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	Author	Broom, M.
	Title	A unified model of dominance hierarchy formation and maintenance			Type	Journal Article
	Year	2002	Publication	Journal of theoretical biology	Abbreviated Journal	J. Theor. Biol.
	Volume	219	Issue	1	Pages	63-72
	Keywords	Animals; Behavior, Animal; Feeding Behavior; Models, Psychological; *Social Dominance; Social Environment
	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.
	Address	Centre for Statistics and Stochastic Modelling, School of Mathematical Sciences, University of Sussex, Falmer, Brighton, BN1 9QH, U.K. m.broom@sussex.ac.uk
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0022-5193	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:12392975			Approved	no
	Call Number	refbase @ user @			Serial	439
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	Author	Viscido, S.V.; Miller, M.; Wethey, D.S.
	Title	The dilemma of the selfish herd: the search for a realistic movement rule			Type	Journal Article
	Year	2002	Publication	Journal of theoretical biology	Abbreviated Journal	J. Theor. Biol.
	Volume	217	Issue	2	Pages	183-194
	Keywords	Animals; Behavior, Animal; Mass Behavior; Models, Biological; *Motor Activity; Predatory Behavior
	Abstract	The selfish herd hypothesis predicts that aggregations form because individuals move toward one another to minimize their own predation risk. The “dilemma of the selfish herd” is that movement rules that are easy for individuals to follow, fail to produce true aggregations, while rules that produce aggregations require individual behavior so complex that one may doubt most animals can follow them. If natural selection at the individual level is responsible for herding behavior, a solution to the dilemma must exist. Using computer simulations, we examined four different movement rules. Relative predation risk was different for all four movement rules (p<0.05). We defined three criteria for measuring the quality of a movement rule. A good movement rule should (a) be statistically likely to benefit an individual that follows it, (b) be something we can imagine most animals are capable of following, and (c) result in a centrally compact flock. The local crowded horizon rule, which allowed individuals to take the positions of many flock-mates into account, but decreased the influence of flock-mates with distance, best satisfied these criteria. The local crowded horizon rule was very sensitive to the animal's perceptive ability. Therefore, the animal's ability to detect its neighbors is an important factor in the dynamics of group formation.
	Address	Department of Biological Sciences, University of South Carolina, Columbia, SC, 29208, USA. viscido@u.washington.edu
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0022-5193	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:12202112			Approved	no
	Call Number	refbase @ user @			Serial	554
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	Author	Viscido, S.V.; Miller, M.; Wethey, D.S.
	Title	The response of a selfish herd to an attack from outside the group perimeter			Type	Journal Article
	Year	2001	Publication	Journal of theoretical biology	Abbreviated Journal	J. Theor. Biol.
	Volume	208	Issue	3	Pages	315-328
	Keywords	Animals; Behavior, Animal; Computer Simulation; Models, Biological; *Movement; Predatory Behavior
	Abstract	According to the selfish herd hypothesis, animals can decrease predation risk by moving toward one another if the predator can appear anywhere and will attack the nearest target. Previous studies have shown that aggregations can form using simple movement rules designed to decrease each animal's Domain of Danger. However, if the predator attacks from outside the group's perimeter, these simple movement rules might not lead to aggregation. To test whether simple selfish movement rules would decrease predation risk for those situations when the predator attacks from outside the flock perimeter, we constructed a computer model that allowed flocks of 75 simulated fiddler crabs to react to one another, and to a predator attacking from 7 m away. We attacked simulated crab flocks with predators of different sizes and attack speeds, and computed relative predation risk after 120 time steps. Final trajectories showed flight toward the center of the flock, but curving away from the predator. Path curvature depended on the predator's size and approach speed. The average crab experienced a greater decrease in predation risk when the predator was small or slow moving. Regardless of the predator's size and speed, however, predation risk always decreased as long as crabs took their flock-mates into account. We conclude that, even when flight away from an external predator occurs, the selfish avoidance of danger can lead to aggregation.
	Address	Department of Biological Sciences, University of South Carolina, Columbia, SC 29208, U.S.A. viscido@u.washington.edu
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0022-5193	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:11207093			Approved	no
	Call Number	refbase @ user @			Serial	555
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	Author	Suzuki, Y.; Toquenaga, Y.
	Title	Effects of information and group structure on evolution of altruism: analysis of two-score model by covariance and contextual analyses			Type	Journal Article
	Year	2005	Publication	Journal of theoretical biology	Abbreviated Journal	J. Theor. Biol.
	Volume	232	Issue	2	Pages	191-201
	Keywords	Altruism; Analysis of Variance; Communication; Cooperative Behavior; Evolution; Game Theory; Group Structure; Humans; Models, Genetic; Models, Psychological; Selection (Genetics); Trust
	Abstract	An altruistic individual has to gamble on cooperation to a stranger because it does not know whether the stranger is trustworthy before direct interaction. Nowak and Sigmund (Nature 393 (1998a) 573; J. Theor. Biol. 194 (1998b) 561) presented a new theoretical framework of indirect reciprocal altruism by image scoring game where all individuals are informed about a partner's behavior from its image score without direct interaction. Interestingly, in a simplified version of the image scoring game, the evolutionarily stability condition for altruism became a similar form of Hamilton's rule, i.e. inequality that the probability of getting correct information is more than the ratio of cost to benefit. Since the Hamilton's rule was derived by evolutionarily stable analysis, the evolutionary meaning of the probability of getting correct information has not been clearly examined in terms of kin and group selection. In this study, we applied covariance analysis to the two-score model for deriving the Hamilton's rule. We confirmed that the probability of getting correct information was proportional to the bias of altruistic interactions caused by using information about a partner's image score. The Hamilton's rule was dependent on the number of game bouts even though the information reduced the risk of cooperation to selfish one at the first encounter. In addition, we incorporated group structure to the two-score model to examine whether the probability of getting correct information affect selection for altruism by group selection. We calculated a Hamilton's rule of group selection by contextual analysis. Group selection is very effective when either the probability of getting correct information or that of future interaction, or both are low. The two Hamilton's rules derived by covariance and contextual analyses demonstrated the effects of information and group structure on the evolution of altruism. We inferred that information about a partner's behavior and group structure can produce flexible pathways for the evolution of altruism.
	Address	Integrative Environmental Sciences, Graduate School of Life and Environmental Sciences, University of Tsukuba, 1-1-1, Ten-Nou-Dai, Tsukuba, Ibaraki 305-8572, Japan. yukari@pe.ies.life.tsukuba.ac.jp
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0022-5193	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15530489			Approved	no
	Call Number	refbase @ user @			Serial	556
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	Author	Hamilton, W.D.
	Title	Geometry for the selfish herd			Type	Journal Article
	Year	1971	Publication	Journal of theoretical biology	Abbreviated Journal	J. Theor. Biol.
	Volume	31	Issue	2	Pages	295-311
	Keywords	Animals; Anura; Behavior, Animal; Breeding; Communication; Evolution; Fear; Metallurgy; Models, Biological; Probability; Snakes; *Spatial Behavior
	Abstract	This paper presents an antithesis to the view that gregarious behaviour is evolved through benefits to the population or species. Following Galton (1871) and Williams (1964) gregarious behaviour is considered as a form of cover-seeking in which each animal tries to reduce its chance of being caught by a predator. It is easy to see how pruning of marginal individuals can maintain centripetal instincts in already gregarious species; some evidence that marginal pruning actually occurs is summarized. Besides this, simply defined models are used to show that even in non-gregarious species selection is likely to favour individuals who stay close to others. Although not universal or unipotent, cover-seeking is a widespread and important element in animal aggregation, as the literature shows. Neglect of the idea has probably followed from a general disbelief that evolution can be dysgenic for a species. Nevertheless, selection theory provides no support for such disbelief in the case of species with outbreeding or unsubdivided populations. The model for two dimensions involves a complex problem in geometrical probability which has relevance also in metallurgy and communication science. Some empirical data on this, gathered from random number plots, is presented as of possible heuristic value.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0022-5193	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:5104951			Approved	no
	Call Number	refbase @ user @			Serial	771
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	Author	Hamilton, W.D.
	Title	The genetical evolution of social behaviour. I			Type	Journal Article
	Year	1964	Publication	Journal of Theoretical Biology	Abbreviated Journal	J. Theor. Biol.
	Volume	7	Issue	1and 2	Pages	1-52
	Keywords	Behavior; Genetics; Humans; *Models, Theoretical
	Abstract	A genetical mathematical model is described which allows for interactions between relatives on one another's fitness. Making use of Wright's Coefficient of Relationship as the measure of the proportion of replica genes in a relative, a quantity is found which incorporates the maximizing property of Darwinian fitness. This quantity is named “inclusive fitness”. Species following the model should tend to evolve behaviour such that each organism appears to be attempting to maximize its inclusive fitness. This implies a limited restraint on selfish competitive behaviour and possibility of limited self-sacrifices. Special cases of the model are used to show (a) that selection in the social situations newly covered tends to be slower than classical selection, (b) how in populations of rather non-dispersive organisms the model may apply to genes affecting dispersion, and (c) how it may apply approximately to competition between relatives, for example, within sibships. Some artificialities of the model are discussed.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0022-5193	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:5875341			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5160
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	Author	Parker, G.A.
	Title	Assessment strategy and the evolution of fighting behaviour			Type	Journal Article
	Year	1974	Publication	Journal of Theoretical Biology	Abbreviated Journal	J. Theor. Biol.
	Volume	47	Issue	1	Pages	223-243
	Keywords
	Abstract	The view is examined that the adaptive value of conventional aspects of fighting behaviour is for assessment of relative RHP (resource holding power) of the combatants. Outcomes of aggressive disputes should be decided by each individual's fitness budget available for expenditure during a fight (determined by the fitness difference between adoption of alternative strategies, escalation or withdrawal without escalation) and on the rate of expenditure of the fitness budget if escalation occurs (determined by the RHPs of the combatants). Thus response thresholds for alternative strategies (“assessments”) will be determined by natural selection on a basis of which opponent is likely to expend its fitness budget first, should escalation occur. This “loser” should retreat (before escalation) and the winner should stay in possession of the resource. Many aggressive decisions depend on whether one is a resource holder, or an attacker. Assuming the RHP of the combatants to be equal, there are many instances of fitness pay-off imbalances between holder and attacker which should weight the dispute outcome in favour of one or other opponent by allowing it a greater expendable fitness budget. Usually the weighting favours the holder; the attacker therefore needs a correspondingly higher RHP before it may be expected to win. This is not invariably the case, and much observed data fits the predictions of this sort of model. If assessments are perfect and budget expenditure rates exactly predictable, then there would never seem to be any case for escalation. Escalation can be explained in terms of injury inflictions (expenditures) occurring as discrete events; i.e. as “bouts” won or lost during fighting. Assessment can give only a probabilistic prediction of the outcome of a bout. A simple model is developed to investigate escalation situations. Each combatant assesses relative RHP; this correlates with an absolute probability of winning the next bout (cabs). The stake played for is infliction of loss of RHP and is determined by the fitness budgets of the opponents. (Each individual plays for the withdrawal of its opponent.) This defines a critical probability of winning (ccrit) for each combatant, above which escalation is the favourable strategy (cabs > ccrit) and below which withdrawal is favourable (cabs < ccrit). Escalation should occur only where cabs-ccrit is positive for both combatants. This model gives predictions compatible with the observations, indicating that RHP loss alone can be adequate to explain withdrawal: escalation behaviour. Withdrawal tendency will be increased by low searching costs. Escalations should be restricted to closely matched RHP opponents if RHP disparity is the major imbalance. Outside the “escalation range” of a given individual, the higher RHP individual wins and the lower one loses (i.e. it should withdraw after conventional display). RHP disparity and holder: attacker imbalance should both interact to shape the observed pattern, though their relative importances will depend on species and situation. In some instances selection may favour immediate withdrawal from an occupied territory even without assessment of RHP.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0022-5193	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	4935
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	Author	Gueron, S.; Levin, S.A.; Rubenstein, D.I.
	Title	The Dynamics of Herds: From Individuals to Aggregations			Type	Journal Article
	Year	1996	Publication	Journal of Theoretical Biology	Abbreviated Journal	J. Theor. Biol.
	Volume	182	Issue	1	Pages	85-98
	Keywords
	Abstract	The dynamic behavior of small herds is investigated by means of simulations of two-dimensional discrete-stochastic models. An individual-based approach is used to relate collective behavior to individual decisions. In our model, the motion of an individual in a herd is assumed to be the combined result of both density-independent and density-dependent decisions, in the latter case based on the influence of surrounding neighbors; assumed decision rules are hierarchical, balancing short range repulsion against long-range attraction. The probability of fragmentation of the model herd depends on parameter values. We explore the variety and characteristics of spatial patterns that develop during migration, for herds that are homogeneous and heterogeneous regarding intrinsic walking speeds. Group integrity can be maintained even in mixed populations, but fragmentation results for these more easily than for a homogeneous herd. Observations of natural populations suggest that animals move away from individuals that intrude too closely into their environment, but are attracted to individuals at a distance. Between these extremes, there appears to be a neutral zone, within which other individuals engender no response. We explore the importance of this neutral zone, and offer evolutionary interpretations. In particular, the neutral zone, if not too large, permits the individual to remain in contact with the herd, while reducing the frequency with which acceleration or deceleration must be undertaken. This offers obvious energetic benefits.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0022-5193	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5253
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	Author	Seyfarth, R.M.
	Title	A model of social grooming among adult female monkeys			Type	Journal Article
	Year	1977	Publication	Journal of Theoretical Biology	Abbreviated Journal	J. Theor. Biol.
	Volume	65	Issue	4	Pages	671-698
	Keywords	Animals; Behavior, Animal; Female; Grooming; Haplorhini/physiology; *Models, Biological; Reproduction; Social Dominance; Time Factors
	Abstract	Grooming networks among adult female monkeys exhibit two similar features across a number of different species. High-ranking animals receive more grooming than others, and the majority of grooming occurs between females of adjacent rank. A theoretical model which duplicates these features is presented, and the properties of the model are used to explain the possible causation and function of female grooming behaviour. The model illustrates how relatively simple principles governing the behaviour of individuals may be used to explain more complex aspects of the social structure of non-human primate groups.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0022-5193	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:406485			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5259
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	Author	Couzin, I.D.; Krause, J.; James, R.; Ruxton, G.D.; Franks, N.R.
	Title	Collective Memory and Spatial Sorting in Animal Groups			Type	Journal Article
	Year	2002	Publication	Journal of Theoretical Biology	Abbreviated Journal	J. Theor. Biol.
	Volume	218	Issue	1	Pages	1-11
	Keywords
	Abstract	We present a self-organizing model of group formation in three-dimensional space, and use it to investigate the spatial dynamics of animal groups such as fish schools and bird flocks. We reveal the existence of major group-level behavioural transitions related to minor changes in individual-level interactions. Further, we present the first evidence for collective memory in such animal groups (where the previous history of group structure influences the collective behaviour exhibited as individual interactions change) during the transition of a group from one type of collective behaviour to another. The model is then used to show how differences among individuals influence group structure, and how individuals employing simple, local rules of thumb, can accurately change their spatial position within a group (e.g. to move to the centre, the front, or the periphery) in the absence of information on their current position within the group as a whole. These results are considered in the context of the evolution and ecological importance of animal groups.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0022-5193	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5310
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