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      How Random Is Social Behaviour? Disentangling Social Complexity through the Study of a Wild House Mouse Population

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          Abstract

          Out of all the complex phenomena displayed in the behaviour of animal groups, many are thought to be emergent properties of rather simple decisions at the individual level. Some of these phenomena may also be explained by random processes only. Here we investigate to what extent the interaction dynamics of a population of wild house mice ( Mus domesticus) in their natural environment can be explained by a simple stochastic model. We first introduce the notion of perceptual landscape, a novel tool used here to describe the utilisation of space by the mouse colony based on the sampling of individuals in discrete locations. We then implement the behavioural assumptions of the perceptual landscape in a multi-agent simulation to verify their accuracy in the reproduction of observed social patterns. We find that many high-level features – with the exception of territoriality – of our behavioural dataset can be accounted for at the population level through the use of this simplified representation. Our findings underline the potential importance of random factors in the apparent complexity of the mice's social structure. These results resonate in the general context of adaptive behaviour versus elementary environmental interactions.

          Author Summary

          From the synchronised beauty of fish schools to the rigorous hierarchy of ant colonies, animals often display awe-inspiring collective behaviour. In recent years, principles of statistical physics have helped to unveil some simple mechanisms behind the emergence of such collective dynamics. Among the most elementary tools used to explain group behaviour are random processes, a typical example being the so-called “random walk”. In this paper, we have developed a framework based on such random assumptions to study the spatial and social structure of a population of wild house mice. We introduce the concept of perceptual landscape to describe the spatial behaviour of animals, whilst including all sensory and social constraints they are subject to: the perceptual landscape effectively maps the environment of animals as they perceive it. By applying our assumptions to a multi-agent model, we are able to reveal that much of the high-level social behaviour observed in the mouse population can indeed be explained through the many interactions of randomly moving individuals. This raises the question of how much of what we often regard as complex natural phenomena may, in fact, be the result of exceedingly simple forces.

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          Analyzing animal movements using Brownian bridges.

          By studying animal movements, researchers can gain insight into many of the ecological characteristics and processes important for understanding population-level dynamics. We developed a Brownian bridge movement model (BBMM) for estimating the expected movement path of an animal, using discrete location data obtained at relatively short time intervals. The BBMM is based on the properties of a conditional random walk between successive pairs of locations, dependent on the time between locations, the distance between locations, and the Brownian motion variance that is related to the animal's mobility. We describe two critical developments that enable widespread use of the BBMM, including a derivation of the model when location data are measured with error and a maximum likelihood approach for estimating the Brownian motion variance. After the BBMM is fitted to location data, an estimate of the animal's probability of occurrence can be generated for an area during the time of observation. To illustrate potential applications, we provide three examples: estimating animal home ranges, estimating animal migration routes, and evaluating the influence of fine-scale resource selection on animal movement patterns.
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            Predator-specific landscapes of fear and resource distribution: effects on spatial range use.

            Although ecologists have long recognized that animal space use is primarily determined by the presence of predators and the distribution of resources, the effects of these two environmental conditions have never been quantified simultaneously in a single spatial model. Here, in a novel approach, predator-specific landscapes of fear are constructed on the basis of behavioral responses of a prey species (vervet monkey; Cercopithecus aethiops), and we show how these can be combined with data on resource distribution to account for the observed variation in intensity of space use. Results from a mixed regressive-spatial regressive analysis demonstrate that ranging behavior can indeed be largely interpreted as an adaptive response to perceived risk of predation by some (but not all) predators and the spatial availability of resources. The theoretical framework behind the model is furthermore such that it can easily be extended to incorporate the effects of additional factors potentially shaping animal range use and thus may be of great value to the study of animal spatial ecology.
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              Global patterns of speciation and diversity.

              In recent years, strikingly consistent patterns of biodiversity have been identified over space, time, organism type and geographical region. A neutral theory (assuming no environmental selection or organismal interactions) has been shown to predict many patterns of ecological biodiversity. This theory is based on a mechanism by which new species arise similarly to point mutations in a population without sexual reproduction. Here we report the simulation of populations with sexual reproduction, mutation and dispersal. We found simulated time dependence of speciation rates, species-area relationships and species abundance distributions consistent with the behaviours found in nature. From our results, we predict steady speciation rates, more species in one-dimensional environments than two-dimensional environments, three scaling regimes of species-area relationships and lognormal distributions of species abundance with an excess of rare species and a tail that may be approximated by Fisher's logarithmic series. These are consistent with dependences reported for, among others, global birds and flowering plants, marine invertebrate fossils, ray-finned fishes, British birds and moths, North American songbirds, mammal fossils from Kansas and Panamanian shrubs. Quantitative comparisons of specific cases are remarkably successful. Our biodiversity results provide additional evidence that species diversity arises without specific physical barriers. This is similar to heavy traffic flows, where traffic jams can form even without accidents or barriers.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS Comput Biol
                PLoS Comput. Biol
                plos
                ploscomp
                PLoS Computational Biology
                Public Library of Science (San Francisco, USA )
                1553-734X
                1553-7358
                November 2012
                November 2012
                29 November 2012
                : 8
                : 11
                : e1002786
                Affiliations
                [1 ]Chair of Systems Design, ETH Zurich, Zurich, Switzerland
                [2 ]Department of Animal Behaviour, Institute of Evolutionary Biology and Environmental Studies, University of Zurich, Zurich, Switzerland
                Pennsylvania State University, United States of America
                Author notes

                The authors have declared that no competing interests exist.

                Analyzed the data: NP. Contributed reagents/materials/analysis tools: NP CJT. Wrote the paper: NP CJT BK FS.

                Article
                PCOMPBIOL-D-12-00456
                10.1371/journal.pcbi.1002786
                3510074
                23209394
                db7114c7-7d56-4b46-ac97-cb9bc89a1a2c
                Copyright @ 2012

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 20 March 2012
                : 3 October 2012
                Page count
                Pages: 11
                Funding
                We acknowledge financial support from the University of Zurich, the former Institute of Zoology, the Institute of Evolutionary Biology and Environmental Studies, and the Swiss National Science Foundation (project 3100A0-120444/1 to Anna Lindholm and Barbara König). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology
                Ecology
                Behavioral Ecology
                Evolutionary Biology
                Animal Behavior
                Mathematics
                Probability Theory
                Stochastic Processes
                Physics
                Statistical Mechanics

                Quantitative & Systems biology
                Quantitative & Systems biology

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