Intimidating rivals
Biologists have long marvelled at how seldom animal conflicts over
food, territories, mates, or other resources escalate to all-out
fighting. Prior to unrestrained aggression, animals often engage in
extended bouts of signalling, making prominant display of their
weapons (e.g. antlers) or running through a reportoire of
highly-stereotyped aggression signals. These displays are presumably
designed to convey information about the aggressive intent and/or the
fighting ability of the participants. But why should these signals be
reliable? Why don't opponents lie, and pretend to be the "tough guy"
all the time? At least three closely related arguments have been
proposed.
First, prominant signals of fighting ability may be impossible, or
impossibly costly, to fake. For example, a full-grown bull elk's rack
of antlers may weigh in excess of 40 pounds - far more than a young,
weak, or sickly individual could hope to carry. Costly traits of this
sort would presumably function in a manner similar to sexual selection
ornaments.
Second, Zahavi (1975) proposed that gestures or displays which put an
individual at a relative disadvantage if attacked, such as exposing a
flank, may serve as reliable signals of fighting ability. If I want to
show an opponent that I am not frightened by him, one approach is to
turn my back and "dare him" to try a sucker punch. Taking a risk like
this can be an honest signal, because a stronger fighter can
afford to send it, whereas a weaker fighter could be knocked out right
away if he tried this tactic.
Third, some signals involved in competitive or aggressive interactions
may be produced without directly imposing a cost or increased risk on
the signaller. These signals - such as the dark throat-patch on a
dominant male sparrow - are called ``conventional signals'' or
``badges or status.'' Such signals are thought to remain honest
because of their indirect consequences: signal receivers will severely
punish those who are caught sending misleading signals. For example,
Rohwer (1977) and Moeller (1987) found that when a sparrow's throat
badge was artificially augmented with a dark dye, making him appear
more dominant than he truly was, other sparrows soon saw through the
deception and vicously attacked the dyed individual.
Primary literature
Geist (1966) and Zahavi (1975) were among the first to consider
unfakable siganls of fighting ability. In the same 1975 paper, Zahavi
also proposed that risks could serve as reliable signals. Adams and
Mesterton-Gibbons (1995) provided a formal model of this scenario;
interestingly, they find that some bluffing is likely at
equilibrium. Rohwer (1975) proposed that variable plumage characters
serve as "badges of status" in several bird species. Enquist (1985)
and Maynard Smith and Harper (1988) published two of the first
game-theoretic models of aggressive signalling. Hurd (1997)
highlighted the distinction between costly signals, risks, and badges
of status.