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class="vector-toc-numb">2</span> <span>Application</span> </div> </a> <button aria-controls="toc-Application-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 Application subsection</span> </button> <ul id="toc-Application-sublist" class="vector-toc-list"> <li id="toc-In_animals" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#In_animals"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>In animals</span> </div> </a> <ul id="toc-In_animals-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Counter-illumination" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Counter-illumination"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Counter-illumination</span> </div> </a> <ul id="toc-Counter-illumination-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Military" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Military"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Military</span> </div> </a> <ul id="toc-Military-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Function" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Function"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Function</span> </div> </a> <button aria-controls="toc-Function-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 Function subsection</span> </button> <ul id="toc-Function-sublist" class="vector-toc-list"> <li id="toc-Flattening_and_background_matching_when_viewed_from_the_side" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Flattening_and_background_matching_when_viewed_from_the_side"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Flattening and background matching when viewed from the side</span> </div> </a> <ul id="toc-Flattening_and_background_matching_when_viewed_from_the_side-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Background_matching_from_above_or_below" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Background_matching_from_above_or_below"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Background matching from above or below</span> </div> </a> <ul id="toc-Background_matching_from_above_or_below-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Outline_obliteration_from_above" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Outline_obliteration_from_above"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Outline obliteration from above</span> </div> </a> <ul id="toc-Outline_obliteration_from_above-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Non-camouflage_theories" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Non-camouflage_theories"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.4</span> <span>Non-camouflage theories</span> </div> </a> <ul id="toc-Non-camouflage_theories-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Evidence" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Evidence"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Evidence</span> </div> </a> <ul id="toc-Evidence-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mechanism" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Mechanism"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Mechanism</span> </div> </a> <ul id="toc-Mechanism-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Reverse_countershading" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Reverse_countershading"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Reverse countershading</span> </div> </a> <ul id="toc-Reverse_countershading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Examples_in_animals" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Examples_in_animals"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Examples in animals</span> </div> </a> <ul id="toc-Examples_in_animals-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Notes" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Notes"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Notes</span> </div> </a> <ul id="toc-Notes-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">10</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Bibliography" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Bibliography"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>Bibliography</span> </div> </a> <button aria-controls="toc-Bibliography-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 Bibliography subsection</span> </button> <ul id="toc-Bibliography-sublist" class="vector-toc-list"> <li id="toc-Pioneering_books" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Pioneering_books"> <div class="vector-toc-text"> <span class="vector-toc-numb">11.1</span> <span>Pioneering books</span> </div> </a> <ul id="toc-Pioneering_books-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-General_reading" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#General_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">11.2</span> <span>General reading</span> </div> </a> <ul id="toc-General_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Journals" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Journals"> <div class="vector-toc-text"> <span class="vector-toc-numb">11.3</span> <span>Journals</span> </div> </a> <ul id="toc-Journals-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label 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mw-list-item"><a href="https://ca.wikipedia.org/wiki/Contraombreig" title="Contraombreig – Catalan" lang="ca" hreflang="ca" data-title="Contraombreig" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Protist%C3%ADn" title="Protistín – Czech" lang="cs" hreflang="cs" data-title="Protistín" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Konterschattierung" title="Konterschattierung – German" lang="de" hreflang="de" data-title="Konterschattierung" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Contracoloraci%C3%B3n" title="Contracoloración – Spanish" lang="es" hreflang="es" data-title="Contracoloración" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Kontra%C5%ADlumo" title="Kontraŭlumo – Esperanto" lang="eo" hreflang="eo" data-title="Kontraŭlumo" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%B6%D8%AF%D8%B3%D8%A7%DB%8C%D9%87" title="ضدسایه – Persian" lang="fa" hreflang="fa" data-title="ضدسایه" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Ombre_invers%C3%A9e" title="Ombre inversée – French" lang="fr" hreflang="fr" data-title="Ombre inversée" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Pembayangan_balik" title="Pembayangan balik – Indonesian" lang="id" hreflang="id" data-title="Pembayangan balik" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%94%D7%A6%D7%9C%D7%9C%D7%AA_%D7%A0%D7%92%D7%93" title="הצללת נגד – Hebrew" lang="he" hreflang="he" data-title="הצללת נגד" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Lawan_penggelapan" title="Lawan penggelapan – Malay" lang="ms" hreflang="ms" data-title="Lawan penggelapan" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Omgekeerde_schaduwwerking" title="Omgekeerde schaduwwerking – Dutch" lang="nl" hreflang="nl" data-title="Omgekeerde schaduwwerking" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%82%AB%E3%82%A6%E3%83%B3%E3%82%BF%E3%83%BC%E3%82%B7%E3%82%A7%E3%83%BC%E3%83%87%E3%82%A3%E3%83%B3%E3%82%B0" title="カウンターシェーディング – Japanese" lang="ja" hreflang="ja" data-title="カウンターシェーディング" 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id="mw-indicator-good-star" class="mw-indicator"><div class="mw-parser-output"><span typeof="mw:File"><a href="/wiki/Wikipedia:Good_articles*" title="This is a good article. Click here for more information."><img alt="This is a good article. Click here for more information." src="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/19px-Symbol_support_vote.svg.png" decoding="async" width="19" height="20" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/29px-Symbol_support_vote.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/39px-Symbol_support_vote.svg.png 2x" data-file-width="180" data-file-height="185" /></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">Camouflage to counteract self-shading</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Tibur%C3%B3n.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/bb/Tibur%C3%B3n.jpg/300px-Tibur%C3%B3n.jpg" decoding="async" width="300" height="300" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/b/bb/Tibur%C3%B3n.jpg 1.5x" data-file-width="303" data-file-height="303" /></a><figcaption>Many animals, such as this <a href="/wiki/Grey_reef_shark" title="Grey reef shark">grey reef shark</a>, are countershaded.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Thayer_Concealing-Coloration_Plate_XII_-_Luna_Caterpillar_a)_in_position_b)_inverted.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Thayer_Concealing-Coloration_Plate_XII_-_Luna_Caterpillar_a%29_in_position_b%29_inverted.jpg/220px-Thayer_Concealing-Coloration_Plate_XII_-_Luna_Caterpillar_a%29_in_position_b%29_inverted.jpg" decoding="async" width="220" height="335" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Thayer_Concealing-Coloration_Plate_XII_-_Luna_Caterpillar_a%29_in_position_b%29_inverted.jpg/330px-Thayer_Concealing-Coloration_Plate_XII_-_Luna_Caterpillar_a%29_in_position_b%29_inverted.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Thayer_Concealing-Coloration_Plate_XII_-_Luna_Caterpillar_a%29_in_position_b%29_inverted.jpg/440px-Thayer_Concealing-Coloration_Plate_XII_-_Luna_Caterpillar_a%29_in_position_b%29_inverted.jpg 2x" data-file-width="1244" data-file-height="1896" /></a><figcaption>Illustration from the artist <a href="/wiki/Abbot_Thayer" class="mw-redirect" title="Abbot Thayer">Abbot Thayer</a>'s 1909 <a href="/wiki/Concealing-Coloration_in_the_Animal_Kingdom" title="Concealing-Coloration in the Animal Kingdom">book on camouflage</a> of a Luna <a href="/wiki/Caterpillar" title="Caterpillar">caterpillar</a> <i><a href="/wiki/Actias_luna" class="mw-redirect" title="Actias luna">Actias luna</a></i><br />a) in position b) inverted.</figcaption></figure> <p><b>Countershading</b>, or <b>Thayer's law</b>, is a method of <a href="/wiki/Camouflage" title="Camouflage">camouflage</a> in which <a href="/wiki/Animal_coloration" title="Animal coloration">an animal's coloration</a> is darker on the top or upper side and lighter on the underside of the body.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This pattern is found in many species of <a href="/wiki/Mammal" title="Mammal">mammals</a>, <a href="/wiki/Reptile" title="Reptile">reptiles</a>, <a href="/wiki/Bird" title="Bird">birds</a>, <a href="/wiki/Fish" title="Fish">fish</a>, and <a href="/wiki/Insect" title="Insect">insects</a>, both in <a href="/wiki/Predator" class="mw-redirect" title="Predator">predators</a> and in <a href="/wiki/Prey" class="mw-redirect" title="Prey">prey</a>. </p><p>When light falls from above on a uniformly coloured three-dimensional object such as a <a href="/wiki/Sphere" title="Sphere">sphere</a>, it makes the upper side appear lighter and the underside darker, grading from one to the other. This pattern of light and shade makes the object appear solid, and therefore easier to detect. The classical form of countershading, discovered in 1909 by the artist <a href="/wiki/Abbott_Handerson_Thayer" title="Abbott Handerson Thayer">Abbott Handerson Thayer</a>, works by counterbalancing the effects of self-shadowing, again typically with grading from dark to light. In theory this could be useful for <a href="/wiki/Military_camouflage" title="Military camouflage">military camouflage</a>, but in practice it has rarely been applied, despite the best efforts of Thayer and, later, in the <a href="/wiki/Second_World_War" class="mw-redirect" title="Second World War">Second World War</a>, of the zoologist <a href="/wiki/Hugh_Cott" class="mw-redirect" title="Hugh Cott">Hugh Cott</a>. </p><p>The precise function of various patterns of animal coloration that have been called countershading has been debated by zoologists such as Hannah Rowland (2009), with the suggestion that there may be multiple functions including flattening and background matching when viewed from the side; background matching when viewed from above or below, implying separate colour schemes for the top and bottom surfaces; outline obliteration from above; and a variety of other largely untested non-camouflage theories. A related mechanism, <a href="/wiki/Counter-illumination" title="Counter-illumination">counter-illumination</a>, adds the creation of light by <a href="/wiki/Bioluminescence" title="Bioluminescence">bioluminescence</a> or lamps to match the actual brightness of a background. Counter-illumination camouflage is common in <a href="/wiki/Marine_organisms" class="mw-redirect" title="Marine organisms">marine organisms</a> such as <a href="/wiki/Squid" title="Squid">squid</a>. It has been studied up to the <a href="/wiki/Prototype" title="Prototype">prototype</a> stage for <a href="/wiki/Military_camouflage" title="Military camouflage">military use</a> in ships and aircraft, but it too has rarely or never been used in warfare. </p><p>The reverse of countershading, with the belly pigmented darker than the back, enhances contrast and so makes animals more conspicuous. It is found in animals that can defend themselves, such as <a href="/wiki/Skunk" title="Skunk">skunks</a>. The pattern is used both in <a href="/wiki/Deimatic_behaviour" title="Deimatic behaviour">startle or deimatic displays</a> and as a <a href="/wiki/Signalling_theory" title="Signalling theory">signal</a> to <a href="/wiki/Aposematism" title="Aposematism">warn off</a> experienced predators. However, animals that habitually live upside-down but lack strong defences, such as the <a href="/wiki/Nile_catfish" class="mw-redirect" title="Nile catfish">Nile catfish</a> and the <a href="/wiki/Luna_moth" title="Luna moth">Luna moth</a> caterpillar, have upside-down countershading for camouflage. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Early_research">Early research</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=1" title="Edit section: Early research"><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:Thayers_ships.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/58/Thayers_ships.jpg/220px-Thayers_ships.jpg" decoding="async" width="220" height="160" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/5/58/Thayers_ships.jpg 1.5x" data-file-width="269" data-file-height="196" /></a><figcaption>Thayer's 1902 patent application. He failed to convince the US Navy.</figcaption></figure> <p>The <a href="/wiki/England" title="England">English</a> <a href="/wiki/Zoologist" class="mw-redirect" title="Zoologist">zoologist</a> <a href="/wiki/Edward_Bagnall_Poulton" title="Edward Bagnall Poulton">Edward Bagnall Poulton</a>, author of <i><a href="/wiki/The_Colours_of_Animals" title="The Colours of Animals">The Colours of Animals</a></i> (1890) discovered the countershading of various insects, including the pupa or <a href="/wiki/Chrysalis" class="mw-redirect" title="Chrysalis">chrysalis</a> of the purple emperor butterfly, <i><a href="/wiki/Apatura_iris" title="Apatura iris">Apatura iris</a></i>,<sup id="cite_ref-Poulton1888_2-0" class="reference"><a href="#cite_note-Poulton1888-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> the caterpillar larvae of the brimstone moth, <i><a href="/wiki/Opisthograptis_luteolata" class="mw-redirect" title="Opisthograptis luteolata">Opisthograptis luteolata</a></i> <sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>a<span class="cite-bracket">]</span></a></sup> and of the peppered moth, <i><a href="/wiki/Biston_betularia" class="mw-redirect" title="Biston betularia">Biston betularia</a></i>.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>b<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Poulton1887_5-0" class="reference"><a href="#cite_note-Poulton1887-5"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> However he did not use the term countershading, nor did he suggest that the effect occurred widely.<sup id="cite_ref-Rowland2009_7-0" class="reference"><a href="#cite_note-Rowland2009-7"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:White_fowl_lacking_countershading.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/03/White_fowl_lacking_countershading.jpg/170px-White_fowl_lacking_countershading.jpg" decoding="async" width="170" height="194" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/03/White_fowl_lacking_countershading.jpg/255px-White_fowl_lacking_countershading.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/03/White_fowl_lacking_countershading.jpg/340px-White_fowl_lacking_countershading.jpg 2x" data-file-width="1447" data-file-height="1652" /></a><figcaption>Thayer's "White fowl, lacking counter-shading, against a flat white cloth."</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Abbott_thayer_countershading.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/76/Abbott_thayer_countershading.jpg/220px-Abbott_thayer_countershading.jpg" decoding="async" width="220" height="148" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/76/Abbott_thayer_countershading.jpg/330px-Abbott_thayer_countershading.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/76/Abbott_thayer_countershading.jpg/440px-Abbott_thayer_countershading.jpg 2x" data-file-width="870" data-file-height="586" /></a><figcaption>A 1917 photograph of a countershading study by Thayer, who became obsessed by the mistaken idea that all animals are countershaded.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>The <a href="/wiki/New_Hampshire" title="New Hampshire">New Hampshire</a> artist <a href="/wiki/Abbott_Handerson_Thayer" title="Abbott Handerson Thayer">Abbott Handerson Thayer</a> was one of the first to study and write about countershading. In his 1909 book <i><a href="/wiki/Concealing-Coloration_in_the_Animal_Kingdom" title="Concealing-Coloration in the Animal Kingdom">Concealing-Coloration in the Animal Kingdom</a></i>, he correctly described and illustrated countershading with photographs and paintings, but wrongly claimed that almost all animals are countershaded.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> For this reason countershading is sometimes called Thayer's law. Thayer wrote: </p> <style data-mw-deduplicate="TemplateStyles:r1244412712">.mw-parser-output .templatequote{overflow:hidden;margin:1em 0;padding:0 32px}.mw-parser-output .templatequotecite{line-height:1.5em;text-align:left;margin-top:0}@media(min-width:500px){.mw-parser-output .templatequotecite{padding-left:1.6em}}</style><blockquote class="templatequote"><p>Animals are painted by Nature darkest on those parts which tend to be most lighted by the sky's light, and <i>vice versa</i>. ... the fact that a vast majority of creatures of the whole animal kingdom wear this gradation, developed to an exquisitely minute degree, and are famous for being hard to see in their homes, speaks for itself.</p><div class="templatequotecite">— <cite>Thayer<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup></cite></div></blockquote> <p>Thayer observed and painted a number of examples, including the Luna moth caterpillar <i><a href="/wiki/Actias_luna" class="mw-redirect" title="Actias luna">Actias luna</a></i>, both in its habitual upside-down feeding position, where its countershading makes it appear flat, and artificially inverted from that position, where sunlight and its inverted countershading combine to make it appear heavily shaded and therefore solid.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Thayer obtained a patent in 1902 to paint warships, both <a href="/wiki/Submarine" title="Submarine">submarines</a> and <a href="/wiki/Warship" title="Warship">surface ships</a>, using countershading,<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> but failed to convince the <a href="/wiki/US_Navy" class="mw-redirect" title="US Navy">US Navy</a> to adopt his ideas.<sup id="cite_ref-Goldstein_13-0" class="reference"><a href="#cite_note-Goldstein-13"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Hugh_Bamford_Cott" class="mw-redirect" title="Hugh Bamford Cott">Hugh Bamford Cott</a> in his 1940 book <i><a href="/wiki/Adaptive_Coloration_in_Animals" title="Adaptive Coloration in Animals">Adaptive Coloration in Animals</a></i> described many instances of countershading, following Thayer in general approach<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> but criticising Thayer's excessive claim ("He says 'All patterns and colors whatsoever of all animals that ever prey or are preyed upon are under certain normal circumstances obliterative.'") that effectively all animals are camouflaged with countershading. Cott called this "Thayer straining the theory to a fantastic extreme".<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>Both Thayer and Cott included in their books photographs of a non-countershaded white cockerel against a white background, to make the point that in Thayer's words "a monochrome object can not be 'obliterated', no matter what its background"<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> or in Cott's words "Colour resemblance alone is not sufficient to afford concealment".<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> Cott explained that </p> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1244412712"><blockquote class="templatequote"><p>Contrary to what might have been expected by any one lacking in artistic perception, the bird appears highly conspicuous, the back looking lighter, and the breast darker, than the background, although in actual fact, back, background and breast are all pure white."<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup></p></blockquote> <div class="mw-heading mw-heading2"><h2 id="Application">Application</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=2" title="Edit section: Application"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="In_animals">In animals</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=3" title="Edit section: In animals"><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:Darga117.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/Darga117.JPG/220px-Darga117.JPG" decoding="async" width="220" height="160" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/Darga117.JPG/330px-Darga117.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/df/Darga117.JPG/440px-Darga117.JPG 2x" data-file-width="1082" data-file-height="789" /></a><figcaption><a href="/wiki/Ibex" title="Ibex">Ibexes</a> are effectively flattened by countershading, making them nearly invisible against a desert background. There are three in the image.</figcaption></figure> <p>Countershading is observed in a wide range of animal groups, both terrestrial, such as <a href="/wiki/Deer" title="Deer">deer</a>, and marine, such as <a href="/wiki/Sharks" class="mw-redirect" title="Sharks">sharks</a>.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> It is the basis of camouflage in both predators and prey.<sup id="cite_ref-cott40_20-0" class="reference"><a href="#cite_note-cott40-20"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> It is used alongside other forms of camouflage including colour matching and disruptive coloration.<sup id="cite_ref-cott40_20-1" class="reference"><a href="#cite_note-cott40-20"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> Among predatory fish, the <a href="/wiki/Mangrove_snapper" title="Mangrove snapper">gray snapper</a>, <i>Lutianus griseus</i>, is effectively flattened by its countershading, while it hunts an "almost invisible" prey, the <a href="/wiki/Atherinomorus" title="Atherinomorus">hardhead silverside</a>, <i>Atherina laticeps</i> which swims over greyish sands.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> Other countershaded marine animals include <a href="/wiki/Blue_shark" title="Blue shark">blue shark</a>, <a href="/wiki/Herring" title="Herring">herring</a>, and <a href="/wiki/Delphinus_delphis" class="mw-redirect" title="Delphinus delphis">dolphin</a>; while fish such as the <a href="/wiki/Scomber_scombrus" class="mw-redirect" title="Scomber scombrus">mackerel</a> and <a href="/wiki/Cobia" title="Cobia">sergeant fish</a> are both countershaded and <a href="/wiki/Patterns_in_nature" title="Patterns in nature">patterned with stripes or spots</a>.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1244412712"><blockquote class="templatequote"><p> It tones the canvas on which are painted the Leopard's spots, the Tiger's stripes ... It is the dress almost universally worn by rodents... It is the essential uniform adopted by Conies, Asses, Antelopes, Deer ... It is repeated extensively among the marsupials ... It provides a basic livery for the great majority of snakes, lizards, and amphibians. Among insects it reaches a fine state of perfection in different caterpillars and grasshoppers. ... It is, however, in rivers, and in the surface waters of the sea, that countershading reaches its maximum development and significance.</p><div class="templatequotecite">— <cite>Hugh Cott<sup id="cite_ref-cott40_20-2" class="reference"><a href="#cite_note-cott40-20"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup></cite></div></blockquote> <p><a href="/wiki/Mesozoic" title="Mesozoic">Mesozoic</a> <a href="/wiki/Marine_reptiles" class="mw-redirect" title="Marine reptiles">marine reptiles</a> had countershading. <a href="/wiki/Fossil" title="Fossil">Fossilised</a> skin pigmented with dark-coloured eumelanin reveals that <a href="/wiki/Ichthyosaur" class="mw-redirect" title="Ichthyosaur">ichthyosaurs</a>, <a href="/wiki/Leatherback_turtle" class="mw-redirect" title="Leatherback turtle">leatherback turtles</a> and <a href="/wiki/Mosasaur" title="Mosasaur">mosasaurs</a> had dark backs and light bellies.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Ornithischian" class="mw-redirect" title="Ornithischian">ornithischian</a> <a href="/wiki/Dinosaur" title="Dinosaur">dinosaur</a> <i><a href="/wiki/Psittacosaurus" title="Psittacosaurus">Psittacosaurus</a></i> similarly appears to have been countershaded, implying that its predators detected their prey by deducing shape from shading. Modelling suggests further that the dinosaur was optimally countershaded for a closed habitat such as a forest.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Counter-illumination">Counter-illumination</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=4" title="Edit section: Counter-illumination"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></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">Main article: <a href="/wiki/Counter-illumination" title="Counter-illumination">Counter-illumination</a></div> <p>Another form of animal camouflage uses <a href="/wiki/Bioluminescence" title="Bioluminescence">bioluminescence</a> to increase the average brightness of an animal to match the brightness of the background.<sup id="cite_ref-Young1977_26-0" class="reference"><a href="#cite_note-Young1977-26"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> This is called <a href="/wiki/Counter-illumination" title="Counter-illumination">counter-illumination</a>. It is common in mid-water <a href="/wiki/Pelagic" class="mw-redirect" title="Pelagic">pelagic</a> fish and invertebrates especially <a href="/wiki/Squid" title="Squid">squid</a>. It makes the counter-illuminated animal practically invisible to predators viewing it from below.<sup id="cite_ref-Young1976_27-0" class="reference"><a href="#cite_note-Young1976-27"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> As such, counter-illumination camouflage can be seen as an extension beyond what countershading can achieve. Where countershading only paints out shadows, counter-illumination can add in actual lights, permitting effective camouflage in changing conditions, including where the background is bright enough to make an animal that is not counter-illuminated appear as a shadow.<sup id="cite_ref-Jones2004_28-0" class="reference"><a href="#cite_note-Jones2004-28"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Military">Military</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=5" title="Edit section: Military"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Countershading, like <a href="/wiki/Counter-illumination" title="Counter-illumination">counter-illumination</a>, has rarely been applied in practice for <a href="/wiki/Military_camouflage" title="Military camouflage">military camouflage</a>, though not because military authorities were unaware of it. Both Abbott Thayer in the <a href="/wiki/First_World_War" class="mw-redirect" title="First World War">First World War</a> and Hugh Cott in the Second World War proposed countershading to their countries' armed forces. They each demonstrated the effectiveness of countershading, without succeeding in persuading their armed forces to adopt the technique, though they influenced military adoption of camouflage in general.<sup id="cite_ref-Goldstein_13-1" class="reference"><a href="#cite_note-Goldstein-13"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p>Cott was a protege of <a href="/wiki/John_Graham_Kerr" title="John Graham Kerr">John Graham Kerr</a> who had quarrelled with <a href="/wiki/Norman_Wilkinson_(artist)" title="Norman Wilkinson (artist)">Norman Wilkinson</a> in the First World War about <a href="/wiki/Dazzle_camouflage" title="Dazzle camouflage">dazzle camouflage</a> for ships. Wilkinson remained influential in 1939 as an inspector of camouflage, so a political argument developed. Cott was invited to camouflage a 12-inch rail-mounted gun, alongside a similar gun camouflaged conventionally. Cott carefully combined disruptive contrast to break up the gun barrel's outlines with countershading to flatten out its appearance as a solid cylinder. The guns were then photographed from the air from various angles, and in Peter Forbes's view "the results were remarkable."<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Cott's gun is "invisible except to the most minute scrutiny by someone who knows exactly where to look and what to look for. The other gun is always highly visible." The authorities hesitated, appearing to be embarrassed by the evidence that Cott was right, and argued that countershading would be too difficult to use as an expert zoologist would be needed to supervise every installation. Cott was posted <a href="/wiki/Middle_East_Command_Camouflage_Directorate" title="Middle East Command Camouflage Directorate">to the Middle East</a>, and Kerr unsuccessfully intervened, pleading for guns to be painted Cott's way and Cott to be brought home.<sup id="cite_ref-Forbes142_30-0" class="reference"><a href="#cite_note-Forbes142-30"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>The Australian zoologist William Dakin in his 1941 book <i>The Art of Camouflage</i> followed Thayer in describing countershading in some detail, and the book was reprinted as a military handbook in 1942. Dakin photographed model birds, much as Thayer and Cott had done, and argued that the shoulders and arms of battledress should be countershaded.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p><p>Countershading was described in the US War Department's 1943 <i>Principles of Camouflage</i>, where after four paragraphs of theory and one on its use in nature, the advice given is that:<sup id="cite_ref-TTT_32-0" class="reference"><a href="#cite_note-TTT-32"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1244412712"><blockquote class="templatequote"><p>Upper surfaces should be painted and textured so as to conform to the color and tone of the surrounding country (background) and the sides graded and toned from this to the <u>white</u> which the under surfaces and parts in shade should be painted.<sup id="cite_ref-TTT_32-1" class="reference"><a href="#cite_note-TTT-32"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup></p></blockquote> <p>Inventors have continued to advocate military usage of countershading, with for example a 2005 US patent for personal camouflage including countershading in the form of "statistical countercoloring" with varying sizes of rounded dark patches on a lighter ground.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p><p>Research by Ariel Tankus and Yehezkel Yeshurun investigating "camouflage breaking", the automated detection of objects such as <a href="/wiki/Tank" title="Tank">tanks</a>, showed that analysing images for <a href="https://en.wiktionary.org/wiki/convex" class="extiw" title="wikt:convex">convexity</a> by looking for graded shadows can "break very strong camouflage, which might delude even human viewers." More precisely, images are searched for places where the gradient of brightness crosses zero, such as the line where a shadow stops becoming darker and starts to become lighter again. The technique defeated camouflage using disruption of edges, but the authors observed that animals with Thayer countershading are using "counter-measures to convexity based detectors", which implied "predators who use convexity based detectors."<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p> <ul class="gallery mw-gallery-nolines center"> <li class="gallerybox" style="width: 205px"> <div class="thumb" style="width: 200px;"><span typeof="mw:File"><a href="/wiki/File:Countershaded_Rail-mounted_Gun_Camouflaged_by_Hugh_Cott_1940.jpg" class="mw-file-description" title="Rail-mounted guns countershaded by Hugh Cott (top) and conventionally camouflaged (middle), August 1940. The British authorities agreed Cott's countershading worked, but refused to adopt it.[28]"><img alt="Rail-mounted guns countershaded by Hugh Cott (top) and conventionally camouflaged (middle), August 1940. The British authorities agreed Cott's countershading worked, but refused to adopt it.[28]" src="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Countershaded_Rail-mounted_Gun_Camouflaged_by_Hugh_Cott_1940.jpg/200px-Countershaded_Rail-mounted_Gun_Camouflaged_by_Hugh_Cott_1940.jpg" decoding="async" width="200" height="143" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Countershaded_Rail-mounted_Gun_Camouflaged_by_Hugh_Cott_1940.jpg/300px-Countershaded_Rail-mounted_Gun_Camouflaged_by_Hugh_Cott_1940.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Countershaded_Rail-mounted_Gun_Camouflaged_by_Hugh_Cott_1940.jpg/400px-Countershaded_Rail-mounted_Gun_Camouflaged_by_Hugh_Cott_1940.jpg 2x" data-file-width="1499" data-file-height="1072" /></a></span></div> <div class="gallerytext">Rail-mounted guns countershaded by <a href="/wiki/Hugh_Cott" class="mw-redirect" title="Hugh Cott">Hugh Cott</a> (top) and conventionally camouflaged (middle), August 1940. The British authorities agreed Cott's countershading worked, but refused to adopt it.<sup id="cite_ref-Forbes142_30-1" class="reference"><a href="#cite_note-Forbes142-30"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup></div> </li> <li class="gallerybox" style="width: 205px"> <div class="thumb" style="width: 200px;"><span typeof="mw:File"><a href="/wiki/File:7.2_inch_howitzer_of_51st_Heavy_Regiment.jpg" class="mw-file-description" title="BL 7.2-inch howitzer with countershaded barrel, September 1944"><img alt="BL 7.2-inch howitzer with countershaded barrel, September 1944" src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f7/7.2_inch_howitzer_of_51st_Heavy_Regiment.jpg/198px-7.2_inch_howitzer_of_51st_Heavy_Regiment.jpg" decoding="async" width="198" height="200" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f7/7.2_inch_howitzer_of_51st_Heavy_Regiment.jpg/297px-7.2_inch_howitzer_of_51st_Heavy_Regiment.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f7/7.2_inch_howitzer_of_51st_Heavy_Regiment.jpg/396px-7.2_inch_howitzer_of_51st_Heavy_Regiment.jpg 2x" data-file-width="792" data-file-height="800" /></a></span></div> <div class="gallerytext"><a href="/wiki/BL_7.2-inch_howitzer" title="BL 7.2-inch howitzer">BL 7.2-inch howitzer</a> with countershaded barrel, September 1944</div> </li> <li class="gallerybox" style="width: 205px"> <div class="thumb" style="width: 200px;"><span typeof="mw:File"><a href="/wiki/File:Sherman_Firefly_9-08-2008_15-05-43.JPG" class="mw-file-description" title="A preserved Sherman Firefly; its gun barrel is countershaded and disruptively patterned to disguise its length."><img alt="A preserved Sherman Firefly; its gun barrel is countershaded and disruptively patterned to disguise its length." src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f8/Sherman_Firefly_9-08-2008_15-05-43.JPG/200px-Sherman_Firefly_9-08-2008_15-05-43.JPG" decoding="async" width="200" height="135" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f8/Sherman_Firefly_9-08-2008_15-05-43.JPG/300px-Sherman_Firefly_9-08-2008_15-05-43.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f8/Sherman_Firefly_9-08-2008_15-05-43.JPG/400px-Sherman_Firefly_9-08-2008_15-05-43.JPG 2x" data-file-width="1520" data-file-height="1027" /></a></span></div> <div class="gallerytext">A preserved <a href="/wiki/Sherman_Firefly" title="Sherman Firefly">Sherman Firefly</a>; its gun barrel is countershaded and <a href="/wiki/Disruptive_coloration" title="Disruptive coloration">disruptively patterned</a> to disguise its length.</div> </li> <li class="gallerybox" style="width: 205px"> <div class="thumb" style="width: 200px;"><span typeof="mw:File"><a href="/wiki/File:Focke-Wulf_Fw_190D-9_outside_USAF.jpg" class="mw-file-description" title="True (graduated from dark to light) countershaded Focke-Wulf Fw 190"><img alt="True (graduated from dark to light) countershaded Focke-Wulf Fw 190" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/67/Focke-Wulf_Fw_190D-9_outside_USAF.jpg/200px-Focke-Wulf_Fw_190D-9_outside_USAF.jpg" decoding="async" width="200" height="133" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/67/Focke-Wulf_Fw_190D-9_outside_USAF.jpg/300px-Focke-Wulf_Fw_190D-9_outside_USAF.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/67/Focke-Wulf_Fw_190D-9_outside_USAF.jpg/400px-Focke-Wulf_Fw_190D-9_outside_USAF.jpg 2x" data-file-width="1800" data-file-height="1196" /></a></span></div> <div class="gallerytext">True (graduated from dark to light) countershaded <a href="/wiki/Focke-Wulf_Fw_190" title="Focke-Wulf Fw 190">Focke-Wulf Fw 190</a></div> </li> </ul> <div class="mw-heading mw-heading2"><h2 id="Function">Function</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=6" title="Edit section: Function"><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:Countershading.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6f/Countershading.svg/330px-Countershading.svg.png" decoding="async" width="330" height="258" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6f/Countershading.svg/495px-Countershading.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6f/Countershading.svg/660px-Countershading.svg.png 2x" data-file-width="952" data-file-height="744" /></a><figcaption>Countershading acts as a form of <a href="/wiki/Camouflage" title="Camouflage">camouflage</a> by 'painting out' the self-shadowing of the body or object. The result is a 'flat' appearance, instead of the 'solid' appearance (with visual convexity) of the body before countershading.</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/List_of_camouflage_methods" title="List of camouflage methods">List of camouflage methods</a></div> <p>Hannah Rowland, reviewing countershading 100 years after Abbott Thayer, observed that countershading, which she defines as "darker pigmentation on those surfaces exposed to the most lighting" is a common but poorly understood aspect of <a href="/wiki/Animal_coloration" title="Animal coloration">animal coloration</a>.<sup id="cite_ref-Rowland2009_7-1" class="reference"><a href="#cite_note-Rowland2009-7"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> She noted there had been "much debate" about how countershading works.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> She considered the evidence for Thayer's theory that this acts as camouflage "by reducing ventral shadowing", and reviewed alternative explanations for countershading.<sup id="cite_ref-Rowland2009_7-2" class="reference"><a href="#cite_note-Rowland2009-7"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>Camouflage theories of countershading, Rowland wrote, include "self-shadow concealment which results in improved background matching when viewed from the side"; "self-shadow concealment that flattens the form when viewed from the side"; "background matching when viewed from above or below"; and "body outline obliteration when viewed from above".<sup id="cite_ref-Rowland2009_7-3" class="reference"><a href="#cite_note-Rowland2009-7"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> These are examined in turn below. </p> <div class="mw-heading mw-heading3"><h3 id="Flattening_and_background_matching_when_viewed_from_the_side">Flattening and background matching when viewed from the side</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=7" title="Edit section: Flattening and background matching when viewed from the side"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1237032888/mw-parser-output/.tmulti">.mw-parser-output .tmulti .multiimageinner{display:flex;flex-direction:column}.mw-parser-output .tmulti .trow{display:flex;flex-direction:row;clear:left;flex-wrap:wrap;width:100%;box-sizing:border-box}.mw-parser-output .tmulti .tsingle{margin:1px;float:left}.mw-parser-output .tmulti .theader{clear:both;font-weight:bold;text-align:center;align-self:center;background-color:transparent;width:100%}.mw-parser-output .tmulti .thumbcaption{background-color:transparent}.mw-parser-output .tmulti .text-align-left{text-align:left}.mw-parser-output .tmulti .text-align-right{text-align:right}.mw-parser-output .tmulti .text-align-center{text-align:center}@media all and (max-width:720px){.mw-parser-output .tmulti .thumbinner{width:100%!important;box-sizing:border-box;max-width:none!important;align-items:center}.mw-parser-output .tmulti .trow{justify-content:center}.mw-parser-output .tmulti .tsingle{float:none!important;max-width:100%!important;box-sizing:border-box;text-align:center}.mw-parser-output .tmulti .tsingle .thumbcaption{text-align:left}.mw-parser-output .tmulti .trow>.thumbcaption{text-align:center}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .tmulti .multiimageinner img{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .tmulti .multiimageinner img{background-color:white}}</style><div class="thumb tmulti tleft"><div class="thumbinner multiimageinner" style="width:258px;max-width:258px"><div class="trow"><div class="tsingle" style="width:127px;max-width:127px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Eastern_Gray_Squirrel_800.jpg" class="mw-file-description"><img alt="Squirrel oriented horizontally" src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f6/Eastern_Gray_Squirrel_800.jpg/125px-Eastern_Gray_Squirrel_800.jpg" decoding="async" width="125" height="107" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f6/Eastern_Gray_Squirrel_800.jpg/188px-Eastern_Gray_Squirrel_800.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f6/Eastern_Gray_Squirrel_800.jpg/250px-Eastern_Gray_Squirrel_800.jpg 2x" data-file-width="800" data-file-height="686" /></a></span></div><div class="thumbcaption">When oriented horizontally, the countershading of the <a href="/wiki/Eastern_gray_squirrel" title="Eastern gray squirrel">eastern gray squirrel</a>, <i>Sciurus carolinensis</i>, helps to "paint out" its ventral shadow.</div></div><div class="tsingle" style="width:127px;max-width:127px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Sciurus_carolinensis-gotigersjf_(cropped).jpg" class="mw-file-description"><img alt="Squirrel oriented vertically" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6b/Sciurus_carolinensis-gotigersjf_%28cropped%29.jpg/125px-Sciurus_carolinensis-gotigersjf_%28cropped%29.jpg" decoding="async" width="125" height="125" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6b/Sciurus_carolinensis-gotigersjf_%28cropped%29.jpg/188px-Sciurus_carolinensis-gotigersjf_%28cropped%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6b/Sciurus_carolinensis-gotigersjf_%28cropped%29.jpg/250px-Sciurus_carolinensis-gotigersjf_%28cropped%29.jpg 2x" data-file-width="1552" data-file-height="1552" /></a></span></div><div class="thumbcaption">When oriented vertically, the squirrel's pale belly is conspicuous rather than camouflaged.</div></div></div></div></div> <p>Cott, like Thayer, argued that countershading would make animals hard to see from the side, as they would "fade into a ghostly elusiveness".<sup id="cite_ref-Cott36_36-0" class="reference"><a href="#cite_note-Cott36-36"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> Rowland notes that Cott is here reviewing Thayer's theory and "reinforcing the view that a gradation in shading would act to eliminate the effects of ventral shadowing."<sup id="cite_ref-Rowland2009_7-4" class="reference"><a href="#cite_note-Rowland2009-7"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Kiltie measured the effect of the countershading of the <a href="/wiki/Eastern_gray_squirrel" title="Eastern gray squirrel">eastern gray squirrel</a>, <i>Sciurus carolinensis</i>, showing that when the squirrel is horizontal the self-shadowing of the belly is partly concealed, but that when the squirrel is vertical (as when climbing a tree trunk) this effect did not occur.<sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p><p>Thayer's original argument, restated by Cott,<sup id="cite_ref-Cott36_36-1" class="reference"><a href="#cite_note-Cott36-36"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> was that nature did the exact opposite with countershading that an artist did with paint when creating the illusion of solid three-dimensionality, namely counteracting the effect of shade to flatten out form. Shading is a powerful cue used by animals in different <a href="/wiki/Phylum" title="Phylum">phyla</a> to identify the shapes of objects. Research with chicks showed that they preferred to peck at grains with shadows falling below them (as if illuminated from above), so both humans and birds may make use of shading as a depth cue.<sup id="cite_ref-Rowland2009_7-5" class="reference"><a href="#cite_note-Rowland2009-7"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Background_matching_from_above_or_below">Background matching from above or below</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=8" title="Edit section: Background matching from above or below"><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:Scomber_scombrus.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c3/Scomber_scombrus.jpg/220px-Scomber_scombrus.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c3/Scomber_scombrus.jpg/330px-Scomber_scombrus.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c3/Scomber_scombrus.jpg/440px-Scomber_scombrus.jpg 2x" data-file-width="2955" data-file-height="2216" /></a><figcaption>The mackerel, <i><a href="/wiki/Scomber_scombrus" class="mw-redirect" title="Scomber scombrus">Scomber scombrus</a></i>, like many <a href="/wiki/Pelagic" class="mw-redirect" title="Pelagic">pelagic</a> fish, is dark above, pale below, camouflaging it against the ocean depths and the bright surface.<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>c<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>A completely different function of animal (and military vehicle) coloration is to camouflage the top and bottom surfaces differently, to match their backgrounds below and above respectively. This was noted, for example, by <a href="/wiki/Frank_Evers_Beddard" title="Frank Evers Beddard">Frank Evers Beddard</a> in 1892: </p> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1244412712"><blockquote class="templatequote"><p>Among <a href="/wiki/Pelagic" class="mw-redirect" title="Pelagic">pelagic</a> fish it is common to find the upper surface dark-coloured and the lower surface white, so that the animal is inconspicuous when seen either from above or below.</p><div class="templatequotecite">— <cite>Frank Evers Beddard<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup></cite></div></blockquote> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Grumman_F6F-5K_Hellcat_(G-50)_AN0622498.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/23/Grumman_F6F-5K_Hellcat_%28G-50%29_AN0622498.jpg/220px-Grumman_F6F-5K_Hellcat_%28G-50%29_AN0622498.jpg" decoding="async" width="220" height="143" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/23/Grumman_F6F-5K_Hellcat_%28G-50%29_AN0622498.jpg/330px-Grumman_F6F-5K_Hellcat_%28G-50%29_AN0622498.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/23/Grumman_F6F-5K_Hellcat_%28G-50%29_AN0622498.jpg/440px-Grumman_F6F-5K_Hellcat_%28G-50%29_AN0622498.jpg 2x" data-file-width="1000" data-file-height="650" /></a><figcaption>Top/bottom countershaded<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>d<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Grumman_F6F_Hellcat" title="Grumman F6F Hellcat">Grumman F6F Hellcat</a></figcaption></figure> <p>Early researchers including <a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a>,<sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> Beddard,<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> Cott<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> and Craik<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> argued that in marine animals including <a href="/wiki/Pelagic" class="mw-redirect" title="Pelagic">pelagic</a> fish such as <a href="/wiki/Marlin" title="Marlin">marlin</a> and <a href="/wiki/Mackerel" title="Mackerel">mackerel</a>, as well as <a href="/wiki/Dolphin" title="Dolphin">dolphins</a>, <a href="/wiki/Shark" title="Shark">sharks</a>, and <a href="/wiki/Penguins" class="mw-redirect" title="Penguins">penguins</a> the upper and lower surfaces are sharply distinct in tone, with a dark upper surface and often a nearly white lower surface. They suggested that when seen from the top, the darker <a href="/wiki/Dorsum_(biology)" class="mw-redirect" title="Dorsum (biology)">dorsal</a> surface of the animal would offer camouflage against the darkness of the deep water below. When seen from below, the lighter ventral area would similarly provide the least possible contrast with the sunlit ocean surface above.<sup id="cite_ref-Rowland2009_7-6" class="reference"><a href="#cite_note-Rowland2009-7"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> There is some evidence for this in birds, where birds that catch fish at a medium depth, rather than at the surface or on the seabed, are more often coloured in this way, and the prey of these birds would see only the underside of the bird.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> Rowland concluded that each possible role for coloration patterns lumped together as "countershading" needs to be evaluated separately, rather than just assuming it functions effectively.<sup id="cite_ref-Rowland2009_7-7" class="reference"><a href="#cite_note-Rowland2009-7"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Outline_obliteration_from_above">Outline obliteration from above</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=9" title="Edit section: Outline obliteration from above"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Rowland (2009) identified an additional mechanism of countershading not previously analysed, namely that a round body such as a cylinder illuminated and seen from above appears to have dark sides. Using a graphics tool, she demonstrated that this effect can be flattened out by countershading. Since predators are known to use edges to identify prey, countershading may therefore, she argues, make prey harder to detect when seen from above.<sup id="cite_ref-Rowland2009_7-8" class="reference"><a href="#cite_note-Rowland2009-7"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Non-camouflage_theories">Non-camouflage theories</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=10" title="Edit section: Non-camouflage theories"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Non-camouflage theories include protection from <a href="/wiki/Ultraviolet" title="Ultraviolet">ultraviolet</a> light; <a href="/wiki/Thermoregulation" title="Thermoregulation">thermoregulation</a>; and protection from abrasion. All three of these "plausible" theories remained largely untested in 2009, according to Rowland.<sup id="cite_ref-Rowland2009_7-9" class="reference"><a href="#cite_note-Rowland2009-7"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Evidence">Evidence</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=11" title="Edit section: Evidence"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Despite demonstrations and examples adduced by Cott and others, little experimental evidence for the effectiveness of countershading was gathered in the century since Thayer's discovery. Experiments in 2009 using artificial prey showed that countershaded objects do have survival benefits<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> and in 2012, a study by William Allen and colleagues showed that countershading in 114 species of <a href="/wiki/Ruminants" class="mw-redirect" title="Ruminants">ruminants</a> closely matched predictions for "self-shadow concealment", the function predicted by Poulton, Thayer and Cott.<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Mechanism">Mechanism</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=12" title="Edit section: Mechanism"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a> has assembled evidence from <a href="/wiki/Embryology" title="Embryology">embryology</a> and <a href="/wiki/Genetics" title="Genetics">genetics</a> to show how <a href="/wiki/Evolution" title="Evolution">evolution</a> has acted at all scales from the whole organism down to individual <a href="/wiki/Gene" title="Gene">genes</a>, <a href="/wiki/Protein" title="Protein">proteins</a> and genetic switches. In the case of countershaded mammals with dark (often brownish) upper parts and lighter (often buff or whitish) under parts, such as in the <a href="/wiki/House_mouse" title="House mouse">house mouse</a>, it is the <a href="/wiki/Agouti_gene" class="mw-redirect" title="Agouti gene"><i>Agouti</i> gene</a> which creates the difference in shading. <i>Agouti</i> encodes for a protein, the <a href="/wiki/Agouti_signalling_peptide" class="mw-redirect" title="Agouti signalling peptide">Agouti signalling peptide</a> (ASP), which specifically inhibits the action of the <a href="/wiki/Melanocortin_1_receptor" title="Melanocortin 1 receptor">Melanocortin 1 receptor</a> (MC1R). In the absence of the Agouti protein, alpha-<a href="/wiki/Melanocyte-stimulating_hormone" title="Melanocyte-stimulating hormone">melanocyte-stimulating hormone</a> stimulates the cells bearing MC1R, <a href="/wiki/Melanocyte" title="Melanocyte">melanocytes</a>, to produce dark <a href="/wiki/Eumelanin" class="mw-redirect" title="Eumelanin">eumelanin</a>, colouring the skin and fur dark brown or black. In the presence of the Agouti protein, the same system produces the lighter-coloured, yellow or red <a href="/wiki/Phaeomelanin" class="mw-redirect" title="Phaeomelanin">phaeomelanin</a>. A genetic switch active in the cells of the <a href="/wiki/Embryo" title="Embryo">embryo</a> that will become the belly skin causes the <i>Agouti</i> gene to become active there, creating the countershading seen in adult mammals.<sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> <span class="anchor" id="Reverse"></span> </p> <div class="mw-heading mw-heading2"><h2 id="Reverse_countershading">Reverse countershading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=13" title="Edit section: Reverse countershading"><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:Honey_badger.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/af/Honey_badger.jpg/220px-Honey_badger.jpg" decoding="async" width="220" height="177" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/af/Honey_badger.jpg/330px-Honey_badger.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/a/af/Honey_badger.jpg 2x" data-file-width="350" data-file-height="282" /></a><figcaption>The <a href="/wiki/Honey_badger" title="Honey badger">honey badger</a> is reverse countershaded, a form of <a href="/wiki/Aposematism" title="Aposematism">aposematism</a> (warning coloration).</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Glaucus_atlanticus_1_cropped.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/21/Glaucus_atlanticus_1_cropped.jpg/170px-Glaucus_atlanticus_1_cropped.jpg" decoding="async" width="170" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/21/Glaucus_atlanticus_1_cropped.jpg/255px-Glaucus_atlanticus_1_cropped.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/21/Glaucus_atlanticus_1_cropped.jpg/340px-Glaucus_atlanticus_1_cropped.jpg 2x" data-file-width="1640" data-file-height="1592" /></a><figcaption>The sea slug <i><a href="/wiki/Glaucus_atlanticus" title="Glaucus atlanticus">Glaucus atlanticus</a></i> swims and is countershaded upside-down.</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Aposematism" title="Aposematism">Aposematism</a> and <a href="/wiki/Advertising_in_biology" title="Advertising in biology">Advertising in biology</a></div> <p>If countershading paints out shadows, the reverse, darkening the belly and lightening the back, would maximise contrast by adding to the natural fall of light. This pattern of <a href="/wiki/Animal_coloration" title="Animal coloration">animal coloration</a> is found in animals such as the <a href="/wiki/Skunk" title="Skunk">skunk</a> and <a href="/wiki/Honey_badger" title="Honey badger">honey badger</a> with strong defences—the offensive stink of the skunk, and the sharp claws, aggressive nature and stink of the honey badger.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> These animals do not run when under attack, but move slowly, often turning to face the danger, and giving <a href="/wiki/Deimatic_behaviour" title="Deimatic behaviour">deimatic or threat displays</a> either to startle inexperienced predators, or as an <a href="/wiki/Aposematism" title="Aposematism">aposematic signal</a>, to warn off experienced ones.<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> </p><p> The caterpillar of the Luna moth, as discovered by Thayer, is in Cott's phrase "countershaded in relation to [its] attitude", i.e. shaded with a light back grading to a dark belly, as is the Nile catfish, <i><a href="/wiki/Synodontis_batensoda" title="Synodontis batensoda">Synodontis batensoda</a></i> for the same reason: these animals (and other caterpillars including <i><a href="/wiki/Automeris_io" title="Automeris io">Automeris io</a></i> and the eyed hawkmoth, <i><a href="/wiki/Smerinthus_ocellatus" title="Smerinthus ocellatus">Smerinthus ocellatus</a></i>) habitually live 'upside down' with the belly uppermost. Similarly in the sea slug <i><a href="/wiki/Glaucus_atlanticus" title="Glaucus atlanticus">Glaucus atlanticus</a></i>, the reverse countershading is associated with inverted habits. These animals are thus employing countershading in the usual way for camouflage.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup></p><div style="clear:both;" class=""></div> <div class="mw-heading mw-heading2"><h2 id="Examples_in_animals">Examples in animals</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=14" title="Edit section: Examples in animals"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul class="gallery mw-gallery-nolines center"> <li class="gallerybox" style="width: 205px"> <div class="thumb" style="width: 200px;"><span typeof="mw:File"><a href="/wiki/File:Tragelaphus_scriptus_(male).jpg" class="mw-file-description" title="Bushbuck, Tragelaphus scriptus, appears almost perfectly even in tone, showing that its countershading has cancelled out its self-shading. The white spots and markings help to disrupt the 'solidity' of the animal further."><img alt="Bushbuck, Tragelaphus scriptus, appears almost perfectly even in tone, showing that its countershading has cancelled out its self-shading. The white spots and markings help to disrupt the 'solidity' of the animal further." src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7a/Tragelaphus_scriptus_%28male%29.jpg/200px-Tragelaphus_scriptus_%28male%29.jpg" decoding="async" width="200" height="150" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7a/Tragelaphus_scriptus_%28male%29.jpg/300px-Tragelaphus_scriptus_%28male%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7a/Tragelaphus_scriptus_%28male%29.jpg/400px-Tragelaphus_scriptus_%28male%29.jpg 2x" data-file-width="2699" data-file-height="2024" /></a></span></div> <div class="gallerytext">Bushbuck, <i><a href="/wiki/Tragelaphus_scriptus" class="mw-redirect" title="Tragelaphus scriptus">Tragelaphus scriptus</a></i>, appears almost perfectly even in tone, showing that its countershading has cancelled out its self-shading. The white spots and markings help to disrupt the 'solidity' of the animal further.</div> </li> <li class="gallerybox" style="width: 205px"> <div class="thumb" style="width: 200px;"><span typeof="mw:File"><a href="/wiki/File:Sylvia_borin_(%C3%96rebro_County).jpg" class="mw-file-description" title="Many birds, such as this garden warbler, Sylvia borin, are countershaded. The lighter belly makes the bird appear almost evenly coloured when seen from the side."><img alt="Many birds, such as this garden warbler, Sylvia borin, are countershaded. The lighter belly makes the bird appear almost evenly coloured when seen from the side." src="//upload.wikimedia.org/wikipedia/commons/thumb/0/07/Sylvia_borin_%28%C3%96rebro_County%29.jpg/200px-Sylvia_borin_%28%C3%96rebro_County%29.jpg" decoding="async" width="200" height="137" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/07/Sylvia_borin_%28%C3%96rebro_County%29.jpg/300px-Sylvia_borin_%28%C3%96rebro_County%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/07/Sylvia_borin_%28%C3%96rebro_County%29.jpg/400px-Sylvia_borin_%28%C3%96rebro_County%29.jpg 2x" data-file-width="1198" data-file-height="823" /></a></span></div> <div class="gallerytext">Many birds, such as this <a href="/wiki/Garden_warbler" title="Garden warbler">garden warbler</a>, <i>Sylvia borin</i>, are countershaded. The lighter belly makes the bird appear almost evenly coloured when seen from the side.</div> </li> <li class="gallerybox" style="width: 205px"> <div class="thumb" style="width: 200px;"><span typeof="mw:File"><a href="/wiki/File:Carolina_anole.jpg" class="mw-file-description" title="The Carolina anole lizard, Anolis carolinensis, is smoothly countershaded."><img alt="The Carolina anole lizard, Anolis carolinensis, is smoothly countershaded." src="//upload.wikimedia.org/wikipedia/commons/thumb/2/23/Carolina_anole.jpg/200px-Carolina_anole.jpg" decoding="async" width="200" height="127" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/2/23/Carolina_anole.jpg 1.5x" data-file-width="300" data-file-height="190" /></a></span></div> <div class="gallerytext">The Carolina anole lizard, <i><a href="/wiki/Carolina_Anole" class="mw-redirect" title="Carolina Anole">Anolis carolinensis</a></i>, is smoothly countershaded.</div> </li> <li class="gallerybox" style="width: 205px"> <div class="thumb" style="width: 200px;"><span typeof="mw:File"><a href="/wiki/File:Adelie_Penguin2.jpg" class="mw-file-description" title="Adelie penguins, Pygoscelis adeliae, are white below and dark above, presumably to enable them to blend with the sea surface when seen from below, and with deep water when seen from above."><img alt="Adelie penguins, Pygoscelis adeliae, are white below and dark above, presumably to enable them to blend with the sea surface when seen from below, and with deep water when seen from above." src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a9/Adelie_Penguin2.jpg/200px-Adelie_Penguin2.jpg" decoding="async" width="200" height="133" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a9/Adelie_Penguin2.jpg/300px-Adelie_Penguin2.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a9/Adelie_Penguin2.jpg/400px-Adelie_Penguin2.jpg 2x" data-file-width="1024" data-file-height="682" /></a></span></div> <div class="gallerytext"><a href="/wiki/Adelie_penguin" class="mw-redirect" title="Adelie penguin">Adelie penguins</a>, <i>Pygoscelis adeliae</i>, are white below and dark above, presumably to enable them to blend with the sea surface when seen from below, and with deep water when seen from above.</div> </li> <li class="gallerybox" style="width: 205px"> <div class="thumb" style="width: 200px;"><span typeof="mw:File"><a href="/wiki/File:Smerinthus_ocellatus_caterpillar_on_apple_tree.jpg" class="mw-file-description" title="The caterpillar larva of the eyed hawkmoth, Smerinthus ocellatus, is reverse countershaded, making it appear flat when upside-down in feeding position."><img alt="The caterpillar larva of the eyed hawkmoth, Smerinthus ocellatus, is reverse countershaded, making it appear flat when upside-down in feeding position." src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d5/Smerinthus_ocellatus_caterpillar_on_apple_tree.jpg/133px-Smerinthus_ocellatus_caterpillar_on_apple_tree.jpg" decoding="async" width="133" height="200" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d5/Smerinthus_ocellatus_caterpillar_on_apple_tree.jpg/199px-Smerinthus_ocellatus_caterpillar_on_apple_tree.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d5/Smerinthus_ocellatus_caterpillar_on_apple_tree.jpg/266px-Smerinthus_ocellatus_caterpillar_on_apple_tree.jpg 2x" data-file-width="2000" data-file-height="3008" /></a></span></div> <div class="gallerytext">The caterpillar larva of the eyed hawkmoth, <i><a href="/wiki/Smerinthus_ocellatus" title="Smerinthus ocellatus">Smerinthus ocellatus</a></i>, is reverse countershaded, making it appear flat when upside-down in feeding position.</div> </li> <li class="gallerybox" style="width: 205px"> <div class="thumb" style="width: 200px;"><span typeof="mw:File"><a href="/wiki/File:Eyed_hawkmoth_larvae_reverse_countershaded.jpg" class="mw-file-description" title="When the eyed hawkmoth caterpillar is turned upright, as here, its countershading adds to the shading caused by sunlight, rather than "painting it out", so its body appears strongly rounded in this position."><img alt="When the eyed hawkmoth caterpillar is turned upright, as here, its countershading adds to the shading caused by sunlight, rather than "painting it out", so its body appears strongly rounded in this position." src="//upload.wikimedia.org/wikipedia/en/thumb/b/b9/Eyed_hawkmoth_larvae_reverse_countershaded.jpg/200px-Eyed_hawkmoth_larvae_reverse_countershaded.jpg" decoding="async" width="200" height="152" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/b/b9/Eyed_hawkmoth_larvae_reverse_countershaded.jpg/300px-Eyed_hawkmoth_larvae_reverse_countershaded.jpg 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/b/b9/Eyed_hawkmoth_larvae_reverse_countershaded.jpg/400px-Eyed_hawkmoth_larvae_reverse_countershaded.jpg 2x" data-file-width="2294" data-file-height="1740" /></a></span></div> <div class="gallerytext">When the eyed hawkmoth caterpillar is turned upright, as here, its countershading adds to the shading caused by sunlight, rather than "painting it out", so its body appears strongly rounded in this position.</div> </li> <li class="gallerybox" style="width: 205px"> <div class="thumb" style="width: 200px;"><span typeof="mw:File"><a href="/wiki/File:Striped_Skunk_(Mephitis_mephitis)_DSC_0030.jpg" class="mw-file-description" title="Striped skunk, Mephitis mephitis, has conspicuous warning coloration with reversed countershading, alerting predators to its powerfully defensive stink."><img alt="Striped skunk, Mephitis mephitis, has conspicuous warning coloration with reversed countershading, alerting predators to its powerfully defensive stink." src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0c/Striped_Skunk_%28Mephitis_mephitis%29_DSC_0030.jpg/200px-Striped_Skunk_%28Mephitis_mephitis%29_DSC_0030.jpg" decoding="async" width="200" height="133" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0c/Striped_Skunk_%28Mephitis_mephitis%29_DSC_0030.jpg/300px-Striped_Skunk_%28Mephitis_mephitis%29_DSC_0030.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0c/Striped_Skunk_%28Mephitis_mephitis%29_DSC_0030.jpg/400px-Striped_Skunk_%28Mephitis_mephitis%29_DSC_0030.jpg 2x" data-file-width="4288" data-file-height="2848" /></a></span></div> <div class="gallerytext">Striped skunk, <i><a href="/wiki/Mephitis_mephitis" class="mw-redirect" title="Mephitis mephitis">Mephitis mephitis</a></i>, has conspicuous <a href="/wiki/Warning_coloration" class="mw-redirect" title="Warning coloration">warning coloration</a> with reversed countershading, alerting predators to its powerfully defensive stink.</div> </li> <li class="gallerybox" style="width: 205px"> <div class="thumb" style="width: 200px;"><span typeof="mw:File"><a href="/wiki/File:Axis_axis_(Nagarhole,_2010).jpg" class="mw-file-description" title="Chital deer, Axis axis. The animals in the background are effectively countershaded with their bodies horizontal, but the upright stag in the foreground is made conspicuous by its light belly. The spotting is disruptive."><img alt="Chital deer, Axis axis. The animals in the background are effectively countershaded with their bodies horizontal, but the upright stag in the foreground is made conspicuous by its light belly. The spotting is disruptive." src="//upload.wikimedia.org/wikipedia/commons/thumb/3/36/Axis_axis_%28Nagarhole%2C_2010%29.jpg/200px-Axis_axis_%28Nagarhole%2C_2010%29.jpg" decoding="async" width="200" height="133" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/36/Axis_axis_%28Nagarhole%2C_2010%29.jpg/300px-Axis_axis_%28Nagarhole%2C_2010%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/36/Axis_axis_%28Nagarhole%2C_2010%29.jpg/400px-Axis_axis_%28Nagarhole%2C_2010%29.jpg 2x" data-file-width="3200" data-file-height="2134" /></a></span></div> <div class="gallerytext">Chital deer, <i><a href="/wiki/Axis_axis" class="mw-redirect" title="Axis axis">Axis axis</a></i>. The animals in the background are effectively countershaded with their bodies horizontal, but the upright stag in the foreground is made conspicuous by its light belly. The spotting is disruptive.</div> </li> </ul> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=15" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><i><a href="/wiki/Synodontis_nigriventris" title="Synodontis nigriventris">Synodontis nigriventris</a></i>, an "upside-down" catfish (with reverse countershading)</li> <li><a href="/wiki/Counterchanging" class="mw-redirect" title="Counterchanging">Counterchanging</a>, a heraldic device of similar appearance</li></ul> <div class="mw-heading mw-heading2"><h2 id="Notes">Notes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=16" title="Edit section: Notes"><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 reflist-lower-alpha"> <div class="mw-references-wrap"><ol class="references"> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text">It was called <i>Rumia crataegata</i> at the time.</span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text">It was called <i>Amphidasis betularia</i> at the time.</span> </li> <li id="cite_note-39"><span class="mw-cite-backlink"><b><a href="#cite_ref-39">^</a></b></span> <span class="reference-text">The mackerel, like many other pelagic fish, is also <a href="/wiki/Silvering_(camouflage)" class="mw-redirect" title="Silvering (camouflage)">camouflaged by silvering</a>, and when seen from above it has a bold <a href="/wiki/Disruptive_pattern" class="mw-redirect" title="Disruptive pattern">disruptive pattern</a>.</span> </li> <li id="cite_note-41"><span class="mw-cite-backlink"><b><a href="#cite_ref-41">^</a></b></span> <span class="reference-text">Colours used are Non-Specular Sea Blue, Intermediate Blue, White.</span> </li> </ol></div></div> <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=Countershading&action=edit&section=17" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239543626"><div class="reflist reflist-columns references-column-width" style="column-width: 22em;"> <ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</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 id="CITEREFArgo2017" class="citation web cs1">Argo, Emily (21 April 2017). <a rel="nofollow" class="external text" href="https://fishionary.fisheries.org/countershading/">"Countershading"</a>. <i>Fishionary</i>. <a href="/wiki/American_Fisheries_Society" title="American Fisheries Society">American Fisheries Society</a><span class="reference-accessdate">. Retrieved <span class="nowrap">17 December</span> 2022</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Fishionary&rft.atitle=Countershading&rft.date=2017-04-21&rft.aulast=Argo&rft.aufirst=Emily&rft_id=https%3A%2F%2Ffishionary.fisheries.org%2Fcountershading%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></span> </li> <li id="cite_note-Poulton1888-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-Poulton1888_2-0">^</a></b></span> <span class="reference-text">Poulton, 1888.</span> </li> <li id="cite_note-Poulton1887-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-Poulton1887_5-0">^</a></b></span> <span class="reference-text">Poulton, 1887.</span> </li> <li id="cite_note-6"><span class="mw-cite-backlink"><b><a href="#cite_ref-6">^</a></b></span> <span class="reference-text">Thayer, 1909. p 22.</span> </li> <li id="cite_note-Rowland2009-7"><span class="mw-cite-backlink">^ <a href="#cite_ref-Rowland2009_7-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Rowland2009_7-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Rowland2009_7-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Rowland2009_7-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Rowland2009_7-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Rowland2009_7-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Rowland2009_7-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-Rowland2009_7-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-Rowland2009_7-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-Rowland2009_7-9"><sup><i><b>j</b></i></sup></a></span> <span class="reference-text">Rowland, 2009.</span> </li> <li id="cite_note-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-8">^</a></b></span> <span class="reference-text">Forbes, 2009. pp. 76–79.</span> </li> <li id="cite_note-9"><span class="mw-cite-backlink"><b><a href="#cite_ref-9">^</a></b></span> <span class="reference-text">Thayer, 1909.</span> </li> <li id="cite_note-10"><span class="mw-cite-backlink"><b><a href="#cite_ref-10">^</a></b></span> <span class="reference-text">Thayer, 1909. pp 14–15.</span> </li> <li id="cite_note-11"><span class="mw-cite-backlink"><b><a href="#cite_ref-11">^</a></b></span> <span class="reference-text">Thayer, 1909. Plate XII.</span> </li> <li id="cite_note-12"><span class="mw-cite-backlink"><b><a href="#cite_ref-12">^</a></b></span> <span class="reference-text"><span><a rel="nofollow" class="external text" href="https://patents.google.com/patent/US715013">U.S. patent 715,013</a></span></span> </li> <li id="cite_note-Goldstein-13"><span class="mw-cite-backlink">^ <a href="#cite_ref-Goldstein_13-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Goldstein_13-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text">Goldstein, 2009, pp. 233–235.</span> </li> <li id="cite_note-14"><span class="mw-cite-backlink"><b><a href="#cite_ref-14">^</a></b></span> <span class="reference-text">Cott, 1940. pp. 35–46.</span> </li> <li id="cite_note-15"><span class="mw-cite-backlink"><b><a href="#cite_ref-15">^</a></b></span> <span class="reference-text">Cott, 1940. pp. 172–173.</span> </li> <li id="cite_note-16"><span class="mw-cite-backlink"><b><a href="#cite_ref-16">^</a></b></span> <span class="reference-text">Thayer, 1909. Caption to Figure 7.</span> </li> <li id="cite_note-17"><span class="mw-cite-backlink"><b><a href="#cite_ref-17">^</a></b></span> <span class="reference-text">Cott, 1940. Caption to Plate 7.</span> </li> <li id="cite_note-18"><span class="mw-cite-backlink"><b><a href="#cite_ref-18">^</a></b></span> <span class="reference-text">Cott, 1940. p. 35.</span> </li> <li id="cite_note-19"><span class="mw-cite-backlink"><b><a href="#cite_ref-19">^</a></b></span> <span class="reference-text">ONR, 2013.</span> </li> <li id="cite_note-cott40-20"><span class="mw-cite-backlink">^ <a href="#cite_ref-cott40_20-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-cott40_20-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-cott40_20-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text">Cott, 1940. p. 40.</span> </li> <li id="cite_note-21"><span class="mw-cite-backlink"><b><a href="#cite_ref-21">^</a></b></span> <span class="reference-text">Cott, 1940. p37.</span> </li> <li id="cite_note-22"><span class="mw-cite-backlink"><b><a href="#cite_ref-22">^</a></b></span> <span class="reference-text">Cott, 1940. p41</span> </li> <li id="cite_note-23"><span class="mw-cite-backlink"><b><a href="#cite_ref-23">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLindgren,_JohanPeter_SjövallRyan_M._CarneyPer_Uvdal2014" class="citation journal cs1">Lindgren, Johan; Peter Sjövall; Ryan M. 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Skin+pigmentation+provides+evidence+of+convergent+melanism+in+extinct+marine+reptiles&rft.volume=506&rft.issue=7489&rft.pages=484-488&rft.date=2014-02&rft_id=info%3Adoi%2F10.1038%2Fnature12899&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A4468035%23id-name%3DS2CID&rft_id=info%3Apmid%2F24402224&rft_id=info%3Abibcode%2F2014Natur.506..484L&rft.au=Lindgren%2C+Johan&rft.au=Peter+Sj%C3%B6vall&rft.au=Ryan+M.+Carney&rft.au=Per+Uvdal&rft.au=Johan+A.+Gren&rft.au=Gareth+Dyke&rft.au=Bo+Pagh+Schultz&rft.au=Matthew+D.+Shawkey&rft.au=Kenneth+R.+Barnes&rft.au=Michael+J.+Polcyn&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></span> </li> <li id="cite_note-24"><span class="mw-cite-backlink"><b><a href="#cite_ref-24">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLindgrenSjövallThielZheng2018" class="citation journal cs1">Lindgren, Johan; Sjövall, Peter; Thiel, Volker; Zheng, Wenxia; Ito, Shosuke; Wakamatsu, Kazumasa; Hauff, Rolf; Kear, Benjamin P.; Engdahl, Anders; Alwmark, Carl; Eriksson, Mats E.; Jarenmark, Martin; Sachs, Sven; Ahlberg, Per E.; Marone, Federica (December 2018). <a rel="nofollow" class="external text" href="https://www.nature.com/articles/s41586-018-0775-X">"Soft-tissue evidence for homeothermy and crypsis in a Jurassic ichthyosaur"</a>. <i>Nature</i>. <b>564</b> (7736): 359–365. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2018Natur.564..359L">2018Natur.564..359L</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fs41586-018-0775-x">10.1038/s41586-018-0775-x</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/1476-4687">1476-4687</a>. <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/30518862">30518862</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:54458324">54458324</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Soft-tissue+evidence+for+homeothermy+and+crypsis+in+a+Jurassic+ichthyosaur&rft.volume=564&rft.issue=7736&rft.pages=359-365&rft.date=2018-12&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A54458324%23id-name%3DS2CID&rft_id=info%3Abibcode%2F2018Natur.564..359L&rft.issn=1476-4687&rft_id=info%3Adoi%2F10.1038%2Fs41586-018-0775-x&rft_id=info%3Apmid%2F30518862&rft.aulast=Lindgren&rft.aufirst=Johan&rft.au=Sj%C3%B6vall%2C+Peter&rft.au=Thiel%2C+Volker&rft.au=Zheng%2C+Wenxia&rft.au=Ito%2C+Shosuke&rft.au=Wakamatsu%2C+Kazumasa&rft.au=Hauff%2C+Rolf&rft.au=Kear%2C+Benjamin+P.&rft.au=Engdahl%2C+Anders&rft.au=Alwmark%2C+Carl&rft.au=Eriksson%2C+Mats+E.&rft.au=Jarenmark%2C+Martin&rft.au=Sachs%2C+Sven&rft.au=Ahlberg%2C+Per+E.&rft.au=Marone%2C+Federica&rft_id=https%3A%2F%2Fwww.nature.com%2Farticles%2Fs41586-018-0775-X&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></span> </li> <li id="cite_note-25"><span class="mw-cite-backlink"><b><a href="#cite_ref-25">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFVintherNichollsLautenschlagerPittman2016" class="citation journal cs1">Vinther, Jakob; Nicholls, Robert; Lautenschlager, Stephan; Pittman, Michael; Kaye, Thomas G.; Rayfield, Emily; Mayr, Gerald; <a href="/wiki/Innes_Cuthill" title="Innes Cuthill">Cuthill, Innes C.</a> (2016). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5049543">"3D Camouflage in an Ornithischian Dinosaur"</a>. <i>Current Biology</i>. <b>26</b> (18): 2456–2462. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.cub.2016.06.065">10.1016/j.cub.2016.06.065</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/PMC5049543">5049543</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/27641767">27641767</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Current+Biology&rft.atitle=3D+Camouflage+in+an+Ornithischian+Dinosaur&rft.volume=26&rft.issue=18&rft.pages=2456-2462&rft.date=2016&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5049543%23id-name%3DPMC&rft_id=info%3Apmid%2F27641767&rft_id=info%3Adoi%2F10.1016%2Fj.cub.2016.06.065&rft.aulast=Vinther&rft.aufirst=Jakob&rft.au=Nicholls%2C+Robert&rft.au=Lautenschlager%2C+Stephan&rft.au=Pittman%2C+Michael&rft.au=Kaye%2C+Thomas+G.&rft.au=Rayfield%2C+Emily&rft.au=Mayr%2C+Gerald&rft.au=Cuthill%2C+Innes+C.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5049543&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></span> </li> <li id="cite_note-Young1977-26"><span class="mw-cite-backlink"><b><a href="#cite_ref-Young1977_26-0">^</a></b></span> <span class="reference-text">Young and Roper, 1977.</span> </li> <li id="cite_note-Young1976-27"><span class="mw-cite-backlink"><b><a href="#cite_ref-Young1976_27-0">^</a></b></span> <span class="reference-text">Young and Roper, 1976.</span> </li> <li id="cite_note-Jones2004-28"><span class="mw-cite-backlink"><b><a href="#cite_ref-Jones2004_28-0">^</a></b></span> <span class="reference-text">Jones, 2004. p. 1151.</span> </li> <li id="cite_note-29"><span class="mw-cite-backlink"><b><a href="#cite_ref-29">^</a></b></span> <span class="reference-text">Forbes, 2009. p. 150.</span> </li> <li id="cite_note-Forbes142-30"><span class="mw-cite-backlink">^ <a href="#cite_ref-Forbes142_30-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Forbes142_30-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text">Forbes, 2009. pp. 142–146, 149–151, 156.</span> </li> <li id="cite_note-31"><span class="mw-cite-backlink"><b><a href="#cite_ref-31">^</a></b></span> <span class="reference-text">Elias, 2011.</span> </li> <li id="cite_note-TTT-32"><span class="mw-cite-backlink">^ <a href="#cite_ref-TTT_32-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-TTT_32-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text">Anon, 1943.</span> </li> <li id="cite_note-33"><span class="mw-cite-backlink"><b><a href="#cite_ref-33">^</a></b></span> <span class="reference-text">Tooley, 2005.</span> </li> <li id="cite_note-34"><span class="mw-cite-backlink"><b><a href="#cite_ref-34">^</a></b></span> <span class="reference-text">Tankus and Yeshurun, 2001.</span> </li> <li id="cite_note-35"><span class="mw-cite-backlink"><b><a href="#cite_ref-35">^</a></b></span> <span class="reference-text">Rowland, 2011.</span> </li> <li id="cite_note-Cott36-36"><span class="mw-cite-backlink">^ <a href="#cite_ref-Cott36_36-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Cott36_36-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text">Cott, 1940. pp 36–37.</span> </li> <li id="cite_note-37"><span class="mw-cite-backlink"><b><a href="#cite_ref-37">^</a></b></span> <span class="reference-text">Kiltie, 1944.</span> </li> <li id="cite_note-38"><span class="mw-cite-backlink"><b><a href="#cite_ref-38">^</a></b></span> <span class="reference-text">Hershberger, 1970.</span> </li> <li id="cite_note-40"><span class="mw-cite-backlink"><b><a href="#cite_ref-40">^</a></b></span> <span class="reference-text">Beddard, 1892. p. 122.</span> </li> <li id="cite_note-42"><span class="mw-cite-backlink"><b><a href="#cite_ref-42">^</a></b></span> <span class="reference-text">Wallace, 1889, p 193</span> </li> <li id="cite_note-43"><span class="mw-cite-backlink"><b><a href="#cite_ref-43">^</a></b></span> <span class="reference-text">Beddard, 1895, p 115</span> </li> <li id="cite_note-44"><span class="mw-cite-backlink"><b><a href="#cite_ref-44">^</a></b></span> <span class="reference-text">Cott, 1940.</span> </li> <li id="cite_note-45"><span class="mw-cite-backlink"><b><a href="#cite_ref-45">^</a></b></span> <span class="reference-text">Craik, 1944.</span> </li> <li id="cite_note-46"><span class="mw-cite-backlink"><b><a href="#cite_ref-46">^</a></b></span> <span class="reference-text">Ruxton 2004.</span> </li> <li id="cite_note-47"><span class="mw-cite-backlink"><b><a href="#cite_ref-47">^</a></b></span> <span class="reference-text">Rowland et al, 2009.</span> </li> <li id="cite_note-48"><span class="mw-cite-backlink"><b><a href="#cite_ref-48">^</a></b></span> <span class="reference-text">Allen et al, 2012.</span> </li> <li id="cite_note-49"><span class="mw-cite-backlink"><b><a href="#cite_ref-49">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCarroll,_Sean_B.2006" class="citation book cs1"><a href="/wiki/Sean_B._Carroll" title="Sean B. Carroll">Carroll, Sean B.</a> (2006). <i>Endless Forms Most Beautiful</i>. <a href="/wiki/Weidenfeld_and_Nicolson" class="mw-redirect" title="Weidenfeld and Nicolson">Weidenfeld and Nicolson</a>. pp. 229–231, 237. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-297-85094-6" title="Special:BookSources/978-0-297-85094-6"><bdi>978-0-297-85094-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Endless+Forms+Most+Beautiful&rft.pages=229-231%2C+237&rft.pub=Weidenfeld+and+Nicolson&rft.date=2006&rft.isbn=978-0-297-85094-6&rft.au=Carroll%2C+Sean+B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></span> </li> <li id="cite_note-50"><span class="mw-cite-backlink"><b><a href="#cite_ref-50">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20141015182936/http://www.umass.edu/newsoffice/article/black-white-and-stinky-explaining-coloration-skunks-and-other-boldly-colored-animals">"Black, White and Stinky: Explaining Coloration in Skunks and Other Boldly Colored Animals"</a>. University of Massachusetts Amherst. 27 May 2011. Archived from <a rel="nofollow" class="external text" href="http://www.umass.edu/newsoffice/article/black-white-and-stinky-explaining-coloration-skunks-and-other-boldly-colored-animals">the original</a> on 15 October 2014<span class="reference-accessdate">. Retrieved <span class="nowrap">19 June</span> 2014</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Black%2C+White+and+Stinky%3A+Explaining+Coloration+in+Skunks+and+Other+Boldly+Colored+Animals&rft.pub=University+of+Massachusetts+Amherst&rft.date=2011-05-27&rft_id=http%3A%2F%2Fwww.umass.edu%2Fnewsoffice%2Farticle%2Fblack-white-and-stinky-explaining-coloration-skunks-and-other-boldly-colored-animals&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></span> </li> <li id="cite_note-51"><span class="mw-cite-backlink"><b><a href="#cite_ref-51">^</a></b></span> <span class="reference-text">Edmunds, 2008.</span> </li> <li id="cite_note-52"><span class="mw-cite-backlink"><b><a href="#cite_ref-52">^</a></b></span> <span class="reference-text">Cott, 1940. p. 43.</span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Bibliography">Bibliography</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=18" title="Edit section: Bibliography"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Pioneering_books">Pioneering books</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=19" title="Edit section: Pioneering books"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeddard,_Frank_Evers1892" class="citation book cs1"><a href="/wiki/Frank_Evers_Beddard" title="Frank Evers Beddard">Beddard, Frank Evers</a> (1892). <a rel="nofollow" class="external text" href="https://archive.org/details/animalcoloratio00beddgoog"><i>Animal coloration; an account of the principal facts and theories relating to the colours and markings of animals</i></a>. Swan Sonnenschein.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Animal+coloration%3B+an+account+of+the+principal+facts+and+theories+relating+to+the+colours+and+markings+of+animals&rft.pub=Swan+Sonnenschein&rft.date=1892&rft.au=Beddard%2C+Frank+Evers&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fanimalcoloratio00beddgoog&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCott,_Hugh_B.1940" class="citation book cs1"><a href="/wiki/Hugh_Cott" class="mw-redirect" title="Hugh Cott">Cott, Hugh B.</a> (1940). <a href="/wiki/Adaptive_Coloration_in_Animals" title="Adaptive Coloration in Animals"><i>Adaptive Coloration in Animals</i></a>. Oxford University Press.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Adaptive+Coloration+in+Animals&rft.pub=Oxford+University+Press&rft.date=1940&rft.au=Cott%2C+Hugh+B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFThayer,_Gerald_H.1909" class="citation book cs1"><a href="/wiki/Abbott_Thayer" class="mw-redirect" title="Abbott Thayer">Thayer, Gerald H.</a> (1909). <i>Concealing-Coloration in the Animal Kingdom. An Exposition of the Laws of Disguise Through Color and Pattern: Being a Summary of Abbott H. Thayer's Discoveries</i>. Macmillan.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Concealing-Coloration+in+the+Animal+Kingdom.+An+Exposition+of+the+Laws+of+Disguise+Through+Color+and+Pattern%3A+Being+a+Summary+of+Abbott+H.+Thayer%27s+Discoveries&rft.pub=Macmillan&rft.date=1909&rft.au=Thayer%2C+Gerald+H.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWallace,_Alfred_Russel1889" class="citation book cs1"><a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Wallace, Alfred Russel</a> (1889). <a rel="nofollow" class="external text" href="https://archive.org/details/darwinismanexpo04wallgoog"><i>Darwinism. An exposition of the theory of natural selection with some of its applications</i></a>. Macmillan.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Darwinism.+An+exposition+of+the+theory+of+natural+selection+with+some+of+its+applications&rft.pub=Macmillan&rft.date=1889&rft.au=Wallace%2C+Alfred+Russel&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fdarwinismanexpo04wallgoog&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></li></ul> <div class="mw-heading mw-heading3"><h3 id="General_reading">General reading</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=20" title="Edit section: General reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBehrens,_Roy_R.2009" class="citation encyclopaedia cs1"><a href="/wiki/Roy_Behrens" title="Roy Behrens">Behrens, Roy R.</a> (2009). Goldstein, E Bruce (ed.). <i>Encyclopedia of Perception, Volume 1</i>. Sage. pp. 233–235.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Encyclopedia+of+Perception%2C+Volume+1&rft.pages=233-235&rft.pub=Sage&rft.date=2009&rft.au=Behrens%2C+Roy+R.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEdmunds,_Malcolm2008" class="citation encyclopaedia cs1">Edmunds, Malcolm (2008). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=i9ITMiiohVQC&pg=PA1173">"Deimatic Behavior"</a>. In Capinera, John L. (ed.). <i>Encyclopedia of Entomology</i>. Springer. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/9781402062421" title="Special:BookSources/9781402062421"><bdi>9781402062421</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Deimatic+Behavior&rft.btitle=Encyclopedia+of+Entomology&rft.pub=Springer&rft.date=2008&rft.isbn=9781402062421&rft.au=Edmunds%2C+Malcolm&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3Di9ITMiiohVQC%26pg%3DPA1173&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFForbes,_Peter2009" class="citation book cs1">Forbes, Peter (2009). <a href="/wiki/Dazzled_and_Deceived:_Mimicry_and_Camouflage" class="mw-redirect" title="Dazzled and Deceived: Mimicry and Camouflage"><i>Dazzled and Deceived: Mimicry and Camouflage</i></a>. Yale.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Dazzled+and+Deceived%3A+Mimicry+and+Camouflage&rft.pub=Yale&rft.date=2009&rft.au=Forbes%2C+Peter&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRowland,_Hannah_M.2011" class="citation book cs1">Rowland, Hannah M. (2011). "The history, theory and evidence for a cryptic function of countershading". In Stevens, Martin; Merilaita, Sami (eds.). <i>Animal Camouflage: Mechanisms and Function</i>. Cambridge University Press.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=The+history%2C+theory+and+evidence+for+a+cryptic+function+of+countershading&rft.btitle=Animal+Camouflage%3A+Mechanisms+and+Function&rft.pub=Cambridge+University+Press&rft.date=2011&rft.au=Rowland%2C+Hannah+M.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRuxton,_Graeme_D.Sherratt,_Thomas_N.Speed,_Michael_P.2004" class="citation book cs1"><a href="/wiki/Graeme_Ruxton" title="Graeme Ruxton">Ruxton, Graeme D.</a>; <a href="/wiki/Thomas_N._Sherratt" title="Thomas N. Sherratt">Sherratt, Thomas N.</a>; Speed, Michael P. (2004). "3. Countershading and counterillumination". <i>Avoiding Attack: The Evolutionary Ecology of Crypsis, Warning Signals and Mimicry</i>. Oxford University Press.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=3.+Countershading+and+counterillumination&rft.btitle=Avoiding+Attack%3A+The+Evolutionary+Ecology+of+Crypsis%2C+Warning+Signals+and+Mimicry&rft.pub=Oxford+University+Press&rft.date=2004&rft.au=Ruxton%2C+Graeme+D.&rft.au=Sherratt%2C+Thomas+N.&rft.au=Speed%2C+Michael+P.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></li></ul> <div class="mw-heading mw-heading3"><h3 id="Journals">Journals</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Countershading&action=edit&section=21" title="Edit section: Journals"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAllen,_William_L.Baddeley,_RolandCuthill,_Innes_C.Scott-Samuel,_Nicholas_E.2012" class="citation journal cs1">Allen, William L.; Baddeley, Roland; <a href="/wiki/Innes_Cuthill" title="Innes Cuthill">Cuthill, Innes C.</a>; Scott-Samuel, Nicholas E. (December 2012). <a rel="nofollow" class="external text" href="https://cronfa.swan.ac.uk/Record/cronfa28000/Download/0028000-762016114031AM.pdf">"A Quantitative Test of the Predicted Relationship between Countershading and Lighting Environment"</a> <span class="cs1-format">(PDF)</span>. <i>The American Naturalist</i>. <b>180</b> (6): 762–776. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1086%2F668011">10.1086/668011</a>. <a href="/wiki/JSTOR_(identifier)" class="mw-redirect" title="JSTOR (identifier)">JSTOR</a> <a rel="nofollow" class="external text" href="https://www.jstor.org/stable/10.1086/668011">10.1086/668011</a>. <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/23149401">23149401</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:28206975">28206975</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+American+Naturalist&rft.atitle=A+Quantitative+Test+of+the+Predicted+Relationship+between+Countershading+and+Lighting+Environment&rft.volume=180&rft.issue=6&rft.pages=762-776&rft.date=2012-12&rft_id=info%3Apmid%2F23149401&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A28206975%23id-name%3DS2CID&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F10.1086%2F668011%23id-name%3DJSTOR&rft_id=info%3Adoi%2F10.1086%2F668011&rft.au=Allen%2C+William+L.&rft.au=Baddeley%2C+Roland&rft.au=Cuthill%2C+Innes+C.&rft.au=Scott-Samuel%2C+Nicholas+E.&rft_id=https%3A%2F%2Fcronfa.swan.ac.uk%2FRecord%2Fcronfa28000%2FDownload%2F0028000-762016114031AM.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAnon1943" class="citation journal cs1">Anon (4 November 1943). <a rel="nofollow" class="external text" href="http://www.lonesentry.com/articles/ttt09/camouflage.html">"Principles of Camouflage"</a>. <i>Tactical and Technical Trends</i>. <b>37</b>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Tactical+and+Technical+Trends&rft.atitle=Principles+of+Camouflage&rft.volume=37&rft.date=1943-11-04&rft.au=Anon&rft_id=http%3A%2F%2Fwww.lonesentry.com%2Farticles%2Fttt09%2Fcamouflage.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBehrens,_Roy2009" class="citation journal cs1"><a href="/wiki/Roy_Behrens" title="Roy Behrens">Behrens, Roy</a> (27 February 2009). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2674083">"Revisiting Abbott Thayer: non-scientific reflections about camouflage in art, war and zoology"</a>. <i><a href="/wiki/Philosophical_Transactions_of_the_Royal_Society_B" title="Philosophical Transactions of the Royal Society B">Philosophical Transactions of the Royal Society B</a></i>. <b>364</b> (1516). 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"Intensity regulation of bioluminescence during countershading in living midwater animals". <i>Fishery Bulletin</i>. <b>75</b> (2): 239–252.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Fishery+Bulletin&rft.atitle=Intensity+regulation+of+bioluminescence+during+countershading+in+living+midwater+animals&rft.volume=75&rft.issue=2&rft.pages=239-252&rft.date=1977&rft.au=Young%2C+R.E.&rft.au=Roper%2C+C.F.E.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACountershading" class="Z3988"></span></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline 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href="/wiki/Template_talk:Camouflage" title="Template talk:Camouflage"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Camouflage" title="Special:EditPage/Template:Camouflage"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Camouflage" style="font-size:114%;margin:0 4em"><a href="/wiki/Camouflage" title="Camouflage">Camouflage</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/List_of_camouflage_methods" title="List of camouflage methods">Methods</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Camouflage" title="Camouflage">Camouflage</a></li> <li><a class="mw-selflink selflink">Countershading</a></li> <li><a href="/wiki/Active_camouflage" title="Active camouflage">Active camouflage</a></li> <li><a href="/wiki/Counter-illumination" title="Counter-illumination">Counter-illumination</a></li> <li><a href="/wiki/Disruptive_coloration" title="Disruptive coloration">Disruptive coloration</a> <ul><li><a href="/wiki/Coincident_disruptive_coloration" title="Coincident disruptive coloration">Coincident disruptive coloration</a></li> <li><a href="/wiki/Disruptive_eye_mask" title="Disruptive eye mask">Disruptive eye mask</a></li></ul></li> <li><a href="/wiki/Distractive_markings" title="Distractive markings">Distractive markings</a></li> <li><a href="/wiki/Motion_camouflage" title="Motion camouflage">Motion camouflage</a></li> <li><a href="/wiki/Multi-scale_camouflage" title="Multi-scale camouflage">Multi-scale camouflage</a></li> <li><a href="/wiki/Multi-spectral_camouflage" title="Multi-spectral camouflage">Multi-spectral camouflage</a></li> <li><a href="/wiki/Self-decoration_camouflage" title="Self-decoration camouflage">Self-decoration</a></li> <li><a href="/wiki/Snow_camouflage" title="Snow camouflage">Snow camouflage</a></li> <li><a href="/wiki/Urban_camouflage" title="Urban camouflage">Urban camouflage</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="5" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Peacock_Flounder_Bothus_mancus_in_Kona_(vertical).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/34/Peacock_Flounder_Bothus_mancus_in_Kona_%28vertical%29.jpg/100px-Peacock_Flounder_Bothus_mancus_in_Kona_%28vertical%29.jpg" decoding="async" width="100" height="322" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/34/Peacock_Flounder_Bothus_mancus_in_Kona_%28vertical%29.jpg/150px-Peacock_Flounder_Bothus_mancus_in_Kona_%28vertical%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/34/Peacock_Flounder_Bothus_mancus_in_Kona_%28vertical%29.jpg/200px-Peacock_Flounder_Bothus_mancus_in_Kona_%28vertical%29.jpg 2x" data-file-width="2000" data-file-height="6438" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">In nature</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Coloration_evidence_for_natural_selection" title="Coloration evidence for natural selection">As evidence for natural selection</a></li> <li><a href="/wiki/Crypsis" title="Crypsis">Crypsis</a></li> <li><a href="/wiki/Decorator_crab" title="Decorator crab">Decorator crab</a></li> <li><a href="/wiki/Flower_mantis" title="Flower mantis">Flower mantis</a></li> <li><a href="/wiki/Mimicry" title="Mimicry">Mimicry</a> <ul><li><a href="/wiki/Batesian_mimicry" title="Batesian mimicry">Batesian</a></li> <li><a href="/wiki/M%C3%BCllerian_mimicry" title="Müllerian mimicry">Müllerian</a></li> <li><a href="/wiki/Aggressive_mimicry" title="Aggressive mimicry">Aggressive</a></li></ul></li> <li><a href="/wiki/Underwater_camouflage" title="Underwater camouflage">Underwater camouflage</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">People</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Early</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Edward_Bagnall_Poulton" title="Edward Bagnall Poulton">Edward Bagnall Poulton</a> <ul><li><i><a href="/wiki/The_Colours_of_Animals" title="The Colours of Animals">The Colours of Animals</a></i></li></ul></li> <li><a href="/wiki/Abbott_Handerson_Thayer" title="Abbott Handerson Thayer">Abbott Handerson Thayer</a> <ul><li><i><a href="/wiki/Concealing-Coloration_in_the_Animal_Kingdom" title="Concealing-Coloration in the Animal Kingdom">Concealing-Coloration in the Animal Kingdom</a></i></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Camoufleurs" class="mw-redirect" title="Camoufleurs">Camoufleurs</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mary_Taylor_Brush" title="Mary Taylor Brush">Mary Taylor Brush</a></li> <li><a href="/wiki/Lucien-Victor_Guirand_de_Sc%C3%A9vola" title="Lucien-Victor Guirand de Scévola">Lucien-Victor Guirand de Scévola</a></li> <li><a href="/wiki/John_Graham_Kerr" title="John Graham Kerr">John Graham Kerr</a></li> <li><a href="/wiki/Norman_Wilkinson_(artist)" title="Norman Wilkinson (artist)">Norman Wilkinson</a></li> <li><a href="/wiki/Everett_Warner" title="Everett Warner">Everett Warner</a></li> <li><a href="/wiki/Leon_Underwood" title="Leon Underwood">Leon Underwood</a></li> <li><a href="/wiki/Johann_Georg_Otto_Schick" class="mw-redirect" title="Johann Georg Otto Schick">Johann Georg Otto Schick</a></li> <li><a href="/wiki/Hugh_B._Cott" title="Hugh B. Cott">Hugh Cott</a> <ul><li><i><a href="/wiki/Adaptive_Coloration_in_Animals" title="Adaptive Coloration in Animals">Adaptive Coloration in Animals</a></i></li></ul></li> <li><a href="/wiki/Geoffrey_Barkas" title="Geoffrey Barkas">Geoffrey Barkas</a></li> <li><a href="/wiki/Timothy_O%27Neill_(camoufleur)" title="Timothy O'Neill (camoufleur)">Timothy O'Neill</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Researchers</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Roy_Behrens" title="Roy Behrens">Roy Behrens</a></li> <li><a href="/wiki/Tim_Caro" title="Tim Caro">Tim Caro</a></li> <li><a href="/wiki/Innes_Cuthill" title="Innes Cuthill">Innes Cuthill</a></li> <li><a href="/wiki/Thomas_N._Sherratt" title="Thomas N. Sherratt">Thomas N. Sherratt</a></li> <li><a href="/wiki/Martin_Stevens_(biologist)" title="Martin Stevens (biologist)">Martin Stevens</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Military_camouflage" title="Military camouflage">Military</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Topics</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Military_camouflage" title="Military camouflage">Military camouflage</a></li> <li><a href="/wiki/Aircraft_camouflage" title="Aircraft camouflage">Aircraft camouflage</a></li> <li><a href="/wiki/Camouflage_clothing_in_Trinidad_and_Tobago" title="Camouflage clothing in Trinidad and Tobago">Camouflage clothing in Trinidad and Tobago</a></li> <li><a href="/wiki/Dazzle_camouflage" title="Dazzle camouflage">Dazzle camouflage</a></li> <li><a href="/wiki/List_of_countries_that_prohibit_camouflage_clothing" title="List of countries that prohibit camouflage clothing">List of countries that prohibit camouflage clothing</a></li> <li><a href="/wiki/Middle_East_Command_Camouflage_Directorate" title="Middle East Command Camouflage Directorate">Middle East Command Camouflage Directorate</a></li> <li><a href="/wiki/Ship_camouflage" title="Ship camouflage">Ship camouflage</a></li> <li><a href="/wiki/World_War_II_ship_camouflage_measures_of_the_United_States_Navy" title="World War II ship camouflage measures of the United States Navy">USN WWII camouflage measures</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/List_of_military_clothing_camouflage_patterns" title="List of military clothing camouflage patterns">Patterns</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Up to WWII</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/German_World_War_II_camouflage_patterns" title="German World War II camouflage patterns">German<br />WWII</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Splittertarnmuster" title="Splittertarnmuster">Splittertarnmuster</a> (1931)</li> <li><a href="/wiki/Platanenmuster" class="mw-redirect" title="Platanenmuster">Platanenmuster</a> (1937)</li> <li><a href="/wiki/Rauchtarnmuster" class="mw-redirect" title="Rauchtarnmuster">Rauchtarnmuster</a> (1939)</li> <li><a href="/wiki/Palmenmuster" class="mw-redirect" title="Palmenmuster">Palmenmuster</a> (c 1941)</li> <li><a href="/wiki/Sumpfmuster" class="mw-redirect" title="Sumpfmuster">Sumpfmuster</a> (1943)</li> <li><a href="/wiki/Erbsenmuster" title="Erbsenmuster">Erbsenmuster</a> (1944)</li> <li><a href="/wiki/Leibermuster" title="Leibermuster">Leibermuster</a> (1945)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Camouflage_tree" title="Camouflage tree">Camouflage tree</a> (1915)</li> <li><a href="/wiki/Lozenge_camouflage" title="Lozenge camouflage">Lozenge</a> (1917 aircraft)</li> <li><a href="/wiki/Telo_mimetico" title="Telo mimetico">Telo mimetico</a> (1929 tent)</li> <li><a href="/wiki/Denison_smock" title="Denison smock">Denison smock</a> (1941)</li> <li><a href="/wiki/Frog_Skin" title="Frog Skin">Frog Skin</a> (1942)</li> <li><a href="/wiki/Ghillie_suit" title="Ghillie suit">Ghillie suit</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Post-war</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Lizard_(camouflage)" title="Lizard (camouflage)">Lizard</a> (1947)</li> <li><a href="/wiki/Strichtarn" title="Strichtarn">Strichtarn</a> (1960)</li> <li><a href="/wiki/Kamuflirovannyi_Letnyi_Maskirovochnyi_Kombinezon" title="Kamuflirovannyi Letnyi Maskirovochnyi Kombinezon">KLMK</a> (1968)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Late 20th<br />century</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Jigsaw_camouflage" title="Jigsaw camouflage">Jigsaw</a> (1958)</li> <li><a href="/wiki/Tiger_stripe_camouflage" title="Tiger stripe camouflage">Tiger stripe</a> (1962)</li> <li><a href="/wiki/Rhodesian_Brushstroke" title="Rhodesian Brushstroke">Rhodesian Brushstroke</a> (1965)</li> <li><a href="/wiki/ERDL_pattern" title="ERDL pattern">ERDL</a> (1967)</li> <li><a href="/wiki/Disruptive_Pattern_Material" title="Disruptive Pattern Material">Disruptive Pattern Material</a> (1969)</li> <li><a href="/wiki/Wz._68_Moro" title="Wz. 68 Moro">wz. 68 Moro</a> (1969)</li> <li><a href="/wiki/Desert_Battle_Dress_Uniform" title="Desert Battle Dress Uniform">Six-Color Desert Pattern (Chocolate Chip)</a> (1981)</li> <li><a href="/wiki/U.S._Woodland" title="U.S. Woodland">U.S. "M81" Woodland</a> (1981)</li> <li><a href="/wiki/Disruptive_Pattern_Camouflage_Uniform" title="Disruptive Pattern Camouflage Uniform">Australian Disruptive Pattern</a> (1982)</li> <li><a href="/wiki/TAZ_83" title="TAZ 83">TAZ 83</a> (1983)</li> <li><a href="/wiki/Dubok_(camouflage)" title="Dubok (camouflage)">Dubok</a> (1984)</li> <li><a href="/wiki/M84_camouflage_pattern" title="M84 camouflage pattern">M84</a> (1984)</li> <li><a href="/wiki/Type_87_(camouflage)" title="Type 87 (camouflage)">Type 87 (China)</a> (1987)</li> <li><a href="/wiki/Wz._89_Puma" title="Wz. 89 Puma">wz. 89 Puma</a> (1989)</li> <li><a href="/wiki/Camouflage_Daguet" title="Camouflage Daguet">Camouflage Daguet</a> (1989)</li> <li><a href="/wiki/M90_(camouflage)" title="M90 (camouflage)">M90</a> (1990)</li> <li><a href="/wiki/Desert_Night_Camouflage" title="Desert Night Camouflage">Desert Night Camouflage</a> (1990)</li> <li><a href="/wiki/Flecktarn" title="Flecktarn">Flecktarn</a> (1990)</li> <li><a href="/wiki/Tropentarn" title="Tropentarn">Tropentarn</a> (1990)</li> <li><a href="/wiki/Desert_Camouflage_Uniform" title="Desert Camouflage Uniform">Desert Camouflage Pattern</a> (1990)</li> <li><a href="/wiki/Camouflage_Central-Europe" title="Camouflage Central-Europe">Camouflage Central-Europe</a> (1991)</li> <li><a href="/wiki/Soldier_2000" title="Soldier 2000">Soldier 2000</a> (1993)</li> <li><a href="/wiki/TAZ_90" title="TAZ 90">TAZ 90</a> (1993)</li> <li><a href="/wiki/Wz._93_Pantera" title="Wz. 93 Pantera">wz. 93 Pantera</a> (1993)</li> <li><a href="/wiki/CADPAT" title="CADPAT">CADPAT</a> (1997)</li> <li><a href="/wiki/M98_camouflage_pattern" title="M98 camouflage pattern">M98</a> (1998)</li> <li><a href="/wiki/Flora_camouflage" title="Flora camouflage">Flora</a> (1998)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">21st<br />century</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/MARPAT" title="MARPAT">MARPAT</a> (2001) (<a href="/wiki/Marine_Corps_Combat_Utility_Uniform" title="Marine Corps Combat Utility Uniform">Marine Corps Combat Utility Uniform</a> (2002))</li> <li><a href="/wiki/MultiCam" title="MultiCam">MultiCam</a> (2002)</li> <li><a href="/wiki/Tactical_Assault_Camouflage" title="Tactical Assault Camouflage">Tactical Assault Camouflage</a> (2004)</li> <li><a href="/wiki/Universal_Camouflage_Pattern" title="Universal Camouflage Pattern">Universal Camouflage Pattern</a> (2004)</li> <li><a href="/wiki/ESTDCU" title="ESTDCU">ESTDCU</a> (2006)</li> <li><a href="/wiki/M05" title="M05">M05</a> (2007)</li> <li><a href="/wiki/Airman_Battle_Uniform" title="Airman Battle Uniform">Airman Battle Uniform</a> (2007)</li> <li><a href="/wiki/Type_07" title="Type 07">Type 07</a> (2007)</li> <li><a href="/wiki/EMR_camouflage" title="EMR camouflage">EMR</a> (2008)</li> <li><a href="/wiki/Multi-Terrain_Pattern" title="Multi-Terrain Pattern">Multi-Terrain Pattern</a> (2010)</li> <li><a href="/wiki/Australian_Multicam_Camouflage_Uniform" title="Australian Multicam Camouflage Uniform">Australian Multicam</a> (2014)</li> <li><a href="/wiki/Hungarian_camouflage_pattern_2015M" title="Hungarian camouflage pattern 2015M">HunCam</a> (2015)</li> <li><a href="/wiki/Operational_Camouflage_Pattern" title="Operational Camouflage Pattern">Operational Camouflage Pattern</a> (2015)</li> <li><a href="/wiki/Netherlands_Fractal_Pattern" title="Netherlands Fractal Pattern">Netherlands Fractal Pattern</a> (2019)</li> <li><a href="/wiki/Xingkong_(camouflage)" title="Xingkong (camouflage)">Xingkong</a> (2019)</li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Technology</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Deployed</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Berberys-R" title="Berberys-R">Berberys-R</a></li> <li><a href="/wiki/Nakidka" title="Nakidka">Nakidka</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Prototypes</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Diffused_lighting_camouflage" title="Diffused lighting camouflage">Diffused lighting camouflage</a> (1941)</li> <li><a href="/wiki/Yehudi_lights" title="Yehudi lights">Yehudi lights</a> (1943)</li> <li><a href="/wiki/Adaptiv" title="Adaptiv">Adaptiv</a> (2011)</li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Dazzled_and_Deceived" title="Dazzled and Deceived">Dazzled and Deceived</a></i></li> <li><a href="/wiki/Stealth_technology" title="Stealth technology">Stealth technology</a> <ul><li><a href="/wiki/Cloaking_device" title="Cloaking device">Cloaking device</a></li> <li><a href="/wiki/Invisibility" title="Invisibility">Invisibility</a></li></ul></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_ecology" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evo_ecol" title="Template:Evo ecol"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evo_ecol" title="Template talk:Evo ecol"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evo_ecol" title="Special:EditPage/Template:Evo ecol"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_ecology" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Patterns of evolution</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Coloration_evidence_for_natural_selection" title="Coloration evidence for natural selection">Coloration evidence for natural selection</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergent evolution</a> <ul><li><a href="/wiki/List_of_examples_of_convergent_evolution" title="List of examples of convergent evolution">examples</a></li></ul></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergent evolution</a></li> <li><a href="/wiki/Paradox_of_the_plankton" title="Paradox of the plankton">Paradox of the plankton</a></li> <li><a href="/wiki/Predator_satiation" title="Predator satiation">Predator satiation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Signals</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a></li> <li><a href="/wiki/Anti-predator_adaptation" title="Anti-predator adaptation">Anti-predator adaptation</a> <ul><li><a href="/wiki/Alarm_signal" title="Alarm signal">Alarm signal</a></li> <li><a href="/wiki/Aposematism" title="Aposematism">Aposematism</a></li> <li><a href="/wiki/Apparent_death" title="Apparent death">Apparent death</a></li> <li><a href="/wiki/Deimatic_behaviour" title="Deimatic behaviour">Deimatic behaviour</a></li> <li><a href="/wiki/Distraction_display" title="Distraction display">Distraction display</a></li></ul></li> <li class="mw-empty-elt"></li> <li><a href="/wiki/Crypsis" title="Crypsis">Crypsis</a></li> <li><a href="/wiki/Camouflage" title="Camouflage">Camouflage</a></li> <li><a href="/wiki/Mimicry" title="Mimicry">Mimicry</a></li> <li><a href="/wiki/Unkenreflex" title="Unkenreflex">Unkenreflex</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Vision_in_animals" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Vision_in_animals" title="Template:Vision in animals"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Vision_in_animals" title="Template talk:Vision in animals"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Vision_in_animals" title="Special:EditPage/Template:Vision in animals"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Vision_in_animals" style="font-size:114%;margin:0 4em">Vision in animals</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Visual_system" title="Visual system">Vision</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bird_vision" title="Bird vision">Birds</a></li> <li><a href="/wiki/Chameleon_vision" title="Chameleon vision">Chameleons</a></li> <li><a href="/wiki/Dinosaur_vision" title="Dinosaur vision">Dinosaurs</a></li> <li><a href="/wiki/Vision_in_fish" title="Vision in fish">Fish</a></li> <li><a href="/wiki/Vision_in_toads" title="Vision in toads">Toads</a></li> <li><a href="/wiki/Mammalian_vision" title="Mammalian vision">Mammals</a> <ul><li><a href="/wiki/Equine_vision" title="Equine vision">horses</a></li> <li><a href="/wiki/Dog_anatomy#Vision" title="Dog anatomy">dogs</a></li> <li><a href="/wiki/Cat_senses#Sight" title="Cat senses">cats</a></li></ul></li></ul> </div></td><td class="noviewer navbox-image" rowspan="5" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Astronotus_ocellatus_-_closeup_(aka).jpg" class="mw-file-description" title="Fish surveying its world"><img alt="Fish surveying its world" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Astronotus_ocellatus_-_closeup_%28aka%29.jpg/140px-Astronotus_ocellatus_-_closeup_%28aka%29.jpg" decoding="async" width="140" height="93" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Astronotus_ocellatus_-_closeup_%28aka%29.jpg/210px-Astronotus_ocellatus_-_closeup_%28aka%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Astronotus_ocellatus_-_closeup_%28aka%29.jpg/280px-Astronotus_ocellatus_-_closeup_%28aka%29.jpg 2x" data-file-width="3008" data-file-height="2000" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Eye" title="Eye">Eyes</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Arthropod_eye" title="Arthropod eye">Arthropod eye</a></li> <li><a href="/wiki/Compound_eye" title="Compound eye">Compound eye</a></li> <li><a href="/wiki/Eagle_eye" title="Eagle eye">Eagle eye</a></li> <li><a href="/wiki/Tapetum_lucidum" title="Tapetum lucidum">Eye shine</a></li> <li><a href="/wiki/Simple_eye_in_invertebrates" title="Simple eye in invertebrates">Simple eye in invertebrates</a></li> <li><a href="/wiki/Mammalian_eye" title="Mammalian eye">Mammalian eye</a> <ul><li><a href="/wiki/Human_eye" title="Human eye">human</a></li></ul></li> <li><a href="/wiki/Mollusc_eye" title="Mollusc eye">Mollusc eye</a> <ul><li><a href="/wiki/Cephalopod_eye" title="Cephalopod eye">cephalopod</a></li> <li><a href="/wiki/Gastropod_eye" class="mw-redirect" title="Gastropod eye">gastropod</a></li></ul></li> <li><a href="/wiki/Holochroal_eye" title="Holochroal eye">Holochroal eye</a></li> <li><a href="/wiki/Parietal_eye" title="Parietal eye">Parietal eye</a></li> <li><a href="/wiki/Schizochroal_eye" class="mw-redirect" title="Schizochroal eye">Schizochroal eye</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution" title="Evolution">Evolution</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">Evolution of the eye</a></li> <li><a href="/wiki/Evolution_of_color_vision" title="Evolution of color vision">Evolution of color vision</a></li> <li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">Evolution of color vision in primates</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Coloration</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Albinism" title="Albinism">Albinism</a></li> <li><a href="/wiki/Animal_coloration" title="Animal coloration">Animal coloration</a></li> <li><a href="/wiki/Aposematism" title="Aposematism">Aposematism</a></li> <li><a href="/wiki/Camouflage" title="Camouflage">Camouflage</a></li> <li><a href="/wiki/Chromatophore" title="Chromatophore">Chromatophore</a></li> <li><a href="/wiki/Counter-illumination" title="Counter-illumination">Counter-illumination</a></li> <li><a class="mw-selflink selflink">Countershading</a></li> <li><a href="/wiki/Crypsis" title="Crypsis">Crypsis</a></li> <li><a href="/wiki/Deimatic_behaviour" title="Deimatic behaviour">Deimatic behaviour</a></li> <li><a href="/wiki/Disruptive_coloration" title="Disruptive coloration">Disruptive coloration</a> <ul><li><a href="/wiki/Coincident_disruptive_coloration" title="Coincident disruptive coloration">coincident</a></li></ul></li> <li><a href="/wiki/Eyespot_(mimicry)" title="Eyespot (mimicry)">Eyespot (mimicry)</a></li> <li><a href="/wiki/Mimicry" title="Mimicry">Mimicry</a></li> <li><a href="/wiki/Structural_coloration" title="Structural coloration">Structural coloration</a></li> <li><a href="/wiki/Underwater_camouflage" title="Underwater camouflage">Underwater camouflage</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related topics</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Sense" title="Sense">Animal senses</a></li> <li><a href="/wiki/Blindness_in_animals" title="Blindness in animals">Blindness in animals</a></li> <li><a href="/wiki/Eyespot_apparatus" title="Eyespot apparatus">Eyespot apparatus</a></li> <li><a href="/wiki/Feature_detection_(nervous_system)" title="Feature detection (nervous system)">Feature detection</a></li> <li><a href="/wiki/Infrared_sensing_in_snakes" title="Infrared sensing in snakes">Infrared sensing in snakes</a></li> <li><a href="/wiki/Monocular_deprivation" title="Monocular deprivation">Monocular deprivation</a></li> <li><a href="/wiki/Ommatidium" title="Ommatidium">Ommatidium</a></li> <li><a href="/wiki/Palpebral_(bone)" title="Palpebral (bone)">Palpebral (bone)</a></li> <li><a href="/wiki/Pseudopupil" title="Pseudopupil">Pseudopupil</a></li> <li><a href="/wiki/Rhopalium" title="Rhopalium">Rhopalium</a></li> <li><a href="/wiki/Underwater_vision" title="Underwater vision">Underwater vision</a></li> <li><a href="/wiki/Visual_perception" title="Visual perception">Visual perception</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Biological_rules" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Biological_rules" title="Template:Biological rules"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Biological_rules" title="Template talk:Biological rules"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Biological_rules" title="Special:EditPage/Template:Biological rules"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Biological_rules" style="font-size:114%;margin:0 4em"><a href="/wiki/Biological_rules" title="Biological rules">Biological rules</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Rules</th><td class="navbox-list-with-group navbox-list navbox-odd vlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allen%27s_rule" title="Allen's rule">Allen's rule</a> <span class="wrap"><span style="font-size:85%;">Shorter appendages in colder climates</span></span></li> <li><a href="/wiki/Bateson%27s_rule" class="mw-redirect" title="Bateson's rule">Bateson's rule</a> <span class="wrap"><span style="font-size:85%;">Extra limbs mirror their neighbours</span></span></li> <li><a href="/wiki/Bergmann%27s_rule" title="Bergmann's rule">Bergmann's rule</a> <span class="wrap"><span style="font-size:85%;">Larger bodies in colder climates</span></span></li> <li><a href="/wiki/Cope%27s_rule" title="Cope's rule">Cope's rule</a> <span class="wrap"><span style="font-size:85%;">Bodies get larger over time</span></span></li> <li><a href="/wiki/Deep-sea_gigantism" title="Deep-sea gigantism">Deep-sea gigantism</a> <span class="wrap"><span style="font-size:85%;">Larger bodies in deep-sea animals</span></span></li> <li><a href="/wiki/Dollo%27s_law_of_irreversibility" title="Dollo's law of irreversibility">Dollo's law</a> <span class="wrap"><span style="font-size:85%;">Loss of complex traits is irreversible</span></span></li> <li><a href="/wiki/Eichler%27s_rule" title="Eichler's rule">Eichler's rule</a> <span class="wrap"><span style="font-size:85%;">Parasites co-vary with their hosts</span></span></li> <li><a href="/wiki/Emery%27s_rule" title="Emery's rule">Emery's rule</a> <span class="wrap"><span style="font-size:85%;">Insect social parasites are often in same genus as their hosts</span></span></li> <li><a href="/wiki/Fahrenholz%27s_rule" class="mw-redirect" title="Fahrenholz's rule">Fahrenholz's rule</a> <span class="wrap"><span style="font-size:85%;">Host and parasite phylogenies become congruent</span></span></li> <li><a href="/wiki/Foster%27s_rule" title="Foster's rule">Foster's rule</a> (<a href="/wiki/Insular_gigantism" class="mw-redirect" title="Insular gigantism">Insular gigantism</a>, <a href="/wiki/Insular_dwarfism" title="Insular dwarfism">Insular dwarfism</a>) <span class="wrap"><span style="font-size:85%;">Small species get larger, large species smaller, after colonizing islands</span></span></li> <li><a href="/wiki/Gause%27s_law" class="mw-redirect" title="Gause's law">Gause's law</a> <span class="wrap"><span style="font-size:85%;">Complete competitors cannot coexist</span></span></li> <li><a href="/wiki/Gloger%27s_rule" title="Gloger's rule">Gloger's rule</a> <span class="wrap"><span style="font-size:85%;">Lighter coloration in colder, drier climates</span></span></li> <li><a href="/wiki/Haldane%27s_rule" title="Haldane's rule">Haldane's rule</a> <span class="wrap"><span style="font-size:85%;">Hybrid sexes that are absent, rare, or sterile, are heterogamic</span></span></li> <li><a href="/wiki/Harrison%27s_rule" title="Harrison's rule">Harrison's rule</a> <span class="wrap"><span style="font-size:85%;">Parasites co-vary in size with their hosts</span></span></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Hamilton's rule</a> <span class="wrap"><span style="font-size:85%;">Genes increase in frequency when relatedness of recipient to actor times benefit to recipient exceeds reproductive cost to actor</span></span></li> <li><a href="/wiki/Kleiber%27s_law" title="Kleiber's law">Kleiber's law</a> <span class="wrap"><span style="font-size:85%;">An animals metabolic rate decreases with its size</span></span></li> <li><a href="/wiki/Willi_Hennig" title="Willi Hennig">Hennig's progression rule</a> <span class="wrap"><span style="font-size:85%;">In cladistics, the most primitive species are found in earliest, central, part of group's area</span></span></li> <li><a href="/wiki/Jarman%E2%80%93Bell_principle" title="Jarman–Bell principle">Jarman–Bell principle</a> <span class="wrap"><span style="font-size:85%;">The correlation between the size of an animal and its diet quality; larger animals can consume lower quality diet</span></span></li> <li><a href="/wiki/Jordan%27s_rule" title="Jordan's rule">Jordan's rule</a> <span class="wrap"><span style="font-size:85%;">Inverse relationship between water temperature and no. of fin rays, vertebrae</span></span></li> <li><a href="/wiki/Lack%27s_principle" title="Lack's principle">Lack's principle</a> <span class="wrap"><span style="font-size:85%;">Birds lay only as many eggs as they can provide food for</span></span></li> <li><a href="/wiki/Rapoport%27s_rule" title="Rapoport's rule">Rapoport's rule</a> <span class="wrap"><span style="font-size:85%;">Latitudinal range increases with latitude</span></span></li> <li><a href="/wiki/Rensch%27s_rule" title="Rensch's rule">Rensch's rule</a> <span class="wrap"><span style="font-size:85%;">Sexual size dimorphism increases with size when males are larger, decreases with size when females are larger</span></span></li> <li><a href="/wiki/Rosa%27s_rule" title="Rosa's rule">Rosa's rule</a> <span class="wrap"><span style="font-size:85%;">Groups evolve from character variation in primitive species to a fixed character state in advanced ones</span></span></li> <li><a href="/wiki/Ivan_Schmalhausen" title="Ivan Schmalhausen">Schmalhausen's law</a> <span class="wrap"><span style="font-size:85%;">A population at limit of tolerance in one aspect is vulnerable to small differences in any other aspect</span></span></li> <li><a class="mw-selflink selflink">Thayer's law</a> <span class="wrap"><span style="font-size:85%;">The top of an animals coloration is darker than the bottom</span></span></li> <li><a href="/wiki/Thorson%27s_rule" title="Thorson's rule">Thorson's rule</a> <span class="wrap"><span style="font-size:85%;">No. of eggs of benthic marine invertebrates decreases with latitude</span></span></li> <li><a href="/wiki/Leigh_Van_Valen" title="Leigh Van Valen">Van Valen's law</a> <span class="wrap"><span style="font-size:85%;">Probability of extinction of a group is constant over time</span></span></li> <li><a href="/wiki/Von_Baer%27s_laws_(embryology)" title="Von Baer's laws (embryology)">von Baer's laws</a> <span class="wrap"><span style="font-size:85%;">Embryos start from a common form and develop into increasingly specialised forms</span></span></li> <li><a href="/wiki/Samuel_Wendell_Williston" title="Samuel Wendell Williston">Williston's law</a> <span class="wrap"><span style="font-size:85%;">Parts in an organism become reduced in number and specialized in function</span></span></li></ul> </div></td><td class="noviewer navbox-image" rowspan="2" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/Bergmann%27s_rule" title="Bergmann's rule"><img alt="Bergmann's rule illustrated with a map and graph" src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b7/Bergmann%27s_Rule.svg/160px-Bergmann%27s_Rule.svg.png" decoding="async" width="160" height="149" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b7/Bergmann%27s_Rule.svg/240px-Bergmann%27s_Rule.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b7/Bergmann%27s_Rule.svg/320px-Bergmann%27s_Rule.svg.png 2x" data-file-width="512" data-file-height="476" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list-with-group navbox-list navbox-even vlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Countergradient_variation" title="Countergradient variation">Countergradient variation</a> <span style="font-size:85%;">Where genetics opposes environment as a factor</span></li> <li><a href="/wiki/Gigantothermy" title="Gigantothermy">Gigantothermy</a> <span style="font-size:85%;">Large ectothermic animals more easily maintain constant body temperature</span></li></ul> </div></td></tr></tbody></table></div> <p class="mw-empty-elt"> </p>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" href="https://en.wikipedia.org/w/index.php?title=Countershading&oldid=1240185831">https://en.wikipedia.org/w/index.php?title=Countershading&oldid=1240185831</a>"</div></div> <div id="catlinks" class="catlinks" data-mw="interface"><div id="mw-normal-catlinks" class="mw-normal-catlinks"><a 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