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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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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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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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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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Cant, M. A., & Field, J. (2005). Helping effort in a dominance hierarchy. Behav. Ecol., 16(4), 708–715.
Abstract: In many cooperatively breeding species, group members form a dominance hierarchy or queue to inherit the position of breeder. Models aimed at understanding individual variation in helping behavior, however, rarely take into account the effect of dominance rank on expected future reproductive success and thus the potential direct fitness costs of helping. Here we develop a kin-selection model of helping behavior in multimember groups in which only the highest ranking individual breeds. Each group member can invest in the dominant's offspring at a cost to its own survivorship. The model predicts that lower ranked subordinates, who have a smaller probability of inheriting the group, should work harder than higher ranked subordinates. This prediction holds regardless of whether the intrinsic mortality rate of subordinates increases or decreases with rank. The prediction does not necessarily hold, however, where the costs of helping are higher for lower ranked individuals: a situation that may be common in vertebrates. The model makes two further testable predictions: that the helping effort of an individual of given rank should be lower in larger groups, and the reproductive success of dominants should be greater where group members are more closely related. Empirical evidence for these predictions is discussed. We argue that the effects of rank on stable helping effort may explain why attempts to correlate individual helping effort with relatedness in cooperatively breeding species have met with limited success.
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Vahl, W. K., Lok, T., van der Meer, J., Piersma, T., & Weissing, F. J. (2005). Spatial clumping of food and social dominance affect interference competition among ruddy turnstones. Behav. Ecol., 16(5), 834–844.
Abstract: In studying the success of foraging animals, studies of interference competition have put emphasis on effects of competitor density, whereas studies of resource defense have focused on the effects of the spatial distribution of food within patches. Very few studies have looked at both factors simultaneously, that is, determined whether the effects of competitor density on foraging success depend on the spatial distribution of food. We studied the behavior and the foraging success of ruddy turnstones (Arenaria interpres) using an experiment in which we varied both the presence of a competitor and the food distribution. Because turnstones may differ strongly in their relative dominance status, we also experimentally varied the foragers' relative dominance status. We found that the presence of a competitor only reduced the foraging success of subordinate birds foraging at the clumped food distribution. At this condition, dominant and subordinate birds differed markedly in their foraging success. Contrary to our expectations, we did not observe more agonistic behavior at the clumped food distribution. This indicates that the amount of agonistic behavior observed may be a bad indicator of interference effects. These findings have specific implications for models of interference competition. Most notably they show that the effects of competitor density on agonistic behavior and foraging success may well depend on the spatial distribution of food and the foragers' relative dominance status. Additionally, our results suggest that social dominance will not be fully understood without considering long-term processes such as the formation and maintenance of social dominance hierarchies.
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Shrader, A. M., Kerley, G. I. H., Kotler, B. P., & Brown, J. S. (2007). Social information, social feding, and competition in group-living goats (Capra hircus). Behav. Ecol., 18(1), 103–107.
Abstract: There are both benefits (e.g., social information) and costs (e.g., intraspecific competition) for individuals foraging in groups. To ascertain how group-foraging goats (Capra hircus) deal with these trade-offs, we asked 1) do goats use social information to make foraging decisions and 2) how do they adjust their intake rate in light of having attracted by other group members? To establish whether goats use social information, we recorded their initial choice of different quality food patches when they were ignorant of patch quality and when they could observe others foraging. After determining that goats use social information, we recorded intake rates while they fed alone and in the presence of potential competitors. Intake rate increased as the number of competitors increased. Interestingly, lone goats achieved an intake rate that was higher than when one competitor was present but similar to when two or more competitors were present. Faster intake rates may allow herbivores to ingest a larger portion of the available food before competing group members arrive at the patch. This however, does not explain the high intake rates achieved when the goats were alone. We provide 2 potential explanations: 1) faster intake rates are a response to greater risk incurred by lone individuals, the loss of social information, and the fear of being left behind by the group and 2) when foraging alone, intake rate is no longer a trade-off between reducing competition and acquiring social information. Thus, individuals are able to feed close to their maximum rate.
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Anderson, C., & Franks, N. R. (2001). Teams in animal societies. Behav. Ecol., 12(5), 534–540.
Abstract: We review the existence of teams in animal societies. Teams have previously been dismissed in all but a tiny minority of insect societies. “Team” is a term not generally used in studies of vertebrates. We propose a new rigorous definition of a team that may be applied to both vertebrate and invertebrate societies. We reconsider what it means to work as a team or group and suggest that there are many more teams in insect societies than previously thought. A team task requires different subtasks to be performed concurrently for successful completion. There is a division of labor within a team. Contrary to previous reviews of teams in social insects, we do not constrain teams to consist of members of different castes and argue that team members may be interchangeable. Consequently, we suggest that a team is simply the set of individuals that performs a team task. We contrast teams with groups and suggest that a group task requires the simultaneous performance and cooperation of two or more individuals for successful completion. In a group, there is no division of labor--each individual performs the same task. We also contrast vertebrate and invertebrate teams and find that vertebrate teams tend to be associated with hunting and are based on individual recognition. Invertebrate teams occur in societies characterized by a great deal of redundancy, and we predict that teams in insect societies are more likely to be found in large polymorphic (“complex”) societies than in small monomorphic (“simple”) societies.
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