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<li id="toc-Origins" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Origins"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Origins</span> </div> </a> <ul id="toc-Origins-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Müller's_mathematical_model" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Müller's_mathematical_model"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Müller's mathematical model</span> </div> </a> <ul id="toc-Müller's_mathematical_model-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Non-deceitful_mimicry" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Non-deceitful_mimicry"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Non-deceitful mimicry</span> </div> </a> <ul id="toc-Non-deceitful_mimicry-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Evolution" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Evolution</span> </div> </a> <button aria-controls="toc-Evolution-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 Evolution subsection</span> </button> <ul id="toc-Evolution-sublist" class="vector-toc-list"> <li id="toc-Aposematism,_camouflage,_and_mimicry" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Aposematism,_camouflage,_and_mimicry"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Aposematism, camouflage, and mimicry</span> </div> </a> <ul id="toc-Aposematism,_camouflage,_and_mimicry-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Selective_advantage" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Selective_advantage"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Selective advantage</span> </div> </a> <ul id="toc-Selective_advantage-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Relationship_to_Batesian_mimicry" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Relationship_to_Batesian_mimicry"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Relationship to Batesian mimicry</span> </div> </a> <ul id="toc-Relationship_to_Batesian_mimicry-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Non-visual_mimicry" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Non-visual_mimicry"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>Non-visual mimicry</span> </div> </a> <ul id="toc-Non-visual_mimicry-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Negative_frequency-dependent_selection" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Negative_frequency-dependent_selection"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.5</span> <span>Negative frequency-dependent selection</span> </div> </a> <ul id="toc-Negative_frequency-dependent_selection-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genetics" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genetics"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.6</span> <span>Genetics</span> </div> </a> <ul id="toc-Genetics-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Two_step_evolution" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Two_step_evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.7</span> <span>Two step evolution</span> </div> </a> <ul id="toc-Two_step_evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Advergence_versus_mutualism" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Advergence_versus_mutualism"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.8</span> <span>Advergence versus mutualism</span> </div> </a> <ul id="toc-Advergence_versus_mutualism-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mimicry_complexes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mimicry_complexes"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.9</span> <span>Mimicry complexes</span> </div> </a> <ul id="toc-Mimicry_complexes-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Taxonomic_range" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Taxonomic_range"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Taxonomic range</span> </div> </a> <ul id="toc-Taxonomic_range-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-In_marketing" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#In_marketing"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>In marketing</span> </div> </a> <ul id="toc-In_marketing-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">5</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">6</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">7</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Sources" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Sources"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Sources</span> </div> </a> <ul id="toc-Sources-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </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 id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Müllerian mimicry</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 16 languages" > <label id="p-lang-btn-label" for="p-lang-btn-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--action-progressive mw-portlet-lang-heading-16" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">16 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Mimetisme_de_M%C3%BCller" title="Mimetisme de Müller – Catalan" lang="ca" hreflang="ca" data-title="Mimetisme de Müller" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cy mw-list-item"><a href="https://cy.wikipedia.org/wiki/Dynwarededd_M%C3%BCelleraidd" title="Dynwarededd Müelleraidd – Welsh" lang="cy" hreflang="cy" data-title="Dynwarededd Müelleraidd" data-language-autonym="Cymraeg" data-language-local-name="Welsh" class="interlanguage-link-target"><span>Cymraeg</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Mimetismo_m%C3%BClleriano" title="Mimetismo mülleriano – Spanish" lang="es" hreflang="es" data-title="Mimetismo mülleriano" 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/Milera_kamuflimito" title="Milera kamuflimito – Esperanto" lang="eo" hreflang="eo" data-title="Milera kamuflimito" 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%AA%D9%82%D9%84%DB%8C%D8%AF_%D9%85%D9%88%D9%84%D8%B1%DB%8C" title="تقلید مولری – Persian" lang="fa" hreflang="fa" data-title="تقلید مولری" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Mimetismo_m%C3%BClleriano" title="Mimetismo mülleriano – Galician" lang="gl" hreflang="gl" data-title="Mimetismo mülleriano" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Mimikri_M%C3%BCller" title="Mimikri Müller – Indonesian" lang="id" hreflang="id" data-title="Mimikri Müller" 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%97%D7%A7%D7%99%D7%99%D7%A0%D7%95%D7%AA_%D7%9E%D7%99%D7%9C%D7%A8%D7%99%D7%99%D7%A0%D7%99%D7%AA" title="חקיינות מילריינית – Hebrew" lang="he" hreflang="he" data-title="חקיינות מילריינית" 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/Kemimikan_M%C3%BCller" title="Kemimikan Müller – Malay" lang="ms" hreflang="ms" data-title="Kemimikan Müller" 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-ja badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://ja.wikipedia.org/wiki/%E3%83%9F%E3%83%A5%E3%83%BC%E3%83%A9%E3%83%BC%E5%9E%8B%E6%93%AC%E6%85%8B" title="ミューラー型擬態 – Japanese" lang="ja" hreflang="ja" data-title="ミューラー型擬態" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Mimetismo_m%C3%BClleriano" title="Mimetismo mülleriano – Portuguese" lang="pt" hreflang="pt" data-title="Mimetismo mülleriano" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9C%D0%B8%D0%BC%D0%B8%D0%BA%D1%80%D0%B8%D1%8F_%D0%9C%D1%8E%D0%BB%D0%BB%D0%B5%D1%80%D0%B0" title="Мимикрия Мюллера – Russian" lang="ru" hreflang="ru" data-title="Мимикрия Мюллера" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/M%C3%BCllers_mimikry" title="Müllers mimikry – Swedish" lang="sv" hreflang="sv" data-title="Müllers mimikry" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9C%D1%96%D0%BC%D1%96%D0%BA%D1%80%D1%96%D1%8F_%D0%9C%D1%8E%D0%BB%D0%BB%D0%B5%D1%80%D0%B0" title="Мімікрія Мюллера – Ukrainian" lang="uk" hreflang="uk" data-title="Мімікрія Мюллера" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/B%E1%BA%AFt_ch%C6%B0%E1%BB%9Bc_ki%E1%BB%83u_M%C3%BCller" title="Bắt chước kiểu Müller – Vietnamese" lang="vi" hreflang="vi" data-title="Bắt chước kiểu Müller" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E7%A9%86%E6%B0%8F%E6%93%AC%E6%85%8B" title="穆氏擬態 – Cantonese" lang="yue" hreflang="yue" data-title="穆氏擬態" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target"><span>粵語</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q1892944#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> </div> </div> </div> </header> <div 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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"><p class="mw-empty-elt"> </p> <div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Mutually beneficial mimicry of strongly defended species</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Heliconius_mimicry.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0a/Heliconius_mimicry.png/220px-Heliconius_mimicry.png" decoding="async" width="220" height="221" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0a/Heliconius_mimicry.png/330px-Heliconius_mimicry.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0a/Heliconius_mimicry.png/440px-Heliconius_mimicry.png 2x" data-file-width="2100" data-file-height="2107" /></a><figcaption>Two examples of Müllerian mimicry in <i><a href="/wiki/Heliconius" title="Heliconius">Heliconius</a></i> butterflies: In this image the top four are forms of <i><a href="/wiki/Heliconius_numata" title="Heliconius numata">Heliconius numata</a>,</i> which mimic species from the genus <i><a href="/wiki/Melinaea" title="Melinaea">Melinaea</a></i>, while the bottom four are <i><a href="/wiki/Heliconius_melpomene" title="Heliconius melpomene">H. melpomene</a></i> (left) and <i><a href="/wiki/Heliconius_erato" title="Heliconius erato">H. erato</a></i> (right), which mimic each other.<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></figcaption></figure> <p><b>Müllerian mimicry</b> is a natural phenomenon in which two or more well-defended <a href="/wiki/Species" title="Species">species</a>, often foul-tasting and sharing common <a href="/wiki/Predator" class="mw-redirect" title="Predator">predators</a>, have come to <a href="/wiki/Mimicry" title="Mimicry">mimic</a> each other's <a href="/wiki/Honest_signal" class="mw-redirect" title="Honest signal">honest</a> <a href="/wiki/Aposematism" title="Aposematism">warning signals</a>, to their <a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">mutual benefit</a>. The benefit to Müllerian mimics is that predators only need one unpleasant encounter with one member of a set of Müllerian mimics, and thereafter avoid all similar coloration, whether or not it belongs to the same species as the initial encounter. It is named after the German naturalist <a href="/wiki/Fritz_M%C3%BCller" title="Fritz Müller">Fritz Müller</a>, who first proposed the concept in 1878, supporting his theory with the first <a href="/wiki/Mathematical_model" title="Mathematical model">mathematical model</a> of <a href="/wiki/Frequency-dependent_selection" title="Frequency-dependent selection">frequency-dependent selection</a>, one of the first such models anywhere in biology.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>a<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>Müllerian mimicry was first identified in tropical <a href="/wiki/Butterfly" title="Butterfly">butterflies</a> that shared colourful wing patterns, but it is found in many groups of insects such as <a href="/wiki/Bumblebee" title="Bumblebee">bumblebees</a>, and other animals such as <a href="/wiki/Poison_dart_frog" title="Poison dart frog">poison frogs</a> and <a href="/wiki/Coral_snake" title="Coral snake">coral snakes</a>. The mimicry need not be visual; for example, many snakes share <a href="/wiki/Hearing_(sense)" class="mw-redirect" title="Hearing (sense)">auditory</a> warning signals. Similarly, the defences involved are not limited to toxicity; anything that tends to deter predators, such as foul taste, sharp spines, or defensive behaviour can make a species unprofitable enough to predators to allow Müllerian mimicry to develop. </p><p>Once a pair of Müllerian mimics has formed, other mimics may join them by advergent evolution (one species changing to conform to the appearance of the pair, rather than mutual <a href="/wiki/Convergent_evolution" title="Convergent evolution">convergence</a>), forming mimicry rings. Large rings are found for example in <a href="/wiki/Dasymutilla" title="Dasymutilla">velvet ants</a>. Since the frequency of mimics is positively correlated with survivability, rarer mimics are likely to adapt to resemble commoner models, favouring both advergence and larger Müllerian mimicry rings. Where mimics are not strongly protected by venom or other defences, <a href="/wiki/Signalling_theory" title="Signalling theory">honest</a> Müllerian mimicry becomes, by degrees, the better-known bluffing of <a href="/wiki/Batesian_mimicry" title="Batesian mimicry">Batesian mimicry</a>. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=1" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Origins">Origins</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=2" title="Edit section: Origins"><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:BatesMimButter.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/63/BatesMimButter.JPG/220px-BatesMimButter.JPG" decoding="async" width="220" height="298" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/6/63/BatesMimButter.JPG 1.5x" data-file-width="300" data-file-height="406" /></a><figcaption>The <a href="/wiki/Viceroy_butterfly" class="mw-redirect" title="Viceroy butterfly">viceroy butterfly</a> (top) appears very similar to the noxious-tasting <a href="/wiki/Monarch_butterfly" title="Monarch butterfly">monarch butterfly</a> (bottom). Although it was for a long time purported to be an example of <a href="/wiki/Batesian_mimicry" title="Batesian mimicry">Batesian mimicry</a>, the viceroy has recently been discovered to be just as unpalatable as the monarch, making this a case of Müllerian mimicry.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup></figcaption></figure> <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">Further information: <a href="/wiki/Coloration_evidence_for_natural_selection" title="Coloration evidence for natural selection">Coloration evidence for natural selection</a></div> <p>Müllerian mimicry was proposed by the <a href="/wiki/Germany" title="Germany">German</a> <a href="/wiki/Zoology" title="Zoology">zoologist</a> and <a href="/wiki/Natural_history" title="Natural history">naturalist</a> <a href="/wiki/Fritz_M%C3%BCller" title="Fritz Müller">Fritz Müller</a> (1821–1897). An early proponent of <a href="/wiki/Evolution" title="Evolution">evolution</a>, Müller offered the first explanation for resemblance between certain <a href="/wiki/Butterflies" class="mw-redirect" title="Butterflies">butterflies</a> that had puzzled the English naturalist <a href="/wiki/Henry_Walter_Bates" title="Henry Walter Bates">Henry Walter Bates</a> in 1862. Bates, like Müller, spent a significant part of his life in <a href="/wiki/Brazil" title="Brazil">Brazil</a>, as described in his book <i><a href="/wiki/The_Naturalist_on_the_River_Amazons" title="The Naturalist on the River Amazons">The Naturalist on the River Amazons</a></i>. Bates conjectured that these abundant and distasteful butterflies might have been caused to resemble each other by their physical environment. Müller had also seen these butterflies first hand, and like Bates had collected specimens, and he proposed a variety of other explanations. One was <a href="/wiki/Sexual_selection" title="Sexual selection">sexual selection</a>, namely that individuals would choose to mate with partners with frequently-seen coloration, such as those resembling other species. However, if as is usual, females are the choosers, then mimicry would be seen in males, but in <a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">sexually dimorphic</a> species, females are more often mimetic.<sup id="cite_ref-Sherratt2008_6-0" class="reference"><a href="#cite_note-Sherratt2008-6"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Another was, as Müller wrote in 1878, that "defended species may evolve a similar appearance so as to share the costs of predator education."<sup id="cite_ref-FOOTNOTEForbes200940–42_7-0" class="reference"><a href="#cite_note-FOOTNOTEForbes200940–42-7"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTERuxtonSpeedSherratt2004116–118_8-0" class="reference"><a href="#cite_note-FOOTNOTERuxtonSpeedSherratt2004116–118-8"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Müller's_mathematical_model"><span id="M.C3.BCller.27s_mathematical_model"></span>Müller's mathematical model</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=3" title="Edit section: Müller's mathematical model"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Mathematical_and_theoretical_biology" title="Mathematical and theoretical biology">Mathematical and theoretical biology</a></div> <p>Müller's 1879 account was one of the earliest uses of a mathematical model in <a href="/wiki/Evolutionary_ecology" title="Evolutionary ecology">evolutionary ecology</a>, and the first exact model of frequency-dependent selection.<sup id="cite_ref-MalletonMuller_9-0" class="reference"><a href="#cite_note-MalletonMuller-9"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Mallet2001_10-0" class="reference"><a href="#cite_note-Mallet2001-10"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Mallet calls Müller's mathematical assumption behind the model "beguilingly simple".<sup id="cite_ref-Mallet2001Interface_11-0" class="reference"><a href="#cite_note-Mallet2001Interface-11"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Müller presumed that the predators had to attack n unprofitable prey in a summer to experience and learn their warning coloration. Calling a<sub>1</sub> and a<sub>2</sub> the total numbers of two unprofitable prey species, Müller then argued that, if the species are completely unalike they each lose n individuals. However, if they resemble each other,<sup id="cite_ref-MalletonMuller_9-1" class="reference"><a href="#cite_note-MalletonMuller-9"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p>then species 1 loses <span class="texhtml"><style data-mw-deduplicate="TemplateStyles:r1214402035">.mw-parser-output .sfrac{white-space:nowrap}.mw-parser-output .sfrac.tion,.mw-parser-output .sfrac .tion{display:inline-block;vertical-align:-0.5em;font-size:85%;text-align:center}.mw-parser-output .sfrac .num{display:block;line-height:1em;margin:0.0em 0.1em;border-bottom:1px solid}.mw-parser-output .sfrac .den{display:block;line-height:1em;margin:0.1em 0.1em}.mw-parser-output .sr-only{border:0;clip:rect(0,0,0,0);clip-path:polygon(0px 0px,0px 0px,0px 0px);height:1px;margin:-1px;overflow:hidden;padding:0;position:absolute;width:1px}</style><span class="sfrac">⁠<span class="tion"><span class="num">a<sub>1</sub>n</span><span class="sr-only">/</span><span class="den">a<sub>1</sub>+a<sub>2</sub></span></span>⁠</span></span> individuals, and species 2 loses <span class="texhtml"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1214402035"><span class="sfrac">⁠<span class="tion"><span class="num">a<sub>2</sub>n</span><span class="sr-only">/</span><span class="den">a<sub>1</sub>+a<sub>2</sub></span></span>⁠</span></span> individuals. </p><p>Species 1 therefore gains <span class="texhtml">n-<link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1214402035"><span class="sfrac">⁠<span class="tion"><span class="num">a<sub>1</sub>n</span><span class="sr-only">/</span><span class="den">a<sub>1</sub>+a<sub>2</sub></span></span>⁠</span> = <span class="texhtml"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1214402035"><span class="sfrac">⁠<span class="tion"><span class="num">a<sub>2</sub>n</span><span class="sr-only">/</span><span class="den">a<sub>1</sub>+a<sub>2</sub></span></span>⁠</span></span></span> and species 2 similarly gains <span class="texhtml"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1214402035"><span class="sfrac">⁠<span class="tion"><span class="num">a<sub>1</sub>n</span><span class="sr-only">/</span><span class="den">a<sub>1</sub>+a<sub>2</sub></span></span>⁠</span></span> in absolute numbers of individuals not killed. </p><p>The proportional gain compared to the total population of species 1 is <span class="texhtml">g<sub>1</sub> = <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1214402035"><span class="sfrac">⁠<span class="tion"><span class="num">a<sub>2</sub>n</span><span class="sr-only">/</span><span class="den">a<sub>1</sub>(a<sub>1</sub>+a<sub>2</sub>)</span></span>⁠</span></span> and similarly for species 2 <span class="texhtml">g<sub>2</sub> = <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1214402035"><span class="sfrac">⁠<span class="tion"><span class="num">a<sub>1</sub>n</span><span class="sr-only">/</span><span class="den">a<sub>2</sub>(a<sub>1</sub>+a<sub>2</sub>)</span></span>⁠</span></span>, giving the per head fitness gain of the mimicry when the predators have been fully educated. </p><p>Hence, Müller concluded, the proportion g1:g2 was <span class="texhtml"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1214402035"><span class="sfrac">⁠<span class="tion"><span class="num">a<sub>2</sub></span><span class="sr-only">/</span><span class="den">a<sub>1</sub></span></span>⁠</span> : <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1214402035"><span class="sfrac">⁠<span class="tion"><span class="num">a<sub>1</sub></span><span class="sr-only">/</span><span class="den">a<sub>2</sub></span></span>⁠</span></span>, which equals <span class="texhtml">a<sub>2</sub><sup>2</sup>:a<sub>1</sub><sup>2</sup></span>, and the rarer species gains far more than the commoner one.<sup id="cite_ref-MalletonMuller_9-2" class="reference"><a href="#cite_note-MalletonMuller-9"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p>The model is an approximation, and assumes the species are equally unprofitable. If one is more distasteful than the other, then the relative gains differ further, the less distasteful species benefiting more (as a square of the relative distastefulness) from the protection afforded by mimicry. This can be thought of as parasitic or quasi-Batesian, the mimic benefiting at the expense of the model. Later models are more complex and take factors such as rarity into account. The assumption of a fixed number n to be attacked is questionable.<sup id="cite_ref-Sherratt2008_6-1" class="reference"><a href="#cite_note-Sherratt2008-6"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Müller also effectively assumed a step function, when a gradual change (a <a href="/wiki/Functional_response" title="Functional response">functional response</a><sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup>) is more plausible.<sup id="cite_ref-Mallet2001Interface_11-1" class="reference"><a href="#cite_note-Mallet2001Interface-11"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Non-deceitful_mimicry">Non-deceitful mimicry</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=4" title="Edit section: Non-deceitful mimicry"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Deception_in_animals" title="Deception in animals">Deception in animals</a></div> <p>Biologists have not always viewed the Müllerian mechanism as mimicry, both because the term was strongly associated with Batesian mimicry, and because no <a href="/wiki/Deception_in_animals" title="Deception in animals">deception</a> was involved—unlike the situation in Batesian mimicry, the aposematic signals given by Müllerian mimics are <a href="/wiki/Honest_signal" class="mw-redirect" title="Honest signal">(unconsciously) honest</a>. Earlier terms, no longer in use, for Müllerian mimicry included "homotypy", "nondeceitful homotypy" and "arithmetic homotypy".<sup id="cite_ref-Pasteur1982_13-0" class="reference"><a href="#cite_note-Pasteur1982-13"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size mw-halign-center" typeof="mw:File/Thumb"><a href="/wiki/File:Batesian_vs_M%C3%BCllerian_mimicry.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a8/Batesian_vs_M%C3%BCllerian_mimicry.svg/440px-Batesian_vs_M%C3%BCllerian_mimicry.svg.png" decoding="async" width="440" height="248" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a8/Batesian_vs_M%C3%BCllerian_mimicry.svg/660px-Batesian_vs_M%C3%BCllerian_mimicry.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a8/Batesian_vs_M%C3%BCllerian_mimicry.svg/880px-Batesian_vs_M%C3%BCllerian_mimicry.svg.png 2x" data-file-width="1280" data-file-height="720" /></a><figcaption><a href="/wiki/Batesian_mimicry" title="Batesian mimicry">Batesian</a> vs Müllerian mimicry: the former is deceptive, the latter honest.</figcaption></figure> <div class="mw-heading mw-heading2"><h2 id="Evolution">Evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=5" title="Edit section: Evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Aposematism,_camouflage,_and_mimicry"><span id="Aposematism.2C_camouflage.2C_and_mimicry"></span>Aposematism, camouflage, and mimicry</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=6" title="Edit section: Aposematism, camouflage, and mimicry"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Aposematism" title="Aposematism">Aposematism</a></div> <p>Müllerian mimicry relies on <a href="/wiki/Aposematism" title="Aposematism">aposematism</a>, or warning signals. Dangerous organisms with these <a href="/wiki/Honest_signals" class="mw-redirect" title="Honest signals">honest signals</a> are avoided by predators, which quickly learn after a bad experience not to pursue the same unprofitable prey again. <a href="/wiki/Learning" title="Learning">Learning</a> is not actually necessary for animals which <a href="/wiki/Instinct" title="Instinct">instinctively</a> avoid certain prey;<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> however, learning from experience is more common.<sup id="cite_ref-EB_15-0" class="reference"><a href="#cite_note-EB-15"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> The underlying concept with predators that learn is that the warning signal makes the harmful organism easier to remember than if it remained as well <a href="/wiki/Camouflage" title="Camouflage">camouflaged</a> as possible. Aposematism and camouflage are in this way opposing concepts, but this does not mean they are mutually exclusive. Many animals remain inconspicuous until threatened, then suddenly employ warning signals, such as startling <a href="/wiki/Eyespot_(mimicry)" title="Eyespot (mimicry)">eyespots</a>, bright colours on their undersides or loud vocalizations. In this way, they enjoy the best of both strategies. These strategies may also be employed differentially throughout development. For instance, <a href="/wiki/Pieris_brassicae" title="Pieris brassicae">large white butterflies</a> are aposematic as <a href="/wiki/Larva" title="Larva">larvae</a>, but are Müllerian mimics once they emerge from development as <a href="/wiki/Imago" title="Imago">adult</a> butterflies.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p><span class="anchor" id="Unprofitability"></span> </p> <div class="mw-heading mw-heading3"><h3 id="Selective_advantage">Selective advantage</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=7" title="Edit section: Selective advantage"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Many different prey of the same predator could all employ their own warning signals, but this would make no sense for any party. If they could all agree on a common warning signal, the predator would have fewer detrimental experiences, and the prey would lose fewer individuals educating it. No such conference needs to take place, as a prey species that just so happens to look a little like an unprofitable<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>b<span class="cite-bracket">]</span></a></sup> species will be safer than its conspecifics, enabling <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> to drive the prey species toward a single warning language. This can lead to the <a href="/wiki/Evolution" title="Evolution">evolution</a> of both <a href="/wiki/Batesian_mimicry" title="Batesian mimicry">Batesian</a> and Müllerian mimicry, depending on whether the mimic is itself unprofitable to its predators, or just a free-rider. Multiple species can join the protective cooperative, expanding the mimicry ring. Müller thus provided an explanation for Bates' paradox; the mimicry was not, in his view, a case of exploitation by one species, but rather a <a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">mutualistic</a> arrangement, though his mathematical model indicated a pronounced asymmetry.<sup id="cite_ref-FOOTNOTERuxtonSpeedSherratt2004116–118_8-1" class="reference"><a href="#cite_note-FOOTNOTERuxtonSpeedSherratt2004116–118-8"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTERuxtonSpeedSherratt2004126_18-0" class="reference"><a href="#cite_note-FOOTNOTERuxtonSpeedSherratt2004126-18"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Mallet2001_10-2" class="reference"><a href="#cite_note-Mallet2001-10"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Relationship_to_Batesian_mimicry">Relationship to Batesian mimicry</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=8" title="Edit section: Relationship to Batesian mimicry"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Batesian_mimicry" title="Batesian mimicry">Batesian mimicry</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Batesian_vs_M%C3%BCllerian_Mimicry.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/93/Batesian_vs_M%C3%BCllerian_Mimicry.svg/400px-Batesian_vs_M%C3%BCllerian_Mimicry.svg.png" decoding="async" width="400" height="212" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/93/Batesian_vs_M%C3%BCllerian_Mimicry.svg/600px-Batesian_vs_M%C3%BCllerian_Mimicry.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/93/Batesian_vs_M%C3%BCllerian_Mimicry.svg/800px-Batesian_vs_M%C3%BCllerian_Mimicry.svg.png 2x" data-file-width="1052" data-file-height="557" /></a><figcaption>Comparison of Batesian and Müllerian mimicry, illustrated with a <a href="/wiki/Hoverfly" title="Hoverfly">hoverfly</a>, a <a href="/wiki/Wasp" title="Wasp">wasp</a> and a <a href="/wiki/Bee" title="Bee">bee</a></figcaption></figure> <p>The Müllerian strategy is usually contrasted with <a href="/wiki/Batesian_mimicry" title="Batesian mimicry">Batesian mimicry</a>, in which one harmless species adopts the appearance of an unprofitable species to gain the advantage of predators' avoidance; Batesian mimicry is thus in a sense <a href="/wiki/Parasite" class="mw-redirect" title="Parasite">parasitic</a> on the model's defences, whereas Müllerian is to mutual benefit. However, because comimics may have differing degrees of protection, the distinction between Müllerian and Batesian mimicry is not absolute, and there can be said to be a spectrum between the two forms.<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> </p><p><a href="/wiki/Viceroy_(butterfly)" title="Viceroy (butterfly)">Viceroy butterflies</a> and <a href="/wiki/Monarch_butterfly" title="Monarch butterfly">monarchs</a> (types of admiral butterfly) are both poisonous Müllerian mimics, though they were long thought to be Batesian. <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">Mitochondrial DNA</a> analysis of admiral butterflies shows that the viceroy is the basal lineage of two western sister species in North America. The variation in wing patterns appears to have preceded the evolution of toxicity, while other species remain non-toxic, refuting the hypothesis that the toxicity of these butterflies is a conserved characteristic from a common ancestor.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Non-visual_mimicry">Non-visual mimicry</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=9" title="Edit section: Non-visual mimicry"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Müllerian mimicry need not involve <a href="/wiki/Visual_perception" title="Visual perception">visual</a> mimicry; it may employ any of the <a href="/wiki/Sense" title="Sense">senses</a>. For example, many snakes share the same <a href="/wiki/Hearing_(sense)" class="mw-redirect" title="Hearing (sense)">auditory</a> warning signals, forming an auditory Müllerian mimicry ring. More than one signal may be shared: snakes can make use of both auditory signals and warning coloration.<sup id="cite_ref-Thalainen_21-0" class="reference"><a href="#cite_note-Thalainen-21"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Negative_frequency-dependent_selection">Negative frequency-dependent selection</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=10" title="Edit section: Negative frequency-dependent selection"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Negative_frequency-dependent_selection" class="mw-redirect" title="Negative frequency-dependent selection">Negative frequency-dependent selection</a></div> <p>There is a <a href="/wiki/Negative_frequency-dependent_selection" class="mw-redirect" title="Negative frequency-dependent selection">negative correlation between the frequency</a> of mimics and the "survivability" of both species involved. This implies that it is reproductively beneficial for both species if the models outnumber the mimics; this increases the negative interactions between predator and prey.<sup id="cite_ref-Thalainen_21-1" class="reference"><a href="#cite_note-Thalainen-21"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Genetics">Genetics</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=11" title="Edit section: Genetics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some insight into the evolution of mimetic color mimicry in Lepidoptera in particular can be seen through the study of the Optix gene. The Optix gene is responsible for the <i><a href="/wiki/Heliconius" title="Heliconius">Heliconius</a></i> butterflies' signature red wing patterns that help it signal to predators that it is toxic. By sharing this coloration with other poisonous red winged butterflies the predator may have pursued previously, the Heliconius butterfly increases its chance of survival through association. By mapping the genome of many related species of <i>Heliconius</i> butterflies "show[s] that the cis-regulatory evolution of a single transcription factor can repeatedly drive the convergent evolution of complex color patterns in distantly related species…".<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> This suggests that the evolution of a <a href="/wiki/Non-coding_DNA" title="Non-coding DNA">non-coding</a> piece of DNA that regulates the transcription of nearby genes can be the reason behind similar <a href="/wiki/Phenotype" title="Phenotype">phenotypic</a> coloration between distant species, making it hard to determine if the trait is <a href="/wiki/Homology_(biology)" title="Homology (biology)">homologous</a> or simply the result of <a href="/wiki/Convergent_evolution" title="Convergent evolution">convergent evolution</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Two_step_evolution">Two step evolution</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=12" title="Edit section: Two step evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>One proposed mechanism for Müllerian mimicry is the "two step hypothesis". This states that a large mutational leap initially establishes an approximate resemblance of the mimic to the model, both species already being aposematic. In a second step, smaller changes establish a closer resemblance. This is only likely to work, however, when a trait is governed by a single gene, and many coloration patterns are certainly controlled by multiple genes.<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> </p> <div class="mw-heading mw-heading3"><h3 id="Advergence_versus_mutualism">Advergence versus mutualism</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=13" title="Edit section: Advergence versus mutualism"><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:Formation_of_M%C3%BCllerian_mimicry_rings.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/17/Formation_of_M%C3%BCllerian_mimicry_rings.svg/330px-Formation_of_M%C3%BCllerian_mimicry_rings.svg.png" decoding="async" width="330" height="344" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/17/Formation_of_M%C3%BCllerian_mimicry_rings.svg/495px-Formation_of_M%C3%BCllerian_mimicry_rings.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/17/Formation_of_M%C3%BCllerian_mimicry_rings.svg/660px-Formation_of_M%C3%BCllerian_mimicry_rings.svg.png 2x" data-file-width="726" data-file-height="756" /></a><figcaption>Formation of Müllerian mimicry rings by a process of advergence of one species or pair of mimics to another, presumably larger or more abundant. Evolution is shown on two axes denoting phenotypes for convenience; in practice there would be any number of dimensions (e.g. coloration features). The model predicts a single mimicry ring in an area, but this is not the case in <i><a href="/wiki/Heliconius" title="Heliconius">Heliconius</a></i> butterflies.<sup id="cite_ref-MalletGilbert1995_24-0" class="reference"><a href="#cite_note-MalletGilbert1995-24"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>The mimic poison frog <i><a href="/wiki/Ranitomeya_imitator" class="mw-redirect" title="Ranitomeya imitator">Ranitomeya (Dendrobates) imitator</a></i> is <a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">polymorphic</a>, with a striped morph that imitates the black and yellow striped morph of <i><a href="/wiki/Ranitomeya_variabilis" class="mw-redirect" title="Ranitomeya variabilis">Ranitomeya variabilis</a></i>, a spotted morph that imitates the largely blue-green highland spotted morph also of <i>R. variabilis</i>, and a banded morph that imitates the red and black banded <i><a href="/wiki/Ranitomeya_summersi" title="Ranitomeya summersi">Ranitomeya summersi</a></i>.<sup id="cite_ref-Sherratt2008_6-2" class="reference"><a href="#cite_note-Sherratt2008-6"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><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> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:R._imitator_Chazuta.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6e/R._imitator_Chazuta.jpg/220px-R._imitator_Chazuta.jpg" decoding="async" width="220" height="156" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6e/R._imitator_Chazuta.jpg/330px-R._imitator_Chazuta.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6e/R._imitator_Chazuta.jpg/440px-R._imitator_Chazuta.jpg 2x" data-file-width="1754" data-file-height="1240" /></a><figcaption>A common morph of <i><a href="/wiki/Ranitomeya_imitator" class="mw-redirect" title="Ranitomeya imitator">Ranitomeya (Dendrobates) imitator</a></i> is <a href="/wiki/Aposematic" class="mw-redirect" title="Aposematic">aposematically</a> striped black and yellow, but in other areas, other morphs imitate differently coloured species.</figcaption></figure> <p><i>R. imitator</i> has thus apparently evolved in separate populations to resemble different targets, i.e. it has changed to resemble (adverged on) those target species, rather than both <i>R. imitator</i> and the other species mutually converging in the way that Müller supposed for tropical butterflies.<sup id="cite_ref-FOOTNOTERuxtonSpeedSherratt2004126–127_26-0" class="reference"><a href="#cite_note-FOOTNOTERuxtonSpeedSherratt2004126–127-26"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>Such advergence may be common. The mechanism was proposed by the entomologist F. A. Dixey in 1909<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> and has remained controversial; the evolutionary biologist <a href="/wiki/James_Mallet" title="James Mallet">James Mallet</a>, reviewing the situation in 2001, suggested that in Müllerian mimicry, advergence may be more common than convergence. In advergent evolution, the mimicking species responds to predation by coming to resemble the model more and more closely. Any initial benefit is thus to the mimic, and there is no implied <a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">mutualism</a>, as there would be with Müller's original convergence theory. However, once model and mimic have become closely similar, some degree of mutual protection becomes likely.<sup id="cite_ref-Mallet2001_10-3" class="reference"><a href="#cite_note-Mallet2001-10"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTERuxtonSpeedSherratt2004126–127_26-1" class="reference"><a href="#cite_note-FOOTNOTERuxtonSpeedSherratt2004126–127-26"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> This theory would predict that all mimicking species in an area should converge on a single pattern of coloration. This does not appear to happen in nature, however, as <i>Heliconius</i> butterflies form multiple Müllerian mimicry rings in a single geographical area. The finding implies that additional evolutionary forces are probably at work.<sup id="cite_ref-MalletGilbert1995_24-1" class="reference"><a href="#cite_note-MalletGilbert1995-24"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Mimicry_complexes">Mimicry complexes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=14" title="Edit section: Mimicry complexes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div style="text-align: center;"><ul class="gallery mw-gallery-packed"> <li class="gallerycaption">Many familiar <a href="/wiki/Bumblebee" title="Bumblebee">bumblebees</a> are Müllerian mimics, with effective <a href="/wiki/Stinger" title="Stinger">stings</a> and similar <a href="/wiki/Warning_coloration" class="mw-redirect" title="Warning coloration">warning coloration</a></li> <li class="gallerybox" style="width: 124.66666666667px"> <div class="thumb" style="width: 122.66666666667px;"><span typeof="mw:File"><a href="/wiki/File:Bombus_terrestris_(Buff-tailed_bumblebee)_-_queen_-_Flickr_-_S._Rae.jpg" class="mw-file-description" title="Bombus terrestris"><img alt="Bombus terrestris" src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b9/Bombus_terrestris_%28Buff-tailed_bumblebee%29_-_queen_-_Flickr_-_S._Rae.jpg/184px-Bombus_terrestris_%28Buff-tailed_bumblebee%29_-_queen_-_Flickr_-_S._Rae.jpg" decoding="async" width="123" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b9/Bombus_terrestris_%28Buff-tailed_bumblebee%29_-_queen_-_Flickr_-_S._Rae.jpg/276px-Bombus_terrestris_%28Buff-tailed_bumblebee%29_-_queen_-_Flickr_-_S._Rae.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b9/Bombus_terrestris_%28Buff-tailed_bumblebee%29_-_queen_-_Flickr_-_S._Rae.jpg/368px-Bombus_terrestris_%28Buff-tailed_bumblebee%29_-_queen_-_Flickr_-_S._Rae.jpg 2x" data-file-width="2134" data-file-height="2090" /></a></span></div> <div class="gallerytext"><i><a href="/wiki/Bombus_terrestris" title="Bombus terrestris">Bombus terrestris</a></i></div> </li> <li class="gallerybox" style="width: 122px"> <div class="thumb" style="width: 120px;"><span typeof="mw:File"><a href="/wiki/File:Bombus_lucorum_-_Centaurea_scabiosa_-_Keila.JPG" class="mw-file-description" title="Bombus lucorum"><img alt="Bombus lucorum" src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Bombus_lucorum_-_Centaurea_scabiosa_-_Keila.JPG/180px-Bombus_lucorum_-_Centaurea_scabiosa_-_Keila.JPG" decoding="async" width="120" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Bombus_lucorum_-_Centaurea_scabiosa_-_Keila.JPG/270px-Bombus_lucorum_-_Centaurea_scabiosa_-_Keila.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Bombus_lucorum_-_Centaurea_scabiosa_-_Keila.JPG/360px-Bombus_lucorum_-_Centaurea_scabiosa_-_Keila.JPG 2x" data-file-width="2626" data-file-height="2626" /></a></span></div> <div class="gallerytext"><i><a href="/wiki/Bombus_lucorum" title="Bombus lucorum">Bombus lucorum</a></i></div> </li> <li class="gallerybox" style="width: 118.66666666667px"> <div class="thumb" style="width: 116.66666666667px;"><span typeof="mw:File"><a href="/wiki/File:Bombus_hortorum_queen_-_Echium_vulgare_-_Keila.jpg" class="mw-file-description" title="Bombus hortorum"><img alt="Bombus hortorum" src="//upload.wikimedia.org/wikipedia/commons/thumb/4/43/Bombus_hortorum_queen_-_Echium_vulgare_-_Keila.jpg/175px-Bombus_hortorum_queen_-_Echium_vulgare_-_Keila.jpg" decoding="async" width="117" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/43/Bombus_hortorum_queen_-_Echium_vulgare_-_Keila.jpg/262px-Bombus_hortorum_queen_-_Echium_vulgare_-_Keila.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/43/Bombus_hortorum_queen_-_Echium_vulgare_-_Keila.jpg/350px-Bombus_hortorum_queen_-_Echium_vulgare_-_Keila.jpg 2x" data-file-width="2952" data-file-height="3037" /></a></span></div> <div class="gallerytext"><i><a href="/wiki/Bombus_hortorum" title="Bombus hortorum">Bombus hortorum</a></i></div> </li> <li class="gallerybox" style="width: 144px"> <div class="thumb" style="width: 142px;"><span typeof="mw:File"><a href="/wiki/File:Tree_bumblebee_(RLs)_(34742598795).jpg" class="mw-file-description" title="Bombus hypnorum"><img alt="Bombus hypnorum" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6f/Tree_bumblebee_%28RLs%29_%2834742598795%29.jpg/213px-Tree_bumblebee_%28RLs%29_%2834742598795%29.jpg" decoding="async" width="142" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6f/Tree_bumblebee_%28RLs%29_%2834742598795%29.jpg/320px-Tree_bumblebee_%28RLs%29_%2834742598795%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6f/Tree_bumblebee_%28RLs%29_%2834742598795%29.jpg/426px-Tree_bumblebee_%28RLs%29_%2834742598795%29.jpg 2x" data-file-width="1870" data-file-height="1579" /></a></span></div> <div class="gallerytext"><i><a href="/wiki/Bombus_hypnorum" title="Bombus hypnorum">Bombus hypnorum</a></i></div> </li> </ul></div> <p>Müllerian mimicry often occurs in clusters of multiple species called rings. Müllerian mimicry is not limited to butterflies, where rings are common; mimicry rings occur among <a href="/wiki/Hymenoptera" title="Hymenoptera">Hymenoptera</a>, such as <a href="/wiki/Bumblebee" title="Bumblebee">bumblebees</a>, and other insects, and among vertebrates including fish and <a href="/wiki/Coral_snake" title="Coral snake">coral snakes</a>. Bumblebees <i><a href="/wiki/Bombus" class="mw-redirect" title="Bombus">Bombus</a></i> are all aposematically coloured in combinations, often stripes, of black, white, yellow, and red; and all their females have stings,<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>c<span class="cite-bracket">]</span></a></sup> so they are certainly unprofitable to predators. There is evidence that several species of bumblebees in each of several areas of the world, namely the American West and East coasts, Western Europe, and Kashmir, have converged or adverged on mutually mimetic coloration patterns. Each of these areas has one to four mimicry rings, with patterns different from those in other areas.<sup id="cite_ref-Mallet2001_10-5" class="reference"><a href="#cite_note-Mallet2001-10"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Velvet_Ant_(Mutillidae)(Id_%3F)_(16821757352).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a5/Velvet_Ant_%28Mutillidae%29%28Id_%3F%29_%2816821757352%29.jpg/220px-Velvet_Ant_%28Mutillidae%29%28Id_%3F%29_%2816821757352%29.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a5/Velvet_Ant_%28Mutillidae%29%28Id_%3F%29_%2816821757352%29.jpg/330px-Velvet_Ant_%28Mutillidae%29%28Id_%3F%29_%2816821757352%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a5/Velvet_Ant_%28Mutillidae%29%28Id_%3F%29_%2816821757352%29.jpg/440px-Velvet_Ant_%28Mutillidae%29%28Id_%3F%29_%2816821757352%29.jpg 2x" data-file-width="5472" data-file-height="3648" /></a><figcaption>Many species of North American velvet ants in the genus <i><a href="/wiki/Dasymutilla" title="Dasymutilla">Dasymutilla</a></i> are involved in mimicry complexes.</figcaption></figure> <p>The relationships among mimics can become complex. For example, the poison fangblenny <i><a href="/wiki/Meiacanthus" title="Meiacanthus">Meiacanthus</a></i> spp. have hollow canines and poison glands, and are avoided by predatory fish. The blenny <i><a href="/wiki/Plagiotremus_townsendi" title="Plagiotremus townsendi">Plagiotremus townsendi</a></i> resembles <i>Meiacanthus</i> and is eaten by a variety of predators, so it is a Batesian mimic in their case: but it is avoided by the lionfish, <i><a href="/wiki/Pterois_volitans" class="mw-redirect" title="Pterois volitans">Pterois volitans</a></i>, making it also a Müllerian mimic.<sup id="cite_ref-FOOTNOTEEdmunds1974127–130_29-0" class="reference"><a href="#cite_note-FOOTNOTEEdmunds1974127–130-29"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p><p>Sets of associated rings are called complexes. Large complexes are known among the North American velvet ants in the genus <i><a href="/wiki/Dasymutilla" title="Dasymutilla">Dasymutilla</a></i>. Out of 351 species examined in one study, 336 had morphological similarities, apparently forming 8 distinct mimetic rings; 65 species in another study appeared to form six rings separable by both morphology and geography.<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Taxonomic_range">Taxonomic range</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=15" title="Edit section: Taxonomic range"><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:Pitohui_kirhocephalus_by_Duperrey_1825-1839.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/bd/Pitohui_kirhocephalus_by_Duperrey_1825-1839.png/170px-Pitohui_kirhocephalus_by_Duperrey_1825-1839.png" decoding="async" width="170" height="172" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/bd/Pitohui_kirhocephalus_by_Duperrey_1825-1839.png/255px-Pitohui_kirhocephalus_by_Duperrey_1825-1839.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/bd/Pitohui_kirhocephalus_by_Duperrey_1825-1839.png/340px-Pitohui_kirhocephalus_by_Duperrey_1825-1839.png 2x" data-file-width="2055" data-file-height="2075" /></a><figcaption><i><a href="/wiki/Pitohui_kirhocephalus" class="mw-redirect" title="Pitohui kirhocephalus">Pitohui kirhocephalus</a></i>, from <a href="/wiki/Louis_Isidore_Duperrey" class="mw-redirect" title="Louis Isidore Duperrey">Duperrey</a>, 1825–1839, appears to be a Müllerian mimic of <i><a href="/wiki/Pitohui_dichrous" class="mw-redirect" title="Pitohui dichrous">Pitohui dichrous</a></i> in some of its plumage types.</figcaption></figure> <p>Müllerian mimicry was discovered and has mainly been researched in insects. However, there is no reason why the mechanism's evolutionary advantages should not be exploited in other groups. There is some evidence that birds in the New Guinea genus <i><a href="/wiki/Pitohui" title="Pitohui">Pitohui</a></i> are Müllerian mimics. <i><a href="/wiki/Pitohui_dichrous" class="mw-redirect" title="Pitohui dichrous">Pitohui dichrous</a></i> and <i><a href="/wiki/Pitohui_kirhocephalus" class="mw-redirect" title="Pitohui kirhocephalus">Pitohui kirhocephalus</a></i> "share a nearly identical colour pattern" where their geographic ranges overlap, but differ elsewhere; they are conspicuous; and they are chemically defended by a powerful <a href="/wiki/Neurotoxic" class="mw-redirect" title="Neurotoxic">neurotoxic</a> <a href="/wiki/Alkaloid" title="Alkaloid">alkaloid</a>, <a href="/wiki/Batrachotoxin" title="Batrachotoxin">batrachotoxin</a>, in their feathers and skin. This combination of facts implies that the populations in these zones of overlap have converged to share honest warning signals.<sup id="cite_ref-DumbacherFleischer2001_32-0" class="reference"><a href="#cite_note-DumbacherFleischer2001-32"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Ictonyx_libyca_multivittata.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/78/Ictonyx_libyca_multivittata.jpg/220px-Ictonyx_libyca_multivittata.jpg" decoding="async" width="220" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/78/Ictonyx_libyca_multivittata.jpg/330px-Ictonyx_libyca_multivittata.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/78/Ictonyx_libyca_multivittata.jpg/440px-Ictonyx_libyca_multivittata.jpg 2x" data-file-width="600" data-file-height="328" /></a><figcaption>Several mammals including the <a href="/wiki/Saharan_striped_polecat" title="Saharan striped polecat">Saharan striped polecat</a>, <i>Ictonyx libycus</i>, are aggressive, aposematically coloured predators. They share black-and-white patterns, suggesting Müllerian mimicry.<sup id="cite_ref-Caro_2005_33-0" class="reference"><a href="#cite_note-Caro_2005-33"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>Many species of <a href="/wiki/Flower" title="Flower">flowers</a> resemble each other but actual mimicry has not been demonstrated.<sup id="cite_ref-Roy1999_34-0" class="reference"><a href="#cite_note-Roy1999-34"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> It has been proposed that spiny plants such as <a href="/wiki/Cactaceae" class="mw-redirect" title="Cactaceae">Cactaceae</a> and <i><a href="/wiki/Agave" title="Agave">Agave</a></i> in the Americas, <i><a href="/wiki/Aloe" title="Aloe">Aloe</a></i>, <i><a href="/wiki/Euphorbia" title="Euphorbia">Euphorbia</a></i>, white-thorned <i><a href="/wiki/Acacia" title="Acacia">Acacia</a></i> in Africa and spiny <a href="/wiki/Asteraceae" title="Asteraceae">Asteraceae</a> of the Mediterranean may form Müllerian mimicry rings, as they are strongly defended, are generally agreed to be aposematic, have similar conspicuous patterns and coloration, and are found in overlapping territories.<sup id="cite_ref-Lev-Yadun2014_35-0" class="reference"><a href="#cite_note-Lev-Yadun2014-35"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p><p>Aposematic <a href="/wiki/Mammal" title="Mammal">mammals</a> in the families <a href="/wiki/Mustelidae" title="Mustelidae">Mustelidae</a>, <a href="/wiki/Viverridae" title="Viverridae">Viverridae</a>, and <a href="/wiki/Herpestidae" class="mw-redirect" title="Herpestidae">Herpestidae</a> have independently evolved conspicuous black-and-white coloration, suggesting that Müllerian mimicry may be involved.<sup id="cite_ref-Caro_2005_33-1" class="reference"><a href="#cite_note-Caro_2005-33"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="In_marketing">In marketing</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=16" title="Edit section: In marketing"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The evolutionary zoologist <a href="/wiki/Thomas_N._Sherratt" title="Thomas N. Sherratt">Thomas N. Sherratt</a> suggests that different types of mimicry occur in <a href="/wiki/Brand" title="Brand">brand</a> and product <a href="/wiki/Marketing" title="Marketing">marketing</a>. He notes that distinctive forms like the <a href="/wiki/Coca-Cola" title="Coca-Cola">Coca-Cola</a> bottle's shape are defended by businesses, whereas rival companies have often imitated such famous motifs to benefit from the investment and reputation of their well-known competitors, constituting Batesian mimicry. Sherratt observes that the <a href="/wiki/Packaging" title="Packaging">packaging</a> of British <a href="/wiki/Supermarket" title="Supermarket">supermarket</a> own brands of <a href="/wiki/Potato_crisps" class="mw-redirect" title="Potato crisps">potato crisps</a> are consistently colour-coded red for the ready-salted variety, green for salt and vinegar, and blue for cheese and onion,<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>d<span class="cite-bracket">]</span></a></sup> across the major chains <a href="/wiki/Sainsbury%27s" title="Sainsbury's">Sainsbury's</a>, <a href="/wiki/Tesco" title="Tesco">Tesco</a>, <a href="/wiki/Asda" title="Asda">Asda</a>, and <a href="/wiki/Waitrose" title="Waitrose">Waitrose</a>. He argues that this sharing of pattern is very unlikely to have arisen by chance, in which case the resemblance is intentionally to inform customers reliably (honest signalling) of what each package contains, to mutual benefit in the manner of Müllerian mimicry.<sup id="cite_ref-Sherratt2008_6-4" class="reference"><a href="#cite_note-Sherratt2008-6"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <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=M%C3%BCllerian_mimicry&action=edit&section=17" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Deception_in_animals" title="Deception in animals">Deception in animals</a></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=M%C3%BCllerian_mimicry&action=edit&section=18" 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-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><a href="/wiki/Thomas_Malthus" class="mw-redirect" title="Thomas Malthus">Thomas Malthus</a>'s use of <a href="/wiki/An_Essay_on_the_Principle_of_Population" title="An Essay on the Principle of Population">tables of numbers illustrating the limits to human population growth</a> is one of the few earlier uses of a mathematical argument that could be called a model.</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">Unprofitability may consist of anything which makes prey not worth a predator's while to eat. Unpalatability on grounds of toxicity or foul taste is a common mechanism, but defences may include sharp spines; an aggressive nature; agility or speed in escape rendering the prey costly to catch; foul smell, and so on.<sup id="cite_ref-Mallet2001_10-1" class="reference"><a href="#cite_note-Mallet2001-10"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup></span> </li> <li id="cite_note-28"><span class="mw-cite-backlink"><b><a href="#cite_ref-28">^</a></b></span> <span class="reference-text">Drones have no sting, but similar patterns, and may (more or less accidentally) benefit from <a href="/wiki/Automimicry" title="Automimicry">automimicry</a> of females of their own species.<sup id="cite_ref-Mallet2001_10-4" class="reference"><a href="#cite_note-Mallet2001-10"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup></span> </li> <li id="cite_note-36"><span class="mw-cite-backlink"><b><a href="#cite_ref-36">^</a></b></span> <span class="reference-text">Sherratt notes that this use of red is shared with Walker's crisps, whereas the uses of blue and green are interchanged with respect to Walker's.<sup id="cite_ref-Sherratt2008_6-3" class="reference"><a href="#cite_note-Sherratt2008-6"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup></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=M%C3%BCllerian_mimicry&action=edit&section=19" 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: 30em;"> <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="CITEREFMeyer2006" class="citation journal cs1">Meyer, A. (2006). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1617347">"Repeating patterns of mimicry"</a>. <i>PLOS Biol</i>. <b>4</b> (10): e341. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1371%2Fjournal.pbio.0040341">10.1371/journal.pbio.0040341</a></span>. <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/PMC1617347">1617347</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/17048984">17048984</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PLOS+Biol&rft.atitle=Repeating+patterns+of+mimicry&rft.volume=4&rft.issue=10&rft.pages=e341&rft.date=2006&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1617347%23id-name%3DPMC&rft_id=info%3Apmid%2F17048984&rft_id=info%3Adoi%2F10.1371%2Fjournal.pbio.0040341&rft.aulast=Meyer&rft.aufirst=A.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1617347&rfr_id=info%3Asid%2Fen.wikipedia.org%3AM%C3%BCllerian+mimicry" class="Z3988"></span></span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMüller1878" class="citation journal cs1"><a href="/wiki/Fritz_M%C3%BCller" title="Fritz Müller">Müller, Fritz</a> (1878). "Ueber die Vortheile der Mimicry bei Schmetterlingen". <i>Zoologischer Anzeiger</i>. <b>1</b>: 54–55.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Zoologischer+Anzeiger&rft.atitle=Ueber+die+Vortheile+der+Mimicry+bei+Schmetterlingen&rft.volume=1&rft.pages=54-55&rft.date=1878&rft.aulast=M%C3%BCller&rft.aufirst=Fritz&rfr_id=info%3Asid%2Fen.wikipedia.org%3AM%C3%BCllerian+mimicry" class="Z3988"></span></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"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMüller1879" class="citation journal cs1"><a href="/wiki/Fritz_M%C3%BCller" title="Fritz Müller">Müller, Fritz</a> (1879). "<i>Ituna</i> and <i>Thyridia</i>; a remarkable case of mimicry in butterflies. (R. Meldola translation)". <i>Proclamations of the Entomological Society of London</i>. <b>1879</b>: 20–29.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proclamations+of+the+Entomological+Society+of+London&rft.atitle=Ituna+and+Thyridia%3B+a+remarkable+case+of+mimicry+in+butterflies.+%28R.+Meldola+translation%29&rft.volume=1879&rft.pages=20-29&rft.date=1879&rft.aulast=M%C3%BCller&rft.aufirst=Fritz&rfr_id=info%3Asid%2Fen.wikipedia.org%3AM%C3%BCllerian+mimicry" class="Z3988"></span></span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRitlandL._P._Brower1991" class="citation journal cs1">Ritland, D.; <a href="/wiki/Lincoln_Brower" title="Lincoln Brower">L. P. Brower</a> (1991). "The viceroy butterfly is not a Batesian mimic". <i><a href="/wiki/Nature_(journal)" title="Nature (journal)">Nature</a></i>. <b>350</b> (6318): 497–498. <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/1991Natur.350..497R">1991Natur.350..497R</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%2F350497a0">10.1038/350497a0</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:28667520">28667520</a>. <q>Viceroys are as unpalatable as monarchs, and significantly more unpalatable than queens from representative Florida populations.</q></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=The+viceroy+butterfly+is+not+a+Batesian+mimic&rft.volume=350&rft.issue=6318&rft.pages=497-498&rft.date=1991&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A28667520%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1038%2F350497a0&rft_id=info%3Abibcode%2F1991Natur.350..497R&rft.aulast=Ritland&rft.aufirst=D.&rft.au=L.+P.+Brower&rfr_id=info%3Asid%2Fen.wikipedia.org%3AM%C3%BCllerian+mimicry" class="Z3988"></span></span> </li> <li id="cite_note-Sherratt2008-6"><span class="mw-cite-backlink">^ <a href="#cite_ref-Sherratt2008_6-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Sherratt2008_6-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Sherratt2008_6-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Sherratt2008_6-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Sherratt2008_6-4"><sup><i><b>e</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSherratt2008" class="citation journal cs1"><a href="/wiki/Thomas_N._Sherratt" title="Thomas N. 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"Floral mimicry: a fascinating yet poorly understood phenomenon". <i>Trends in Plant Science</i>. <b>4</b> (8): 325–330. <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%2FS1360-1385%2899%2901445-4">10.1016/S1360-1385(99)01445-4</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/10431223">10431223</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Trends+in+Plant+Science&rft.atitle=Floral+mimicry%3A+a+fascinating+yet+poorly+understood+phenomenon&rft.volume=4&rft.issue=8&rft.pages=325-330&rft.date=1999&rft_id=info%3Adoi%2F10.1016%2FS1360-1385%2899%2901445-4&rft_id=info%3Apmid%2F10431223&rft.aulast=Roy&rft.aufirst=B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AM%C3%BCllerian+mimicry" class="Z3988"></span></span> </li> <li id="cite_note-Lev-Yadun2014-35"><span class="mw-cite-backlink"><b><a href="#cite_ref-Lev-Yadun2014_35-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLev-Yadun2014" class="citation journal cs1">Lev-Yadun, Simcha (2014). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2688291">"Müllerian mimicry in aposematic spiny plants"</a>. <i>Plant Signaling & Behavior</i>. <b>4</b> (6): 482–483. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.4161%2Fpsb.4.6.8848">10.4161/psb.4.6.8848</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/PMC2688291">2688291</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/19816137">19816137</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Plant+Signaling+%26+Behavior&rft.atitle=M%C3%BCllerian+mimicry+in+aposematic+spiny+plants&rft.volume=4&rft.issue=6&rft.pages=482-483&rft.date=2014&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2688291%23id-name%3DPMC&rft_id=info%3Apmid%2F19816137&rft_id=info%3Adoi%2F10.4161%2Fpsb.4.6.8848&rft.aulast=Lev-Yadun&rft.aufirst=Simcha&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2688291&rfr_id=info%3Asid%2Fen.wikipedia.org%3AM%C3%BCllerian+mimicry" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Sources">Sources</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=20" title="Edit section: Sources"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Wikisource-logo.svg/38px-Wikisource-logo.svg.png" decoding="async" width="38" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Wikisource-logo.svg/57px-Wikisource-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Wikisource-logo.svg/76px-Wikisource-logo.svg.png 2x" data-file-width="410" data-file-height="430" /></span></span></div> <div class="side-box-text plainlist"><a href="/wiki/Wikisource" title="Wikisource">Wikisource</a> has original text related to this article: <div style="margin-left: 10px;"><b><a href="https://en.wikisource.org/wiki/Mimicry_in_Butterflies" class="extiw" title="wikisource:Mimicry in Butterflies">Mimicry in Butterflies</a></b></div></div></div> </div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEdmunds1974" class="citation book cs1">Edmunds, M. (1974). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/defenceinanimals0000edmu"><i>Defence in Animals</i></a></span>. Longmans. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-582-44132-3" title="Special:BookSources/978-0-582-44132-3"><bdi>978-0-582-44132-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Defence+in+Animals&rft.pub=Longmans&rft.date=1974&rft.isbn=978-0-582-44132-3&rft.aulast=Edmunds&rft.aufirst=M.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fdefenceinanimals0000edmu&rfr_id=info%3Asid%2Fen.wikipedia.org%3AM%C3%BCllerian+mimicry" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFForbes2009" 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 University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-300-17896-8" title="Special:BookSources/978-0-300-17896-8"><bdi>978-0-300-17896-8</bdi></a>.</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+University+Press&rft.date=2009&rft.isbn=978-0-300-17896-8&rft.aulast=Forbes&rft.aufirst=Peter&rfr_id=info%3Asid%2Fen.wikipedia.org%3AM%C3%BCllerian+mimicry" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRuxtonSpeedSherratt2004" class="citation book cs1"><a href="/wiki/Graeme_Ruxton" title="Graeme Ruxton">Ruxton, Graeme</a>; Speed, M. P.; <a href="/wiki/Thomas_N._Sherratt" title="Thomas N. Sherratt">Sherratt, T. N.</a> (2004). <i>Avoiding Attack. The Evolutionary Ecology of Crypsis, Warning Signals and Mimicry</i>. Oxford University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-852860-9" title="Special:BookSources/978-0-19-852860-9"><bdi>978-0-19-852860-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Avoiding+Attack.+The+Evolutionary+Ecology+of+Crypsis%2C+Warning+Signals+and+Mimicry&rft.pub=Oxford+University+Press&rft.date=2004&rft.isbn=978-0-19-852860-9&rft.aulast=Ruxton&rft.aufirst=Graeme&rft.au=Speed%2C+M.+P.&rft.au=Sherratt%2C+T.+N.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AM%C3%BCllerian+mimicry" class="Z3988"></span> Chapters 9 and 11 provide an overview.</li></ul> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=M%C3%BCllerian_mimicry&action=edit&section=21" title="Edit section: Further 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="CITEREFWickler1968" class="citation book cs1"><a href="/wiki/Wolfgang_Wickler" title="Wolfgang Wickler">Wickler, Wolfgang</a> (1968). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/mimicryinplantsa00wick"><i>Mimicry in Plants and Animals</i></a></span>. McGraw-Hill. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-07-070100-7" title="Special:BookSources/978-0-07-070100-7"><bdi>978-0-07-070100-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Mimicry+in+Plants+and+Animals&rft.pub=McGraw-Hill&rft.date=1968&rft.isbn=978-0-07-070100-7&rft.aulast=Wickler&rft.aufirst=Wolfgang&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fmimicryinplantsa00wick&rfr_id=info%3Asid%2Fen.wikipedia.org%3AM%C3%BCllerian+mimicry" class="Z3988"></span> Especially chapters 7 and 8.</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 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display</a></li> <li><a href="/wiki/Automimicry" title="Automimicry">Automimicry</a></li> <li><a href="/wiki/Batesian_mimicry" title="Batesian mimicry">Batesian</a> <ul><li><a href="/wiki/Locomotor_mimicry" title="Locomotor mimicry">Locomotor</a></li></ul></li> <li><a href="/wiki/Brood_parasitism" title="Brood parasitism">Brood</a></li> <li><a href="/wiki/Chemical_mimicry" title="Chemical mimicry">Chemical</a></li> <li><a href="/wiki/Emsleyan_mimicry" title="Emsleyan mimicry">Emsleyan/Mertensian</a></li> <li><a href="/wiki/Eyespot_(mimicry)" title="Eyespot (mimicry)">Eyespot</a></li> <li><a class="mw-selflink selflink">Müllerian</a></li> <li><a href="/wiki/Sexual_mimicry" title="Sexual mimicry">Sexual</a></li> <li><a href="/wiki/Mimicry_in_vertebrates" title="Mimicry in vertebrates">In vertebrates</a></li> <li><a href="/wiki/Wasmannian_mimicry" class="mw-redirect" title="Wasmannian mimicry">Wasmannian</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="3" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Batesplate_ArM.jpg" class="mw-file-description" title="Batesian mimicry in butterflies"><img alt="Batesian mimicry in butterflies" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/95/Batesplate_ArM.jpg/70px-Batesplate_ArM.jpg" decoding="async" width="70" height="60" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/95/Batesplate_ArM.jpg/105px-Batesplate_ArM.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/95/Batesplate_ArM.jpg/140px-Batesplate_ArM.jpg 2x" data-file-width="1200" data-file-height="1024" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Mimicry_in_plants" title="Mimicry in plants">In plants</a></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/Mimicry_in_plants#Bakerian" title="Mimicry in plants">Bakerian</a></li> <li><a href="/wiki/Mimicry_in_plants#Dodsonian" title="Mimicry in plants">Dodsonian</a></li> <li><a href="/wiki/Gilbertian_mimicry" title="Gilbertian mimicry">Gilbertian</a></li> <li><a href="/wiki/Pouyannian_mimicry" title="Pouyannian mimicry">Pouyannian</a></li> <li><a href="/wiki/Vavilovian_mimicry" title="Vavilovian mimicry">Vavilovian</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 hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Anti-predator_adaptation" title="Anti-predator adaptation">Anti-predator adaptation</a></li> <li><a href="/wiki/Animal_communication" title="Animal communication">Animal communication</a></li> <li><a href="/wiki/Aposematism" title="Aposematism">Aposematism</a></li> <li><a href="/wiki/Camouflage" title="Camouflage">Camouflage</a>/<a href="/wiki/Crypsis" title="Crypsis">Crypsis</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Co-evolution</a></li> <li><a href="/wiki/Animal_coloration" title="Animal coloration">Animal coloration</a></li> <li><a href="/wiki/Community_(ecology)" title="Community (ecology)">Community ecology</a></li> <li><a href="/wiki/Coloration_evidence_for_natural_selection" title="Coloration evidence for natural selection">Coloration evidence for natural selection</a></li> <li><i><a href="/wiki/Dazzled_and_Deceived" title="Dazzled and Deceived">Dazzled and Deceived</a></i></li> <li><a href="/wiki/Deception_in_animals" title="Deception in animals">Deception in animals</a></li> <li><a href="/wiki/Deimatic_behaviour" title="Deimatic behaviour">Deimatic behaviour</a></li> <li><a href="/wiki/Mimicry#Evolution" title="Mimicry">Mimicry#Evolution</a></li> <li><a href="/wiki/Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></li> <li><a href="/wiki/Frequency-dependent_selection" title="Frequency-dependent selection">Frequency-dependent selection</a></li> <li><a href="/wiki/Phagomimicry" title="Phagomimicry">Phagomimicry</a></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a></li> <li><a href="/wiki/Underwater_camouflage" title="Underwater camouflage">Underwater camouflage</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="3"><div> <ul><li> <span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, 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