Fighting for space: How do weaker individuals compete?

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.