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728:. 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. 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 sexually monomorphic species (that is, species where the sexes look outwardly similar). It appears to inhibit or redirect aggression, as it is typically the dominant bird that initiates the behaviour.
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allopreening of another bird's body rather than head and neck) increase with group size. Evidence suggests this type of allopreening reduces social tension, and thus plays an important role in group cohesion. More dominant birds receive far more body allopreening services than do lower-ranked birds, and lower-ranked birds initiate far more body allopreening bouts than do their higher-ranked flock mates. Body allopreening is only reciprocal when done between members of a mated pair; otherwise, the dominant bird reciprocates in fewer than 10% of the instances.
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301:. This alters the colour of the eggs (darkening them) but there is also evidence that the bacteria may help to protect the developing chicks. Other studies have shown that removing or restricting access to the uropygial gland typically results in a higher bacterial parasite load on the plumage, though not necessarily of feather-degrading bacterial species. Preen oil may play a part in protecting at least some species from some internal parasites; a study of the incidence of
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activities â dislodged when the bird brushes up against vegetation, for instance, or when it interacts with another bird during fighting or mating. Preening may involve two kinds of bill actions: nibbling (or mandibulating) while working the feather from base to tip, or stroking with the bill either open or closed. The nibbling action is the one used most often; it is more effective than stroking for applying preen oil, removing
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459:) have comb-like serrations on the claw (a pectinate claw) of the middle toe which may aid in scratch-preening. Some species stretch their leg over their lowered wing to reach their head (known as "indirect" scratching), while others extend their leg between their wing and their body (known as "direct" scratching). There is some evidence that the method used by a species may be related to its ecology. For instance,
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considerable time in their efforts, they do not use proper techniques to groom effectively and may do a poor job overall as a result. Displaced feathers can cause birds considerable trouble; such feathers might become damaged, could interrupt the smooth flow of air over a flying bird, or might allow the bird's body heat to escape. Preening allows a bird to reposition such displaced feathers. There is evidence that
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functions are not mutually exclusive. Evidence suggests that different species may participate for different reasons, and that those reasons may change depending on the season and the individuals involved. In most cases, allopreening involves members of the same species, although some cases of interspecific allopreening are known; the vast majority of these involve
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members in a social species. Such behaviour may assist in effective grooming, in the recognition of individuals (mates or potential sexual partners), or in reducing or redirecting potential aggressive tendencies in social species. Most allopreening is confined to the head and neck, smaller efforts being directed towards other parts of the body.
434:, rejoining unzipped barbules, and rearranging feathers. The stroking action is typically done in the direction the feathers lie, with the bill either opened or closed. Stroking is used to apply preen oil, as well as to dry and smooth plumage. Grebes stroke more vigorously with an open bill â a behaviour known as "stropping".
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85:, and so are vital to its survival. Because of this, birds spend considerable time each day maintaining their feathers, primarily through preening. Several actions make up preening behaviour. Birds fluff up and shake their feathers, which helps to "rezip" feather barbules that have become unhooked. Using their
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feathers which continually break down into a fine dust that the birds apply to their contour feathers while preening. These powder down feathers may be scattered throughout the bird's plumage or concentrated into dense patches. As well as helping to waterproof and preserve the bird's feathers, powder
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Preening is a maintenance behaviour used by all birds to care for their feathers. It is an innate behaviour; birds are born knowing the basics, but there is a learned component. Birds that are hand-reared without access to a role model have abnormalities in their preening behaviours. Despite spending
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and waterproofing, and rapidly leads to the bird becoming chilled. If waterbirds are exposed, they can lose both buoyancy and the ability to fly; this means they must swim constantly to stay warm and afloat (if they cannot reach land), and eventually die of exhaustion. While preening in an effort to
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of eggs (by the removal of eggs from their nest) responded by preening and nest building â both displacement activities. When all three eggs in their regular clutch were removed, the gulls showed a significant increase in the amount of time they spent preening. The conflict between two incompatible
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Birds cannot use their beaks to apply preen oil to their own heads. Instead, many use their feet in an action called scratch-preening. Once they have gathered preen oil on their beak, they scrape a foot across their bill to transfer the oil, and then scratch the oil into the feathers on their head.
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that can occur when birds are subjected to two conflicting drives. Though primarily an individual function, preening can be a social activity involving two or more birds â a behaviour known as allopreening. In general, allopreening occurs either between two members of a mated pair or between flock
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Because feathers are critical to a bird's survival â contributing to insulation, waterproofing and aerodynamic flight â birds spend a great deal of time maintaining them. When resting, birds may preen at least once an hour. Studies on multiple species have shown that they spend an average of more
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Preening may help to send sexual signals to potential mates because plumage colouration (which can be altered by the act of preening) can reliably reflect the health or "quality" of its bearer. In some species, preen oil is used to cosmetically colour the plumage. During the breeding season, the
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Several hypotheses have been advanced to explain the behaviour: that it assists in effective grooming, that it assists in recognition of individuals (mates or potential sexual partners), and that it assists in social communication, reducing or redirecting potential aggressive tendencies. These
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has been documented. Birds seeking allopreening adopt specific, ritualised postures to signal so; they may fluff their feathers out or put their heads down. Captive birds of social species that normally live in flocks, such as parrots, will regularly solicit preening from their human owners.
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branching from that shaft. Pennaceous feathers also have much smaller barbules branching from the entire length of each barb; these barbules have tiny hooks along their length, which interlock with the hooks of neighbouring barbules. Barbules can become unhooked as a result of a bird's daily
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that engaged in allopreening were much less likely to fight. Since fights often lead to eggs or chicks being knocked off breeding cliffs, fewer fights led to greater breeding success for allopreening neighbours. Among social flocks of green wood hoopoes, rates of body allopreening (that is,
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Preen oil helps to maintain the waterproofing of a bird's plumage. Though the oil does not provide any direct waterproofing agent, it helps to extend the life of the feather â including the microscopic structures (the barbs and barbules) which interlock to create the waterproof barrier.
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261:, which opens above the base of the tail feathers and secretes a substance containing fatty acids, water, and waxes. The bird gathers this substance on its bill and applies it to its feathers. The gland is generally larger (in relation to body size) in waterbirds, including
705:, 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.
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or disease-causing organisms during preening can lead to problems ranging from liver and kidney damage to pneumonia and disease transmission. Injury and infection can cause overpreening in caged birds, as can confining a bird with a dominant or aggressive cage mate.
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of both sexes will lift a wing so that the brightly coloured speculum is showing, then will place their bill behind the speculum as if preening it. Courtship preening is more conspicuous than is preening for feather maintenance, using more stereotypical movements.
773:, as well as liver and kidney damage. Studies done with black guillemots showed that even small amounts of ingested oil caused the birds physiological distress. It interfered with the foraging efficiency of adults and decreased the growth rates of young birds.
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tend to be overwing scratchers, while those that spend significant time on the ground are typically underwing scratchers. In general, preening takes place while the bird is perched, on the ground, or swimming, but some of the more aerial species (including
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use their whole heads to stroke, in a motion referred to as "wiping". Birds regularly fluff up their plumage and repeatedly shake their bodies while preening. Experiments have shown that the shaking action can "rezip" a majority of split feather barbules.
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Williams, Joseph B.; Siegfried, W. Roy; Milton, Suzanne J.; Adams, Nigel J.; Dean, W. R. J.; du
Plessis, Morne A. & Jackson, Sue (March 1993). "Field metabolism, water requirements, and foraging behavior of wild ostriches in the Namib".
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and petrels. However, studies have found no clear correlation between the size of the gland and the amount of time a species spends in the water; it is not consistently largest in those species that spend the most time in the water.
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Fully grown feathers are essentially dead structures, so it is vital that birds have some way to protect and lubricate them. Otherwise, age and exposure cause them to become brittle. To facilitate that care, many bird species have a
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at the base of the filoplumes only fire when contour feathers are displaced or the filoplume moves. Preening enables birds to remove dirt and parasites from their plumage, and assists in the waterproofing of feathers. During
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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. It may also help to remove
495:, and can either precede or follow these other behaviours. All birds typically preen after bathing. Groups of birds sometimes all groom individually at the same time. This has been seen in species ranging from herons to
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spent 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. Some
297:. Female hoopoes transfer preen oil onto their brood patches and eggs, which results in the transfer of bacteria as well. Preen oil and bacteria are rubbed into microscopic pits on the surface of the eggs during
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Karunakaran, Athira
Cheruplackal; Murugkar, Harshad V.; Kumar, Manoj; Nagarajan, Shanmugasundaram; Tosh, Chakradhar; Pathak, Anubha; Rajendrakumar, Arunraj Mekhemadhom; Agarwal, Rajesh Kumar (May 2019).
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Delogu, Mauro; De Marco, Maria A.; Di Trani, Livia; Raffini, Elisabetta; Cotti, Claudia; Puzelli, Simona; Ostanello, Fabio; Webster, Robert G.; Cassone, Antonio & Donatelli, Isabella (25 June 2010).
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Although preening is primarily an individual behaviour, some species indulge in allopreening, one individual preening another. It is not particularly common among birds, though species from at least 43
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found that uninfected birds had larger uropygial glands and higher antimicrobial activity in those glands than infected birds did. There is even evidence that the foul-smelling preen oil of hoopoes and
528:, contains a pink colourant which does the same. The heads of these birds typically show little pink, because of the difficulty of reaching those areas with preen oil. The yellow feathers of the
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MartĂnez-GarcĂa, Ăngela; Soler, Juan J.; RodrĂguez-Ruano, Sonia M.; MartĂnez-Bueno, Manuel; MartĂn-Platero, Antonio Manuel; JuĂĄrez-GarcĂa, Natalia & MartĂn-Vivaldi, Manuel (November 2015).
784:, for instance, meaning that birds that preen infected partners might become infected themselves. Even preening its own body may expose a bird to pathogens. There is evidence that water-borne
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PĂ©rez-RodrĂguez, Lorenzo; Mougeot, Francois & Bortolotti, Gary R. (2011-07-01). "The effects of preen oils and soiling on the UVâvisible reflectance of carotenoid-pigmented feathers".
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480:) preen while flying. Many birds have a slight overhang at the tip of their upper mandible. Experiments suggest that this allows birds to apply shearing forces that kill the flat-bodied
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virus is "captured" by the preen oil on feathers, providing a possible route for infection. The ingestion of parasites during preening may result in infection; the tick-borne disease
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than 9% of each day on maintenance behaviours, preening occupying over 92% of that time, though this figure can be significantly higher. Studies found that some
810:) can help to eliminate the behaviour. Confining a bird with an incompatible or very dominant cage mate can lead to excessive allopreening, which can result in
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Preen oil plays a role in reducing the presence of parasitic organisms, such as feather-degrading bacteria, lice and fungi, on a bird's feathers. One study of
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Necker, Reinhold (May 1985). "Observations on the function of a slowly-adapting mechanoreceptor associated with filoplumes in the feathered skin of pigeons".
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CzirjĂĄk, GĂĄbor ĂrpĂĄd; Pap, PĂ©ter LĂĄszlĂł; VĂĄgĂĄsi, Csongor IstvĂĄn; Giraudeau, Mathieu; MureĆan, Cosmin; Mirleau, Pascal & Heeb, Philipp (February 2013).
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preen when they have been alarmed by a potential predator or when they have had an aggressive encounter with a neighbouring bird, for instance. Fighting
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Most allopreening is done between the two members of a mated pair, and the activity appears to play an important role in strengthening and maintaining
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are known to engage in the mutual activity. Most allopreening activity concentrates on the head and neck, a lesser amount being directed towards the
802:, particularly parrots, sometimes overpreen in response to being exposed to strong scents (such as nicotine or air fresheners) or as a result of
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Allopreening appears to reduce the incidence of conflict between members of some colonially living or colonially nesting species. Neighbouring
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by several species, particularly ducks; such preening is typically designed to draw attention to a modified structure (such as the sail-shaped
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Tinbergen, N. (March 1952). ""Derived" activities; their causation, biological significance, origin, and emancipation during evolution".
2000:
Delhey, Kaspar; Peters, Anne & Kempenaers, Bart (2007-01-01). "Cosmetic coloration in birds: occurrence, function, and evolution".
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Clayton, Dale H.; Moyer, Brett R.; Bush, Sarah E.; Jones, Tony G.; Gardiner, David W.; Rhodes, Barry B. & Goller, Franz (2005).
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Elphick, Chris & Dunning, John B. Jr. (2001). "Behaviour". In
Elphick, Chris; Dunning, John B. Jr. & Sibley, David (eds.).
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Van Iersel, J. J. A. & Bol, A. C. Angela (January 1958). "Preening of two tern species: a study on displacement activities".
1517:"Volume and antimicrobial activity of secretions of the uropygial gland are correlated with malaria infection in house sparrows"
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2832:"Survivability of highly pathogenic avian influenza virus (H5N1) in naturally preened duck feathers at different temperatures"
806:. Reducing exposure to the offending odour, or treating the underlying cause of the neuropathy (such as injury, infection, or
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from those hard-to-reach areas. Allopreening is most common among species that are regularly in close physical contact due to
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Ruiz-RodrĂguez, M.; Valdivia, E.; Soler, Juan J.; MartĂn-Vivaldi, M.; MartĂn-Platero, A. M. & MartĂnez-Bueno, M. (2009).
568:. In some cases, it is done in place of another activity that birds are strongly motivated, but unable, to do. In one study,
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becomes red-orange, imparting a pink flush to the bird's plumage. The preen oil of several gull and tern species, including
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Gilbert, Lucy; Jones, Linda D.; Laurenson, M. Karen; Gould, Ernie A.; Reid, Hugh W. & Hudson, Peter J. (7 May 2004).
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or social behaviours, where such contact allows for easier transfer of ectoparasites between individuals. In one study,
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2335:"Allopreening in birds is associated with parental cooperation over offspring care and stable pair bonds across years"
2289:"Dermatological aspects of displacement activity: attention to the body surface as a substitute for "fight or flight""
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Giraudeau, M.; CzirjĂĄk, G.Ă.; Duval, C.; Bretagnolle, V.; Gutierrez, C.; Guillon, N. & Heeb, P. (January 2013).
228:, which are not dependent on their feathers for flight, spend far less time on maintenance behaviours. One study of
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Rowley, Ian (1997). "Family
Cacatuidae (Cockatoos)". In del Hoyo, Josep; Elliott, Andrew; Sargatal, Jordi (eds.).
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Magallanes, Sergio; Pape MĂžller, Anders; GarcĂa-Longoria, Luz; de Lope, Florentino & Marzal, Alfonso (2016).
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Zhao, Jing-Shan; Zhanga, Jiayue; Zhao, Yuping; Zhanga, Zhaodong & Godefroit, Pascal (February 2020).
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1661:"Shaking the wings and preening feathers with the beak help a bird to recover its ruffled feather vane"
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Soler, Juan J.; MartĂn-Vivaldi, M.; Peralta-SĂĄnchez, J. M.; Arco, L.; JuĂĄrez-GarcĂa-Pelayo, N. (2014).
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branching from those barbs. The barbules' hooks interlock as shown to provide strength and flexibility.
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drives, such as incubating and escape, can lead a bird to engage in displacement activities. Nesting
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156:, which had the same meaning. This usage was combined with the Scottish and northern English dialect
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Lewis, Sue; Roberts, Gilbert; Harris, Mike P.; Prigmore, Carina & Wanless, Sarah (August 2007).
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This article is about a maintenance behaviour in birds. For similar behaviour in other animals, see
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clean themselves, they may ingest harmfully large amounts of the petroleum. Ingested oil can cause
664:. A few species are known to allopreen other areas, including the rump, tail, belly and underwing.
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species spent 15% of daylight hours during the breeding season preening, while another showed that
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Longer-necked birds may rub their head directly on their uropygial gland. Some species (including
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Stettenheim, Peter R. (August 2000). "The integumentary morphology of modern birdsâan overview".
1407:"Preen gland removal increases plumage bacterial load but not that of feather-degrading bacteria"
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Docherty, Douglas E.; Long, Renee
Romaine; Griffin, Kathryn M. & Saito, Emi K. (June 2004).
1816:"Records of flight preening and related aerial activities in birds, particularly the black tern"
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Allopreening may facilitate disease transmission between infected and non-infected individuals.
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Preening is often done in association with other maintenance behaviours, including bathing,
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meaning "pierce" or "pin", due to the "pricking" action of the bird's beak during preening.
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1208:"Symbiotic bacteria living in the hoopoe's uropygial gland prevent feather degradation"
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and her ducklings demonstrate the shaking, nibbling and stroking movements of preening.
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Moynihan, M. (January 1953). "Some displacement activities of the black-headed gull".
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in check. Feathers contribute significantly to a bird's insulation, waterproofing and
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2881:"Ticks need not bite their red grouse hosts to infect them with louping ill virus"
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Ehrlich, Paul R.; Dobkin, David S.; Wheye, Darryl & Pimms, Stuart L. (1994).
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34:) draws individual feathers through its beak, realigning and re-interlocking the
3313:"Veterinary Models of Compulsive Self-grooming: Parallels with Trichotillamania"
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Moon-Fandli, Alice M.; Dodman, Nicholas A. & O'Sullivan, Richard L. (1999).
1470:"Effect of preen oil on plumage bacteria: an experimental test with the mallard"
2486:"Dual function of allopreening in the cooperatively breeding green woodhoopoe,
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Stray
Feathers: Reflections on the Structure, Behaviour and Evolution of Birds
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Kenny, Elspeth; Birkhead, Tim R. & Green, Jonathan P. (JulyâAugust 2017).
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increases the UV reflectance of its feathers. Ritualised preening is used in
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found that they spent less than 1% of their time engaged in such behaviours.
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2772:"Can preening contribute to influenza A virus infection in wild waterbirds?"
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While most species have a preen gland, the organ is missing in the ratites (
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was created from a revision of this article dated 18 July 2023
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1300:
3315:. In Stein, Dan J.; Christenson, Gary A. & Hollander, Eric (eds.).
3162:
Hailman, Jack P. (1985). "Behavior". In
Pettingill, Olin Sewall (ed.).
2848:
2571:
2545:"Interspecific allopreening between crested caracara and black vulture"
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2436:
Harrison, C. J. O. (July 1965). "Allopreening as
Agonistic Behaviour".
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Philosophical
Transactions of the Royal Society B: Biological Sciences
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are also cosmetically coloured during preening. The preen oil of the
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3036:. In del Hoyo, Josep; Elliott, Andrew & Sargatal, Jordi (eds.).
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2013:
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3042:. Vol. 1: Ostrich to Ducks. Barcelona, Spain: Lynx Edicions.
1341:"Hoopoes color their eggs with antimicrobial uropygial secretions"
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underneath. Both feather types have a central shaft with narrower
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1685:
1628:
672:, though at least one instance of mutual grooming between a wild
669:
557:
496:
473:
435:
413:
A bird's plumage is primarily made up of two feather types: firm
349:
229:
182:
101:
74:
62:
3399:. Vol. 4: Sandgrouse to Cuckoos. Barcelona: Lynx Edicions.
3190:
2062:
2044:
1467:
925:
3310:
3078:
Ehrlich, Paul R.; Dobkin, David S. & Wheye, Darryl (1988).
3060:
Nests, Eggs, and
Incubation: New ideas about avian reproduction
2484:
Radford, Andrew N. & Du Plessis, Morné A. (December 2006).
1640:
860:
781:
744:
361:
357:
315:
225:
194:, specialised feathers buried under a bird's outer covering of
93:
and distribute this oil through their feathers. They draw each
2158:
1163:
1161:
1159:
1157:
1155:
1153:
248:, typically use their beak to gather preen gland oil from the
3181:
Huxley, Julian; Hardy, A. C. & Ford, E. B., eds. (1954).
2768:
2629:
448:
266:
204:
150:
1095:
198:, help to signal when contour feathers have been displaced.
3057:
Deeming, D. Charles & Reynolds, S. James, eds. (2015).
2650:"Fitness increases with partner and neighbour allopreening"
1743:
10.1668/0003-1569(2000)040[0461:timomb]2.0.co;2
1150:
952:
940:
698:
262:
216:
86:
58:
3269:
Lovette, Irby C. & Fitzpatrick, John W., eds. (2016).
2878:
2647:
2617:
1978:
1976:
1974:
1404:
780:
has been found in the feather pulp of several species of
329:
2711:
2583:
2581:
515:
ritually preen their distinctive orange "sail" feathers.
3096:
2885:
Proceedings of the Royal Society B: Biological Sciences
2699:
2188:
2176:
1965:
1862:
Proceedings of the Royal Society B: Biological Sciences
3015:(3rd ed.). Oxford, UK: Blackwood Publishing Ltd.
2605:
1999:
1971:
1947:
1700:
1571:
1140:
1138:
1136:
866:
207:, birds remove the sheaths from around their emerging
2929:
2714:"Corvidae feather pulp and West Nile virus detection"
2593:
2578:
2399:
Forsman, Eric D. & Wight, Howard M. (July 1979).
2375:
1855:
1559:
2965:
Campbell, Bruce & Lack, Elizabeth, eds. (1985).
1658:
697:
that frequently allopreened had significantly fewer
364:. Some species that lack a preen gland instead have
137:
to mean the tidying of a bird's feathers dates from
3275:(3rd ed.). Oxford, UK: John Wiley & Sons.
2941:
1268:"Preening as a vehicle for key bacteria in hoopoes"
1133:
1040:"Preening and associated comfort behavior in birds"
3268:
3140:
3098:
3077:
3031:
2966:
2332:
2050:
1694:
1634:
964:
934:
796:if the bird consumes a tick carrying the disease.
701:on their heads and necks than those that did not.
97:through their bill, nibbling it from base to tip.
3380:. Princeton, NJ, US: Princeton University Press.
3166:(5th ed.). Orlando, FL, US: Academic Press.
2483:
913:
381:A shaft runs down the middle of the feather with
3517:
2543:Ng, David & Jasperson, Bruce D. (May 1984).
1180:Montalti, Diego & SalibiĂĄn, Alfredo (2000).
1179:
988:Cotgreave, Peter & Clayton, Dale H. (1994).
987:
277:showed that the presence of symbiotic bacteria (
3420:. Cambridge, MA, US: Harvard University Press.
3180:
3056:
2635:
2401:"Allopreening in owls: what are its functions?"
2164:
1259:
369:down can give a metallic sheen to the plumage.
3508:Splendid fairy-wrens preening and allopreening
3319:. Washington, DC: American Psychiatric Press.
3119:
1772:Nice, Margaret M. & Schantz, W.E. (1959).
958:
946:
656:and an even smaller percentage applied to the
61:that involves the use of the beak to position
3212:. Boston, MA, US: Houghton Mifflin Harcourt.
3122:The Sibley Guide to Bird Life & Behaviour
2243:
2994:. Johannesburg, South Africa: Jacana Media.
2964:
2398:
1646:
1167:
983:
981:
979:
572:which were prevented from incubating a full
3354:
2822:
2623:
2542:
2431:
2429:
1904:
1771:
1718:
1712:
2479:
2477:
2475:
2394:
2392:
2390:
1508:
1047:Annals of the New York Academy of Sciences
100:Over time, some elements of preening have
3209:Peterson Reference Guide to Bird Behavior
2912:
2847:
2805:
2795:
2745:
2673:
2358:
2304:
2121:
2115:
1995:
1993:
1991:
1930:
1905:Bush, Sarah E.; Clayton, Dale H. (2018).
1881:
1732:
1676:
1542:
1532:
1461:
1299:
1226:
976:
861:Moon-Fandli, Dodman & O'Sullivan 1999
838:. Oxford University Press. Archived from
389:branching from the main shaft and hooked
141:. It appears to be a variant of the word
3485:, and does not reflect subsequent edits.
3468:
3289:
3247:
3029:
3012:Essentials of Avian Medicine and Surgery
2705:
2611:
2435:
2426:
2328:
2326:
2324:
2286:
2280:
2200:
2194:
1595:"The function of powder downs in herons"
1577:
1182:"Uropygial gland size and avian habitat"
1173:
1089:
872:
743:
707:
618:
602:
506:
400:
376:
239:
167:
25:
3375:
3355:Olsen, Penny & Joseph, Leo (2011).
3205:
3161:
3105:. Oxford, UK: Oxford University Press.
3063:. Oxford, UK: Oxford University Press.
2987:
2872:
2472:
2387:
1982:
1953:
1849:
1813:
1807:
1706:
1589:
1583:
1565:
1398:
589:will break off their battles to preen.
3518:
3413:
3394:
3361:. Collingwood, VIC, Australia: CSIRO.
3296:. Ames, IA, US: Blackwell Publishing.
3248:Loon, Rael & Loon, HĂ©lĂšne (2005).
3226:
2935:
2762:
2599:
2587:
2381:
1988:
1037:
1031:
884:
502:
409:is used as a comb for facial preening.
3008:
2947:
2641:
2321:
2056:
1332:
878:
739:
548:) or distinctive colour (such as the
3333:
3233:. Hoboken, NJ, US: Wiley Blackwell.
3185:. London: George Allan & Unwin.
3138:
3082:. New York, NY: Simon and Schuster.
2293:Canadian Medical Association Journal
1898:
1774:"Head-scratching movements in birds"
1765:
1652:
1199:
1144:
970:
919:
712:Allopreening can reduce or redirect
607:Allopreening may help to strengthen
405:The specialized pectinate claw of a
30:When preening, a bird (such as this
3378:The Princeton Encyclopedia of Birds
3254:. Cape Town, South Africa: Struik.
3164:Ornithology in Laboratory and Field
3147:(3rd ed.). New York: Freeman.
2836:Transboundary and Emerging Diseases
2494:Behavioral Ecology and Sociobiology
2237:
2065:Behavioral Ecology and Sociobiology
1814:Goodwin, Robert E. (October 1959).
887:Journal of Comparative Physiology A
631:, may solicit preening from humans.
108:preening has become a part of some
13:
3455:
3397:Handbook of the Birds of the World
3039:Handbook of the Birds of the World
2536:
1907:"Anti-parasite behaviour of birds"
1067:10.1111/j.1749-6632.1988.tb38594.x
824:
372:
73:that have become separated, clean
14:
3547:
3436:
3290:Luescher, Andrew U., ed. (2006).
2973:. Carlton, UK: T and A D Poyser.
145:; one now-obsolete definition of
91:a gland at the base of their tail
3467:
3376:Perrins, Christopher M. (2009).
2051:Ehrlich, Dobkin & Wheye 1988
1695:Ehrlich, Dobkin & Wheye 1988
1635:Ehrlich, Dobkin & Wheye 1988
836:Oxford English Living Dictionary
3417:Sociobiology: The New Synthesis
3230:Handbook of Exotic Pet Medicine
2287:Mitchell, J. C. (18 May 1968).
2124:The Quarterly Review of Biology
1215:Journal of Experimental Biology
592:
564:Preening may be performed as a
2957:
935:Lovette & Fitzpatrick 2016
716:within a flock, as with these
16:Maintenance behaviour of birds
1:
3009:Coles, Brian H., ed. (2007).
2165:Huxley, Hardy & Ford 1954
817:
756:If birds are exposed to some
163:
149:meant "anoint", based on the
104:to have secondary functions.
89:, they gather preen oil from
3337:Understanding Bird Behaviour
3124:. London: Christopher Helm.
2797:10.1371/journal.pone.0011315
2718:Emerging Infectious Diseases
1678:10.1016/j.matdes.2019.108410
1486:10.1016/j.beproc.2012.08.001
417:on the surface, with softer
415:vaned or pennaceous feathers
235:
128:
112:, for example. It is also a
7:
3414:Wilson, Edward O. (2000) .
3227:Kubiak, Marie, ed. (2021).
2636:Deeming & Reynolds 2015
10:
3552:
3101:The Birdwatcher's Handbook
3030:de Juana, Eduardo (1992).
2991:Beat about the Bush: Birds
1038:Delius, J. D. (May 1988).
959:Elphick & Dunning 2001
947:Elphick & Dunning 2001
596:
18:
3293:Manual of Parrot Behavior
2514:10.1007/s00265-006-0253-6
2085:10.1007/s00265-011-1153-y
1534:10.1186/s13071-016-1512-7
1431:10.1007/s00114-012-1005-2
1368:10.1007/s00114-014-1201-3
1292:10.1007/s00248-015-0636-1
3272:Handbook of Bird Biology
2988:Carnaby, Trevor (2008).
1647:Campbell & Lack 1985
1168:Campbell & Lack 1985
808:heavy metal intoxication
511:During courtship, drake
293:did this by releasing a
259:preen or uropygial gland
3251:Birds: The Inside Story
3139:Gill, Frank B. (2007).
2624:Olsen & Joseph 2011
2450:10.1163/156853965x00011
2258:10.1163/156853958x00037
2215:10.1163/156853953X00041
2002:The American Naturalist
1521:Parasites & Vectors
1189:Ornitologia Neotropical
1009:10.1163/156853994X00424
748:Oiled birds, like this
611:in species such as the
3463:
3443:Listen to this article
3340:. London: Bloomsbury.
3334:Moss, Stephen (2015).
3206:Kricher, John (2020).
3183:Evolution as a Process
2897:10.1098/rsbl.2003.0147
2738:10.3201/eid1005.030825
2666:10.1098/rsbl.2007.0258
1923:10.1098/rstb.2017.0196
1874:10.1098/rspb.2004.3036
1665:Materials & Design
792:can be transmitted to
753:
721:
632:
616:
516:
410:
398:
286:Bacillus licheniformis
253:
186:
43:
3462:
3080:The Birder's Handbook
2969:A Dictionary of Birds
2488:Phoeniculus purpureus
2351:10.1093/beheco/arx078
1474:Behavioural Processes
747:
711:
622:
606:
597:Further information:
566:displacement activity
510:
491:, sunning, oiling or
404:
380:
291:Enterococcus faecalis
280:Enterococcus faecalis
243:
180:
114:displacement activity
53:maintenance behaviour
29:
3494:More spoken articles
3033:"Class Aves (Birds)"
2612:Loon & Loon 2005
657:
649:
641:
549:
422:
390:
382:
246:two-barred crossbill
133:The use of the word
66:
65:, interlock feather
50:
35:
3502:Barred owl preening
2891:(Suppl 4): S202-5.
2788:2010PLoSO...511315D
2730:2004EIDis..10..907D
2700:Ehrlich et al. 1994
2506:2006BEcoS..61..221R
2189:Ehrlich et al. 1994
2177:Ehrlich et al. 1994
2077:2011BEcoS..65.1425P
1966:Ehrlich et al. 1994
1649:, pp. 102â103.
1423:2013NW....100..145C
1411:Naturwissenschaften
1360:2014NW....101..697S
1348:Naturwissenschaften
1284:2015MicEc..70.1024M
1111:1993Ecol...74..390W
1059:1988NYASA.525...40D
629:black-capped conure
522:great white pelican
503:Secondary functions
463:that are primarily
139:Late Middle English
3464:
2849:10.1111/tbed.13148
2654:Biological Letters
2339:Behavioral Ecology
1917:(1751): 20170196.
1721:American Zoologist
1591:Wetmore, Alexander
1228:10.1242/jeb.031336
899:10.1007/BF00610731
760:, such as leaking
754:
740:Potential problems
722:
718:great woodswallows
703:Green wood hoopoes
633:
617:
587:European starlings
570:black-headed gulls
538:courtship displays
517:
461:New World warblers
411:
399:
314:may help to repel
254:
187:
110:courtship displays
83:aerodynamic flight
44:
3460:
3427:978-0-674-00089-6
3406:978-84-87334-22-1
3387:978-0-691-14070-4
3368:978-0-643-09493-2
3347:978-1-4729-1206-0
3326:978-0-88048-759-7
3303:978-0-8138-2749-0
3282:978-1-118-29105-4
3261:978-1-77007-151-3
3240:978-1-119-38994-1
3219:978-1-328-78736-1
3173:978-0-12-552455-1
3154:978-0-7167-4983-7
3112:978-0-19-858407-0
3089:978-0-671-62133-9
3070:978-0-19-871866-6
3049:978-84-87334-10-8
3022:978-1-4051-5755-1
3001:978-1-77009-241-9
2980:978-0-85661-039-4
2008:(S1): S145âS158.
1868:(1565): 811â817.
1272:Microbial Ecology
1221:(22): 3621â3626.
842:on August 6, 2017
790:louping ill virus
733:common guillemots
695:Macaroni penguins
520:preen oil of the
348:birds, including
244:Birds, like this
178:
21:Personal grooming
3543:
3484:
3482:
3471:
3470:
3461:
3451:
3449:
3444:
3431:
3410:
3391:
3372:
3351:
3330:
3317:Trichotillomania
3307:
3286:
3265:
3244:
3223:
3202:
3177:
3158:
3146:
3135:
3116:
3104:
3093:
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2984:
2972:
2951:
2945:
2939:
2933:
2927:
2926:
2916:
2876:
2870:
2869:
2851:
2842:(3): 1306â1313.
2826:
2820:
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2809:
2799:
2766:
2760:
2759:
2749:
2709:
2703:
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2345:(4): 1142â1148.
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2071:(7): 1425â1435.
2060:
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1997:
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1278:(4): 1024â1033.
1263:
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1230:
1212:
1203:
1197:
1196:
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923:
917:
911:
910:
882:
876:
870:
864:
858:
852:
851:
849:
847:
828:
812:feather plucking
678:crested caracara
661:
653:
645:
613:Laysan albatross
553:
534:Bohemian waxwing
426:
394:
386:
275:Eurasian hoopoes
211:while preening.
200:Mechanoreceptors
196:contour feathers
179:
70:
54:
39:
3551:
3550:
3546:
3545:
3544:
3542:
3541:
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3516:
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3486:
3480:
3478:
3475:This audio file
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2564:10.2307/1367047
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2397:
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2299:(20): 962â964.
2285:
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1835:10.2307/4082320
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1793:10.2307/4081811
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1734:10.1.1.559.1172
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1614:10.2307/1362391
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1119:10.2307/1939301
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883:
879:
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867:
859:
855:
845:
843:
830:
829:
825:
820:
786:avian influenza
778:West Nile virus
766:heat regulation
742:
663:
655:
647:
627:, such as this
601:
599:Social grooming
595:
556:) on the bird.
555:
505:
428:
396:
388:
375:
373:Preening action
250:uropygial gland
238:
168:
166:
131:
95:contour feather
72:
56:
41:
24:
17:
12:
11:
5:
3549:
3539:
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3528:
3512:
3511:
3505:
3487:
3473:
3466:
3454:
3441:
3440:
3438:
3437:External links
3435:
3433:
3432:
3426:
3411:
3405:
3392:
3386:
3373:
3367:
3352:
3346:
3331:
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3308:
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3287:
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3266:
3260:
3245:
3239:
3224:
3218:
3203:
3178:
3172:
3159:
3153:
3136:
3131:978-0713662504
3130:
3117:
3111:
3094:
3088:
3075:
3069:
3054:
3048:
3027:
3021:
3006:
3000:
2985:
2979:
2961:
2959:
2956:
2953:
2952:
2940:
2938:, p. 175.
2928:
2871:
2821:
2761:
2724:(5): 907â909.
2704:
2702:, p. 225.
2689:
2660:(4): 386â389.
2640:
2628:
2626:, p. 249.
2616:
2604:
2602:, p. 257.
2592:
2590:, p. 209.
2577:
2558:(2): 214â215.
2535:
2500:(2): 221â230.
2471:
2444:(3): 161â208.
2425:
2414:(3): 525â531.
2386:
2384:, p. 208.
2374:
2320:
2279:
2236:
2193:
2181:
2169:
2167:, p. 242.
2157:
2136:10.1086/398642
2114:
2055:
2043:
2014:10.1086/510095
1987:
1985:, p. 118.
1970:
1968:, p. 357.
1958:
1956:, p. 358.
1946:
1897:
1848:
1829:(4): 521â523.
1806:
1787:(3): 339â342.
1764:
1727:(4): 461â477.
1711:
1709:, p. 214.
1699:
1697:, p. 543.
1684:
1651:
1639:
1637:, p. 311.
1627:
1608:(5): 168â170.
1582:
1570:
1558:
1507:
1460:
1417:(2): 145â151.
1397:
1354:(9): 697â705.
1331:
1258:
1198:
1172:
1170:, p. 102.
1149:
1147:, p. 102.
1132:
1105:(2): 390â404.
1088:
1030:
1003:(3): 171â187.
975:
973:, p. 104.
963:
951:
939:
937:, p. 129.
924:
912:
893:(3): 391â394.
877:
875:, p. 198.
865:
853:
822:
821:
819:
816:
741:
738:
594:
591:
530:great hornbill
513:mandarin ducks
504:
501:
374:
371:
307:house sparrows
237:
234:
165:
162:
130:
127:
15:
9:
6:
4:
3:
2:
3548:
3537:
3534:
3532:
3529:
3527:
3526:Bird behavior
3524:
3523:
3521:
3514:
3509:
3506:
3503:
3500:
3499:
3495:
3491:
3476:
3429:
3423:
3419:
3418:
3412:
3408:
3402:
3398:
3393:
3389:
3383:
3379:
3374:
3370:
3364:
3360:
3359:
3353:
3349:
3343:
3339:
3338:
3332:
3328:
3322:
3318:
3314:
3309:
3305:
3299:
3295:
3294:
3288:
3284:
3278:
3274:
3273:
3267:
3263:
3257:
3253:
3252:
3246:
3242:
3236:
3232:
3231:
3225:
3221:
3215:
3211:
3210:
3204:
3200:
3196:
3192:
3188:
3184:
3179:
3175:
3169:
3165:
3160:
3156:
3150:
3145:
3144:
3137:
3133:
3127:
3123:
3118:
3114:
3108:
3103:
3102:
3095:
3091:
3085:
3081:
3076:
3072:
3066:
3062:
3061:
3055:
3051:
3045:
3041:
3040:
3034:
3028:
3024:
3018:
3014:
3013:
3007:
3003:
2997:
2993:
2992:
2986:
2982:
2976:
2971:
2970:
2963:
2962:
2950:, p. 46.
2949:
2944:
2937:
2932:
2924:
2920:
2915:
2910:
2906:
2902:
2898:
2894:
2890:
2886:
2882:
2875:
2867:
2863:
2859:
2855:
2850:
2845:
2841:
2837:
2833:
2825:
2817:
2813:
2808:
2803:
2798:
2793:
2789:
2785:
2782:(6): e11315.
2781:
2777:
2773:
2765:
2757:
2753:
2748:
2743:
2739:
2735:
2731:
2727:
2723:
2719:
2715:
2708:
2701:
2696:
2694:
2685:
2681:
2676:
2671:
2667:
2663:
2659:
2655:
2651:
2644:
2638:, p. 94.
2637:
2632:
2625:
2620:
2614:, p. 36.
2613:
2608:
2601:
2596:
2589:
2584:
2582:
2573:
2569:
2565:
2561:
2557:
2553:
2546:
2539:
2531:
2527:
2523:
2519:
2515:
2511:
2507:
2503:
2499:
2495:
2491:
2489:
2480:
2478:
2476:
2467:
2463:
2459:
2455:
2451:
2447:
2443:
2439:
2432:
2430:
2421:
2417:
2413:
2409:
2402:
2395:
2393:
2391:
2383:
2378:
2370:
2366:
2361:
2356:
2352:
2348:
2344:
2340:
2336:
2329:
2327:
2325:
2316:
2312:
2307:
2302:
2298:
2294:
2290:
2283:
2275:
2271:
2267:
2263:
2259:
2255:
2251:
2247:
2240:
2232:
2228:
2224:
2220:
2216:
2212:
2208:
2204:
2197:
2191:, p. 37.
2190:
2185:
2179:, p. 49.
2178:
2173:
2166:
2161:
2153:
2149:
2145:
2141:
2137:
2133:
2129:
2125:
2118:
2110:
2106:
2102:
2098:
2094:
2090:
2086:
2082:
2078:
2074:
2070:
2066:
2059:
2053:, p. 58.
2052:
2047:
2039:
2035:
2031:
2027:
2023:
2019:
2015:
2011:
2007:
2003:
1996:
1994:
1992:
1984:
1979:
1977:
1975:
1967:
1962:
1955:
1950:
1942:
1938:
1933:
1928:
1924:
1920:
1916:
1912:
1908:
1901:
1893:
1889:
1884:
1879:
1875:
1871:
1867:
1863:
1859:
1852:
1844:
1840:
1836:
1832:
1828:
1824:
1817:
1810:
1802:
1798:
1794:
1790:
1786:
1782:
1775:
1768:
1760:
1756:
1752:
1748:
1744:
1740:
1735:
1730:
1726:
1722:
1715:
1708:
1703:
1696:
1691:
1689:
1679:
1674:
1670:
1666:
1662:
1655:
1648:
1643:
1636:
1631:
1623:
1619:
1615:
1611:
1607:
1603:
1596:
1592:
1586:
1580:, p. 40.
1579:
1578:de Juana 1992
1574:
1568:, p. 37.
1567:
1562:
1554:
1550:
1545:
1540:
1535:
1530:
1526:
1522:
1518:
1511:
1503:
1499:
1495:
1491:
1487:
1483:
1479:
1475:
1471:
1464:
1456:
1452:
1448:
1444:
1440:
1436:
1432:
1428:
1424:
1420:
1416:
1412:
1408:
1401:
1393:
1389:
1385:
1381:
1377:
1373:
1369:
1365:
1361:
1357:
1353:
1349:
1342:
1335:
1327:
1323:
1319:
1315:
1311:
1307:
1302:
1297:
1293:
1289:
1285:
1281:
1277:
1273:
1269:
1262:
1254:
1250:
1246:
1242:
1238:
1234:
1229:
1224:
1220:
1216:
1209:
1202:
1194:
1190:
1183:
1176:
1169:
1164:
1162:
1160:
1158:
1156:
1154:
1146:
1141:
1139:
1137:
1128:
1124:
1120:
1116:
1112:
1108:
1104:
1100:
1092:
1084:
1080:
1076:
1072:
1068:
1064:
1060:
1056:
1052:
1048:
1041:
1034:
1026:
1022:
1018:
1014:
1010:
1006:
1002:
998:
991:
984:
982:
980:
972:
967:
961:, p. 57.
960:
955:
949:, p. 58.
948:
943:
936:
931:
929:
922:, p. 71.
921:
916:
908:
904:
900:
896:
892:
888:
881:
874:
873:Luescher 2006
869:
863:, p. 68.
862:
857:
841:
837:
833:
827:
823:
815:
813:
809:
805:
801:
797:
795:
791:
787:
783:
779:
774:
772:
767:
763:
759:
751:
746:
737:
734:
729:
727:
719:
715:
710:
706:
704:
700:
696:
692:
688:
687:ectoparasites
682:
679:
675:
674:black vulture
671:
665:
662:
654:
646:
639:
630:
626:
621:
614:
610:
605:
600:
590:
588:
584:
580:
575:
571:
567:
562:
559:
554:
547:
546:mandarin duck
544:of the drake
543:
539:
535:
531:
527:
523:
514:
509:
500:
498:
494:
490:
485:
483:
479:
475:
471:
466:
462:
458:
454:
450:
446:
440:
437:
433:
432:ectoparasites
427:
420:
419:down feathers
416:
408:
403:
395:
387:
379:
370:
367:
363:
359:
355:
351:
347:
343:
339:
335:
332:, ostriches,
331:
326:
322:
320:
317:
313:
308:
304:
303:avian malaria
300:
296:
292:
288:
287:
282:
281:
276:
271:
268:
264:
260:
251:
247:
242:
233:
231:
227:
222:
218:
212:
210:
206:
201:
197:
193:
184:
161:
159:
155:
152:
148:
144:
140:
136:
126:
123:
120:Ingestion of
118:
115:
111:
107:
103:
98:
96:
92:
88:
84:
80:
79:ectoparasites
76:
71:
64:
60:
55:
48:
40:
33:
28:
22:
3513:
3416:
3396:
3377:
3357:
3336:
3316:
3292:
3271:
3250:
3229:
3208:
3182:
3163:
3142:
3121:
3100:
3079:
3059:
3037:
3011:
2990:
2968:
2943:
2931:
2888:
2884:
2874:
2839:
2835:
2824:
2779:
2775:
2764:
2721:
2717:
2707:
2657:
2653:
2643:
2631:
2619:
2607:
2595:
2555:
2551:
2538:
2497:
2493:
2487:
2441:
2437:
2411:
2407:
2377:
2342:
2338:
2296:
2292:
2282:
2249:
2245:
2239:
2209:(1): 58â80.
2206:
2202:
2196:
2184:
2172:
2160:
2127:
2123:
2117:
2093:10261/143995
2068:
2064:
2058:
2046:
2005:
2001:
1983:Kricher 2020
1961:
1954:Carnaby 2008
1949:
1914:
1910:
1900:
1865:
1861:
1851:
1826:
1822:
1809:
1784:
1780:
1767:
1724:
1720:
1714:
1707:Hailman 1985
1702:
1668:
1664:
1654:
1642:
1630:
1605:
1601:
1585:
1573:
1566:Perrins 2009
1561:
1524:
1520:
1510:
1477:
1473:
1463:
1414:
1410:
1400:
1351:
1347:
1334:
1301:10261/123119
1275:
1271:
1261:
1218:
1214:
1201:
1192:
1188:
1175:
1102:
1098:
1091:
1053:(1): 40â55.
1050:
1046:
1033:
1000:
996:
966:
954:
942:
915:
890:
886:
880:
868:
856:
844:. Retrieved
840:the original
835:
826:
798:
775:
755:
730:
723:
683:
666:
634:
593:Allopreening
583:common terns
563:
518:
486:
482:feather lice
476:, terns and
453:frigatebirds
441:
412:
327:
323:
312:wood hoopoes
290:
284:
278:
272:
255:
221:common loons
213:
209:pin feathers
188:
157:
153:
146:
142:
134:
132:
119:
99:
46:
45:
3143:Ornithology
2958:Works cited
2936:Kubiak 2021
2600:Rowley 1997
2588:Wilson 2000
2382:Wilson 2000
2252:(1): 1â87.
2130:(1): 1â32.
814:or injury.
800:Caged birds
750:surf scoter
676:and a wild
625:pet parrots
542:secondaries
526:Ross's gull
478:albatrosses
457:pratincoles
455:, owls and
366:powder down
354:woodpeckers
344:) and some
334:cassowaries
295:bacteriocin
77:, and keep
3520:Categories
3510:on YouTube
3504:on YouTube
3490:Audio help
3481:2023-07-18
2948:Coles 2007
2552:The Condor
1671:: 108410.
1602:The Condor
1195:: 297â306.
818:References
804:neuropathy
794:red grouse
758:pollutants
726:pair bonds
714:aggression
609:pair bonds
497:blackbirds
299:incubation
192:filoplumes
164:Importance
122:pollutants
106:Ritualised
2905:0962-8452
2458:1568-539X
2438:Behaviour
2266:1568-539X
2246:Behaviour
2223:0005-7959
2203:Behaviour
2101:0340-5443
2022:0003-0147
1759:198156620
1751:0003-1569
1729:CiteSeerX
1439:1432-1904
1376:0028-1042
1310:1432-184X
1237:0022-0949
1145:Gill 2007
1017:0005-7959
997:Behaviour
971:Gill 2007
920:Moss 2015
771:pneumonia
762:petroleum
445:nightjars
319:predators
316:mammalian
236:Preen oil
230:ostriches
129:Etymology
57:found in
3531:Feathers
3492: ·
3191:54001781
2923:15252984
2866:76664136
2858:30861310
2816:20593026
2776:PLOS ONE
2756:15200828
2684:17550875
2530:43724298
2522:25511576
2369:29622926
2152:31957387
2144:14930222
2109:38405658
2038:29592388
2030:19426089
1941:29866911
1892:15888414
1593:(1920).
1553:27114098
1502:21076057
1494:22940115
1455:15209444
1447:23288399
1384:25011415
1318:26078039
1245:19880722
1083:17744188
846:6 August
691:flocking
670:icterids
638:families
579:Sandwich
558:Mallards
552:speculum
474:swallows
465:arboreal
436:Penguins
407:barn owl
393:barbules
356:, a few
350:bustards
346:neognath
69:barbules
63:feathers
47:Preening
38:barbules
32:red lory
3536:Hygiene
3479: (
3450:minutes
3199:1434718
2914:1810039
2807:2892510
2784:Bibcode
2747:3323200
2726:Bibcode
2675:2390679
2572:1367047
2502:Bibcode
2466:4533105
2420:4085549
2408:The Auk
2360:5873249
2315:5657176
2306:1924139
2274:4532894
2231:4532768
2073:Bibcode
1932:6000146
1883:1599863
1843:4082320
1823:The Auk
1801:4081811
1781:The Auk
1622:1362391
1544:4845389
1527:: 232.
1480:: 1â5.
1419:Bibcode
1392:4278863
1356:Bibcode
1326:8342661
1280:Bibcode
1253:9884724
1127:1939301
1107:Bibcode
1099:Ecology
1075:2839072
1055:Bibcode
1025:4535237
907:8499915
832:"Preen"
489:dusting
362:pigeons
358:parrots
226:ratites
183:mallard
102:evolved
75:plumage
3424:
3403:
3384:
3365:
3344:
3323:
3300:
3279:
3258:
3237:
3216:
3197:
3189:
3170:
3151:
3128:
3109:
3086:
3067:
3046:
3019:
2998:
2977:
2921:
2911:
2903:
2864:
2856:
2814:
2804:
2754:
2744:
2682:
2672:
2570:
2528:
2520:
2464:
2456:
2418:
2367:
2357:
2313:
2303:
2272:
2264:
2229:
2221:
2150:
2142:
2107:
2099:
2036:
2028:
2020:
1939:
1929:
1890:
1880:
1841:
1799:
1757:
1749:
1731:
1620:
1551:
1541:
1500:
1492:
1453:
1445:
1437:
1390:
1382:
1374:
1324:
1316:
1308:
1251:
1243:
1235:
1125:
1081:
1073:
1023:
1015:
905:
782:corvid
660:flanks
652:mantle
644:breast
574:clutch
493:anting
470:swifts
449:herons
267:grebes
154:ungere
2862:S2CID
2568:JSTOR
2548:(PDF)
2526:S2CID
2518:JSTOR
2462:JSTOR
2416:JSTOR
2404:(PDF)
2270:JSTOR
2227:JSTOR
2148:S2CID
2105:S2CID
2034:S2CID
1839:JSTOR
1819:(PDF)
1797:JSTOR
1777:(PDF)
1755:S2CID
1618:JSTOR
1598:(PDF)
1498:S2CID
1451:S2CID
1388:S2CID
1344:(PDF)
1322:S2CID
1249:S2CID
1211:(PDF)
1185:(PDF)
1123:JSTOR
1079:S2CID
1043:(PDF)
1021:JSTOR
993:(PDF)
903:S2CID
699:ticks
623:Some
425:barbs
385:barbs
342:kiwis
338:rheas
263:terns
205:moult
158:preen
151:Latin
147:prune
143:prune
135:preen
87:beaks
59:birds
49:is a
3422:ISBN
3401:ISBN
3382:ISBN
3363:ISBN
3342:ISBN
3321:ISBN
3298:ISBN
3277:ISBN
3256:ISBN
3235:ISBN
3214:ISBN
3195:OCLC
3187:LCCN
3168:ISBN
3149:ISBN
3126:ISBN
3107:ISBN
3084:ISBN
3065:ISBN
3044:ISBN
3017:ISBN
2996:ISBN
2975:ISBN
2919:PMID
2901:ISSN
2854:PMID
2812:PMID
2752:PMID
2680:PMID
2454:ISSN
2365:PMID
2311:PMID
2262:ISSN
2219:ISSN
2140:PMID
2097:ISSN
2026:PMID
2018:ISSN
1937:PMID
1888:PMID
1747:ISSN
1549:PMID
1490:PMID
1443:PMID
1435:ISSN
1380:PMID
1372:ISSN
1314:PMID
1306:ISSN
1241:PMID
1233:ISSN
1071:PMID
1013:ISSN
848:2017
648:and
581:and
360:and
340:and
217:gull
2909:PMC
2893:doi
2889:271
2844:doi
2802:PMC
2792:doi
2742:PMC
2734:doi
2670:PMC
2662:doi
2560:doi
2510:doi
2446:doi
2355:PMC
2347:doi
2301:PMC
2254:doi
2211:doi
2132:doi
2089:hdl
2081:doi
2010:doi
2006:169
1927:PMC
1919:doi
1915:373
1878:PMC
1870:doi
1866:272
1831:doi
1789:doi
1739:doi
1673:doi
1669:187
1610:doi
1539:PMC
1529:doi
1482:doi
1427:doi
1415:100
1364:doi
1352:101
1296:hdl
1288:doi
1223:doi
1219:212
1115:doi
1063:doi
1051:525
1005:doi
1001:131
895:doi
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