Social structure in a colonial mammal: unravelling hidden structural layers and their foundations by network analysis

The processes governing social interactions and the resulting relationships among members of colonially organized animals are largely unknown. Using network analytical tools we investigated fundamental components of social structure in a highly fluid fission–fusion society. We constructed a social network and studied the relative importance of sex and age class, fine-scale site fidelity and male territory distribution in a breeding colony of the Galapagos sealion, Zalophus wollebaeki. The social system was partitioned into a hierarchy with at least three levels. The outermost level of organization was given by a single social network, where all individuals of the population were interconnected. This level split into communities and further into cliques. Sex and age class influenced social structure at the population level, whereas fine-scale site fidelity explained most of the structure found at the community level. None of these assortment variables could account for clique structure, which might therefore be explained in terms of individual preferences, genetic relatedness or a combination of both. Male territories did not form the basic unit of social structure, but appeared simply to be superimposed on the structural backbone formed by females and young.

[1]  M. Newman,et al.  Mixing patterns in networks. , 2002, Physical review. E, Statistical, nonlinear, and soft matter physics.

[2]  J. Hoogland,et al.  Population genetics meets behavioral ecology. , 1996, Trends in ecology & evolution.

[3]  W. Hamilton,et al.  The Evolution of Cooperation , 1984 .

[4]  C. D. Gelatt,et al.  Optimization by Simulated Annealing , 1983, Science.

[5]  C. Barnard Animal Behaviour: Mechanism, Development, Function and Evolution , 2003 .

[6]  F. Trillmich,et al.  The mating systems of pinnipeds and marine iguanas : convergent evolution of polygyny , 1984 .

[7]  Peter M. Kappeler,et al.  Evolution of Primate Social Systems , 2002, International Journal of Primatology.

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

[9]  J. Godin,et al.  Association patterns and shoal fidelity in the three–spined stickleback , 2002, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[10]  R. Harcourt Factors affecting early mortality in the South American fur seal (Arctocephalus australis) in Peru: density‐related effects and predation , 1992 .

[11]  M. Slatkin,et al.  Fifty years of evolution: essays in honour of John Maynard Smith. , 1987, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[12]  Levy Bj,et al.  Perinatal behavior of northern elephant seal females and their young. , 1973 .

[13]  Lars Bejder,et al.  Testing association patterns: issues arising and extensions , 2005, Animal Behaviour.

[14]  H. Whitehead Analysing animal social structure , 1997, Animal Behaviour.

[15]  D. Tautz,et al.  Galápagos and Californian sea lions are separate species: Genetic analysis of the genus Zalophus and its implications for conservation management , 2007, Frontiers in Zoology.

[16]  John W. Pepper,et al.  Sex Differences in Patterns of Association Among Indian Ocean Bottlenose Dolphins , 1992 .

[17]  G. Kauermann,et al.  Males in the shade: habitat use and sexual segregation in the Galápagos sea lion (Zalophus californianus wollebaeki) , 2005, Behavioral Ecology and Sociobiology.

[18]  R. Bon,et al.  Unexplained sexual segregation in polygamous ungulates: a defense of an ontogenetic approach , 1996, Behavioural Processes.

[19]  G. Bartholomew A MODEL FOR THE EVOLUTION OF PINNIPED POLYGYNY , 1970, Evolution; international journal of organic evolution.

[20]  F. Galimberti,et al.  Relatedness and site fidelity at the southern elephant seal, Mirounga leonina, breeding colony in the Falkland Islands , 2006, Animal Behaviour.

[21]  F. Strayer,et al.  Genèse De La Ségrégation Sexuelle Et Différences Comportementales Chez Des Enfants D'Age Préscolaire , 1991 .

[22]  M. Newman,et al.  Finding community structure in very large networks. , 2004, Physical review. E, Statistical, nonlinear, and soft matter physics.

[23]  F. Trillmich,et al.  Parent–offspring and sibling conflict in Galápagos fur seals and sea lions , 2007, Behavioral Ecology and Sociobiology.

[24]  I. Gordon,et al.  Are social factors sufficient to explain sexual segregation in ungulates? , 2005, Animal Behaviour.

[25]  Ronald J. Schusterman,et al.  Kin recognition in captive California sea lions (Zalophus californianus). , 1990 .

[26]  D. Renouf The Behaviour of Pinnipeds , 1990, Springer Netherlands.

[27]  T. Härkönen,et al.  Spatial structure of harbour seal populations and the implications thereof , 2001 .

[28]  M E J Newman,et al.  Finding and evaluating community structure in networks. , 2003, Physical review. E, Statistical, nonlinear, and soft matter physics.

[29]  G. Orians,et al.  Familiar neighbors enhance breeding success in birds. , 1989, Proceedings of the National Academy of Sciences of the United States of America.

[30]  Richard E. Michod,et al.  The Evolution of Cooperation in Spatially Heterogeneous Populations , 1996, The American Naturalist.

[31]  Edward H. Miller,et al.  Communication in pinnipeds, with special reference to non-acoustic signalling , 1991 .

[32]  D. Lusseau,et al.  Quantifying the influence of sociality on population structure in bottlenose dolphins. , 2006, The Journal of animal ecology.

[33]  Roger L. Gentry,et al.  Northern Fur Seal Behavior and Ecology@@@Behavior and Ecology of the Northern fur Seal , 1998 .

[34]  Richard James,et al.  Social networks in the guppy (Poecilia reticulata) , 2004, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[35]  Jochen B. W. Wolf,et al.  Beyond habitat requirements: individual fine-scale site fidelity in a colony of the Galapagos sea lion (Zalophus wollebaeki) creates conditions for social structuring , 2007, Oecologia.

[36]  R. Gentry The Development of Social Behavior Through Play in the Steller Sea Lion , 1974 .

[37]  S. Twiss,et al.  Reproductive performance links to fine-scale spatial patterns of female grey seal relatedness , 2001, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[38]  Mark E. J. Newman,et al.  The Structure and Function of Complex Networks , 2003, SIAM Rev..

[39]  P. Redman,et al.  Breeding site choice fails to explain interannual associations of female grey seals , 2005, Behavioral Ecology and Sociobiology.

[40]  R. Guimerà,et al.  Modularity from fluctuations in random graphs and complex networks. , 2004, Physical review. E, Statistical, nonlinear, and soft matter physics.

[41]  T. Clutton‐Brock,et al.  The Evolution of Parental Care , 2019 .

[42]  J. Krause,et al.  Assortative interactions and social networks in fish , 2005, Oecologia.

[43]  M. Fenton,et al.  Analysis of Spix's disc-winged bat association patterns and roosting home ranges reveal a novel social structure among bats , 2004, Animal Behaviour.

[44]  K. Ruckstuhl,et al.  Sexual segregation in vertebrates : ecology of the two sexes , 2006 .

[45]  S. Twiss,et al.  Dispersion and site fidelity of breeding male grey seals (Halichoevus grypus) on North Rona, Scotland , 1994 .

[46]  P. Redman,et al.  Philopatry, Site Fidelity and Local Kin Associations within Grey Seal Breeding Colonies , 2000 .

[47]  W. Getz,et al.  The socioecology of elephants: analysis of the processes creating multitiered social structures , 2005, Animal Behaviour.