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Packer, C.; Pusey, A.E.
Cooperation and competition within coalitions of male lions: kin selection or game theory?
1982  Nature (296): 740-742

Male lions form cooperative coalitions which compete against other coalitions for exclusive access to female groups. This cooperation and the apparently low level of intra-coalition competition over oestrus females, have been considered to be due to the close genetic relatedness of the males in the coalition. However, we now present evidence that breeding coalitions of male lions include non-relatives much more commonly than was generally supposed, that intra-coalition competition over females is widespread and that kinship is not the primary factor determining levels of competition.

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