The dynamics of group formation in large mammalian herbivores: an analysis in the European roe deer

In large mammalian herbivores, herd size usually increases with openness of the environment, but the proximate mechanisms that underlie this phenomenon remain poorly known. We investigated the dynamics of group formation in a population of roe deer, Capreolus capreolus, living in open cultivated plain. Our results show the high spontaneous instability of groups. The probability of fission per unit time increased approximately as the square of group size, and the sizes of the groups resulting from a splitting-up followed a uniform distribution. Attraction between groups was the main cause of fusion and was at work over distances of more than 200 m, far exceeding the perception radius of roe deer in forest. In addition, when not yet separated by than more than 200 m, two groups resulting from a single splitting-up had a high probability of remerging and restoring the parent group. Our results are thus consistent with the assumption that in large herbivores group size increases with habitat openness, not because of a change in individual's behaviour, but because groups are unstable and any increase of the distance at which animals perceive one another enhances the rate at which groups merge.

[1]  R. Putman The natural history of deer , 1988 .

[2]  A. Frid Vigilance by female Dall's sheep: interactions between predation risk factors , 1997, Animal Behaviour.

[3]  L. Conradt,et al.  Could asynchrony in activity between the sexes cause intersexual social segregation in ruminants? , 1998, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[4]  Jean-François Gerard,et al.  Sexual segregation in fallow deer: are mixed-sex groups especially unstable because of asynchrony between the sexes? , 2006, Comptes rendus biologies.

[5]  Jane Uhd Jepsen,et al.  Modelling roe deer (Capreolus capreolus) in a gradient of forest fragmentation: behavioural plasticity and choice of cover , 2004 .

[6]  H. Pulliam,et al.  On the advantages of flocking. , 1973, Journal of theoretical biology.

[7]  J. Vincent,et al.  Evolution saisonnière de la taille des groupes chez le chevreuil en milieu forestier , 1983 .

[8]  S. Lovari,et al.  The biology and management of mountain ungulates , 1986 .

[9]  T. Caraco,et al.  Social Foraging Theory , 2018 .

[10]  C. Johnson Variations in Group Size and Composition in Red and Western Grey Kangaroos, Macropus rufus (Desmarest ) and M. fulignosus (Desmarest) , 1983 .

[11]  Robin I. M. Dunbar,et al.  Sexual segregation among feral goats: testing between alternative hypotheses , 2006, Animal Behaviour.

[12]  J. Gerard,et al.  Instability and diurnal variation in size of winter groupings of field roe deer , 1998, Revue d'Écologie (La Terre et La Vie).

[13]  E. Milner‐Gulland,et al.  The ecology and management of the Saiga antelope in Kazakhstan , 1998 .

[14]  J. Vincent,et al.  Flexibilité de l'organisation sociale du chevreuil en fonction des caractéristiques de l'environnement , 1987 .

[15]  C. Southwell Variability in Grouping in the Eastern Grey Kangaroo, Macropus giganteus I. Group Density and Group Size , 1984 .

[16]  D. Fraser,et al.  The behaviour of Ungulates and its relation to management , 1975 .

[17]  J. Gerard,et al.  Herd Size in Large Herbivores: Encoded in the Individual or Emergent? , 2002, The Biological Bulletin.

[18]  A. Schaal Influence de l’environnement sur les composantes du groupe social chez le Daim Cervus (Dama) dama L. , 1982, Revue d'Écologie (La Terre et La Vie).

[19]  J. Peek,et al.  Dynamics of Moose Aggregations in Alaska, Minnesota, and Montana , 1974 .

[20]  C. S. José,et al.  Grouping in roe deer: an effect of habitat openness or cover distribution? , 1997 .

[21]  J. Gerard,et al.  Group dynamics and local population density dependence of group size in the Pyrenean chamois, Rupicapra pyrenaica , 2008, Animal Behaviour.

[22]  SIMON BENHAMOU,et al.  Path integration in dogs , 1998, Animal Behaviour.

[23]  R. Estes,et al.  The Behavior Guide to African Mammals: Including Hoofed Mammals, Carnivores, Primates , 1991 .

[24]  M. Murray Structure of association in impala, Aepyceros melampus , 1981, Behavioral Ecology and Sociobiology.

[25]  J. Gerard,et al.  Large group formation in European Roe deer : an adaptive feature ? , 1995 .

[26]  P. Evans Habitat preferences of ungulates in closed savanna of central Africa , 1979 .

[27]  W. Hamilton Geometry for the selfish herd. , 1971, Journal of theoretical biology.

[28]  Pulliam Hr,et al.  On the advantages of flocking , 1973 .

[29]  S. Gueron,et al.  The dynamics of group formation. , 1995, Mathematical biosciences.

[30]  A. Pusey The behavior guide to African mammals: by Richard Despard Estes, University of California Press, 1991. $75.00 hbk (xxii + 611 pages) ISBN 0 520 05831 3 , 1992 .

[31]  D. Cox,et al.  Analysis of Survival Data. , 1986 .

[32]  E. Bonabeau,et al.  Possible universality in the size distribution of fish schools. , 1995, Physical review. E, Statistical physics, plasmas, fluids, and related interdisciplinary topics.

[33]  Do the sexes tend to segregate in roe deer in agricultural environments? An analysis of group composition , 2006 .

[34]  T. Clutton‐Brock,et al.  Red Deer: Behavior and Ecology of Two Sexes , 1992 .

[35]  Paul Martin,et al.  Measuring behaviour: Preface to second edition , 1993 .

[36]  C. Barrette The size of Axis deer fluid groups in Wilpattu national park, Sri Lanka , 1991 .

[37]  J. Berger Group size, foraging, and antipredator ploys: An analysis of bighorn sheep decisions , 1978, Behavioral Ecology and Sociobiology.

[38]  J. Gerard,et al.  Inter-individual associations and social structure of a mouflon population (Ovis orientalis musimon) , 1995, Behavioural Processes.

[39]  T. Pitcher,et al.  When fish shoals meet: outcomes for evolution and fisheries , 2003 .

[40]  Stuart A. Altmann,et al.  Estes, R. D. The Behavior Guide to African Mammals, Including Hoofed Mammals, Carnivores and Primates. University of California Press, Berkeley, California , 1992 .

[41]  W. Franklin,et al.  Social Organization and Home Range of Roosevelt Elk , 1975 .

[42]  W. Bresiński Grouping tendencies in roe deer under agrocenosis conditions , 1982 .

[43]  Simon Benhamou,et al.  FAD: Fish Aggregating Device or Fish Attracting Device? A new analysis of yellowfin tuna movements around floating objects , 2004, Animal Behaviour.

[44]  Peter Wirtz,et al.  Group Sizes of Antelopes in an East African National Park , 1983 .

[45]  T. J. Roper,et al.  Activity synchrony and social cohesion: a fission-fusion model , 2000, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[46]  Simon Benhamou,et al.  Homing in green turtles Chelonia mydas: oceanic currents act as a constraint rather than as an information source , 2006 .

[47]  J. Gerard,et al.  Spontaneous Emergence of a Relationship between Habitat Openness and Mean Group Size and its Possible Evolutionary Consequences in Large Herbivores , 1995 .

[48]  SIMON BENHAMOU,et al.  No evidence for cognitive mapping in rats , 1996, Animal Behaviour.

[49]  David R. Anderson,et al.  Model selection and multimodel inference : a practical information-theoretic approach , 2003 .

[50]  David Sloan Wilson,et al.  Living in Groups.Oxford Series in Ecology and Evolution.ByJens Krauseand, Graeme D Ruxton.Oxford and New York: Oxford University Press. $95.00 (hardcover); $45.00 (paper). xiv + 210 p; ill.; author and general indexes. ISBN: 0–19–850817–4 (hc); 0–19–850818–2 (pb). 2002. , 2003 .

[51]  M. Stüwe,et al.  Organization of roe deer (Capreolus capreolus) in an open field habitat , 1984 .

[52]  S. Benhamou How to reliably estimate the tortuosity of an animal's path: straightness, sinuosity, or fractal dimension? , 2004, Journal of theoretical biology.

[53]  G. Caughley Social Organization and Daily Activity of the Red Kangaroo and the Grey Kangaroo , 1964 .

[54]  P. Jarman,et al.  The Social Organisation of Antelope in Relation To Their Ecology , 1974 .

[55]  R. R. Krausz Living in Groups , 2013 .

[56]  P. Banks Predation-sensitive grouping and habitat use by eastern grey kangaroos: a field experiment , 2001, Animal Behaviour.

[57]  Simon Benhamou,et al.  Detecting an orientation component in animal paths when the preferred direction is individual-dependent. , 2006, Ecology.

[58]  David R. Anderson,et al.  Model Selection and Multimodel Inference , 2003 .

[59]  A theoretical study of the socioecology of ungulates. II. A dynamic programming study of the stochastic formulation. , 1992, Theoretical population biology.

[60]  J. Gerard,et al.  Probable exaptations within the distributed' herd , 1993 .