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Dominance Hierarchies

Dominance Hierarchies

It all started with hens. Norwegian scientist Thorleif Schjelderup-Ebbe


wondered how peace was kept in their flocks, and conducted a series of
experiments to find out. He discovered that things were tranquil only in
established flocks -- ones in which each hen knew its place. And the hens
learned their places in fights over chicken feed. Once a hen had been bested in
a squabble, it henceforth would defer to the victor. Each hen knew whom it
could dominate, and by whom it would be dominated. A "peck order" was
thus established in the flock and functioned to maintain social stability. Hens
can recognize many other hens and can remember their dominance status
relative to each. One hen demonstrated the ability to recognize 27 other
individuals belonging to four different flocks. As you would expect, the birds
at the top of the peck order benefited both by increased access to food and by
avoidance of injuries (even bullies can get hurt in fights). The birds at the
bottom, while having to wait until those higher up had eaten their fill, at least
were not subjected to continuous fights that they were likely to lose.

In all but polyandrous species of birds (those where one female mates with
more than one male), males are normally dominant over females. That is
especially true in the early stages of pair formation, although in some buntings
and other finches and in some gulls, a reversal of dominance, with the male
becoming subordinate, reportedly occurs after the pair bond is established. In
experiments, females have been able to assume higher ranks in dominance
hierarchies after receiving injections of male hormones and by dyeing their
feathers to resemble male plumage. Results of such experiments varied from
species to species; doses of male hormone increased the social status of female
European Chaffinches and Japanese Quail, but not of starlings.

It is usually assumed that a high position in a dominance hierarchy increases


the chances of survival and also increases reproductive output. In species
where males display on leks, the dominant male generally holds the best
territory on the lek and successfully copulates with the most females. But there
is relatively little evidence in monogamous species that dominant individuals
have a better chance of survival than subordinates.

A study of juvenile Song Sparrows on an island near Victoria, British


Columbia, showed that dominance relationships at millet-provisioned feeders
during summer were reflected in chances for surviving the subsequent winter
and establishing a territory the following spring. In two consecutive years,
dominant males showed 35 and 22 percent better survival than subordinate
males. For dominant females the equivalent figures were 32 and 33 percent.
The effects on successful settlement of territories were similar for both sexes --
dominant individuals were more likely to settle into territories than
subordinates.

The same study showed that sex and age were the major determinants of
dominance. Males more frequently dominated females than vice versa; males
tended to win about 60 percent of their encounters with females at the feeders,
females about 40 percent. Time since hatching had a strong effect on the
success of encounters, even when only 24 days separated the oldest juvenile
from the youngest. In contrast, no effect of body size on dominance was
detected; a small bird was as likely to dominate a larger one as the reverse.
Overall, it appeared that next after sex and age, previous experience in
encounters was the most important factor in achieving dominance. Thus, early
nesting would appear to be the best reproductive strategy for Song Sparrows.
Young produced early in the season will gain the most experience and are the
most likely to survive to reproduce. But if nesting is started too early,
offspring may be killed by food shortage or late winter storms. Undoubtedly,
there is a fine ecological line between reaping the advantages of early nesting
and suffering the consequences of premature breeding.

In another study of Yellow Warblers, however, Michael Studd and Raleigh


Robertson of Queens University in Ontario discovered that the nestlings of
subordinate birds grew just as fast as those of dominant birds. At least by that
measure (to be sure, not an exhaustive one), natural selection was not
operating in favor of dominant birds. It has been suggested that in many
circumstances being dominant is no more advantageous than being
subordinate. For example, a dominant male may obtain a territory which is
more resource-rich than that of a subordinate, but it may have to spend much
more energy to protect it. Therefore, costs and benefits may more or less
balance one another at each level in a dominance hierarchy, and evolution
may favor the maintenance of the hierarchy itself, rather than just those near
the top (which could lead to the disappearance of differences in dominance).
Such a situation in which selection leads to the coexistence of several different
modes of behavior has been called by British evolutionist John Maynard Smith
a "mixed evolutionarily stable strategy."

SEE: Bird Badges; Natural Selection; Visual Displays.

Copyright ® 1988 by Paul R. Ehrlich, David S. Dobkin, and Darryl Wheye.

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