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insulation, waterproofing and aerodynamic flight � birds spend a great deal of time
maintaining them. Studies on multiple species have shown that birds spend an
average of more than 9% of each day on maintenance behaviours, with more than 92%
of that time spent preening. However, this number can be significantly higher. One
study found that some gull species spend 15% of their daylight hours preening
during the breeding season, for example, and another showed common loons spend
upwards of 25% of their day preening. In most of the studied species where the
bird's sex could be determined in the field, males spent more time preening than
females, though this was reversed in ducks.
There is some evidence that allopreening may help to keep in good condition those
feathers that a bird cannot easily reach by itself; allopreening activities tend to
focus on the head and neck, for example.[34][35] It may also help to remove
ectoparasites from those hard-to-reach areas. Allopreening is most common among
species that are regularly in close physical contact due to flocking or social
behaviours,[29] and such contact allows for easier transfer of ectoparasites
between individuals.[36] In one study, Macaroni penguins that frequently
allopreened had significantly fewer ticks on their heads and necks than those that
did not.[37] Green wood hoopoes, a flocking species with a complex hierarchy, show
similar frequencies of initiating and reciprocating allopreening of the head and
neck regardless of social status, time of year or group size, which suggests that
such activity is primarily related to feather hygiene.[32]
Most allopreening is done between the two members of a mated pair,[9] and the
activity appears to play an important role in strengthening and maintaining pair
bonds. It is more common in species where both parents help to raise the offspring,
and correlates with an increased likelihood that partners will remain together for
successive breeding seasons.[36] Allopreening often features as part of the
"greeting ceremony" between the members of a pair in species such as albatrosses
and penguins, where partners may be separated for a relatively long period of time,
and is far more common among species that are sexually monomorphic (that is,
species where the sexes look outwardly similar). It appears to inhibit or sublimate
aggression, as it is typically the dominant bird that initiates the behaviour.[31]