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Hemelrijk, C. K. (2000). Towards the integration of social dominance and spatial structure. Anim. Behav., 59(5), 1035–1048.
Abstract: My aim was to show how individual-oriented (or artificial life) models may provide an integrative background for the development of theories about dominance by including effects of spatial structure. Dominance interactions are thought to serve two different, contrasting functions: acquisition of high rank and reduction of aggression. The model I present consists of a homogeneous virtual world inhabited by artificial agents whose actions are restricted to grouping and dominance interactions in which the effects of winning and losing are self-reinforcing. The two functions are implemented as strategies to initiate dominance interactions and the intensity of aggression and dominance perception (direct or memory based) are varied experimentally. Behaviour is studied by recording the same behavioural units as in real animals. Ranks appear to differentiate more clearly at high than at low intensity of aggression and also more in the case of direct than of memory-based rank perception. Strong differentiation of rank produces a cascade of unexpected effects that differ depending on which function is implemented: for instance, a decline in aggression, spatial centrality of dominants and a correlation between rank and aggression. Insight into the origination of these self-organized patterns leads to new hypotheses for the study of the social behaviour of real animals.
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Hemelrijk, C. K. (2002). Despotic societies, sexual attraction and the emergence of male 'tolerance': an agent-based model. Behaviour, 139(6), 729–747.
Abstract: During the period when females are sexually attractive – but only then – males of certain species of primates, such as chimpanzees, allow females access to resources. Because males are usually dominant over females, such male 'tolerance' is explained as a special, reproductive strategy to gain access to females. In this paper a simpler hypothesis is proposed on the basis of an individual-based model (called DomWorld): male 'tolerance' towards females arises in 'despotic' artificial societies as a kind of 'respectful timidity', because sexual attraction automatically increases female dominance over males as a side-effect. The model consists in a homogeneous, virtual world with agents that group and perform dominance-interactions in which the effects of victory and defeat are self-reinforcing. The artificial sexes differ in that VirtualMales have a higher intensity of aggression, they start with a greater capacity to win conflicts than VirtualFemales and they are especially attracted to the opposite sex during certain periods, whereas VirtualFemales are not. I shall explain how the introduction into DomWorld of the attraction of VirtualMales by VirtualFemales leads to female dominance, why it does so only in despotic, but not in egalitarian societies, and how it leads to other phenomena that are relevant to the study of primate behaviour.
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Hemelrijk, C. K. (1990). Models of, and tests for, reciprocity, unidirectionality and other social interaction patterns at a group level. Anim. Behav., 39(6), 1013–1029.
Abstract: Research on reciprocity is impaired by confusing definitions and often wrongly used statistical tests. Here, two models of the mechanism on which reciprocity may be based are discussed and an initial step towards a new fremework for its analysis is presented. A distinction is made between reciprocity and interchange. In the case of reciprocity, for one kind of act the same kind is received in return. In interchange, however, two different kinds of acts are bartered. Three types of reciprocity/interchange in social actions among all pairs of group-members are distinguished ([`]qualitative', [`]relative' and [`]absolute') on the basis of the precision of the reciprocity/interchange. Permutation procedures for association between matrices (such as the Mantel Z and two other newly derived tests) are used as a statistical test for detecting reciprocity/interchange. A rough comparison of the power of the two new tests is included. The tests can be applied to all kinds of group-living animals and to all sorts of social behaviour. The distinction between the three types of reciprocity/interchange and the matching statistical methods are also useful for defining and detecting other patterns in social interactions, like unidirectionality and associations between different kinds of social behaviour. The influence on social interactions of variables like dominance rank, age and sex can be analysed in the three forms by testing correlations between invented matrices which represent the influence of these variables (the so-called hypothesis matrices) and social interaction matrices. These methods are extended for two categories of individuals, thus allowing the investigation of, for example, reciprocity between males and females. The methods are illustrated with examples of coalition formation and grooming behaviour among captive chimpanzees, Pan troglodytes.
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Hemelrijk, C. K., Wantia, J.,, & Isler, K. (2008). Female Dominance over Males in Primates: Self-Organisation and Sexual Dimorphism. PLoS ONE, 3(7), e2678.
Abstract: The processes that underlie the formation of the dominance hierarchy in a group are since long under debate. Models of self-organisation suggest that dominance hierarchies develop by the self-reinforcing effects of winning and losing fights (the so-called winner-loser effect), but according to ‘the prior attribute hypothesis’, dominance hierarchies develop from pre-existing individual differences, such as in body mass. In the present paper, we investigate the relevance of each of these two theories for the degree of female dominance over males. We investigate this in a correlative study in which we compare female dominance between groups of 22 species throughout the primate order. In our study female dominance may range from 0 (no female dominance) to 1 (complete female dominance). As regards ‘the prior attribute hypothesis’, we expected a negative correlation between female dominance over males and species-specific sexual dimorphism in body mass. However, to our surprise we found none (we use the method of independent contrasts). Instead, we confirm the self-organisation hypothesis: our model based on the winner-loser effect predicts that female dominance over males increases with the percentage of males in the group. We confirm this pattern at several levels in empirical data (among groups of a single species and between species of the same genus and of different ones). Since the winner-loser effect has been shown to work in many taxa including humans, these results may have broad implications.3
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Hemelrijk, C. K. (2002). Understanding Social Behaviour with the Help of Complexity Science (Invited Article). Ethology, 108(8), 655–671.
Abstract: Abstract In the study of complexity, a new kind of explanation has been developed for social behaviour. It shows how patterns of social behaviour can arise as a side-effect of the interaction of individuals with their social or physical environment (e.g. by self-organization). This development may influence our ideas about the direct causation and evolution of social behaviour. Furthermore, it may influence our theories about the integration of different traits. This new method has been made possible by the increase in computing power. It is now applied in many areas of science, such as physics, chemistry, sociology and economics. However, in zoology and anthropology it is still rare. The major aim of this paper is to make this method more generally accepted among behavioural scientists.
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Hemelrijk, C. K. (2002). Self-Organization and Natural Selection in the Evolution of Complex Despotic Societies. Biol Bull, 202(3), 283–288.
Abstract: Differences between related species are usually explained as separate adaptations produced by individual selection. I discuss in this paper how related species, which differ in many respects, may evolve by a combination of individual selection, self-organization, and group-selection, requiring an evolutionary adaptation of only a single trait. In line with the supposed evolution of despotic species of macaques, we take as a starting point an ancestral species that is egalitarian and mildly aggressive. We suppose it to live in an environment with abundant food and we put the case that, if food becomes scarce and more clumped, natural selection at the level of the individual will favor individuals with a more intense aggression (implying, for instance, biting and fierce fighting). Using an individual-centered model, called DomWorld, I show what happens when the intensity of aggression increases. In DomWorld, group life is represented by artificial individuals that live in a homogeneous world. Individuals are extremely simple: all they do is flock together and, upon meeting one another, they may perform dominance interactions in which the effects of winning and losing are self-reinforcing. When the intensity of aggression in the model is increased, a complex feedback between the hierarchy and spatial structure results; via self-organization, this feedback causes the egalitarian society to change into a despotic one. The many differences between the two types of artificial society closely correspond to those between despotic and egalitarian macaques in the real world. Given that, in the model, the organization changes as a side effect of the change of one single trait proper to an egalitarian society, in the real world a despotic society may also have arisen as a side effect of the mutation of a single trait of an egalitarian species. If groups with different intensities of aggression evolve in this way, they will also have different gradients of hierarchy. When food is scarce, groups with the steepest hierarchy may have the best chance to survive, because at least a small number of individuals in such a group may succeed in producing offspring, whereas in egalitarian societies every individual is at risk of being insufficiently fed to reproduce. Therefore, intrademic group selection (selection within an interbreeding group) may have contributed to the evolution of despotic societies. N1 -
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Hemelrijk, C. K., & Hildenbrandt, H. (2008). Self-Organized Shape and Frontal Density of Fish Schools. Ethology, 114(3), 245–254.
Abstract: Abstract Models of swarming (based on avoidance, alignment and attraction) produce patterns of behaviour also seen in schools of fish. However, the significance of such similarities has been questioned, because some model assumptions are unrealistic [e.g. speed in most models is constant with random error, the perception is global and the size of the schools that have been studied is small (up to 128 individuals)]. This criticism also applies to our former model, in which we demonstrated the emergence of two patterns of spatial organization, i.e. oblong school form and high frontal density, which are supposed to function as protection against predators. In our new model we respond to this criticism by making the following improvements: individuals have a preferred ‘cruise speed’ from which they can deviate in order to avoid others or to catch up with them. Their range of perception is inversely related to density, with which we take into account that high density limits the perception of others that are further away. Swarm sizes range from 10 to 2000 individuals. The model is three-dimensional. Further, we show that the two spatial patterns (oblong shape and high frontal density) emerge by self-organization as a side-effect of coordination at two speeds (of two or four body lengths per second) for schools of sizes above 20. Our analysis of the model leads to the development of a new set of hypotheses. If empirical data confirm these hypotheses, then in a school of real fish these patterns may arise as a side-effect of their coordination in the same way as in the model.
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Puga-Gonzalez, I., Hildenbrandt, H., & Hemelrijk, C. K. (2009). Emergent Patterns of Social Affiliation in Primates, a Model. PLoS Comput Biol, 5(12), e1000630.
Abstract: Author Summary
<p>Individual primates distribute their affiliative behaviour (such as grooming) in complex patterns among their group members. For instance, they reciprocate grooming, direct it more to partners the higher the partner's rank, use it to reconcile fights and do so in particular with partners that are more valuable. For several types of patterns (such as reconciliation and exchange), a separate theory based on specific cognitive processes has been developed (such as individual recordkeeping, a tendency to exchange, selective attraction to the former opponent, and estimation of the value of a relationship). It is difficult to imagine how these separate theories can all be integrated scientifically and how these processes can be combined in the animal's mind. To solve this problem, we first surveyed the empirical patterns and then we developed an individual-based model (called GrooFiWorld) in which individuals group, compete and groom. The grooming rule is based on grooming out of fear of defeat and on the anxiety reducing effects of grooming. We show that in this context this rule alone can explain many of the patterns of affiliation as well as the differences between egalitarian and despotic species. Our model can be used as a null model to increase our understanding of affiliative patterns of primates.</p> |
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Hildenbrandt, H., Carere, C., & Hemelrijk, C. K. (2010). Self-organized aerial displays of thousands of starlings: a model. Behav. Ecol., 21(6), 1349–1359.
Abstract: Through combining theoretical models and empirical data, complexity science has increased our understanding of social behavior of animals, in particular of social insects, primates, and fish. What are missing are studies of collective behavior of huge swarms of birds. Recently detailed empirical data have been collected of the swarming maneuvers of large flocks of thousands of starlings (Sturnus vulgaris) at their communal sleeping site (roost). Their flocking maneuvers are of dazzling complexity in their changes in density and flock shape, but the processes underlying them are still a mystery. Recent models show that flocking may arise by self-organization from rules of co-ordination with nearby neighbors, but patterns in these models come nowhere near the complexity of those of the real starlings. The question of this paper, therefore, is whether such complex patterns can emerge by self-organization. In our computer model, called StarDisplay, we combine the usual rules of co-ordination based on separation, attraction, and alignment with specifics of starling behavior: 1) simplified aerodynamics of flight, especially rolling during turning, 2) movement above a “roosting area” (sleeping site), and 3) the low fixed number of interaction neighbors (i.e., the topological range). Our model generates patterns that resemble remarkably not only qualitative but also quantitative empirical data collected in Rome through video recordings and position measurements by stereo photography. Our results provide new insights into the mechanisms underlying complex flocking maneuvers of starlings and other birds.
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Krueger, K., Flauger, B., Farmer, K., & Hemelrijk, C. (2014). Movement initiation in groups of feral horses. Behav. Process., 103, 91–101.
Abstract: Abstract Herds of ungulates, flocks of birds, swarms of insects and schools of fish move in coordinated groups. Computer models show that only one or very few animals are needed to initiate and direct movement. To investigate initiation mechanisms further, we studied two ways in which movement can be initiated in feral horses: herding, and departure from the group. We examined traits affecting the likelihood of a horse initiating movement i.e. social rank, affiliative relationships, spatial position, and social network. We also investigated whether group members join a movement in dominance rank order. Our results show that whereas herding is exclusive to alpha males, any group member may initiate movement by departure. Social bonds, the number of animals interacted with, and the spatial position were not significantly associated with movement initiation. We did not find movement initiation by departure to be exclusive to any type of individual. Instead we find evidence for a limited form of distributed leadership, with higher ranking animals being followed more often.
Keywords: Horse; Equus ferus caballus; Distributed leadership; Herding; Departure; Rank
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