Fighting for space: How do weaker individuals compete?

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Animals fight for territories and larger, stronger individuals usually win. How then, in fiddler crabs, do small males and weaponless females defend their territories against larger males? I will investigate four contexts in which strength alone may not determine fighting success. I predict that: (a) positional and motivational advantages allow territory owners to repel intruders; (b) weak individuals compete with stronger neighbours by 'nagging' rather than by winning fights; (c) residents form coalitions to fend off intruders; and d) females, who lack the large claws of males, employ novel alternative tactics to maintain territories.This study will test important theories of territoriality and in so doing make a major contribution to behavioural ecology.