eavesdropping had relatively little effect on bystander-loser contest dynamics, bystanders were less prone to initiate aggression and win against losers that had escalated in the witnessed bout. Thus, bystanders appear to preferentially retain and utilize information gained about potentially dangerous opponents (winners or persistent losers). Our data lend clear support for the importance of eavesdropping in visually based aggressive signalling systems.
Keywords: *Aggression; Animals; *Behavior, Animal; *Cyprinodontiformes; Female; Male
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Peake, T. M., Terry, A. M. R., McGregor, P. K., & Dabelsteen, T. (2002). Do great tits assess rivals by combining direct experience with information gathered by eavesdropping? Proc Biol Sci, 269(1503), 1925–1929.
Abstract: Animals frequently use signals that travel further than the spacing between individuals. For every intended recipient of a given signal there are likely to be many other individuals that receive information. Eavesdropping on signalling interactions between other individuals provides a relatively cost-free method of assessing future opponents or mates. Male great tits (Parus major) extract relative information from such interactions between individuals unknown to them. Here, we show that male great tits can take information gathering a stage further and obtain more information about a previously unencountered intruder, by the hitherto unknown capability of combining information gathered by eavesdropping with that derived from their own direct interaction with an individual. Prior experience with an intruder (A) was achieved by subjecting a focal male to different levels of intrusion simulated using interactive playback. This intruder (A) then took part in a simulated interaction with an unknown male (B) outside the territorial boundary of the focal males. In response to subsequent intrusion by the second male (B), focal males showed low song output in response to males that had lost to a male that the subject was able to beat. Males of known high quality, or those about which information was ambiguous, elicited a high level of song output by focal males. We discuss the implications of this finding for the evolution of communication and social behaviour.
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Dzieweczynski, T. L., Eklund, A. C., & Rowland, W. J. (2006). Male 11-ketotestosterone levels change as a result of being watched in Siamese fighting fish, Betta splendens. Gen Comp Endocrinol, 147(2), 184–189.
Abstract: This study investigated the effects of nesting status and the presence of an audience on 11-ketotestosterone (11KT) levels in male Siamese fighting fish, Betta splendens. Prior studies have demonstrated that both nesting status, an indicator of territory-holding power and reproductive state, and the sex of a conspecific audience lead to differences in male behavior during aggressive encounters. Since behavioral changes have already been demonstrated, we chose to investigate whether 11KT levels were also influenced by nesting status and audience presence as 11KT both stimulates, and is stimulated by, reproductive and aggressive behaviors in male teleosts. Male 11KT levels were measured from water samples taken from containers holding fish both before and after interaction. Males interacted under three treatment conditions: no audience, female audience, and male audience. Within these treatments were two nest paradigms: both males had nests or neither male had a nest. 11KT levels varied depending on nesting status and audience type. In general, 11KT levels were lower in interacting males when a female audience was present or when males had nests. Overall, 11KT showed increases or decreases as aggression increased or decreased, as shown by already established behavioral findings [see Dzieweczynski T.L., Green T.M., Earley R.L., Rowland W.J., 2005. Audience effect is context dependent in Siamese fighting fish, Betta splendens. Behav. Ecol. 16, 1025-1030; Doutrelant, C., McGregor, P.K., Oliveira, R.F., 2001. Effect of an audience on intrasexual communication in male Siamese fighting fish (Betta splendens). Behav. Ecol. 12, 283-286.]. Our results suggest that 11KT levels are influenced by reproductive status, as indicated by nest ownership, and audience presence and are most likely modulated by territorial behavior and social environment.
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Drent, P. J., van Oers, K., & van Noordwijk, A. J. (2003). Realized heritability of personalities in the great tit (Parus major). Proc Biol Sci, 270(1510), 45–51.
Abstract: Behaviour under conditions of mild stress shows consistent patterns in all vertebrates: exploratory behaviour, boldness, aggressiveness covary in the same way. The existence of highly consistent individual variation in these behavioural strategies, also referred to as personalities or coping styles, allows us to measure the behaviour under standardized conditions on birds bred in captivity, link the standardized measurements to the behaviour under natural conditions and measure natural selection in the field. We have bred the great tit (Parus major), a classical model species for the study of behaviour under natural conditions, in captivity. Here, we report a realized heritability of 54 +/- 5% for early exploratory behaviour, based on four generations of bi-directional artificial selection. In addition to this, we measured hand-reared juveniles and their wild-caught parents in the laboratory. The heritability found in the mid-offspring-mid-parent regression was significantly different from zero. We have thus established the presence of considerable amounts of genetic variation for personality types in a wild bird.
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Grosenick, L., Clement, T. S., & Fernald, R. D. (2007). Fish can infer social rank by observation alone. Nature, 445(7126), 429–432.
Abstract: Transitive inference (TI) involves using known relationships to deduce unknown ones (for example, using A > B and B > C to infer A > C), and is thus essential to logical reasoning. First described as a developmental milestone in children, TI has since been reported in nonhuman primates, rats and birds. Still, how animals acquire and represent transitive relationships and why such abilities might have evolved remain open problems. Here we show that male fish (Astatotilapia burtoni) can successfully make inferences on a hierarchy implied by pairwise fights between rival males. These fish learned the implied hierarchy vicariously (as 'bystanders'), by watching fights between rivals arranged around them in separate tank units. Our findings show that fish use TI when trained on socially relevant stimuli, and that they can make such inferences by using indirect information alone. Further, these bystanders seem to have both spatial and featural representations related to rival abilities, which they can use to make correct inferences depending on what kind of information is available to them. Beyond extending TI to fish and experimentally demonstrating indirect TI learning in animals, these results indicate that a universal mechanism underlying TI is unlikely. Rather, animals probably use multiple domain-specific representations adapted to different social and ecological pressures that they encounter during the course of their natural lives.
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