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[o]" accesskey="o"><span class="vector-icon mw-ui-icon-logIn mw-ui-icon-wikimedia-logIn"></span> <span>Log in</span></a></li> </ul> </div> </div> <div id="p-user-menu-anon-editor" class="vector-menu mw-portlet mw-portlet-user-menu-anon-editor" > <div class="vector-menu-heading"> Pages for logged out editors <a href="/wiki/Help:Introduction" aria-label="Learn more about editing"><span>learn more</span></a> </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="pt-anoncontribs" class="mw-list-item"><a href="/wiki/Special:MyContributions" title="A list of edits made from this IP address [y]" accesskey="y"><span>Contributions</span></a></li><li id="pt-anontalk" class="mw-list-item"><a href="/wiki/Special:MyTalk" title="Discussion about edits from this IP address [n]" accesskey="n"><span>Talk</span></a></li> </ul> </div> </div> </div> </div> </nav> </div> </header> </div> <div class="mw-page-container"> <div class="mw-page-container-inner"> <div 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data-event-name="pinnable-header.vector-toc.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-toc.unpin">hide</button> </div> <ul class="vector-toc-contents" id="mw-panel-toc-list"> <li id="toc-mw-content-text" class="vector-toc-list-item vector-toc-level-1"> <a href="#" class="vector-toc-link"> <div class="vector-toc-text">(Top)</div> </a> </li> <li id="toc-Etymology" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Etymology"> <div class="vector-toc-text"> <span class="vector-toc-numb">1</span> <span>Etymology</span> </div> </a> <ul id="toc-Etymology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Importance" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Importance"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Importance</span> </div> </a> <ul id="toc-Importance-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Preen_oil" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Preen_oil"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Preen oil</span> </div> </a> <ul id="toc-Preen_oil-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Preening_action" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Preening_action"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Preening action</span> </div> </a> <ul id="toc-Preening_action-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Secondary_functions" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Secondary_functions"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Secondary functions</span> </div> </a> <ul id="toc-Secondary_functions-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Allopreening" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Allopreening"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Allopreening</span> </div> </a> <ul id="toc-Allopreening-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Potential_problems" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Potential_problems"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Potential problems</span> </div> </a> <ul id="toc-Potential_problems-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>References</span> </div> </a> <button aria-controls="toc-References-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle References subsection</span> </button> <ul id="toc-References-sublist" class="vector-toc-list"> <li id="toc-Works_cited" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Works_cited"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.1</span> <span>Works cited</span> </div> </a> <ul id="toc-Works_cited-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>External links</span> </div> </a> <ul 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title="This is a featured article. Click here for more information."><img alt="Featured article" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e7/Cscr-featured.svg/20px-Cscr-featured.svg.png" decoding="async" width="20" height="19" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e7/Cscr-featured.svg/30px-Cscr-featured.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e7/Cscr-featured.svg/40px-Cscr-featured.svg.png 2x" data-file-width="466" data-file-height="443" /></a></span></div></div> <div id="mw-indicator-spoken-icon" class="mw-indicator"><div class="mw-parser-output"><span typeof="mw:File"><a href="/wiki/File:En-Preening-article.ogg" title="Listen to this article"><img alt="Listen to this article" src="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/Sound-icon.svg/20px-Sound-icon.svg.png" decoding="async" width="20" height="15" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/Sound-icon.svg/30px-Sound-icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/47/Sound-icon.svg/40px-Sound-icon.svg.png 2x" data-file-width="128" data-file-height="96" /></a></span></div></div> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Maintenance behaviour of birds</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">This article is about a maintenance behaviour in birds. For similar behaviour in other animals, see <a href="/wiki/Personal_grooming" title="Personal grooming">Personal grooming</a>.</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Red_Lory_(Eos_bornea)-6.jpg" class="mw-file-description"><img alt="The head and back of a short-billed red parrot, which is chewing on the tip of a black and red feather" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/81/Red_Lory_%28Eos_bornea%29-6.jpg/170px-Red_Lory_%28Eos_bornea%29-6.jpg" decoding="async" width="170" height="212" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/81/Red_Lory_%28Eos_bornea%29-6.jpg/255px-Red_Lory_%28Eos_bornea%29-6.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/81/Red_Lory_%28Eos_bornea%29-6.jpg/340px-Red_Lory_%28Eos_bornea%29-6.jpg 2x" data-file-width="683" data-file-height="850" /></a><figcaption>When preening, a bird (such as this <a href="/wiki/Red_lory" title="Red lory">red lory</a>) draws individual feathers through its beak, realigning and re-interlocking the <dfn><a href="/wiki/Glossary_of_bird_terms#barbules" title="Glossary of bird terms"><span title="See entry at: Glossary of bird terms § barbules" style="color:inherit;" class="glossary-link">barbules</span></a></dfn>.</figcaption></figure> <p><b>Preening</b> is a <dfn><a href="/wiki/Glossary_of_bird_terms#maintenance_behaviour" title="Glossary of bird terms"><span title="See entry at: Glossary of bird terms § maintenance behaviour" style="color:inherit;" class="glossary-link">maintenance behaviour</span></a></dfn> found in <a href="/wiki/Bird" title="Bird">birds</a> that involves the use of the beak to position <a href="/wiki/Feather" title="Feather">feathers</a>, interlock feather <dfn><a href="/wiki/Glossary_of_bird_terms#barbules" title="Glossary of bird terms"><span title="See entry at: Glossary of bird terms § barbules" style="color:inherit;" class="glossary-link">barbules</span></a></dfn> that have become separated, clean <a href="/wiki/Plumage" title="Plumage">plumage</a>, and keep <a href="/wiki/Ectoparasites" class="mw-redirect" title="Ectoparasites">ectoparasites</a> in check. Feathers contribute significantly to a bird's insulation, waterproofing and <a href="/wiki/Bird_flight" title="Bird flight">aerodynamic flight</a>, 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 <a href="/wiki/Beak" title="Beak">beaks</a>, they gather preen oil from <a href="/wiki/Uropygial_gland" title="Uropygial gland">a gland at the base of their tail</a> and distribute this oil through their feathers. They draw each <a href="/wiki/Contour_feather" class="mw-redirect" title="Contour feather">contour feather</a> through their bill, nibbling it from base to tip. </p><p>Over time, some elements of preening have <a href="/wiki/Evolved" class="mw-redirect" title="Evolved">evolved</a> to have secondary functions. <a href="/wiki/Ritualization" title="Ritualization">Ritualised</a> preening has become a part of some <a href="/wiki/Courtship_display" title="Courtship display">courtship displays</a>, for example. It is also a <a href="/wiki/Displacement_activity" title="Displacement activity">displacement activity</a> 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 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. </p><p>Ingestion of <a href="/wiki/Pollutant" title="Pollutant">pollutants</a> 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. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Etymology">Etymology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Preening&action=edit&section=1" title="Edit section: Etymology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The use of the word <i>preen</i> to mean the tidying of a bird's feathers dates from <a href="/wiki/Late_Middle_English" class="mw-redirect" title="Late Middle English">Late Middle English</a>. It appears to be a variant of the word <i>prune</i>; one now-obsolete definition of <i>prune</i> meant "anoint", based on the <a href="/wiki/Latin" title="Latin">Latin</a> <i>ungere</i>, which had the same meaning. This usage was combined with the Scottish and northern English dialect <i>preen</i> meaning "pierce" or "pin", due to the "pricking" action of the bird's beak during preening.<sup id="cite_ref-OED_1-0" class="reference"><a href="#cite_note-OED-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Importance">Importance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Preening&action=edit&section=2" title="Edit section: Importance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><span><video id="mwe_player_0" poster="//upload.wikimedia.org/wikipedia/commons/thumb/0/03/Mallard_and_ducklings_preening_%2881284%29.webm/220px--Mallard_and_ducklings_preening_%2881284%29.webm.jpg" controls="" preload="none" data-mw-tmh="" class="mw-file-element" width="220" height="124" data-durationhint="64" data-mwtitle="Mallard_and_ducklings_preening_(81284).webm" data-mwprovider="wikimediacommons" resource="/wiki/File:Mallard_and_ducklings_preening_(81284).webm"><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/0/03/Mallard_and_ducklings_preening_%2881284%29.webm/Mallard_and_ducklings_preening_%2881284%29.webm.480p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="480p.vp9.webm" data-width="854" data-height="480" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/0/03/Mallard_and_ducklings_preening_%2881284%29.webm/Mallard_and_ducklings_preening_%2881284%29.webm.720p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="720p.vp9.webm" data-width="1280" data-height="720" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/0/03/Mallard_and_ducklings_preening_%2881284%29.webm/Mallard_and_ducklings_preening_%2881284%29.webm.1080p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="1080p.vp9.webm" data-width="1920" data-height="1080" /><source src="//upload.wikimedia.org/wikipedia/commons/0/03/Mallard_and_ducklings_preening_%2881284%29.webm" type="video/webm; codecs="vp9, opus"" data-width="1920" data-height="1080" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/0/03/Mallard_and_ducklings_preening_%2881284%29.webm/Mallard_and_ducklings_preening_%2881284%29.webm.144p.mjpeg.mov" type="video/quicktime" data-transcodekey="144p.mjpeg.mov" data-width="256" data-height="144" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/0/03/Mallard_and_ducklings_preening_%2881284%29.webm/Mallard_and_ducklings_preening_%2881284%29.webm.240p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="240p.vp9.webm" data-width="426" data-height="240" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/0/03/Mallard_and_ducklings_preening_%2881284%29.webm/Mallard_and_ducklings_preening_%2881284%29.webm.360p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="360p.vp9.webm" data-width="640" data-height="360" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/0/03/Mallard_and_ducklings_preening_%2881284%29.webm/Mallard_and_ducklings_preening_%2881284%29.webm.360p.webm" type="video/webm; codecs="vp8, vorbis"" data-transcodekey="360p.webm" data-width="640" data-height="360" /></video></span><figcaption>A <a href="/wiki/Mallard" title="Mallard">mallard</a> and her ducklings demonstrate the shaking, nibbling and stroking movements of preening.</figcaption></figure> <p>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.<sup id="cite_ref-FOOTNOTEMoon-FandliDodmanO'Sullivan199968_2-0" class="reference"><a href="#cite_note-FOOTNOTEMoon-FandliDodmanO'Sullivan199968-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Despite spending considerable time in their efforts, they do not use proper techniques to groom effectively and may do a poor job overall as a result.<sup id="cite_ref-FOOTNOTELuescher2006198_3-0" class="reference"><a href="#cite_note-FOOTNOTELuescher2006198-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> 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 <a href="/wiki/Filoplume" class="mw-redirect" title="Filoplume">filoplumes</a>, specialised feathers buried under a bird's outer covering of <a href="/wiki/Contour_feather" class="mw-redirect" title="Contour feather">contour feathers</a>, help to signal when contour feathers have been displaced. <a href="/wiki/Mechanoreceptor" title="Mechanoreceptor">Mechanoreceptors</a> at the base of the filoplumes only fire when contour feathers are displaced or the filoplume moves.<sup id="cite_ref-Necker_4-0" class="reference"><a href="#cite_note-Necker-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Preening enables birds to remove dirt and parasites from their plumage, and assists in the waterproofing of feathers.<sup id="cite_ref-FOOTNOTEMoss201571_5-0" class="reference"><a href="#cite_note-FOOTNOTEMoss201571-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTELovetteFitzpatrick2016129_6-0" class="reference"><a href="#cite_note-FOOTNOTELovetteFitzpatrick2016129-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> During <a href="/wiki/Moult" class="mw-redirect" title="Moult">moult</a>, birds remove the sheaths from around their emerging <a href="/wiki/Pin_feather" title="Pin feather">pin feathers</a> while preening.<sup id="cite_ref-FOOTNOTEElphickDunning200158_7-0" class="reference"><a href="#cite_note-FOOTNOTEElphickDunning200158-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p>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.<sup id="cite_ref-FOOTNOTEElphickDunning200157_8-0" class="reference"><a href="#cite_note-FOOTNOTEElphickDunning200157-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> When resting, birds may preen at least once an hour.<sup id="cite_ref-FOOTNOTEGill2007104_9-0" class="reference"><a href="#cite_note-FOOTNOTEGill2007104-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Studies on multiple species have shown that they spend an average of more than 9% of each day on maintenance behaviours, preening occupying over 92% of that time, though this figure can be significantly higher.<sup id="cite_ref-Cotgreave_10-0" class="reference"><a href="#cite_note-Cotgreave-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Studies found that some <a href="/wiki/Gull" title="Gull">gull</a> species spent 15% of daylight hours during the breeding season preening, while another showed that <a href="/wiki/Common_loon" title="Common loon">common loons</a> spent upwards of 25% of their day preening.<sup id="cite_ref-Cotgreave_10-1" class="reference"><a href="#cite_note-Cotgreave-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Delius_11-0" class="reference"><a href="#cite_note-Delius-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> 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.<sup id="cite_ref-Cotgreave_10-2" class="reference"><a href="#cite_note-Cotgreave-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Some <a href="/wiki/Ratite" title="Ratite">ratites</a>, which are not dependent on their feathers for flight, spend far less time on maintenance behaviours. One study of <a href="/wiki/Ostrich" title="Ostrich">ostriches</a> found that they spent less than 1% of their time engaged in such behaviours.<sup id="cite_ref-Williams_12-0" class="reference"><a href="#cite_note-Williams-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Preen_oil">Preen oil</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Preening&action=edit&section=3" title="Edit section: Preen oil"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:White-winged_Crossbill_Uropygial.JPG" class="mw-file-description"><img alt="A reddish-headed bird puts its beak on a small nipple at the base of its brownish-grey tail." src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4d/White-winged_Crossbill_Uropygial.JPG/220px-White-winged_Crossbill_Uropygial.JPG" decoding="async" width="220" height="160" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4d/White-winged_Crossbill_Uropygial.JPG/330px-White-winged_Crossbill_Uropygial.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4d/White-winged_Crossbill_Uropygial.JPG/440px-White-winged_Crossbill_Uropygial.JPG 2x" data-file-width="3225" data-file-height="2344" /></a><figcaption>Birds, like this <a href="/wiki/Two-barred_crossbill" title="Two-barred crossbill">two-barred crossbill</a>, typically use their beak to gather preen gland oil from the <a href="/wiki/Uropygial_gland" title="Uropygial gland">uropygial gland</a>.</figcaption></figure> <p>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.<sup id="cite_ref-FOOTNOTEGill2007102_13-0" class="reference"><a href="#cite_note-FOOTNOTEGill2007102-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> To facilitate that care, many bird species have a <a href="/wiki/Uropygial_gland" title="Uropygial gland">preen or uropygial gland</a>, 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.<sup id="cite_ref-FOOTNOTECampbellLack1985102_14-0" class="reference"><a href="#cite_note-FOOTNOTECampbellLack1985102-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> The gland is generally larger (in relation to body size) in waterbirds, including <a href="/wiki/Tern" title="Tern">terns</a>, <a href="/wiki/Grebe" title="Grebe">grebes</a> 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.<sup id="cite_ref-Montalti_15-0" class="reference"><a href="#cite_note-Montalti-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p>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.<sup id="cite_ref-FOOTNOTEGill2007102_13-1" class="reference"><a href="#cite_note-FOOTNOTEGill2007102-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> One study of <a href="/wiki/Eurasian_hoopoe" title="Eurasian hoopoe">Eurasian hoopoes</a> showed that the presence of symbiotic bacteria (<i><a href="/wiki/Enterococcus_faecalis" title="Enterococcus faecalis">Enterococcus faecalis</a></i>) in their preen oil inhibited the growth of the feather-degrading bacteria <i><a href="/wiki/Bacillus_licheniformis" title="Bacillus licheniformis">Bacillus licheniformis</a></i>. <i>Enterococcus faecalis</i> did this by releasing a <a href="/wiki/Bacteriocin" title="Bacteriocin">bacteriocin</a>.<sup id="cite_ref-Ruiz_16-0" class="reference"><a href="#cite_note-Ruiz-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> 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 <a href="/wiki/Egg_incubation" title="Egg incubation">incubation</a>. This alters the colour of the eggs (darkening them) but there is also evidence that the bacteria may help to protect the developing chicks.<sup id="cite_ref-Martínez-García_17-0" class="reference"><a href="#cite_note-Martínez-García-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Soler_18-0" class="reference"><a href="#cite_note-Soler-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> 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.<sup id="cite_ref-Czirják_19-0" class="reference"><a href="#cite_note-Czirják-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Giraudeau_20-0" class="reference"><a href="#cite_note-Giraudeau-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> Preen oil may play a part in protecting at least some species from some internal parasites; a study of the incidence of <a href="/wiki/Avian_malaria" title="Avian malaria">avian malaria</a> in <a href="/wiki/House_sparrow" title="House sparrow">house sparrows</a> found that uninfected birds had larger uropygial glands and higher antimicrobial activity in those glands than infected birds did.<sup id="cite_ref-Magallanes_21-0" class="reference"><a href="#cite_note-Magallanes-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> There is even evidence that the foul-smelling preen oil of hoopoes and <a href="/wiki/Wood_hoopoe" title="Wood hoopoe">wood hoopoes</a> may help to repel <a href="/wiki/Mammal" title="Mammal">mammalian</a> <a href="/wiki/Predator" class="mw-redirect" title="Predator">predators</a>.<sup id="cite_ref-FOOTNOTEGill2007102_13-2" class="reference"><a href="#cite_note-FOOTNOTEGill2007102-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>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.<sup id="cite_ref-FOOTNOTELovetteFitzpatrick2016129_6-1" class="reference"><a href="#cite_note-FOOTNOTELovetteFitzpatrick2016129-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>While most species have a preen gland, the organ is missing in the ratites (<a href="/wiki/Emu" title="Emu">emu</a>, ostriches, <a href="/wiki/Cassowary" title="Cassowary">cassowaries</a>, <a href="/wiki/Rhea_(bird)" title="Rhea (bird)">rheas</a> and <a href="/wiki/Kiwi_(bird)" title="Kiwi (bird)">kiwis</a>) and some <a href="/wiki/Neognath" class="mw-redirect" title="Neognath">neognath</a> birds, including <a href="/wiki/Bustard" title="Bustard">bustards</a>, <a href="/wiki/Woodpecker" title="Woodpecker">woodpeckers</a>, a few <a href="/wiki/Parrot" title="Parrot">parrots</a> and <a href="/wiki/Pigeon" class="mw-redirect" title="Pigeon">pigeons</a>.<sup id="cite_ref-FOOTNOTEPerrins200937_22-0" class="reference"><a href="#cite_note-FOOTNOTEPerrins200937-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEde_Juana199240_23-0" class="reference"><a href="#cite_note-FOOTNOTEde_Juana199240-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Some species that lack a preen gland instead have <a href="/wiki/Powder_down" class="mw-redirect" title="Powder down">powder down</a> feathers which continually break down into a fine dust that the birds apply to their contour feathers while preening.<sup id="cite_ref-Wetmore_24-0" class="reference"><a href="#cite_note-Wetmore-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> 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 down can give a metallic sheen to the plumage.<sup id="cite_ref-FOOTNOTEEhrlichDobkinWheye1988311_25-0" class="reference"><a href="#cite_note-FOOTNOTEEhrlichDobkinWheye1988311-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Preening_action">Preening action</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Preening&action=edit&section=4" title="Edit section: Preening action"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Feather_zipping_microstructure.svg" class="mw-file-description"><img alt="A diagram showing a series of increasingly closer views of feather structure, to show how the microscopic elements interact" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6b/Feather_zipping_microstructure.svg/220px-Feather_zipping_microstructure.svg.png" decoding="async" width="220" height="156" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6b/Feather_zipping_microstructure.svg/330px-Feather_zipping_microstructure.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6b/Feather_zipping_microstructure.svg/440px-Feather_zipping_microstructure.svg.png 2x" data-file-width="1052" data-file-height="744" /></a><figcaption>A shaft runs down the middle of the feather with <dfn><a href="/wiki/Glossary_of_bird_terms#barbs" title="Glossary of bird terms"><span title="See entry at: Glossary of bird terms § barbs" style="color:inherit;" class="glossary-link">barbs</span></a></dfn> branching from the main shaft and hooked <dfn><a href="/wiki/Glossary_of_bird_terms#barbules" title="Glossary of bird terms"><span title="See entry at: Glossary of bird terms § barbules" style="color:inherit;" class="glossary-link">barbules</span></a></dfn> branching from those barbs. The barbules' hooks interlock as shown to provide strength and flexibility.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Barn_Owl_Pectinate_Claw.jpg" class="mw-file-description"><img alt="Close-up of a talon with a pointed tip and small grooves along one edge" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Barn_Owl_Pectinate_Claw.jpg/220px-Barn_Owl_Pectinate_Claw.jpg" decoding="async" width="220" height="308" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Barn_Owl_Pectinate_Claw.jpg/330px-Barn_Owl_Pectinate_Claw.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Barn_Owl_Pectinate_Claw.jpg/440px-Barn_Owl_Pectinate_Claw.jpg 2x" data-file-width="1989" data-file-height="2784" /></a><figcaption>The specialized pectinate claw of a <a href="/wiki/Barn-owl" class="mw-redirect" title="Barn-owl">barn owl</a> is used as a comb for facial preening.</figcaption></figure> <p>A bird's plumage is primarily made up of two feather types: firm <a href="/wiki/Pennaceous" class="mw-redirect" title="Pennaceous">vaned or pennaceous feathers</a> on the surface, with softer <a href="/wiki/Down_feather" title="Down feather">down feathers</a> underneath. Both feather types have a central shaft with narrower <dfn><a href="/wiki/Glossary_of_bird_terms#barbs" title="Glossary of bird terms"><span title="See entry at: Glossary of bird terms § barbs" style="color:inherit;" class="glossary-link">barbs</span></a></dfn> 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 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 <a href="/wiki/Ectoparasite" class="mw-redirect" title="Ectoparasite">ectoparasites</a>, rejoining unzipped barbules, and rearranging feathers.<sup id="cite_ref-FOOTNOTECampbellLack1985102_14-1" class="reference"><a href="#cite_note-FOOTNOTECampbellLack1985102-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> 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".<sup id="cite_ref-FOOTNOTECampbellLack1985102–103_26-0" class="reference"><a href="#cite_note-FOOTNOTECampbellLack1985102–103-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Penguin" title="Penguin">Penguins</a> use their whole heads to stroke, in a motion referred to as "wiping".<sup id="cite_ref-FOOTNOTECampbellLack1985102_14-2" class="reference"><a href="#cite_note-FOOTNOTECampbellLack1985102-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> 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.<sup id="cite_ref-Zhao_27-0" class="reference"><a href="#cite_note-Zhao-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p><p>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.<sup id="cite_ref-FOOTNOTEEhrlichDobkinWheye1988543_28-0" class="reference"><a href="#cite_note-FOOTNOTEEhrlichDobkinWheye1988543-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Longer-necked birds may rub their head directly on their uropygial gland.<sup id="cite_ref-FOOTNOTEHailman1985214_29-0" class="reference"><a href="#cite_note-FOOTNOTEHailman1985214-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> Some species (including <a href="/wiki/Nightjar" title="Nightjar">nightjars</a>, <a href="/wiki/Heron" title="Heron">herons</a>, <a href="/wiki/Frigatebird" title="Frigatebird">frigatebirds</a>, owls and <a href="/wiki/Pratincole" title="Pratincole">pratincoles</a>) have comb-like serrations on the claw (a pectinate claw) of the middle toe which may aid in scratch-preening.<sup id="cite_ref-Stettenheim_30-0" class="reference"><a href="#cite_note-Stettenheim-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> 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).<sup id="cite_ref-FOOTNOTECampbellLack1985102_14-3" class="reference"><a href="#cite_note-FOOTNOTECampbellLack1985102-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Nice_31-0" class="reference"><a href="#cite_note-Nice-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> There is some evidence that the method used by a species may be related to its ecology. For instance, <a href="/wiki/New_World_warbler" title="New World warbler">New World warblers</a> that are primarily <a href="https://en.wiktionary.org/wiki/arboreal" class="extiw" title="wikt:arboreal">arboreal</a> tend to be overwing scratchers, while those that spend significant time on the ground are typically underwing scratchers.<sup id="cite_ref-FOOTNOTEEhrlichDobkinWheye1988543_28-1" class="reference"><a href="#cite_note-FOOTNOTEEhrlichDobkinWheye1988543-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> In general, preening takes place while the bird is perched, on the ground, or swimming, but some of the more aerial species (including <a href="/wiki/Swift_(bird)" title="Swift (bird)">swifts</a>, <a href="/wiki/Swallow" title="Swallow">swallows</a>, terns and <a href="/wiki/Albatross" title="Albatross">albatrosses</a>) preen while flying.<sup id="cite_ref-Goodwin_32-0" class="reference"><a href="#cite_note-Goodwin-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> 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 <a href="/wiki/Feather_lice" class="mw-redirect" title="Feather lice">feather lice</a>; the removal of the bill tip caused an increase in feather lice due to ineffective preening.<sup id="cite_ref-Clayton_33-0" class="reference"><a href="#cite_note-Clayton-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bush_34-0" class="reference"><a href="#cite_note-Bush-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p><p>Preening is often done in association with other maintenance behaviours, including bathing, <a href="/wiki/Dust_bathing" title="Dust bathing">dusting</a>, sunning, oiling or <a href="/wiki/Anting_(bird_activity)" class="mw-redirect" title="Anting (bird activity)">anting</a>, and can either precede or follow these other behaviours.<sup id="cite_ref-FOOTNOTECarnaby2008358_35-0" class="reference"><a href="#cite_note-FOOTNOTECarnaby2008358-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> All birds typically preen after bathing.<sup id="cite_ref-FOOTNOTEEhrlichDobkinWheyePimms1994357_36-0" class="reference"><a href="#cite_note-FOOTNOTEEhrlichDobkinWheyePimms1994357-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> Groups of birds sometimes all groom individually at the same time. This has been seen in species ranging from herons to <a href="/wiki/Icterid" title="Icterid">blackbirds</a>.<sup id="cite_ref-FOOTNOTEKricher2020118_37-0" class="reference"><a href="#cite_note-FOOTNOTEKricher2020118-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Secondary_functions">Secondary functions</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Preening&action=edit&section=5" title="Edit section: Secondary functions"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Mandarin.duck.arp.jpg" class="mw-file-description"><img alt="A rusty-brown, black and white duck with large orange "sail" feathers sticking up from its back floats on water." src="//upload.wikimedia.org/wikipedia/commons/thumb/5/51/Mandarin.duck.arp.jpg/220px-Mandarin.duck.arp.jpg" decoding="async" width="220" height="157" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/51/Mandarin.duck.arp.jpg/330px-Mandarin.duck.arp.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/51/Mandarin.duck.arp.jpg/440px-Mandarin.duck.arp.jpg 2x" data-file-width="1770" data-file-height="1264" /></a><figcaption>During courtship, drake <a href="/wiki/Mandarin_duck" title="Mandarin duck">mandarin ducks</a> ritually preen their distinctive orange "sail" feathers.</figcaption></figure> <p>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.<sup id="cite_ref-Delhey_38-0" class="reference"><a href="#cite_note-Delhey-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> In some species, preen oil is used to cosmetically colour the plumage. During the breeding season, the preen oil of the <a href="/wiki/Great_white_pelican" title="Great white pelican">great white pelican</a> becomes red-orange, imparting a pink flush to the bird's plumage.<sup id="cite_ref-Delhey_38-1" class="reference"><a href="#cite_note-Delhey-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> The preen oil of several gull and tern species, including <a href="/wiki/Ross%27s_gull" title="Ross's gull">Ross's gull</a>, 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.<sup id="cite_ref-FOOTNOTEEhrlichDobkinWheye198858_39-0" class="reference"><a href="#cite_note-FOOTNOTEEhrlichDobkinWheye198858-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> The yellow feathers of the <a href="/wiki/Great_hornbill" title="Great hornbill">great hornbill</a> are also cosmetically coloured during preening.<sup id="cite_ref-Delhey_38-2" class="reference"><a href="#cite_note-Delhey-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> The preen oil of the <a href="/wiki/Bohemian_waxwing" title="Bohemian waxwing">Bohemian waxwing</a> increases the UV reflectance of its feathers.<sup id="cite_ref-Pérez_40-0" class="reference"><a href="#cite_note-Pérez-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> Ritualised preening is used in <a href="/wiki/Courtship_display" title="Courtship display">courtship displays</a> by several species, particularly ducks; such preening is typically designed to draw attention to a modified structure (such as the sail-shaped <a href="/wiki/Flight_feather#Secondaries" title="Flight feather">secondaries</a> of the drake <a href="/wiki/Mandarin_duck" title="Mandarin duck">mandarin duck</a>) or distinctive colour (such as the <dfn><a href="/wiki/Glossary_of_bird_terms#speculum" title="Glossary of bird terms"><span title="See entry at: Glossary of bird terms § speculum" style="color:inherit;" class="glossary-link">speculum</span></a></dfn>) on the bird.<sup id="cite_ref-Tinbergen_41-0" class="reference"><a href="#cite_note-Tinbergen-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEHuxleyHardyFord1954242_42-0" class="reference"><a href="#cite_note-FOOTNOTEHuxleyHardyFord1954242-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Mallard" title="Mallard">Mallards</a> 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.<sup id="cite_ref-FOOTNOTEEhrlichDobkinWheyePimms199449_43-0" class="reference"><a href="#cite_note-FOOTNOTEEhrlichDobkinWheyePimms199449-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> Courtship preening is more conspicuous than is preening for feather maintenance, using more stereotypical movements.<sup id="cite_ref-FOOTNOTEEhrlichDobkinWheyePimms199437_44-0" class="reference"><a href="#cite_note-FOOTNOTEEhrlichDobkinWheyePimms199437-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p><p>Preening may be performed as a <a href="/wiki/Displacement_activity" title="Displacement activity">displacement activity</a>. In some cases, it is done in place of another activity that birds are strongly motivated, but unable, to do. In one study, <a href="/wiki/Black-headed_gull" title="Black-headed gull">black-headed gulls</a> which were prevented from incubating a full <a href="/wiki/Clutch_(eggs)" title="Clutch (eggs)">clutch</a> 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.<sup id="cite_ref-Moynihan_45-0" class="reference"><a href="#cite_note-Moynihan-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> The conflict between two incompatible drives, such as incubating and escape, can lead a bird to engage in displacement activities. Nesting <a href="/wiki/Sandwich_tern" title="Sandwich tern">Sandwich</a> and <a href="/wiki/Common_tern" title="Common tern">common terns</a> preen when they have been alarmed by a potential predator or when they have had an aggressive encounter with a neighbouring bird, for instance.<sup id="cite_ref-Van_46-0" class="reference"><a href="#cite_note-Van-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> Fighting <a href="/wiki/European_starling" class="mw-redirect" title="European starling">European starlings</a> will break off their battles to preen.<sup id="cite_ref-Mitchell_47-0" class="reference"><a href="#cite_note-Mitchell-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Allopreening">Allopreening</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Preening&action=edit&section=6" title="Edit section: Allopreening"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Social_grooming" title="Social grooming">Social grooming</a></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Starr_080606-6754_Ipomoea_pes-caprae_subsp._brasiliensis.jpg" class="mw-file-description"><img alt="Two black and white birds lie on their bellies on a sandy beach; one nuzzles the other's head with the tip of its long, yellow bill." src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6a/Starr_080606-6754_Ipomoea_pes-caprae_subsp._brasiliensis.jpg/220px-Starr_080606-6754_Ipomoea_pes-caprae_subsp._brasiliensis.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6a/Starr_080606-6754_Ipomoea_pes-caprae_subsp._brasiliensis.jpg/330px-Starr_080606-6754_Ipomoea_pes-caprae_subsp._brasiliensis.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6a/Starr_080606-6754_Ipomoea_pes-caprae_subsp._brasiliensis.jpg/440px-Starr_080606-6754_Ipomoea_pes-caprae_subsp._brasiliensis.jpg 2x" data-file-width="2816" data-file-height="2112" /></a><figcaption>Allopreening may help to strengthen <a href="/wiki/Pair_bond" title="Pair bond">pair bonds</a> in species such as the <a href="/wiki/Laysan_albatross" title="Laysan albatross">Laysan albatross</a>.<sup id="cite_ref-Kenny_48-0" class="reference"><a href="#cite_note-Kenny-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup></figcaption></figure> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Black_capped_parakeet_receiving_scritches.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0b/Black_capped_parakeet_receiving_scritches.png/220px-Black_capped_parakeet_receiving_scritches.png" decoding="async" width="220" height="293" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0b/Black_capped_parakeet_receiving_scritches.png/330px-Black_capped_parakeet_receiving_scritches.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0b/Black_capped_parakeet_receiving_scritches.png/440px-Black_capped_parakeet_receiving_scritches.png 2x" data-file-width="3024" data-file-height="4032" /></a><figcaption>Some <a href="/wiki/Companion_parrot" title="Companion parrot">pet parrots</a>, such as this <a href="/wiki/Black-capped_conure" class="mw-redirect" title="Black-capped conure">black-capped conure</a>, may solicit preening from humans.</figcaption></figure> <p>Although preening is primarily an individual behaviour, some species indulge in allopreening, one individual preening another.<sup id="cite_ref-FOOTNOTECampbellLack1985102_14-4" class="reference"><a href="#cite_note-FOOTNOTECampbellLack1985102-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> It is not particularly common among birds, though species from at least 43 <a href="/wiki/Family_(biology)" title="Family (biology)">families</a> are known to engage in the mutual activity.<sup id="cite_ref-FOOTNOTEWilson2000208_49-0" class="reference"><a href="#cite_note-FOOTNOTEWilson2000208-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Forsman_50-0" class="reference"><a href="#cite_note-Forsman-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> Most allopreening activity concentrates on the head and neck, a lesser amount being directed towards the <dfn><a href="/wiki/Glossary_of_bird_terms#breast" title="Glossary of bird terms"><span title="See entry at: Glossary of bird terms § breast" style="color:inherit;" class="glossary-link">breast</span></a></dfn> and <dfn><a href="/wiki/Glossary_of_bird_terms#mantle" title="Glossary of bird terms"><span title="See entry at: Glossary of bird terms § mantle" style="color:inherit;" class="glossary-link">mantle</span></a></dfn> and an even smaller percentage applied to the <dfn><a href="/wiki/Glossary_of_bird_terms#flanks" title="Glossary of bird terms"><span title="See entry at: Glossary of bird terms § flanks" style="color:inherit;" class="glossary-link">flanks</span></a></dfn>. A few species are known to allopreen other areas, including the rump, tail, belly and underwing.<sup id="cite_ref-Harrison_51-0" class="reference"><a href="#cite_note-Harrison-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> </p><p>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.<sup id="cite_ref-Forsman_50-1" class="reference"><a href="#cite_note-Forsman-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> These functions are not mutually exclusive.<sup id="cite_ref-Radford_52-0" class="reference"><a href="#cite_note-Radford-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> Evidence suggests that different species may participate for different reasons, and that those reasons may change depending on the season and the individuals involved.<sup id="cite_ref-Forsman_50-2" class="reference"><a href="#cite_note-Forsman-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> In most cases, allopreening involves members of the same species, although some cases of interspecific allopreening are known; the vast majority of these involve <a href="/wiki/Icterid" title="Icterid">icterids</a>, though at least one instance of mutual grooming between a wild <a href="/wiki/Black_vulture" title="Black vulture">black vulture</a> and a wild <a href="/wiki/Crested_caracara" title="Crested caracara">crested caracara</a> has been documented.<sup id="cite_ref-Ng_53-0" class="reference"><a href="#cite_note-Ng-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> Birds seeking allopreening adopt specific, ritualised postures to signal so; they may fluff their feathers out or put their heads down.<sup id="cite_ref-FOOTNOTEWilson2000209_54-0" class="reference"><a href="#cite_note-FOOTNOTEWilson2000209-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> Captive birds of social species that normally live in flocks, such as parrots, will regularly solicit preening from their human owners.<sup id="cite_ref-FOOTNOTERowley1997257_55-0" class="reference"><a href="#cite_note-FOOTNOTERowley1997257-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> </p><p>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.<sup id="cite_ref-FOOTNOTELoonLoon200536_56-0" class="reference"><a href="#cite_note-FOOTNOTELoonLoon200536-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEOlsenJoseph2011249_57-0" class="reference"><a href="#cite_note-FOOTNOTEOlsenJoseph2011249-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> It may also help to remove <a href="/wiki/Ectoparasite" class="mw-redirect" title="Ectoparasite">ectoparasites</a> from those hard-to-reach areas. Allopreening is most common among species that are regularly in close physical contact due to <a href="/wiki/Flock_(birds)" title="Flock (birds)">flocking</a> or social behaviours, where such contact allows for easier transfer of ectoparasites between individuals.<sup id="cite_ref-Kenny_48-1" class="reference"><a href="#cite_note-Kenny-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEWilson2000209_54-1" class="reference"><a href="#cite_note-FOOTNOTEWilson2000209-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> In one study, <a href="/wiki/Macaroni_penguin" title="Macaroni penguin">Macaroni penguins</a> that frequently allopreened had significantly fewer <a href="/wiki/Tick" title="Tick">ticks</a> on their heads and necks than those that did not.<sup id="cite_ref-FOOTNOTEDeemingReynolds201594_58-0" class="reference"><a href="#cite_note-FOOTNOTEDeemingReynolds201594-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Green_wood_hoopoe" title="Green wood hoopoe">Green wood hoopoes</a>, 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.<sup id="cite_ref-Radford_52-1" class="reference"><a href="#cite_note-Radford-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Great_Woodswallow_group.jpg" class="mw-file-description"><img alt="Five small black-and-white birds perch on a branch; three of them are preening themselves, and a fourth is preening the fifth one's head." src="//upload.wikimedia.org/wikipedia/commons/thumb/0/03/Great_Woodswallow_group.jpg/220px-Great_Woodswallow_group.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/03/Great_Woodswallow_group.jpg/330px-Great_Woodswallow_group.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/03/Great_Woodswallow_group.jpg/440px-Great_Woodswallow_group.jpg 2x" data-file-width="1024" data-file-height="768" /></a><figcaption>Allopreening can reduce or redirect <a href="/wiki/Agonistic_behaviour" title="Agonistic behaviour">aggression</a> within a flock, as with these <a href="/wiki/Great_woodswallow" title="Great woodswallow">great woodswallows</a>.</figcaption></figure> <p>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 <a href="/wiki/Pair_bond" title="Pair bond">pair bonds</a>.<sup id="cite_ref-FOOTNOTECampbellLack1985102_14-5" class="reference"><a href="#cite_note-FOOTNOTECampbellLack1985102-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> 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.<sup id="cite_ref-Kenny_48-2" class="reference"><a href="#cite_note-Kenny-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> 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.<sup id="cite_ref-Harrison_51-1" class="reference"><a href="#cite_note-Harrison-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> </p><p>Allopreening appears to reduce the incidence of conflict between members of some colonially living or colonially nesting species. Neighbouring <a href="/wiki/Common_guillemot" class="mw-redirect" title="Common guillemot">common guillemots</a> 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.<sup id="cite_ref-Lewis_59-0" class="reference"><a href="#cite_note-Lewis-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> Among social flocks of green wood hoopoes, rates of body allopreening (that is, 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.<sup id="cite_ref-Radford_52-2" class="reference"><a href="#cite_note-Radford-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Potential_problems">Potential problems</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Preening&action=edit&section=7" title="Edit section: Potential problems"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Oiled_bird_3.jpg" class="mw-file-description"><img alt="A duck coated in oil sits on a stone wall with its head turned to the side and the tip of its bill resting on its feathers." src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3f/Oiled_bird_3.jpg/220px-Oiled_bird_3.jpg" decoding="async" width="220" height="194" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3f/Oiled_bird_3.jpg/330px-Oiled_bird_3.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3f/Oiled_bird_3.jpg/440px-Oiled_bird_3.jpg 2x" data-file-width="2440" data-file-height="2152" /></a><figcaption>Oiled birds, like this <a href="/wiki/Surf_scoter" title="Surf scoter">surf scoter</a>, can quickly ingest harmful or fatal amounts of oil during their attempts to preen.</figcaption></figure> <p>If birds are exposed to some <a href="/wiki/Pollutant" title="Pollutant">pollutants</a>, such as leaking <a href="/wiki/Petroleum" title="Petroleum">petroleum</a>, they can quickly lose the preen oil from their feathers. This causes a loss of <a href="/wiki/Thermoregulation" title="Thermoregulation">heat regulation</a> and waterproofing, and rapidly leads to the bird becoming chilled.<sup id="cite_ref-FOOTNOTEKricher2020118_37-1" class="reference"><a href="#cite_note-FOOTNOTEKricher2020118-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> 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.<sup id="cite_ref-FOOTNOTEEhrlichDobkinWheyePimms1994225_60-0" class="reference"><a href="#cite_note-FOOTNOTEEhrlichDobkinWheyePimms1994225-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> While preening in an effort to clean themselves, they may ingest harmfully large amounts of the petroleum.<sup id="cite_ref-FOOTNOTEKricher2020118_37-2" class="reference"><a href="#cite_note-FOOTNOTEKricher2020118-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> Ingested oil can cause <a href="/wiki/Pneumonia" title="Pneumonia">pneumonia</a>, 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.<sup id="cite_ref-FOOTNOTEEhrlichDobkinWheyePimms1994225_60-1" class="reference"><a href="#cite_note-FOOTNOTEEhrlichDobkinWheyePimms1994225-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> </p><p>Allopreening may facilitate disease transmission between infected and non-infected individuals. <a href="/wiki/West_Nile_virus" title="West Nile virus">West Nile virus</a> has been found in the feather pulp of several species of <a href="/wiki/Corvid" class="mw-redirect" title="Corvid">corvid</a>, for instance, meaning that birds that preen infected partners might become infected themselves.<sup id="cite_ref-Docherty_61-0" class="reference"><a href="#cite_note-Docherty-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> Even preening its own body may expose a bird to pathogens. There is evidence that water-borne <a href="/wiki/Avian_influenza" title="Avian influenza">avian influenza</a> virus is "captured" by the preen oil on feathers, providing a possible route for infection.<sup id="cite_ref-Delogu_62-0" class="reference"><a href="#cite_note-Delogu-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Karunakaran_63-0" class="reference"><a href="#cite_note-Karunakaran-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> The ingestion of parasites during preening may result in infection; the tick-borne disease <a href="/wiki/Louping_ill_virus" class="mw-redirect" title="Louping ill virus">louping ill virus</a> can be transmitted to <a href="/wiki/Red_grouse" title="Red grouse">red grouse</a> if the bird consumes a tick carrying the disease.<sup id="cite_ref-Gilbert_64-0" class="reference"><a href="#cite_note-Gilbert-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Cagebird" class="mw-redirect" title="Cagebird">Caged birds</a>, particularly parrots, sometimes overpreen in response to being exposed to strong scents (such as nicotine or air fresheners) or as a result of <a href="/wiki/Neuropathy" class="mw-redirect" title="Neuropathy">neuropathy</a>. Reducing exposure to the offending odour, or treating the underlying cause of the neuropathy (such as injury, infection, or <a href="/wiki/Toxic_heavy_metal" title="Toxic heavy metal">heavy metal intoxication</a>) can help to eliminate the behaviour.<sup id="cite_ref-FOOTNOTEKubiak2021175_65-0" class="reference"><a href="#cite_note-FOOTNOTEKubiak2021175-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> Confining a bird with an incompatible or very dominant cage mate can lead to excessive allopreening, which can result in <a href="/wiki/Feather_plucking" class="mw-redirect" title="Feather plucking">feather plucking</a> or injury.<sup id="cite_ref-FOOTNOTEColes200746_66-0" class="reference"><a href="#cite_note-FOOTNOTEColes200746-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Preening&action=edit&section=8" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-OED-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-OED_1-0">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20170806142240/https://en.oxforddictionaries.com/definition/preen">"Preen"</a>. <i>Oxford English Living Dictionary</i>. Oxford University Press. Archived from <a rel="nofollow" class="external text" href="https://en.oxforddictionaries.com/definition/preen">the original</a> on August 6, 2017<span class="reference-accessdate">. Retrieved <span class="nowrap">6 August</span> 2017</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Oxford+English+Living+Dictionary&rft.atitle=Preen&rft_id=https%3A%2F%2Fen.oxforddictionaries.com%2Fdefinition%2Fpreen&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></span> </li> <li id="cite_note-FOOTNOTEMoon-FandliDodmanO'Sullivan199968-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-FOOTNOTEMoon-FandliDodmanO'Sullivan199968_2-0">^</a></b></span> <span class="reference-text"><a href="#CITEREFMoon-FandliDodmanO'Sullivan1999">Moon-Fandli, Dodman & O'Sullivan 1999</a>, p. 68.</span> </li> <li id="cite_note-FOOTNOTELuescher2006198-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-FOOTNOTELuescher2006198_3-0">^</a></b></span> <span class="reference-text"><a href="#CITEREFLuescher2006">Luescher 2006</a>, p. 198.</span> </li> <li id="cite_note-Necker-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-Necker_4-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNecker1985" class="citation journal cs1">Necker, Reinhold (May 1985). 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(7 May 2004). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1810039">"Ticks need not bite their red grouse hosts to infect them with louping ill virus"</a>. <i>Proceedings of the Royal Society B: Biological Sciences</i>. <b>271</b> (Suppl 4): S202-5. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frsbl.2003.0147">10.1098/rsbl.2003.0147</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0962-8452">0962-8452</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1810039">1810039</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/15252984">15252984</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+Royal+Society+B%3A+Biological+Sciences&rft.atitle=Ticks+need+not+bite+their+red+grouse+hosts+to+infect+them+with+louping+ill+virus&rft.volume=271&rft.issue=Suppl+4&rft.pages=S202-5&rft.date=2004-05-07&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1810039%23id-name%3DPMC&rft.issn=0962-8452&rft_id=info%3Apmid%2F15252984&rft_id=info%3Adoi%2F10.1098%2Frsbl.2003.0147&rft.aulast=Gilbert&rft.aufirst=Lucy&rft.au=Jones%2C+Linda+D.&rft.au=Laurenson%2C+M.+Karen&rft.au=Gould%2C+Ernie+A.&rft.au=Reid%2C+Hugh+W.&rft.au=Hudson%2C+Peter+J.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1810039&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></span> </li> <li id="cite_note-FOOTNOTEKubiak2021175-65"><span class="mw-cite-backlink"><b><a href="#cite_ref-FOOTNOTEKubiak2021175_65-0">^</a></b></span> <span class="reference-text"><a href="#CITEREFKubiak2021">Kubiak 2021</a>, p. 175.</span> </li> <li id="cite_note-FOOTNOTEColes200746-66"><span class="mw-cite-backlink"><b><a href="#cite_ref-FOOTNOTEColes200746_66-0">^</a></b></span> <span class="reference-text"><a href="#CITEREFColes2007">Coles 2007</a>, p. 46.</span> </li> </ol></div></div> <div class="mw-heading mw-heading3"><h3 id="Works_cited">Works cited</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Preening&action=edit&section=9" title="Edit section: Works cited"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin refbegin-columns references-column-width" style="column-width: 30em"> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCampbellLack1985" class="citation book cs1">Campbell, Bruce & Lack, Elizabeth, eds. (1985). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/dictionaryofbird0000unse"><i>A Dictionary of Birds</i></a></span>. Carlton, UK: T and A D Poyser. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-85661-039-4" title="Special:BookSources/978-0-85661-039-4"><bdi>978-0-85661-039-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=A+Dictionary+of+Birds&rft.place=Carlton%2C+UK&rft.pub=T+and+A+D+Poyser&rft.date=1985&rft.isbn=978-0-85661-039-4&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fdictionaryofbird0000unse&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCarnaby2008" class="citation book cs1">Carnaby, Trevor (2008). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=hh11gLdZyDgC&pg=PA358"><i>Beat about the Bush: Birds</i></a>. Johannesburg, South Africa: Jacana Media. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-77009-241-9" title="Special:BookSources/978-1-77009-241-9"><bdi>978-1-77009-241-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Beat+about+the+Bush%3A+Birds&rft.place=Johannesburg%2C+South+Africa&rft.pub=Jacana+Media&rft.date=2008&rft.isbn=978-1-77009-241-9&rft.aulast=Carnaby&rft.aufirst=Trevor&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3Dhh11gLdZyDgC%26pg%3DPA358&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFColes2007" class="citation book cs1">Coles, Brian H., ed. (2007). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=Y__okjurvl8C&pg=PA46"><i>Essentials of Avian Medicine and Surgery</i></a> (3rd ed.). Oxford, UK: Blackwood Publishing Ltd. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4051-5755-1" title="Special:BookSources/978-1-4051-5755-1"><bdi>978-1-4051-5755-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Essentials+of+Avian+Medicine+and+Surgery&rft.place=Oxford%2C+UK&rft.edition=3rd&rft.pub=Blackwood+Publishing+Ltd.&rft.date=2007&rft.isbn=978-1-4051-5755-1&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DY__okjurvl8C%26pg%3DPA46&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFde_Juana1992" class="citation encyclopaedia cs1 cs1-prop-long-vol">de Juana, Eduardo (1992). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/handbookofbirdso0001unse">"Class Aves (Birds)"</a></span>. In del Hoyo, Josep; Elliott, Andrew & Sargatal, Jordi (eds.). <i><a href="/wiki/Handbook_of_the_Birds_of_the_World" title="Handbook of the Birds of the World">Handbook of the Birds of the World</a></i>. Vol. 1: Ostrich to Ducks. Barcelona, Spain: Lynx Edicions. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-84-87334-10-8" title="Special:BookSources/978-84-87334-10-8"><bdi>978-84-87334-10-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Class+Aves+%28Birds%29&rft.btitle=Handbook+of+the+Birds+of+the+World&rft.place=Barcelona%2C+Spain&rft.pub=Lynx+Edicions&rft.date=1992&rft.isbn=978-84-87334-10-8&rft.aulast=de+Juana&rft.aufirst=Eduardo&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fhandbookofbirdso0001unse&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDeemingReynolds2015" class="citation book cs1">Deeming, D. Charles & Reynolds, S. James, eds. (2015). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=w3yECgAAQBAJ&pg=PA94"><i>Nests, Eggs, and Incubation: New ideas about avian reproduction</i></a>. Oxford, UK: Oxford University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-871866-6" title="Special:BookSources/978-0-19-871866-6"><bdi>978-0-19-871866-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Nests%2C+Eggs%2C+and+Incubation%3A+New+ideas+about+avian+reproduction&rft.place=Oxford%2C+UK&rft.pub=Oxford+University+Press&rft.date=2015&rft.isbn=978-0-19-871866-6&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3Dw3yECgAAQBAJ%26pg%3DPA94&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEhrlichDobkinWheye1988" class="citation book cs1">Ehrlich, Paul R.; Dobkin, David S. & Wheye, Darryl (1988). <i>The Birder's Handbook</i>. New York, NY: Simon and Schuster. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-671-62133-9" title="Special:BookSources/978-0-671-62133-9"><bdi>978-0-671-62133-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Birder%27s+Handbook&rft.place=New+York%2C+NY&rft.pub=Simon+and+Schuster&rft.date=1988&rft.isbn=978-0-671-62133-9&rft.aulast=Ehrlich&rft.aufirst=Paul+R.&rft.au=Dobkin%2C+David+S.&rft.au=Wheye%2C+Darryl&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEhrlichDobkinWheyePimms1994" class="citation book cs1">Ehrlich, Paul R.; Dobkin, David S.; Wheye, Darryl & Pimms, Stuart L. (1994). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/birdwatchershand00ehrl"><i>The Birdwatcher's Handbook</i></a></span>. Oxford, UK: Oxford University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-858407-0" title="Special:BookSources/978-0-19-858407-0"><bdi>978-0-19-858407-0</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Birdwatcher%27s+Handbook&rft.place=Oxford%2C+UK&rft.pub=Oxford+University+Press&rft.date=1994&rft.isbn=978-0-19-858407-0&rft.aulast=Ehrlich&rft.aufirst=Paul+R.&rft.au=Dobkin%2C+David+S.&rft.au=Wheye%2C+Darryl&rft.au=Pimms%2C+Stuart+L.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fbirdwatchershand00ehrl&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFElphickDunning2001" class="citation book cs1">Elphick, Chris & Dunning, John B. Jr. (2001). "Behaviour". In Elphick, Chris; Dunning, John B. Jr. & Sibley, David (eds.). <i>The Sibley Guide to Bird Life & Behaviour</i>. London: Christopher Helm. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0713662504" title="Special:BookSources/978-0713662504"><bdi>978-0713662504</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Behaviour&rft.btitle=The+Sibley+Guide+to+Bird+Life+%26+Behaviour&rft.place=London&rft.pub=Christopher+Helm&rft.date=2001&rft.isbn=978-0713662504&rft.aulast=Elphick&rft.aufirst=Chris&rft.au=Dunning%2C+John+B.+Jr.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGill2007" class="citation book cs1">Gill, Frank B. (2007). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/OrnithologyThirdEditionFrankB.GillW.H.Freeman200671mb"><i>Ornithology</i></a></span> (3rd ed.). New York: Freeman. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-7167-4983-7" title="Special:BookSources/978-0-7167-4983-7"><bdi>978-0-7167-4983-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Ornithology&rft.place=New+York&rft.edition=3rd&rft.pub=Freeman&rft.date=2007&rft.isbn=978-0-7167-4983-7&rft.aulast=Gill&rft.aufirst=Frank+B.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2FOrnithologyThirdEditionFrankB.GillW.H.Freeman200671mb&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHailman1985" class="citation book cs1">Hailman, Jack P. (1985). "Behavior". In Pettingill, Olin Sewall (ed.). <i>Ornithology in Laboratory and Field</i> (5th ed.). Orlando, FL, US: Academic Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-12-552455-1" title="Special:BookSources/978-0-12-552455-1"><bdi>978-0-12-552455-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Behavior&rft.btitle=Ornithology+in+Laboratory+and+Field&rft.place=Orlando%2C+FL%2C+US&rft.edition=5th&rft.pub=Academic+Press&rft.date=1985&rft.isbn=978-0-12-552455-1&rft.aulast=Hailman&rft.aufirst=Jack+P.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHuxleyHardyFord1954" class="citation book cs1">Huxley, Julian; Hardy, A. C. & Ford, E. B., eds. (1954). <i>Evolution as a Process</i>. London: George Allan & Unwin. <a href="/wiki/LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/54001781">54001781</a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/1434718">1434718</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Evolution+as+a+Process&rft.place=London&rft.pub=George+Allan+%26+Unwin&rft.date=1954&rft_id=info%3Aoclcnum%2F1434718&rft_id=info%3Alccn%2F54001781&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKricher2020" class="citation book cs1">Kricher, John (2020). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=Lce_DwAAQBAJ&pg=PA118"><i>Peterson Reference Guide to Bird Behavior</i></a>. Boston, MA, US: Houghton Mifflin Harcourt. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-328-78736-1" title="Special:BookSources/978-1-328-78736-1"><bdi>978-1-328-78736-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Peterson+Reference+Guide+to+Bird+Behavior&rft.place=Boston%2C+MA%2C+US&rft.pub=Houghton+Mifflin+Harcourt&rft.date=2020&rft.isbn=978-1-328-78736-1&rft.aulast=Kricher&rft.aufirst=John&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DLce_DwAAQBAJ%26pg%3DPA118&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKubiak2021" class="citation book cs1">Kubiak, Marie, ed. (2021). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=4YL4DwAAQBAJ&pg=PA175"><i>Handbook of Exotic Pet Medicine</i></a>. Hoboken, NJ, US: Wiley Blackwell. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-119-38994-1" title="Special:BookSources/978-1-119-38994-1"><bdi>978-1-119-38994-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Handbook+of+Exotic+Pet+Medicine&rft.place=Hoboken%2C+NJ%2C+US&rft.pub=Wiley+Blackwell&rft.date=2021&rft.isbn=978-1-119-38994-1&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3D4YL4DwAAQBAJ%26pg%3DPA175&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLoonLoon2005" class="citation book cs1">Loon, Rael & Loon, Hélène (2005). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=rhBbDwAAQBAJ&pg=PA36"><i>Birds: The Inside Story</i></a>. Cape Town, South Africa: Struik. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-77007-151-3" title="Special:BookSources/978-1-77007-151-3"><bdi>978-1-77007-151-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Birds%3A+The+Inside+Story&rft.place=Cape+Town%2C+South+Africa&rft.pub=Struik&rft.date=2005&rft.isbn=978-1-77007-151-3&rft.aulast=Loon&rft.aufirst=Rael&rft.au=Loon%2C+H%C3%A9l%C3%A8ne&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DrhBbDwAAQBAJ%26pg%3DPA36&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLovetteFitzpatrick2016" class="citation book cs1">Lovette, Irby C. & Fitzpatrick, John W., eds. (2016). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=OGyQDAAAQBAJ&pg=PA129"><i>Handbook of Bird Biology</i></a> (3rd ed.). Oxford, UK: John Wiley & Sons. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-118-29105-4" title="Special:BookSources/978-1-118-29105-4"><bdi>978-1-118-29105-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Handbook+of+Bird+Biology&rft.place=Oxford%2C+UK&rft.edition=3rd&rft.pub=John+Wiley+%26+Sons&rft.date=2016&rft.isbn=978-1-118-29105-4&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DOGyQDAAAQBAJ%26pg%3DPA129&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLuescher2006" class="citation book cs1">Luescher, Andrew U., ed. (2006). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=GmRPyX1a9hMC"><i>Manual of Parrot Behavior</i></a>. Ames, IA, US: Blackwell Publishing. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-8138-2749-0" title="Special:BookSources/978-0-8138-2749-0"><bdi>978-0-8138-2749-0</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Manual+of+Parrot+Behavior&rft.place=Ames%2C+IA%2C+US&rft.pub=Blackwell+Publishing&rft.date=2006&rft.isbn=978-0-8138-2749-0&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DGmRPyX1a9hMC&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMoon-FandliDodmanO'Sullivan1999" class="citation book cs1">Moon-Fandli, Alice M.; Dodman, Nicholas A. & O'Sullivan, Richard L. (1999). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=59ZNHpio5eEC&pg=PA68">"Veterinary Models of Compulsive Self-grooming: Parallels with Trichotillamania"</a>. In Stein, Dan J.; Christenson, Gary A. & Hollander, Eric (eds.). <i>Trichotillomania</i>. Washington, DC: American Psychiatric Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-88048-759-7" title="Special:BookSources/978-0-88048-759-7"><bdi>978-0-88048-759-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Veterinary+Models+of+Compulsive+Self-grooming%3A+Parallels+with+Trichotillamania&rft.btitle=Trichotillomania&rft.place=Washington%2C+DC&rft.pub=American+Psychiatric+Press&rft.date=1999&rft.isbn=978-0-88048-759-7&rft.aulast=Moon-Fandli&rft.aufirst=Alice+M.&rft.au=Dodman%2C+Nicholas+A.&rft.au=O%27Sullivan%2C+Richard+L.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3D59ZNHpio5eEC%26pg%3DPA68&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMoss2015" class="citation book cs1">Moss, Stephen (2015). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=hbE-CgAAQBAJ&pg=PA71"><i>Understanding Bird Behaviour</i></a>. London: Bloomsbury. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4729-1206-0" title="Special:BookSources/978-1-4729-1206-0"><bdi>978-1-4729-1206-0</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Understanding+Bird+Behaviour&rft.place=London&rft.pub=Bloomsbury&rft.date=2015&rft.isbn=978-1-4729-1206-0&rft.aulast=Moss&rft.aufirst=Stephen&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DhbE-CgAAQBAJ%26pg%3DPA71&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFOlsenJoseph2011" class="citation book cs1">Olsen, Penny & Joseph, Leo (2011). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=Jsm5AoA4qfcC&pg=PA249"><i>Stray Feathers: Reflections on the Structure, Behaviour and Evolution of Birds</i></a>. Collingwood, VIC, Australia: CSIRO. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-643-09493-2" title="Special:BookSources/978-0-643-09493-2"><bdi>978-0-643-09493-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Stray+Feathers%3A+Reflections+on+the+Structure%2C+Behaviour+and+Evolution+of+Birds&rft.place=Collingwood%2C+VIC%2C+Australia&rft.pub=CSIRO&rft.date=2011&rft.isbn=978-0-643-09493-2&rft.aulast=Olsen&rft.aufirst=Penny&rft.au=Joseph%2C+Leo&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DJsm5AoA4qfcC%26pg%3DPA249&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPerrins2009" class="citation book cs1">Perrins, Christopher M. (2009). <i>The Princeton Encyclopedia of Birds</i>. Princeton, NJ, US: Princeton University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-691-14070-4" title="Special:BookSources/978-0-691-14070-4"><bdi>978-0-691-14070-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Princeton+Encyclopedia+of+Birds&rft.place=Princeton%2C+NJ%2C+US&rft.pub=Princeton+University+Press&rft.date=2009&rft.isbn=978-0-691-14070-4&rft.aulast=Perrins&rft.aufirst=Christopher+M.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRowley1997" class="citation book cs1 cs1-prop-long-vol">Rowley, Ian (1997). "Family Cacatuidae (Cockatoos)". In del Hoyo, Josep; Elliott, Andrew; Sargatal, Jordi (eds.). <i>Handbook of the Birds of the World</i>. Vol. 4: Sandgrouse to Cuckoos. Barcelona: Lynx Edicions. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-84-87334-22-1" title="Special:BookSources/978-84-87334-22-1"><bdi>978-84-87334-22-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Family+Cacatuidae+%28Cockatoos%29&rft.btitle=Handbook+of+the+Birds+of+the+World&rft.place=Barcelona&rft.pub=Lynx+Edicions&rft.date=1997&rft.isbn=978-84-87334-22-1&rft.aulast=Rowley&rft.aufirst=Ian&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWilson2000" class="citation book cs1">Wilson, Edward O. (2000) [1975]. <a rel="nofollow" class="external text" href="https://books.google.com/books?id=v7lV9tz8fXAC&pg=PA208"><i>Sociobiology: The New Synthesis</i></a>. Cambridge, MA, US: Harvard University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-674-00089-6" title="Special:BookSources/978-0-674-00089-6"><bdi>978-0-674-00089-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Sociobiology%3A+The+New+Synthesis&rft.place=Cambridge%2C+MA%2C+US&rft.pub=Harvard+University+Press&rft.date=2000&rft.isbn=978-0-674-00089-6&rft.aulast=Wilson&rft.aufirst=Edward+O.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3Dv7lV9tz8fXAC%26pg%3DPA208&rfr_id=info%3Asid%2Fen.wikipedia.org%3APreening" class="Z3988"></span></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Preening&action=edit&section=10" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235611614">.mw-parser-output .spoken-wikipedia{border:1px solid #a2a9b1;background-color:var(--background-color-interactive-subtle,#f8f9fa);margin:0.5em 0;padding:0.2em;line-height:1.5em;font-size:90%}.mw-parser-output .spoken-wikipedia-header{text-align:center}.mw-parser-output .spoken-wikipedia-listen-to{font-weight:bold}.mw-parser-output .spoken-wikipedia-files{text-align:center;margin-top:10px;margin-bottom:0.4em}.mw-parser-output .spoken-wikipedia-icon{float:left;margin-left:5px;margin-top:10px}.mw-parser-output .spoken-wikipedia-disclaimer{margin-left:60px;margin-top:10px;font-size:95%;line-height:1.4em}.mw-parser-output .spoken-wikipedia-footer{margin-top:10px;text-align:center}@media(min-width:720px){.mw-parser-output .spoken-wikipedia{width:20em;float:right;clear:right;margin-left:1em}}</style><div class="spoken-wikipedia noprint haudio"><div class="spoken-wikipedia-header"><span class="spoken-wikipedia-listen-to">Listen to this article</span> (<span class="duration"><span class="min">22</span> minutes</span>)</div><div class="spoken-wikipedia-files"><figure class="mw-halign-center" typeof="mw:File"><span><audio id="mwe_player_1" controls="" preload="none" data-mw-tmh="" class="mw-file-element" width="200" style="width:200px;" data-durationhint="1324" data-mwtitle="En-Preening-article.ogg" data-mwprovider="wikimediacommons"><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/d/de/En-Preening-article.ogg/En-Preening-article.ogg.mp3" type="audio/mpeg" data-transcodekey="mp3" data-width="0" data-height="0" /><source src="//upload.wikimedia.org/wikipedia/commons/d/de/En-Preening-article.ogg" type="audio/ogg; 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