…carnivores has been a focus for theoretical models (e.g. Macdonald 1983; Carr and Macdonald 1986; Bacon et al. 1991) and empirical studies (e.g. Kruuk 1978; Kruuk and Parish 1982; Mills 1989; Mills and Gorman 1997; Geffen et al. 1992; Patterson and Messier 2001; Valenzuela and Macdonald 2002).