, 2006) and spotted hyenas (Holekamp & Smale, 2000), increased levels of competition between females can extend back into adolescence and early development. For example, in meerkats, competitive interactions between adolescents are more frequent between females than between males (Clutton-Brock, 2009b) while, in spotted hyenas, siblicide (which occurs when resources are at short supply) is more frequent between females than between males or litters of mixed sex (Hofer & East, 1997, 2008; James & Hofer, 1999). As yet, detailed studies of fighting tactics have been almost totally confined find more to studies of males. However, accounts of fights between females suggest

that their distribution and duration coincide with the predictions of theoretical models: fights appear to be most frequent and intense where the benefits of winning or the costs of losing are large, and longest when the resource holding power (RHP; Parker, 1974) of contestants is approximately similar. There are probably several reasons why physical attacks are usually less frequent and less intense in females than in males (Andersson, 1980).

First, the fitness benefits associated with the resources at stake are greater Selleckchem Galunisertib in males than in females, as a consequence of both increased variance in reproductive success and of contrasts in Bateman gradients (Kokko, Klug & Jennions, 2012). Second, a lesser number of individuals commonly compete simultaneously for the same resources as a result of biases in the operational sex ratio (Emlen & Oring, 1977). Third, risks associated with escalated fights may frequently be higher for females than for males, as they may entail fatal injuries for dependent offspring: for example, territorial fights among females frequently result in infant deaths in ring-tailed lemurs (Jolly et al., 2000) and, in several species, lactating females

MCE rarely engage in aggressive interactions (Wasser & Starling, 1988; Huchard & Cowlishaw, 2011). Finally, as a result of female philopatry, females are frequently competing with relatives, whereas males are typically competing with unrelated individuals. In addition, philopatry can allow females to control the presence or development of potential rivals, so that threats between individuals of approximately equal RHP are less common than among males (Clutton-Brock, 2009b; Clutton-Brock et al., 2010). While conflicts between females sometimes lead to direct fighting, the majority of aggressive interactions between group members involve threats rather than physical attacks (Andersson, 1980). For example, in studies of vervet monkeys, although maternal interventions occurred in less than 4% of juvenile interactions, maternal dominance rank predicted the outcome of up to 85.5% of all dyadic aggressive interactions between juveniles and 94.