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Broom, M. (2002). A unified model of dominance hierarchy formation and maintenance. J. Theor. Biol., 219(1), 63–72.
Abstract: In many different species it is common for animals to spend large portions of their lives in groups. Such groups need to divide available resources amongst the individuals they contain and this is often achieved by means of a dominance hierarchy. Sometimes hierarchies are stable over a long period of time and new individuals slot into pre-determined positions, but there are many situations where this is not so and a hierarchy is formed out of a group of individuals meeting for the first time. There are several different models both of the formation of such dominance hierarchies and of already existing hierarchies. These models often treat the two phases as entirely separate, whereas in reality, if there is a genuine formation phase to the hierarchy, behaviour in this phase will be governed by the rewards available, which in turn depends upon how the hierarchy operates once it has been formed. This paper describes a method of unifying models of these two distinct phases, assuming that the hierarchy formed is stable. In particular a framework is introduced which allows a variety of different models of each of the two parts to be used in conjunction with each other, thus enabling a wide range of situations to be modelled. Some examples are given to show how this works in practice.
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Nicol, C. J. (2006). How animals learn from each other. Appl. Anim. Behav. Sci., 100(1-2), 58–63.
Abstract: This paper explores ways by which animals may learn from one another, using examples drawn mostly from the chicken, an animal for which social learning is likely to be less dangerous than individual learning. In early life, the behaviour of the hen is important in encouraging chicks to peck at edible items. Maternal display not only attracts chicks to profitable food items, but also redirects their attention away from harmful or non-profitable items. Older chicks can enhance their foraging success by observing the behaviour of conspecifics within their own social group. Hens have been trained to perform a novel behaviour (key-pecking for food) by observation of a trained demonstrator bird. Moreover, observers learnt most from watching dominant demonstrators. Thus the ability to learn from others is not `fixed', but depends on the context and the social identity of both the observer and the demonstrator.
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Rands, S. A., Cowlishaw, G., Pettifor, R. A., Rowcliffe, J. M., & Johnstone, R. A. (2008). The emergence of leaders and followers in foraging pairs when the qualities of individuals differ. BMC Evol Biol, 8, 51.
Abstract: BACKGROUND: Foraging in groups offers animals a number of advantages, such as increasing their likelihood of finding food or detecting and avoiding predators. In order for a group to remain together, there has to be some degree of coordination of behaviour and movement between its members (which may in some cases be initiated by a decision-making leader, and in other cases may emerge as an underlying property of the group). For example, behavioural synchronisation is a phenomenon where animals within a group initiate and then continue to conduct identical behaviours, and has been characterised for a wide range of species. We examine how a pair of animals should behave using a state-dependent approach, and ask what conditions are likely to lead to behavioural synchronisation occurring, and whether one of the individuals is more likely to act as a leader. RESULTS: The model we describe considers how the energetic gain, metabolic requirements and predation risks faced by the individuals affect measures of their energetic state and behaviour (such as the degree of behavioural synchronisation seen within the pair, and the value to an individual of knowing the energetic state of its colleague). We explore how predictable changes in these measures are in response to changes in physiological requirements and predation risk. We also consider how these measures should change when the members of the pair are not identical in their metabolic requirements or their susceptibility to predation. We find that many of the changes seen in these measures are complex, especially when asymmetries exist between the members of the pair. CONCLUSION: Analyses are presented that demonstrate that, although these general patterns are robust, care needs to be taken when considering the effects of individual differences, as the relationship between individual differences and the resulting qualitative changes in behaviour may be complex. We discuss how these results are related to experimental observations, and how the model and its predictions could be extended.
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Saayman, G. S. (1971). Behaviour of the adult males in a troop of free-ranging Chacma baboons (Papio ursinus). Folia Primatol (Basel), 15(1), 36–57.
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Heitor, F., & Vicente, L. (2010). Dominance relationships and patterns of aggression in a bachelor group of Sorraia horses (Equus caballus). J. Ethol., 28(1), 35–44.
Abstract: Abstract The influence of individual factors on dominance rank and the relationship between rank distance and patterns of aggression predicted by models of evolutionarily stable strategies (ESS) of animal conflict were investigated in a managed bachelor group of Sorraia horses, Equus caballus. The group was composed of four to six stallions 3- to 12-years-old during the study period. The dominance hierarchy was significantly linear and rank was not related to age, weight, height or aggressiveness. Frequency and intensity of agonistic interactions were low, but higher-ranking stallions did not receive lower aggressiveness than lower-ranking stallions. There was some evidence that dominance relationships were more contested among close-ranking stallions, as predicted. Agonistic-related interactions among close-ranking stallions served similar functions to those among distant-ranking stallions, but the latter interacted more frequently than expected for access to resting sites and/or resting partners. Therefore, we found some evidence that agonistic-related interactions among distant-ranking stallions play a larger role in providing access to valuable and defendable resources than those among close-ranking stallions. Nevertheless, the fact that space to escape from aggression was limited and breeding access was independent from dominance rank may have reduced the benefits relative to costs of aggression and therefore limited the occurrence of contests over dominance and resources.
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Schooening, B. (1998). Ethology of the horse. Prakt. Tierarzt, 79(6 Suppl.), 25–28.
Abstract: The paper starts with a short introduction/definition about ethology and the used methods in this scientific field, giving special examples for horses and about how their “normal behaviour” is measured. The behaviour repertoire of horses is described in a brief outline with special emphasis on their social systems and hierarchies and the problem of dominance, especially in interaction with humans. Schlütersche GmbH & Co. KG, Verlag und Druckerei.
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Fujita, K., Kuroshima, H., & Masuda, T. (2002). Do tufted capuchin monkeys (Cebus apella) spontaneously deceive opponents? A preliminary analysis of an experimental food-competition contest between monkeys. Anim. Cogn., 5(1), 19–25.
Abstract: A new laboratory procedure which allows the study of deceptive behavior in nonhuman primates is described. Pairs of tufted capuchin monkeys faced each other in a food-competition contest. Two feeder boxes were placed between the monkeys. A piece of food was placed in one of the boxes. The subordinate individual was able to see the food and to open the box to obtain the bait. A dominant male was unable to see the food or to open the box but was able to take the food once the box was opened by the subordinate. In experiment 1, two of four subordinate monkeys spontaneously started to open the unbaited box first with increasing frequency. Experiment 2 confirmed that this “deceptive” act was not due to a drop in the rate of reinforcement caused by the usurping dominant male, under the situation in which food sometimes automatically dropped from the opened box. In experiment 3, two subordinate monkeys were rerun in the same situation as experiment 1. One of them showed some recovery of the “deceptive” act but the other did not; instead the latter tended to position himself on the side where there was no food before he started to open the box. Although the results do not clearly indicate spontaneous deception, we suggest that operationally defined spontaneous deceptive behaviors in monkeys can be analyzed with experimental procedures such as those used here.
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Houpt, K. A., & Wolski, T. R. (1980). Stability of equine hierarchies and the prevention of dominance related aggression. Equine Vet J, 12(1), 15–18.
Abstract: The dominance hierarchy of a herd of 10 Thoroughbred mares was determined twice, at an interval of 18 months, using paired feeding tests. Each mare's rank was correlated significantly between the 2 tests. This indicated that the hierarchy within the herd was stable. The offspring of dominant and subordinate mares were also tested for dominance in their own age groups. The offspring of dominant mares tended to be near the top of the hierarchy while those of middle and low ranking mares were not consistently found in the middle or bottom of their own hierarchies. Paired feeding tests were carried out on 8 ponies. During tests the time that each pony spent eating and the ponies' aggressive interactions were recorded. Two situations were used. Each pony-pair was tested when both ponies were in the same paddock and also when they were separated by a rail fence. The subordinate ponies spent significantly more time eating and the domonant pony was significantly less aggressive, when the pony-pair was separated by a fence than when they were in one paddock. It was concluded that the dominance hierarchies of adult horse groups changed very little over time and that the foals of dominant mares will tend to be dominant in their own age groups. Management practices can be used to reduce aggression and consequent injury that may arise in group feeding situations.
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Feist, J. D., & McCullough, D. R. (1975). Reproduction in feral horses. J Reprod Fertil Suppl, (23), 13–18.
Abstract: A behavioural study of feral horses was conducted on the Pryor Mountain Wild Horse Range in the western United States. All 270 horses on the Range were identified individually. The sex ratio was nearly balanced. Foal to adult female ratio was 43-2:100. Morality was concentrated among foals and old horses. Horses were organized as forty-four harem groups each with a dominant stallion, one to two immature stallions, one to three immature mares, one to three adult mares and their yearling and foal offspring, and 23 bachelor groups of one to eight stallions. Harem groups were quite stable year-round because of dominance and leadership by the stallions and group fidelity by mares and their offsring. Most changes occurred during the breeding season and involved immature females. Defeat of dominant stallions was infrequent. Immature males were tolerated because of their submissive behaviour. Bachelor stallion groups were inherently unstable. Mares came into heat after foaling in May/June, and were mated by harem stallions only.
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Hirsch, B. T. (2007). Costs and benefits of within-group spatial position: a feeding competition model. Q Rev Biol, 82(1), 9–27.
Abstract: An animal's within-group spatial position has several important fitness consequences. Risk of predation, time spent engaging in antipredatory behavior and feeding competition can all vary with respect to spatial position. Previous research has found evidence that feeding rates are higher at the group edge in many species, but these studies have not represented the entire breadth of dietary diversity and ecological situations faced by many animals. In particular the presence of concentrated, defendable food patches can lead to increased feeding rates by dominants in the center of the group that are able to monopolize or defend these areas. To fully understand the tradeoffs of within-group spatial position in relation to a variety of factors, it is important to be able to predict where individuals should preferably position themselves in relation to feeding rates and food competition. A qualitative model is presented here to predict how food depletion time, abundance of food patches within a group, and the presence of prior knowledge of feeding sites affect the payoffs of different within-group spatial positions for dominant and subordinate animals. In general, when feeding on small abundant food items, individuals at the front edge of the group should have higher foraging success. When feeding on slowly depleted, rare food items, dominants will often have the highest feeding rates in the center of the group. Between these two extreme points of a continuum, an individual's optimal spatial position is predicted to be influenced by an additional combination of factors, such as group size, group spread, satiation rates, and the presence of producer-scrounger tactics.
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