ferociously, for an oestrual female (Schaller et al., 1985). It has been proposed that male hyperaggression in captivity may be an expression of a species-typical behaviour with no outlet and, thus, is misdirected toward females (see Chapter 12). The result is that in the ex situ environment, and in the absence of access to same-sex competitors, a male will often fight with a female, even one in oestrus. Thus it makes sense that a behaviourally aggressive female is more competent at handling an aggressive male, and perhaps breeding is less successful when only one of the animals is aggressive. In both cases there are many anecdotal reports of either shy males or shy females being so aggressively pursued by a cantankerous conspecific that mating is impossible. Being wild born also conferred a distinctive advantage, presumably appropriate aggressive behaviours are learned from the dam, and they better equip an individual for future agonistic encounters, including during mating.
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