Preening - Knowledge

615: 389: 252: 739:. 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. 720: 38: 631: 519: 747:

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.

413: 756: 312:. 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 180: 441:

activities – 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

3480: 470:) 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, 201:

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.

445:, 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 – a behaviour known as "stropping". 183: 96:, 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 182: 187: 186: 181: 188: 379:

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 – 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 – 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 – contributing to insulation, waterproofing and aerodynamic flight – 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 – including the microscopic structures (the barbs and barbules) which interlock to create the waterproof barrier.

185: 272:, 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 716:, 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. 135:

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.

784:, 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. 478:

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

506:, 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 234:

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

308:. 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 3469: 184: 2840:

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

539:, 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 1277:

Martínez-García, Ángela; Soler, Juan J.; Rodríguez-Ruano, Sonia M.; Martínez-Bueno, Manuel; Martín-Platero, Antonio Manuel; Juárez-García, Natalia & Martín-Vivaldi, Manuel (November 2015).

795:, 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 1050: 2074:

Pérez-Rodríguez, Lorenzo; Mougeot, Francois & Bortolotti, Gary R. (2011-07-01). "The effects of preen oils and soiling on the UV–visible reflectance of carotenoid-pigmented feathers".

3468: 491:) 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 799:

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

821:) 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 284:

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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Czirják, Gábor Árpád; Pap, Péter László; Vágási, Csongor István; Giraudeau, Mathieu; Mureşan, 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

813:, particularly parrots, sometimes overpreen in response to being exposed to strong scents (such as nicotine or air fresheners) or as a result of 742:

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".

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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".

1528:"Volume and antimicrobial activity of secretions of the uropygial gland are correlated with malaria infection in house sparrows" 842: 2843:"Survivability of highly pathogenic avian influenza virus (H5N1) in naturally preened duck feathers at different temperatures" 817:. Reducing exposure to the offending odour, or treating the underlying cause of the neuropathy (such as injury, infection, or 700:

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-Rodríguez, M.; Valdivia, E.; Soler, Juan J.; Martín-Vivaldi, M.; Martín-Platero, A. M. & Martínez-Bueno, M. (2009).

579:. In some cases, it is done in place of another activity that birds are strongly motivated, but unable, to do. In one study, 535:

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,

3140: 3049: 2346:"Allopreening in birds is associated with parental cooperation over offspring care and stable pair bonds across years" 2300:"Dermatological aspects of displacement activity: attention to the body surface as a substitute for "fight or flight"" 62: 3152: 1479:

Giraudeau, M.; Czirják, G.Á.; Duval, C.; Bretagnolle, V.; Gutierrez, C.; Guillon, N. & Heeb, P. (January 2013).

239:, which are not dependent on their feathers for flight, spend far less time on maintenance behaviours. One study of 2555: 1605: 3406:

Rowley, Ian (1997). "Family Cacatuidae (Cockatoos)". In del Hoyo, Josep; Elliott, Andrew; Sargatal, Jordi (eds.).

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Magallanes, Sergio; Pape Møller, Anders; García-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).

561: 402: 78: 47: 1672:"Shaking the wings and preening feathers with the beak help a bird to recover its ruffled feather vane" 1350:

Soler, Juan J.; Martín-Vivaldi, M.; Peralta-Sánchez, J. M.; Arco, L.; Juárez-García-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

434: 394: 167:, which had the same meaning. This usage was combined with the Scottish and northern English dialect 2659:

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

675:. A few species are known to allopreen other areas, including the rump, tail, belly and underwing. 230:

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 birds—an overview".

1418:"Preen gland removal increases plumage bacterial load but not that of feather-degrading bacteria" 2723:

Docherty, Douglas E.; Long, Renee Romaine; Griffin, Kathryn M. & Saito, Emi K. (June 2004).

1827:"Records of flight preening and related aerial activities in birds, particularly the black tern" 1218: 787:

Allopreening may facilitate disease transmission between infected and non-infected individuals.

3536: 1739: 503: 296: 3426: 3367: 3281: 3239: 3218: 3000: 495:; the removal of the bill tip caused an increase in feather lice due to ineffective preening. 3346: 3323: 3260: 3069: 3021: 576: 290: 124: 630: 498:

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.

3500: 2794: 2736: 2512: 2083: 1429: 1366: 1290: 1117: 1065: 552: 256: 8: 724: 639: 532: 149: 2798: 2740: 2516: 2087: 1433: 1370: 1294: 1121: 1069: 3043: 2978: 2924: 2891: 2872: 2817: 2782: 2757: 2724: 2685: 2660: 2578: 2536: 2528: 2472: 2426: 2370: 2345: 2316: 2299: 2280: 2237: 2158: 2115: 2044: 1942: 1917: 1893: 1868: 1849: 1807: 1765: 1628: 1554: 1527: 1508: 1461: 1398: 1332: 1259: 1219:"Symbiotic bacteria living in the hoopoe's uropygial gland prevent feather degradation" 1133: 1089: 1077: 1031: 913: 196:

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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Ehrlich, Paul R.; Dobkin, David S.; Wheye, Darryl & Pimms, Stuart L. (1994).

2807: 1688: 1671: 1496: 1276: 796: 788: 775:, they can quickly lose the preen oil from their feathers. This causes a loss of 609: 388: 285: 269: 260: 210: 206: 105: 101: 45:) draws individual feathers through its beak, realigning and re-interlocking the 3324:"Veterinary Models of Compulsive Self-grooming: Parallels with Trichotillamania" 3322:

Moon-Fandli, Alice M.; Dodman, Nicholas A. & O'Sullivan, Richard L. (1999).

1481:"Effect of preen oil on plumage bacteria: an experimental test with the mallard" 2497:"Dual function of allopreening in the cooperatively breeding green woodhoopoe, 540: 536: 309: 3518: 3512: 3369:

Stray Feathers: Reflections on the Structure, Behaviour and Evolution of Birds

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Kenny, Elspeth; Birkhead, Tim R. & Green, Jonathan P. (July–August 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.

116: 89: 2783:"Can preening contribute to influenza A virus infection in wild waterbirds?" 2460: 2361: 2268: 2225: 1278: 1019: 865: 475: 339:

While most species have a preen gland, the organ is missing in the ratites (

294:) in their preen oil inhibited the growth of the feather-degrading bacteria 3209: 2933: 2907: 2868: 2826: 2766: 2748: 2694: 2676: 2379: 2154: 2040: 1951: 1933: 1902: 1884: 1563: 1525: 1504: 1457: 1394: 1328: 1255: 697: 499: 492: 480: 442: 429: 2325: 1085: 719: 412: 37: 2706: 2704: 1869:"Adaptive significance of avian beak morphology for ectoparasite control" 1216: 760: 593: 463: 376: 352: 348: 322: 305: 231: 219: 93: 3488:

was created from a revision of this article dated 18 July 2023

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are also cosmetically coloured during preening. The preen oil of the

488: 344: 202: 132: 3047:. In del Hoyo, Josep; Elliott, Andrew & Sargatal, Jordi (eds.). 2574: 1845: 1803: 1701: 1699: 1624: 1129: 518: 435: 395: 2146: 2024: 1480: 941: 939: 810: 455: 417: 356: 329: 42: 3053:. Vol. 1: Ostrich to Ducks. Barcelona, Spain: Lynx Edicions. 1352:"Hoopoes color their eggs with antimicrobial uropygial secretions" 432:

underneath. Both feather types have a central shaft with narrower

2839: 1696: 1639: 683:, though at least one instance of mutual grooming between a wild 680: 568: 507: 484: 446: 424:

A bird's plumage is primarily made up of two feather types: firm

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Radford, Andrew N. & Du Plessis, Morné A. (December 2006).

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and distribute this oil through their feathers. They draw each

2169: 1174: 1172: 1170: 1168: 1166: 1164: 259:, typically use their beak to gather preen gland oil from the 3192:

Huxley, Julian; Hardy, A. C. & Ford, E. B., eds. (1954).

2779: 2640: 459: 277: 215: 161: 1106: 209:, help to signal when contour feathers have been displaced. 3068:

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has been found in the feather pulp of several species of

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ritually preen their distinctive orange "sail" feathers.

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Proceedings of the Royal Society B: Biological Sciences

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3026:(3rd ed.). Oxford, UK: Blackwood Publishing Ltd. 2616: 2010: 1982: 1958: 1711: 1582: 1151: 1149: 1147: 877: 218:, birds remove the sheaths from around their emerging 2940: 2725:"Corvidae feather pulp and West Nile virus detection" 2604: 2589: 2410:

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that frequently allopreened had significantly fewer

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to mean the tidying of a bird's feathers dates from

3286:(3rd ed.). Oxford, UK: John Wiley & Sons. 2952: 1279:"Preening as a vehicle for key bacteria in hoopoes" 1144: 1051:"Preening and associated comfort behavior in birds" 3279: 3151: 3109: 3088: 3042: 2977: 2343: 2061: 1705: 1645: 975: 945: 807:if the bird consumes a tick carrying the disease. 712:on their heads and necks than those that did not. 108:through their bill, nibbling it from base to tip. 3391:. Princeton, NJ, US: Princeton University Press. 3177:(5th ed.). Orlando, FL, US: Academic Press. 2494: 924: 392:A shaft runs down the middle of the feather with 3528: 2554:Ng, David & Jasperson, Bruce D. (May 1984). 1191:Montalti, Diego & Salibián, Alfredo (2000). 1190: 999:Cotgreave, Peter & Clayton, Dale H. (1994). 998: 288:showed that the presence of symbiotic bacteria ( 3431:. Cambridge, MA, US: Harvard University Press. 3191: 3067: 2646: 2412:"Allopreening in owls: what are its functions?" 2175: 1270: 380:down can give a metallic sheen to the plumage. 3519:Splendid fairy-wrens preening and allopreening 3330:. Washington, DC: American Psychiatric Press. 3130: 1783:Nice, Margaret M. & Schantz, W.E. (1959). 969: 957: 667:and an even smaller percentage applied to the 72:that involves the use of the beak to position 3223:. Boston, MA, US: Houghton Mifflin Harcourt. 3133:The Sibley Guide to Bird Life & Behaviour 2254: 3005:. Johannesburg, South Africa: Jacana Media. 2975: 2409: 1657: 1178: 994: 992: 990: 583:which were prevented from incubating a full 3365: 2833: 2634: 2553: 2442: 2440: 1915: 1782: 1729: 1723: 2490: 2488: 2486: 2405: 2403: 2401: 1519: 1058:Annals of the New York Academy of Sciences 111:Over time, some elements of preening have 3220:Peterson Reference Guide to Bird Behavior 2923: 2858: 2816: 2806: 2756: 2684: 2369: 2315: 2132: 2126: 2006: 2004: 2002: 1941: 1916:Bush, Sarah E.; Clayton, Dale H. (2018). 1892: 1743: 1687: 1553: 1543: 1472: 1310: 1237: 987: 872:Moon-Fandli, Dodman & O'Sullivan 1999 849:. Oxford University Press. Archived from 400:branching from the main shaft and hooked 152:. It appears to be a variant of the word 3496:, and does not reflect subsequent edits. 3479: 3300: 3258: 3040: 3023:Essentials of Avian Medicine and Surgery 2716: 2622: 2446: 2437: 2339: 2337: 2335: 2297: 2291: 2211: 2205: 1606:"The function of powder downs in herons" 1588: 1193:"Uropygial gland size and avian habitat" 1184: 1100: 883: 754: 718: 629: 613: 517: 411: 387: 250: 178: 36: 3386: 3366:Olsen, Penny & Joseph, Leo (2011). 3216: 3172: 3116:. Oxford, UK: Oxford University Press. 3074:. Oxford, UK: Oxford University Press. 2998: 2883: 2483: 2398: 1993: 1964: 1860: 1824: 1818: 1717: 1600: 1594: 1576: 1409: 600:will break off their battles to preen. 14: 3529: 3424: 3405: 3372:. Collingwood, VIC, Australia: CSIRO. 3307:. Ames, IA, US: Blackwell Publishing. 3259:Loon, Rael & Loon, Hélène (2005). 3237: 2946: 2773: 2610: 2598: 2392: 1999: 1048: 1042: 895: 513: 420:is used as a comb for facial preening. 3019: 2958: 2652: 2332: 2067: 1343: 889: 750: 559:) or distinctive colour (such as the 3344: 3244:. Hoboken, NJ, US: Wiley Blackwell. 3196:. London: George Allan & Unwin. 3149: 3093:. New York, NY: Simon and Schuster. 2304:Canadian Medical Association Journal 1909: 1785:"Head-scratching movements in birds" 1776: 1663: 1210: 1155: 981: 930: 723:Allopreening can reduce or redirect 618:Allopreening may help to strengthen 416:The specialized pectinate claw of a 41:When preening, a bird (such as this 3389:The Princeton Encyclopedia of Birds 3265:. Cape Town, South Africa: Struik. 3175:Ornithology in Laboratory and Field 3158:(3rd ed.). New York: Freeman. 2847:Transboundary and Emerging Diseases 2505:Behavioral Ecology and Sociobiology 2248: 2076:Behavioral Ecology and Sociobiology 1825:Goodwin, Robert E. (October 1959). 898:Journal of Comparative Physiology A 642:, may solicit preening from humans. 119:preening has become a part of some 24: 3466: 3408:Handbook of the Birds of the World 3050:Handbook of the Birds of the World 2547: 1918:"Anti-parasite behaviour of birds" 1078:10.1111/j.1749-6632.1988.tb38594.x 835: 383: 84:that have become separated, clean 25: 3558: 3447: 3301:Luescher, Andrew U., ed. (2006). 2984:. Carlton, UK: T and A D Poyser. 156:; one now-obsolete definition of 102:a gland at the base of their tail 3478: 3387:Perrins, Christopher M. (2009). 2062:Ehrlich, Dobkin & Wheye 1988 1706:Ehrlich, Dobkin & Wheye 1988 1646:Ehrlich, Dobkin & Wheye 1988 847:Oxford English Living Dictionary 3428:Sociobiology: The New Synthesis 3241:Handbook of Exotic Pet Medicine 2298:Mitchell, J. C. (18 May 1968). 2135:The Quarterly Review of Biology 1226:Journal of Experimental Biology 603: 575:Preening may be performed as a 2968: 946:Lovette & Fitzpatrick 2016 727:within a flock, as with these 27:Maintenance behaviour of birds 13: 1: 3020:Coles, Brian H., ed. (2007). 2176:Huxley, Hardy & Ford 1954 828: 767:If birds are exposed to some 174: 160:meant "anoint", based on the 115:to have secondary functions. 100:, they gather preen oil from 3348:Understanding Bird Behaviour 3135:. London: Christopher Helm. 2808:10.1371/journal.pone.0011315 2729:Emerging Infectious Diseases 1689:10.1016/j.matdes.2019.108410 1497:10.1016/j.beproc.2012.08.001 428:on the surface, with softer 426:vaned or pennaceous feathers 246: 139: 123:, for example. It is also a 7: 3425:Wilson, Edward O. (2000) . 3238:Kubiak, Marie, ed. (2021). 2647:Deeming & Reynolds 2015 10: 3563: 3112:The Birdwatcher's Handbook 3041:de Juana, Eduardo (1992). 3002:Beat about the Bush: Birds 1049:Delius, J. D. (May 1988). 970:Elphick & Dunning 2001 958:Elphick & Dunning 2001 607: 29: 3304:Manual of Parrot Behavior 2525:10.1007/s00265-006-0253-6 2096:10.1007/s00265-011-1153-y 1545:10.1186/s13071-016-1512-7 1442:10.1007/s00114-012-1005-2 1379:10.1007/s00114-014-1201-3 1303:10.1007/s00248-015-0636-1 3283:Handbook of Bird Biology 2999:Carnaby, Trevor (2008). 1658:Campbell & Lack 1985 1179:Campbell & Lack 1985 819:heavy metal intoxication 522:During courtship, drake 304:did this by releasing a 270:preen or uropygial gland 3262:Birds: The Inside Story 3150:Gill, Frank B. (2007). 2635:Olsen & Joseph 2011 2461:10.1163/156853965x00011 2269:10.1163/156853958x00037 2226:10.1163/156853953X00041 2013:The American Naturalist 1532:Parasites & Vectors 1200:Ornitologia Neotropical 1020:10.1163/156853994X00424 759:Oiled birds, like this 622:in species such as the 3474: 3454:Listen to this article 3351:. London: Bloomsbury. 3345:Moss, Stephen (2015). 3217:Kricher, John (2020). 3194:Evolution as a Process 2908:10.1098/rsbl.2003.0147 2749:10.3201/eid1005.030825 2677:10.1098/rsbl.2007.0258 1934:10.1098/rstb.2017.0196 1885:10.1098/rspb.2004.3036 1676:Materials & Design 803:can be transmitted to 764: 732: 643: 627: 527: 421: 409: 297:Bacillus licheniformis 264: 197: 54: 3473: 3091:The Birder's Handbook 2980:A Dictionary of Birds 2499:Phoeniculus purpureus 2362:10.1093/beheco/arx078 1485:Behavioural Processes 758: 722: 633: 617: 608:Further information: 577:displacement activity 521: 502:, sunning, oiling or 415: 391: 302:Enterococcus faecalis 291:Enterococcus faecalis 254: 191: 125:displacement activity 64:maintenance behaviour 40: 3505:More spoken articles 3044:"Class Aves (Birds)" 2623:Loon & Loon 2005 668: 660: 652: 560: 433: 401: 393: 257:two-barred crossbill 144:The use of the word 77: 76:, interlock feather 61: 46: 3513:Barred owl preening 2902:(Suppl 4): S202-5. 2799:2010PLoSO...511315D 2741:2004EIDis..10..907D 2711:Ehrlich et al. 1994 2517:2006BEcoS..61..221R 2200:Ehrlich et al. 1994 2188:Ehrlich et al. 1994 2088:2011BEcoS..65.1425P 1977:Ehrlich et al. 1994 1660:, pp. 102–103. 1434:2013NW....100..145C 1422:Naturwissenschaften 1371:2014NW....101..697S 1359:Naturwissenschaften 1295:2015MicEc..70.1024M 1122:1993Ecol...74..390W 1070:1988NYASA.525...40D 640:black-capped conure 533:great white pelican 514:Secondary functions 474:that are primarily 150:Late Middle English 3475: 2860:10.1111/tbed.13148 2665:Biological Letters 2350:Behavioral Ecology 1928:(1751): 20170196. 1732:American Zoologist 1602:Wetmore, Alexander 1239:10.1242/jeb.031336 910:10.1007/BF00610731 771:, such as leaking 765: 751:Potential problems 733: 729:great woodswallows 714:Green wood hoopoes 644: 628: 598:European starlings 581:black-headed gulls 549:courtship displays 528: 472:New World warblers 422: 410: 325:may help to repel 265: 198: 121:courtship displays 94:aerodynamic flight 55: 3471: 3438:978-0-674-00089-6 3417:978-84-87334-22-1 3398:978-0-691-14070-4 3379:978-0-643-09493-2 3358:978-1-4729-1206-0 3337:978-0-88048-759-7 3314:978-0-8138-2749-0 3293:978-1-118-29105-4 3272:978-1-77007-151-3 3251:978-1-119-38994-1 3230:978-1-328-78736-1 3184:978-0-12-552455-1 3165:978-0-7167-4983-7 3123:978-0-19-858407-0 3100:978-0-671-62133-9 3081:978-0-19-871866-6 3060:978-84-87334-10-8 3033:978-1-4051-5755-1 3012:978-1-77009-241-9 2991:978-0-85661-039-4 2019:(S1): S145–S158. 1879:(1565): 811–817. 1283:Microbial Ecology 1232:(22): 3621–3626. 853:on August 6, 2017 801:louping ill virus 744:common guillemots 706:Macaroni penguins 531:preen oil of the 359:birds, including 255:Birds, like this 189: 32:Personal grooming 16:(Redirected from 3554: 3495: 3493: 3482: 3481: 3472: 3462: 3460: 3455: 3442: 3421: 3402: 3383: 3362: 3341: 3328:Trichotillomania 3318: 3297: 3276: 3255: 3234: 3213: 3188: 3169: 3157: 3146: 3127: 3115: 3104: 3085: 3064: 3046: 3037: 3016: 2995: 2983: 2962: 2956: 2950: 2944: 2938: 2937: 2927: 2887: 2881: 2880: 2862: 2853:(3): 1306–1313. 2837: 2831: 2830: 2820: 2810: 2777: 2771: 2770: 2760: 2720: 2714: 2708: 2699: 2698: 2688: 2656: 2650: 2644: 2638: 2632: 2626: 2620: 2614: 2608: 2602: 2596: 2587: 2586: 2560: 2551: 2545: 2544: 2492: 2481: 2480: 2444: 2435: 2434: 2416: 2407: 2396: 2390: 2384: 2383: 2373: 2356:(4): 1142–1148. 2341: 2330: 2329: 2319: 2295: 2289: 2288: 2252: 2246: 2245: 2209: 2203: 2197: 2191: 2185: 2179: 2173: 2167: 2166: 2130: 2124: 2123: 2082:(7): 1425–1435. 2071: 2065: 2059: 2053: 2052: 2008: 1997: 1991: 1980: 1974: 1968: 1962: 1956: 1955: 1945: 1913: 1907: 1906: 1896: 1864: 1858: 1857: 1831: 1822: 1816: 1815: 1789: 1780: 1774: 1773: 1747: 1727: 1721: 1715: 1709: 1703: 1694: 1693: 1691: 1667: 1661: 1655: 1649: 1643: 1637: 1636: 1610: 1598: 1592: 1586: 1580: 1574: 1568: 1567: 1557: 1547: 1523: 1517: 1516: 1476: 1470: 1469: 1413: 1407: 1406: 1356: 1347: 1341: 1340: 1314: 1289:(4): 1024–1033. 1274: 1268: 1267: 1241: 1223: 1214: 1208: 1207: 1197: 1188: 1182: 1176: 1159: 1153: 1142: 1141: 1104: 1098: 1097: 1055: 1046: 1040: 1039: 1005: 996: 985: 979: 973: 967: 961: 955: 949: 943: 934: 928: 922: 921: 893: 887: 881: 875: 869: 863: 862: 860: 858: 839: 823:feather plucking 689:crested caracara 672: 664: 656: 624:Laysan albatross 564: 545:Bohemian waxwing 437: 405: 397: 286:Eurasian hoopoes 222:while preening. 211:Mechanoreceptors 207:contour feathers 190: 81: 65: 50: 21: 3562: 3561: 3557: 3556: 3555: 3553: 3552: 3551: 3527: 3526: 3509: 3508: 3497: 3491: 3489: 3486:This audio file 3483: 3476: 3467: 3464: 3458: 3457: 3453: 3450: 3445: 3439: 3418: 3399: 3380: 3359: 3338: 3315: 3294: 3273: 3252: 3231: 3185: 3166: 3143: 3124: 3101: 3082: 3061: 3034: 3013: 2992: 2971: 2966: 2965: 2957: 2953: 2945: 2941: 2888: 2884: 2838: 2834: 2778: 2774: 2721: 2717: 2709: 2702: 2657: 2653: 2645: 2641: 2633: 2629: 2621: 2617: 2609: 2605: 2597: 2590: 2575:10.2307/1367047 2558: 2552: 2548: 2493: 2484: 2445: 2438: 2414: 2408: 2399: 2391: 2387: 2342: 2333: 2310:(20): 962–964. 2296: 2292: 2253: 2249: 2210: 2206: 2198: 2194: 2186: 2182: 2174: 2170: 2131: 2127: 2072: 2068: 2060: 2056: 2009: 2000: 1992: 1983: 1975: 1971: 1963: 1959: 1914: 1910: 1865: 1861: 1846:10.2307/4082320 1829: 1823: 1819: 1804:10.2307/4081811 1787: 1781: 1777: 1745:10.1.1.559.1172 1728: 1724: 1716: 1712: 1704: 1697: 1668: 1664: 1656: 1652: 1644: 1640: 1625:10.2307/1362391 1608: 1599: 1595: 1587: 1583: 1575: 1571: 1524: 1520: 1477: 1473: 1414: 1410: 1354: 1348: 1344: 1275: 1271: 1221: 1215: 1211: 1195: 1189: 1185: 1177: 1162: 1154: 1145: 1130:10.2307/1939301 1105: 1101: 1053: 1047: 1043: 1003: 997: 988: 980: 976: 968: 964: 956: 952: 944: 937: 929: 925: 894: 890: 882: 878: 870: 866: 856: 854: 841: 840: 836: 831: 797:avian influenza 789:West Nile virus 777:heat regulation 753: 674: 666: 658: 638:, such as this 612: 610:Social grooming 606: 567:) on the bird. 566: 516: 439: 407: 399: 386: 384:Preening action 261:uropygial gland 249: 179: 177: 142: 106:contour feather 83: 67: 52: 35: 28: 23: 22: 18:Preening (bird) 15: 12: 11: 5: 3560: 3550: 3549: 3544: 3539: 3523: 3522: 3516: 3498: 3484: 3477: 3465: 3452: 3451: 3449: 3448:External links 3446: 3444: 3443: 3437: 3422: 3416: 3403: 3397: 3384: 3378: 3363: 3357: 3342: 3336: 3319: 3313: 3298: 3292: 3277: 3271: 3256: 3250: 3235: 3229: 3214: 3189: 3183: 3170: 3164: 3147: 3142:978-0713662504 3141: 3128: 3122: 3105: 3099: 3086: 3080: 3065: 3059: 3038: 3032: 3017: 3011: 2996: 2990: 2972: 2970: 2967: 2964: 2963: 2951: 2949:, p. 175. 2939: 2882: 2832: 2772: 2735:(5): 907–909. 2715: 2713:, p. 225. 2700: 2671:(4): 386–389. 2651: 2639: 2637:, p. 249. 2627: 2615: 2613:, p. 257. 2603: 2601:, p. 209. 2588: 2569:(2): 214–215. 2546: 2511:(2): 221–230. 2482: 2455:(3): 161–208. 2436: 2425:(3): 525–531. 2397: 2395:, p. 208. 2385: 2331: 2290: 2247: 2204: 2192: 2180: 2178:, p. 242. 2168: 2147:10.1086/398642 2125: 2066: 2054: 2025:10.1086/510095 1998: 1996:, p. 118. 1981: 1979:, p. 357. 1969: 1967:, p. 358. 1957: 1908: 1859: 1840:(4): 521–523. 1817: 1798:(3): 339–342. 1775: 1738:(4): 461–477. 1722: 1720:, p. 214. 1710: 1708:, p. 543. 1695: 1662: 1650: 1648:, p. 311. 1638: 1619:(5): 168–170. 1593: 1581: 1569: 1518: 1471: 1428:(2): 145–151. 1408: 1365:(9): 697–705. 1342: 1269: 1209: 1183: 1181:, p. 102. 1160: 1158:, p. 102. 1143: 1116:(2): 390–404. 1099: 1041: 1014:(3): 171–187. 986: 984:, p. 104. 974: 962: 950: 948:, p. 129. 935: 923: 904:(3): 391–394. 888: 886:, p. 198. 876: 864: 833: 832: 830: 827: 752: 749: 605: 602: 541:great hornbill 524:mandarin ducks 515: 512: 385: 382: 318:house sparrows 248: 245: 176: 173: 141: 138: 26: 9: 6: 4: 3: 2: 3559: 3548: 3545: 3543: 3540: 3538: 3537:Bird behavior 3535: 3534: 3532: 3525: 3520: 3517: 3514: 3511: 3510: 3506: 3502: 3487: 3440: 3434: 3430: 3429: 3423: 3419: 3413: 3409: 3404: 3400: 3394: 3390: 3385: 3381: 3375: 3371: 3370: 3364: 3360: 3354: 3350: 3349: 3343: 3339: 3333: 3329: 3325: 3320: 3316: 3310: 3306: 3305: 3299: 3295: 3289: 3285: 3284: 3278: 3274: 3268: 3264: 3263: 3257: 3253: 3247: 3243: 3242: 3236: 3232: 3226: 3222: 3221: 3215: 3211: 3207: 3203: 3199: 3195: 3190: 3186: 3180: 3176: 3171: 3167: 3161: 3156: 3155: 3148: 3144: 3138: 3134: 3129: 3125: 3119: 3114: 3113: 3106: 3102: 3096: 3092: 3087: 3083: 3077: 3073: 3072: 3066: 3062: 3056: 3052: 3051: 3045: 3039: 3035: 3029: 3025: 3024: 3018: 3014: 3008: 3004: 3003: 2997: 2993: 2987: 2982: 2981: 2974: 2973: 2961:, p. 46. 2960: 2955: 2948: 2943: 2935: 2931: 2926: 2921: 2917: 2913: 2909: 2905: 2901: 2897: 2893: 2886: 2878: 2874: 2870: 2866: 2861: 2856: 2852: 2848: 2844: 2836: 2828: 2824: 2819: 2814: 2809: 2804: 2800: 2796: 2793:(6): e11315. 2792: 2788: 2784: 2776: 2768: 2764: 2759: 2754: 2750: 2746: 2742: 2738: 2734: 2730: 2726: 2719: 2712: 2707: 2705: 2696: 2692: 2687: 2682: 2678: 2674: 2670: 2666: 2662: 2655: 2649:, p. 94. 2648: 2643: 2636: 2631: 2625:, p. 36. 2624: 2619: 2612: 2607: 2600: 2595: 2593: 2584: 2580: 2576: 2572: 2568: 2564: 2557: 2550: 2542: 2538: 2534: 2530: 2526: 2522: 2518: 2514: 2510: 2506: 2502: 2500: 2491: 2489: 2487: 2478: 2474: 2470: 2466: 2462: 2458: 2454: 2450: 2443: 2441: 2432: 2428: 2424: 2420: 2413: 2406: 2404: 2402: 2394: 2389: 2381: 2377: 2372: 2367: 2363: 2359: 2355: 2351: 2347: 2340: 2338: 2336: 2327: 2323: 2318: 2313: 2309: 2305: 2301: 2294: 2286: 2282: 2278: 2274: 2270: 2266: 2262: 2258: 2251: 2243: 2239: 2235: 2231: 2227: 2223: 2219: 2215: 2208: 2202:, p. 37. 2201: 2196: 2190:, p. 49. 2189: 2184: 2177: 2172: 2164: 2160: 2156: 2152: 2148: 2144: 2140: 2136: 2129: 2121: 2117: 2113: 2109: 2105: 2101: 2097: 2093: 2089: 2085: 2081: 2077: 2070: 2064:, p. 58. 2063: 2058: 2050: 2046: 2042: 2038: 2034: 2030: 2026: 2022: 2018: 2014: 2007: 2005: 2003: 1995: 1990: 1988: 1986: 1978: 1973: 1966: 1961: 1953: 1949: 1944: 1939: 1935: 1931: 1927: 1923: 1919: 1912: 1904: 1900: 1895: 1890: 1886: 1882: 1878: 1874: 1870: 1863: 1855: 1851: 1847: 1843: 1839: 1835: 1828: 1821: 1813: 1809: 1805: 1801: 1797: 1793: 1786: 1779: 1771: 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995: 993: 991: 983: 978: 972:, p. 57. 971: 966: 960:, p. 58. 959: 954: 947: 942: 940: 933:, p. 71. 932: 927: 919: 915: 911: 907: 903: 899: 892: 885: 884:Luescher 2006 880: 874:, p. 68. 873: 868: 852: 848: 844: 838: 834: 826: 824: 820: 816: 812: 808: 806: 802: 798: 794: 790: 785: 783: 778: 774: 770: 762: 757: 748: 745: 740: 738: 730: 726: 721: 717: 715: 711: 707: 703: 699: 698:ectoparasites 693: 690: 686: 685:black vulture 682: 676: 673: 665: 657: 650: 641: 637: 632: 625: 621: 616: 611: 601: 599: 595: 591: 586: 582: 578: 573: 570: 565: 558: 557:mandarin duck 555:of the drake 554: 550: 546: 542: 538: 534: 525: 520: 511: 509: 505: 501: 496: 494: 490: 486: 482: 477: 473: 469: 465: 461: 457: 451: 448: 444: 443:ectoparasites 438: 431: 430:down feathers 427: 419: 414: 406: 398: 390: 381: 378: 374: 370: 366: 362: 358: 354: 350: 346: 343:, ostriches, 342: 337: 333: 331: 328: 324: 319: 315: 314:avian malaria 311: 307: 303: 299: 298: 293: 292: 287: 282: 279: 275: 271: 262: 258: 253: 244: 242: 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