Dugatkin, L. A. (2001). Bystander effects and the structure of dominance hierarchies. Behav. Ecol., 12(3), 348–352.
Abstract: Prior modeling work has found that pure winner and loser effects (i.e., changing the estimation of your own fighting ability as a function of direct prior experience) can have important consequences for hierarchy formation. Here these models are extended to incorporate “bystander effects.” When bystander effects are in operation, observers (i.e., bystanders) of aggressive interactions change their assessment of the protagonists' fighting abilities (depending on who wins and who loses). Computer simulations demonstrate that when bystander winner effects alone are at play, groups have a clear omega (bottom-ranking individual), while the relative position of other group members remains difficult to determine. When only bystander loser effects are in operation, wins and losses are randomly distributed throughout a group (i.e., no discernible hierarchy). When pure and bystander winner effects are jointly in place, a linear hierarchy, in which all positions (i.e., {alpha} to {delta} when N = 4) are clearly defined, emerges. Joint pure and bystander loser effects produce the same result. In principle one could test the predictions from the models developed here in a straightforward comparative study. Hopefully, the results of this model will spur on such studies in the future.
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Mesterton-Gibbons, M., & Dugatkin, L. A. (1995). Toward a theory of dominance hierarchies: effects of assessment, group size, and variation in fighting ability. Behav. Ecol., 6(4), 416–423.
Abstract: We introduce assessment to the analysis of dominance hierarchies by exploring the effect of an evolutionarily stable fighting rule when there is variation in resource holding potential (RHP) and RHP is not a perfectly reliable predictor of the outcome of a fight. With assessment, the probability of a linear hierarchy decreases with group size but can remain appreciable for groups of up to seven or eight individuals, whereas it decreases virtually to zero if there is no assessment. The probability of a hierarchy that correlates perfectly with RHP is low unless group size is small.
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Cameron, E. Z.,, Linklater, W. L.,, Stafford, K. J.,, & Minot, E. O.,. (2003). Social grouping and maternal behaviour in feral horses (Equus caballus): the influence of males on maternal protectiveness. Behav. Ecol. Sociobiol., 53(2), 92–101.
Abstract: The risk of infant injury or mortality influences maternal behaviour, particularly protectiveness. Mares are found in bands with a single stallion or bands with more than one stallion in which paternity is less certain. We investigated maternal behaviour in relation to band type. Mares in bands with more than one stallion were more protective of their foals, particularly when stallions and foals approached one another. The rate of aggression between the stallion and foal was a significant predictor of maternal protectiveness, and mare protectiveness was significantly correlated with reduced reproductive success in the subsequent year. Mares that changed band types with a foal at foot, or had their band type experimentally altered, were more protective of their foal in multi-stallion bands than they were in single-stallion bands. Equids are unusual amongst ungulates in that infanticide and feticide have been reported. Both occur where paternity has been uncertain, and equid social structure is similar to other species in which infanticide has been reported. Stallions benefit from infanticide as the mare has greater reproductive success in the subsequent year. Stallion aggression is a significant modifier of mare behaviour and maternal effort, probably due to the risk of infanticide.
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Naguib, M., Amrhein, V., & Kunc, H. P. (2004). Effects of territorial intrusions on eavesdropping neighbors: communication networks in nightingales. Behav. Ecol., 15(6), 1011–1015.
Abstract: Animal communication often occurs in communication networks in which multiple signalers and receivers are within signaling range of each other. In such networks, individuals can obtain information on the quality and motivation of territorial neighbors by eavesdropping on their signaling interactions. In songbirds, extracting information from interactions involving neighbors is thought to be an important factor in the evolution of strategies of territory defense. In a playback experiment with radio-tagged nightingales Luscinia megarhynchos we here demonstrate that territorial males use their familiar neighbors' performance in a vocal interaction with an unfamiliar intruder as a standard for their own response. Males were attracted by a vocal interaction between their neighbor and a simulated stranger and intruded into the neighbor's territory. The more intensely the neighbor had interacted with playback, the earlier the intrusions were made, indicating that males eavesdropped on the vocal contest involving a neighbor. However, males never intruded when we had simulated by a second playback that the intruder had retreated and sang outside the neighbor's territory. These results suggest that territorial males use their neighbors' singing behavior as an early warning system when territorial integrity is threatened. Simultaneous responses by neighboring males towards unfamiliar rivals are likely to be beneficial to the individuals in maintaining territorial integrity.
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Dugatkin, L. A., & Earley, R. L. (2003). Group fusion: the impact of winner, loser, and bystander effects on hierarchy formation in large groups. Behav. Ecol., 14(3), 367–373.
Abstract: We present the results of a series of computer simulations that examined the impact of winner, loser, and bystander effects on hierarchy formation in fused groups. These effects and their implications for hierarchy structure and aggressive interactions were first examined in small four-member groups. Subsequent to this, the two small groups were fused into a single larger group. Further interactions took place in this fused group, generating a new hierarchy. Our models demonstrate clearly that winner, loser, and bystander effects strongly influence both the structure and types of interactions that emerge from the fusion of smaller groups. Four conditions produced results in which the same general patterns were uncovered in pre- and postfusion groups: (1) winner effects alone, (2) bystander loser effects alone, (3) winner and bystander winner effects operating simultaneously, and (4) all four effects in play simultaneously. Outside this parameter space, hierarchy structure and the nature of aggressive interactions differed in pre- and postfusion groups. When only loser effects were in play, one of the two clear alphas from the prefused groups dropped in rank in the eight-member fused group. When bystander winner effects were in play, it was difficult to rank any of the eight individuals in the fused group, and players interacted almost exclusively with those that were not in their original four-member group. When loser and bystander loser effects operated simultaneously, two top-ranking individuals emerged in the fused groups, but the relative rank of the other players was difficult to assign.
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FitzGibbon, C. D. (1994). The costs and benefits of predator inspection behaviour in Thomson's gazelles. Behav. Ecol. Sociobiol., 34(2), 139–148.
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Dugatkin, L. A. (1992). Tendency to inspect predators predicts mortality risk in the guppy (Poecilia reticulata). Behav. Ecol., 3(2), 124–127.
Abstract: Although predator inspection behavior in fishes has become a model system for examining game theoretical strategies such as Tit for Tat, the direct costs of inspection behavior have not been quantified. To begin quantifying such costs, I conducted an experiment that examined mortality due to predation as a function of predator inspection in the guppy (Poecilia reticulata). Before being subjected to a “survivorship” experiment, guppies were assayed for their tendency to inspect a predator. Groups were then composed of six guppies that differed in their tendency to inspect. These groups were placed into a pool containing a predator, and survivorship of guppies with different inspection tendencies was noted 36 and 60 h later. Results indicate that individuals that display high degrees of inspection behavior suffer greater mortality than their noninspecting shoalmates.
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Kavaliers, M., Colwell, D. D., & Choleris, E. (2005). Kinship, familiarity and social status modulate social learning about “micropredators” (biting flies) in deer mice. Behav. Ecol. Sociobiol., 58(1), 60–71.
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Berger, J. (1977). Organizational systems and dominance in feral horses in the Grand Canyon. Behav. Ecol. Sociobiol., 2(2), 131–146.
Abstract: 1. Several aspects of the behavioral ecology of feral horses (Equus caballus) were studied in Grand Canyon, Arizona, USA. Most bands contained three to five horses that included one stallion and his harem. Males that did not obtain a harem remained solitary. Throughout the study bands remained stable in composition.
2. Home ranges for all bands decreased in size in successive warm months, probably due to increased ambient temperature and drought. This resulted in greater utilization of spring areas that led to increased interband confrontation and agonistic display.
3. Territoriality was not observed in individual horses or bands, but bands hierarchial in both inter- and intraband structures. Interband stallion dominance was reinforced through posturing and fighting. Intraband hierarchies, as determined by dominance coefficients, were independent of individual size in three of four bands.
4. Indexes of nervousness (NER), calculated while horses were drinking, showed that stallions were less nervous than mares. A low NER was correlated with individuals leading toward drinking areas, whereas a high NER existed in individuals initiating flight although no single horse acted consistently as a leader.
5. Diurnal activity patterns were correlated with ambient temperatures.
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Dugatkin, L. A. (1997). Winner and loser effects and the structure of dominance hierarchies. Behav. Ecol., 8(6), 583–587.
Abstract: In the literature on dominance hierarchies, “winner” and “loser” effects usually are denned as an increased probability of winning at time T, bated on victories at time T-l, T-2, etc, and an increased probability of losing at time T, based on losing at T-1, T-2, etc., respectively. Despite some early theoretical work on winner and loser effects, these factors and how they affect the structure of dominance hierarchies have not been examined in detail. I developed a computer simulation to examine winner and loser effects when such effects are independent of one another (as well as when they interact) and when combatants assess each other's resource-holding power. When winner effects alone were important, a hierarchy in which all individuals held an unambiguous rank was found. When only loser effects were important, a dear alpha individual always emerged, but the rank of others in the group was often unclear because of the scarcity of aggressive interactions. Increasing winner effects for a given value of the loser effect increase the number of individuals with unambiguous positions in a hierarchy and the converse is true for increasing the value of the loser effect for a given winner effect Although winner and loser effects have been documented in a number of species, no study has documented both winner and loser effects (using some controlled, pairwise testing system) and the detailed nature of behavioral interactions when individuals are in groups. I hope the results of this model will spur such studies in the future.
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