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J. theor. Biol. (1971) 31, 295-311
Geometry for the Selfish Herd
W. D. HAMILTON
Department of Zoology,
Imperial College, London, S. W.7, England
(Received 28 September 1970)
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 un-
subdivided populations.
The model for two dimensions involves a complex problem in geo-
metrical probability which has relevance a
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