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class="vector-toc-numb">2.1</span> <span>Type I functional response</span> </div> </a> <ul id="toc-Type_I_functional_response-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Type_II_functional_response" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Type_II_functional_response"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Type II functional response</span> </div> </a> <ul id="toc-Type_II_functional_response-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Structure_of_networks" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Structure_of_networks"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Structure of networks</span> </div> </a> <ul id="toc-Structure_of_networks-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Humans" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Humans"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Humans</span> </div> </a> <button aria-controls="toc-Humans-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 Humans subsection</span> </button> <ul id="toc-Humans-sublist" class="vector-toc-list"> <li id="toc-Intestinal_microbiota" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Intestinal_microbiota"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Intestinal microbiota</span> </div> </a> <ul id="toc-Intestinal_microbiota-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Evolution_of_mutualism" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Evolution_of_mutualism"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Evolution of mutualism</span> </div> </a> <button aria-controls="toc-Evolution_of_mutualism-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 of mutualism subsection</span> </button> <ul id="toc-Evolution_of_mutualism-sublist" class="vector-toc-list"> <li id="toc-Evolution_by_type" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Evolution_by_type"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Evolution by type</span> </div> </a> <ul id="toc-Evolution_by_type-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mutualism_breakdown" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mutualism_breakdown"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Mutualism breakdown</span> </div> </a> <ul id="toc-Mutualism_breakdown-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Measuring_and_defining_mutualism" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Measuring_and_defining_mutualism"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Measuring and defining mutualism</span> </div> </a> <ul id="toc-Measuring_and_defining_mutualism-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">7</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_references" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_references"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Further references</span> </div> </a> <ul id="toc-Further_references-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">10</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 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</div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Mutualism (biology)</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 54 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-54" 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">54 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%AA%D9%82%D8%A7%D9%8A%D8%B6" title="تقايض – Arabic" lang="ar" hreflang="ar" data-title="تقايض" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%9C%D1%83%D1%82%D1%83%D0%B0%D0%BB%D1%96%D0%B7%D0%BC" title="Мутуалізм – Belarusian" lang="be" hreflang="be" data-title="Мутуалізм" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target"><span>Беларуская</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9C%D1%83%D1%82%D1%83%D0%B0%D0%BB%D0%B8%D0%B7%D1%8A%D0%BC" title="Мутуализъм – Bulgarian" lang="bg" hreflang="bg" data-title="Мутуализъм" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Mutualizam" title="Mutualizam – Bosnian" lang="bs" hreflang="bs" data-title="Mutualizam" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Mutualisme_(biologia)" title="Mutualisme (biologia) – Catalan" lang="ca" hreflang="ca" data-title="Mutualisme (biologia)" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Mutualismus" title="Mutualismus – Czech" lang="cs" hreflang="cs" data-title="Mutualismus" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Mutualisme_(%C3%B8kologi)" title="Mutualisme (økologi) – Danish" lang="da" hreflang="da" data-title="Mutualisme (økologi)" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Mutualismus_(Biologie)" title="Mutualismus (Biologie) – German" lang="de" hreflang="de" data-title="Mutualismus (Biologie)" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Mutualism" title="Mutualism – Estonian" lang="et" hreflang="et" data-title="Mutualism" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%92%CE%B9%CE%BF%CE%BB%CE%BF%CE%B3%CE%B9%CE%BA%CE%AE_%CE%B1%CE%BC%CE%BF%CE%B9%CE%B2%CE%B1%CE%B9%CF%8C%CF%84%CE%B7%CF%84%CE%B1" title="Βιολογική αμοιβαιότητα – Greek" lang="el" hreflang="el" data-title="Βιολογική αμοιβαιότητα" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Mutualismo_(biolog%C3%ADa)" title="Mutualismo (biología) – Spanish" lang="es" hreflang="es" data-title="Mutualismo (biología)" 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/Mutualismo" title="Mutualismo – Esperanto" lang="eo" hreflang="eo" data-title="Mutualismo" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Mutualismo_(biologia)" title="Mutualismo (biologia) – Basque" lang="eu" hreflang="eu" data-title="Mutualismo (biologia)" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%87%D9%85%D8%B2%DB%8C%D8%B3%D8%AA%DB%8C_%D8%AF%D9%88%D8%B3%D9%88%DB%8C%D9%87" title="همزیستی دوسویه – Persian" lang="fa" hreflang="fa" data-title="همزیستی دوسویه" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Mutualisme_(biologie)" title="Mutualisme (biologie) – French" lang="fr" hreflang="fr" data-title="Mutualisme (biologie)" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Comhara%C3%ADocht" title="Comharaíocht – Irish" lang="ga" hreflang="ga" data-title="Comharaíocht" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Mutualismo" title="Mutualismo – Galician" lang="gl" hreflang="gl" data-title="Mutualismo" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%83%81%EB%A6%AC_%EA%B3%B5%EC%83%9D" title="상리 공생 – Korean" lang="ko" hreflang="ko" data-title="상리 공생" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D5%84%D5%B8%D6%82%D5%BF%D5%B8%D6%82%D5%A1%D5%AC%D5%AB%D5%A6%D5%B4" title="Մուտուալիզմ – Armenian" lang="hy" hreflang="hy" data-title="Մուտուալիզմ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%B8%E0%A4%B9%E0%A5%8B%E0%A4%AA%E0%A4%95%E0%A4%BE%E0%A4%B0%E0%A4%BF%E0%A4%A4%E0%A4%BE" title="सहोपकारिता – Hindi" lang="hi" hreflang="hi" data-title="सहोपकारिता" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Mutualizam" title="Mutualizam – Croatian" lang="hr" hreflang="hr" data-title="Mutualizam" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Mutualisme" title="Mutualisme – Indonesian" lang="id" hreflang="id" data-title="Mutualisme" 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-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Samhj%C3%A1lp" title="Samhjálp – Icelandic" lang="is" hreflang="is" data-title="Samhjálp" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target"><span>Íslenska</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Mutualismo" title="Mutualismo – Italian" lang="it" hreflang="it" data-title="Mutualismo" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%94%D7%93%D7%93%D7%99%D7%95%D7%AA" title="הדדיות – Hebrew" lang="he" hreflang="he" data-title="הדדיות" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%9C%D1%83%D1%82%D1%83%D0%B0%D0%BB%D0%B8%D0%B7%D0%BC" title="Мутуализм – Kazakh" lang="kk" hreflang="kk" data-title="Мутуализм" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ht mw-list-item"><a href="https://ht.wikipedia.org/wiki/Mityalis" title="Mityalis – Haitian Creole" lang="ht" hreflang="ht" data-title="Mityalis" data-language-autonym="Kreyòl ayisyen" data-language-local-name="Haitian Creole" class="interlanguage-link-target"><span>Kreyòl ayisyen</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Mutualismus" title="Mutualismus – Latin" lang="la" hreflang="la" data-title="Mutualismus" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Mutualizmas" title="Mutualizmas – Lithuanian" lang="lt" hreflang="lt" data-title="Mutualizmas" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Mutualizmus_(biol%C3%B3gia)" title="Mutualizmus (biológia) – Hungarian" lang="hu" hreflang="hu" data-title="Mutualizmus (biológia)" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%9C%D1%83%D1%82%D1%83%D0%B0%D0%BB%D0%B8%D0%B7%D0%B0%D0%BC_(%D0%B1%D0%B8%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%98%D0%B0)" title="Мутуализам (биологија) – Macedonian" lang="mk" hreflang="mk" data-title="Мутуализам (биологија)" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%B8%E0%B4%B9%E0%B5%8B%E0%B4%AA%E0%B4%95%E0%B4%BE%E0%B4%B0%E0%B4%BF%E0%B4%A4" title="സഹോപകാരിത – Malayalam" lang="ml" hreflang="ml" data-title="സഹോപകാരിത" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Kesalingan_(biologi)" title="Kesalingan (biologi) – Malay" lang="ms" hreflang="ms" data-title="Kesalingan (biologi)" 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-mni mw-list-item"><a href="https://mni.wikipedia.org/wiki/%EA%AF%80%EA%AF%A5%EA%AF%A1%EA%AF%82%EA%AF%A8%EA%AF%9E_%EA%AF%83%EA%AF%81%EA%AF%A6%EA%AF%9F_%EA%AF%87%EA%AF%A6%EA%AF%A1%EA%AF%84%EA%AF%A5%EA%AF%A1%EA%AF%85%EA%AF%84_(%EA%AF%8A%EA%AF%8B%EA%AF%A5%EA%AF%A2%EA%AF%84%EA%AF%A5%EA%AF%9F%EA%AF%84_%EA%AF%85%EA%AF%A9%EA%AF%85%EA%AF%82%EA%AF%A3%EA%AF%9F)" title="ꯀꯥꯡꯂꯨꯞ ꯃꯁꯦꯟ ꯇꯦꯡꯄꯥꯡꯅꯄ (ꯊꯋꯥꯢꯄꯥꯟꯄ ꯅꯩꯅꯂꯣꯟ) – Manipuri" lang="mni" hreflang="mni" data-title="ꯀꯥꯡꯂꯨꯞ ꯃꯁꯦꯟ ꯇꯦꯡꯄꯥꯡꯅꯄ (ꯊꯋꯥꯢꯄꯥꯟꯄ ꯅꯩꯅꯂꯣꯟ)" data-language-autonym="ꯃꯤꯇꯩ ꯂꯣꯟ" data-language-local-name="Manipuri" class="interlanguage-link-target"><span>ꯃꯤꯇꯩ ꯂꯣꯟ</span></a></li><li class="interlanguage-link interwiki-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%9C%D1%83%D1%82%D1%83%D0%B0%D0%BB%D0%B8%D0%B7%D0%BC_(%D0%B1%D0%B8%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8)" title="Мутуализм (биологи) – Mongolian" lang="mn" hreflang="mn" data-title="Мутуализм (биологи)" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target"><span>Монгол</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Mutualisme" title="Mutualisme – Dutch" lang="nl" hreflang="nl" data-title="Mutualisme" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E7%9B%B8%E5%88%A9%E5%85%B1%E7%94%9F" title="相利共生 – Japanese" lang="ja" hreflang="ja" data-title="相利共生" 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-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Mutualisme_(biologi)" title="Mutualisme (biologi) – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Mutualisme (biologi)" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Mutualisme" title="Mutualisme – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Mutualisme" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Mutualizm" title="Mutualizm – Polish" lang="pl" hreflang="pl" data-title="Mutualizm" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Mutualismo_(biologia)" title="Mutualismo (biologia) – Portuguese" lang="pt" hreflang="pt" data-title="Mutualismo (biologia)" 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%D1%83%D1%82%D1%83%D0%B0%D0%BB%D0%B8%D0%B7%D0%BC" title="Мутуализм – Russian" lang="ru" hreflang="ru" data-title="Мутуализм" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Mutualism_(biology)" title="Mutualism (biology) – Simple English" lang="en-simple" hreflang="en-simple" data-title="Mutualism (biology)" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Mutualizem" title="Mutualizem – Slovenian" lang="sl" hreflang="sl" data-title="Mutualizem" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D8%B3%D9%88%D9%88%D8%AF%DA%AF%DB%86%DA%95%DA%A9%DB%8E" title="سوودگۆڕکێ – Central Kurdish" lang="ckb" hreflang="ckb" data-title="سوودگۆڕکێ" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%9C%D1%83%D1%82%D1%83%D0%B0%D0%BB%D0%B8%D0%B7%D0%B0%D0%BC_(%D0%B1%D0%B8%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%98%D0%B0)" title="Мутуализам (биологија) – Serbian" lang="sr" hreflang="sr" data-title="Мутуализам (биологија)" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Mutualizam" title="Mutualizam – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Mutualizam" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Mutualismi_(biologia)" title="Mutualismi (biologia) – Finnish" lang="fi" hreflang="fi" data-title="Mutualismi (biologia)" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Mutualism_(biologi)" title="Mutualism (biologi) – Swedish" lang="sv" hreflang="sv" data-title="Mutualism (biologi)" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Mutualizm_(biyoloji)" title="Mutualizm (biyoloji) – Turkish" lang="tr" hreflang="tr" data-title="Mutualizm (biyoloji)" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9C%D1%83%D1%82%D1%83%D0%B0%D0%BB%D1%96%D0%B7%D0%BC" title="Мутуалізм – Ukrainian" lang="uk" hreflang="uk" data-title="Мутуалізм" 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/Quan_h%E1%BB%87_h%E1%BB%97_tr%E1%BB%A3" title="Quan hệ hỗ trợ – Vietnamese" lang="vi" hreflang="vi" data-title="Quan hệ hỗ trợ" 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/%E4%BA%92%E5%88%A9%E5%85%B1%E7%94%9F" title="互利共生 – Cantonese" lang="yue" hreflang="yue" data-title="互利共生" data-language-autonym="粵語" data-language-local-name="Cantonese" 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0.5em;text-align:center}.mw-parser-output .ambox .mbox-imageright{border:none;padding:2px 0.5em 2px 0;text-align:center}.mw-parser-output .ambox .mbox-empty-cell{border:none;padding:0;width:1px}.mw-parser-output .ambox .mbox-image-div{width:52px}@media(min-width:720px){.mw-parser-output .ambox{margin:0 10%}}@media print{body.ns-0 .mw-parser-output .ambox{display:none!important}}</style><table class="box-Merge_from plainlinks metadata ambox ambox-move" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0f/Mergefrom.svg/50px-Mergefrom.svg.png" decoding="async" width="50" height="20" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0f/Mergefrom.svg/75px-Mergefrom.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0f/Mergefrom.svg/100px-Mergefrom.svg.png 2x" data-file-width="50" data-file-height="20" /></span></span></div></td><td class="mbox-text"><div class="mbox-text-span">It has been suggested that <i><a href="/wiki/Protocooperation" title="Protocooperation">Protocooperation</a></i> be <a href="/wiki/Wikipedia:Merging" title="Wikipedia:Merging">merged</a> into this article. (<a href="/wiki/Talk:Mutualism_(biology)" title="Talk:Mutualism (biology)">Discuss</a>)<small><i> Proposed since November 2024.</i></small></div></td></tr></tbody></table> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">This article is about the biological term. For the economic theory and other uses, see <a href="/wiki/Mutualism_(disambiguation)" class="mw-redirect mw-disambig" title="Mutualism (disambiguation)">Mutualism (disambiguation)</a>.</div> <p class="mw-empty-elt"> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Hummingbird_hawkmoth_a.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4b/Hummingbird_hawkmoth_a.jpg/220px-Hummingbird_hawkmoth_a.jpg" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4b/Hummingbird_hawkmoth_a.jpg/330px-Hummingbird_hawkmoth_a.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4b/Hummingbird_hawkmoth_a.jpg/440px-Hummingbird_hawkmoth_a.jpg 2x" data-file-width="1568" data-file-height="1568" /></a><figcaption><a href="/wiki/Hummingbird_hawkmoth" class="mw-redirect" title="Hummingbird hawkmoth">Hummingbird hawkmoth</a> drinking from <i><a href="/wiki/Dianthus" title="Dianthus">Dianthus</a></i>, with <a href="/wiki/Pollination" title="Pollination">pollination</a> being a classic example of mutualism</figcaption></figure> <p><b>Mutualism</b> describes the ecological <a href="/wiki/Biological_interaction" title="Biological interaction">interaction</a> between two or more <a href="/wiki/Species" title="Species">species</a> where each species has a net benefit.<sup id="cite_ref-Bronstein15_1-0" class="reference"><a href="#cite_note-Bronstein15-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Mutualism is a common type of <a href="/wiki/Ecology" title="Ecology">ecological</a> interaction. Prominent examples are: </p> <ul><li>the nutrient exchange between vascular plants and <a href="/wiki/Mycorrhizae" class="mw-redirect" title="Mycorrhizae">mycorrhizal fungi</a>,</li> <li>the <a href="/wiki/Fertilisation" title="Fertilisation">fertilization</a> of flowering plants by <a href="/wiki/Pollinators" class="mw-redirect" title="Pollinators">pollinators</a>,</li> <li>the ways plants use fruits and <a href="/wiki/Edible" title="Edible">edible</a> seeds to encourage animal aid in <a href="/wiki/Seed_dispersal" title="Seed dispersal">seed dispersal</a>, and</li> <li>the way <a href="/wiki/Coral" title="Coral">corals</a> become photosynthetic with the help of the microorganism <a href="/wiki/Zooxanthellae" title="Zooxanthellae">zooxanthellae</a>.</li></ul> <p>Mutualism can be contrasted with <a href="/wiki/Interspecific_competition" title="Interspecific competition">interspecific competition</a>, in which each species experiences <i>reduced</i> fitness, and <a href="/wiki/Cheating_(biology)" title="Cheating (biology)">exploitation</a>, and with <a href="/wiki/Parasitism" title="Parasitism">parasitism</a>, in which one species benefits at the expense of the other.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> However, mutualism may evolve from interactions that began with imbalanced benefits, such as <a href="/wiki/Parasitism" title="Parasitism">parasitism</a>.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>The term <i>mutualism</i> was introduced by <a href="/wiki/Pierre-Joseph_van_Beneden" title="Pierre-Joseph van Beneden">Pierre-Joseph van Beneden</a> in his 1876 book <i>Animal Parasites and Messmates</i> to mean "mutual aid among species".<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>Mutualism is often <a href="/wiki/Conflated" class="mw-redirect" title="Conflated">conflated</a> with two other types of ecological phenomena: cooperation and <a href="/wiki/Symbiosis" title="Symbiosis">symbiosis</a>. <a href="/wiki/Cooperation" title="Cooperation">Cooperation</a> most commonly refers to increases in fitness through within-species (intraspecific) interactions, although it has been used (especially in the past) to refer to mutualistic interactions, and it is sometimes used to refer to mutualistic interactions that are not obligate.<sup id="cite_ref-Bronstein15_1-1" class="reference"><a href="#cite_note-Bronstein15-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Symbiosis" title="Symbiosis">Symbiosis</a> involves two species living in close physical contact over a long period of their existence and may be mutualistic, <a href="/wiki/Parasitism" title="Parasitism">parasitic</a>, or <a href="/wiki/Commensalism" title="Commensalism">commensal</a>, so <a href="/wiki/Symbiosis" title="Symbiosis">symbiotic relationships</a> are not always mutualistic, and mutualistic interactions are not always symbiotic. Despite a different definition between mutualism and symbiosis, they have been largely used interchangeably in the past, and confusion on their use has persisted.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>Mutualism plays a key part in <a href="/wiki/Ecology" title="Ecology">ecology</a> and <a href="/wiki/Evolution" title="Evolution">evolution</a>. For example, mutualistic interactions are vital for terrestrial <a href="/wiki/Ecosystem" title="Ecosystem">ecosystem</a> function as: </p> <ul><li>about 80% of land plants species rely on <a href="/wiki/Mycorrhizal" class="mw-redirect" title="Mycorrhizal">mycorrhizal</a> relationships with <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a> to provide them with inorganic compounds and trace elements.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup></li> <li>estimates of tropical rainforest plants with seed dispersal mutualisms with animals range at least from 70% to 93.5%.<sup id="cite_ref-Jordano2000_8-0" class="reference"><a href="#cite_note-Jordano2000-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> In addition, mutualism is thought to have driven the evolution of much of the biological diversity we see, such as <a href="/wiki/Flower" title="Flower">flower</a> forms (important for <a href="/wiki/Pollination" title="Pollination">pollination</a> mutualisms) and <a href="/wiki/Co-evolution" class="mw-redirect" title="Co-evolution">co-evolution</a> between groups of species.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup></li></ul> <p>A prominent example of pollination mutualism is with bees and flowering plants. Bees use these plants as their food source with pollen and nectar. In turn, they transfer pollen to other nearby flowers, inadvertently allowing for cross-pollination. Cross-pollination has become essential in plant reproduction and fruit/seed production. The bees get their nutrients from the plants, and allow for successful fertilization of plants, demonstrating a mutualistic relationship between two seemingly-unlike species. </p><p>Mutualism has also been linked to major <a href="/wiki/Evolution" title="Evolution">evolutionary</a> events, such as the evolution of the eukaryotic cell (<a href="/wiki/Symbiogenesis" title="Symbiogenesis">symbiogenesis</a>) and the colonization of land by plants in association with <a href="/wiki/Mycorrhiza" title="Mycorrhiza">mycorrhizal</a> fungi. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Types">Types</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Mutualism_(biology)&action=edit&section=1" title="Edit section: Types"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Resource-resource_relationships">Resource-resource relationships</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Mutualism_(biology)&action=edit&section=2" title="Edit section: Resource-resource relationships"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Mutualistic relationships can be thought of as a form of "<a href="/wiki/Biology" title="Biology">biological</a> barter"<sup id="cite_ref-Ollerton06_10-0" class="reference"><a href="#cite_note-Ollerton06-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> in <a href="/wiki/Mycorrhizal" class="mw-redirect" title="Mycorrhizal">mycorrhizal</a> associations between plant <a href="/wiki/Root" title="Root">roots</a> and <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a>, with the plant providing <a href="/wiki/Carbohydrates" class="mw-redirect" title="Carbohydrates">carbohydrates</a> to the <a href="/wiki/Fungus" title="Fungus">fungus</a> in return for primarily <a href="/wiki/Phosphate" title="Phosphate">phosphate</a> but also <a href="/wiki/Nitrogenous" class="mw-redirect" title="Nitrogenous">nitrogenous</a> compounds. Other examples include <a href="/wiki/Rhizobia" title="Rhizobia">rhizobia</a> bacteria that fix nitrogen for <a href="/wiki/Leguminous" class="mw-redirect" title="Leguminous">leguminous</a> plants (family Fabaceae) in return for energy-containing <a href="/wiki/Carbohydrates" class="mw-redirect" title="Carbohydrates">carbohydrates</a>.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Metabolite exchange between multiple mutualistic species of <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> has also been observed in a process known as <a href="/wiki/Syntrophy" title="Syntrophy">cross-feeding</a>.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Service-resource_relationships">Service-resource relationships</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Mutualism_(biology)&action=edit&section=3" title="Edit section: Service-resource relationships"><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:Impala_mutualim_with_birds_wide.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Impala_mutualim_with_birds_wide.jpg/220px-Impala_mutualim_with_birds_wide.jpg" decoding="async" width="220" height="151" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Impala_mutualim_with_birds_wide.jpg/330px-Impala_mutualim_with_birds_wide.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/09/Impala_mutualim_with_birds_wide.jpg/440px-Impala_mutualim_with_birds_wide.jpg 2x" data-file-width="2231" data-file-height="1531" /></a><figcaption>The <a href="/wiki/Red-billed_oxpecker" title="Red-billed oxpecker">red-billed oxpecker</a> eats ticks on the <a href="/wiki/Impala" title="Impala">impala</a>'s coat, in a <a href="/wiki/Cleaning_symbiosis" title="Cleaning symbiosis">cleaning symbiosis</a>.</figcaption></figure> <p>Service-resource relationships are common. Three important types are <a href="/wiki/Pollination" title="Pollination">pollination</a>, <a href="/wiki/Cleaning" title="Cleaning">cleaning</a> symbiosis, and <a href="/wiki/Seed_dispersal" title="Seed dispersal">zoochory</a>. </p><p>In <a href="/wiki/Pollination" title="Pollination">pollination</a>, a plant trades food resources in the form of <a href="/wiki/Nectar" title="Nectar">nectar</a> or <a href="/wiki/Pollen" title="Pollen">pollen</a> for the service of pollen dispersal. However, daciniphilous <i><a href="/wiki/Bulbophyllum" title="Bulbophyllum">Bulbophyllum</a></i> orchid species trade <a href="/wiki/Sex_pheromone" title="Sex pheromone">sex pheromone</a> precursor or booster components via floral <a href="/wiki/Synomone" class="mw-redirect" title="Synomone">synomones</a>/attractants in a true mutualistic interactions with males of <a href="/wiki/Dacini" class="mw-redirect" title="Dacini">Dacini</a> fruit flies (Diptera: Tephritidae: Dacinae).<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Phagophile" class="mw-redirect" title="Phagophile">Phagophiles</a> feed (resource) on <a href="/wiki/Ectoparasite" class="mw-redirect" title="Ectoparasite">ectoparasites</a>, thereby providing anti-pest service, as in <a href="/wiki/Cleaning_symbiosis" title="Cleaning symbiosis">cleaning symbiosis</a>. <i><a href="/wiki/Elacatinus" title="Elacatinus">Elacatinus</a></i> and <i><a href="/wiki/Gobiosoma" title="Gobiosoma">Gobiosoma</a></i>, genera of <a href="/wiki/Goby" title="Goby">gobies</a>, feed on ectoparasites of their clients while cleaning them.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Zoochory" class="mw-redirect" title="Zoochory">Zoochory</a> is the dispersal of the seeds of plants by animals. This is similar to pollination in that the plant produces food resources (for example, fleshy fruit, overabundance of seeds) for animals that disperse the seeds (service). Plants may advertise these resources using colour<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> and a variety of other fruit characteristics, e.g., scent. Fruit of the <a href="/wiki/Aardvark_cucumber" class="mw-redirect" title="Aardvark cucumber">aardvark cucumber</a> <i>(Cucumis humifructus)</i> is buried so deeply that the plant is solely reliant upon the <a href="/wiki/Aardvark#Head" title="Aardvark">aardvark's keen sense of smell</a> to detect its ripened fruit, extract, consume and then scatter its seeds;<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> <i>C. humifructus's</i> geographical range is thus restricted to that of the aardvark's. </p><p>Another type is <a href="/wiki/Ant" title="Ant">ant</a> protection of <a href="/wiki/Aphids" class="mw-redirect" title="Aphids">aphids</a>, where the aphids trade <a href="/wiki/Sugar" title="Sugar">sugar</a>-rich <a href="/wiki/Honeydew_(secretion)" title="Honeydew (secretion)">honeydew</a> (a by-product of their mode of feeding on plant <a href="/wiki/Sap" title="Sap">sap</a>) in return for defense against <a href="/wiki/Predator" class="mw-redirect" title="Predator">predators</a> such as <a href="/wiki/Ladybug" class="mw-redirect" title="Ladybug">ladybugs</a>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (June 2024)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="Service-service_relationships">Service-service relationships</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Mutualism_(biology)&action=edit&section=4" title="Edit section: Service-service relationships"><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:Common_clownfish_curves_dnsmpl.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/70/Common_clownfish_curves_dnsmpl.jpg/220px-Common_clownfish_curves_dnsmpl.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/70/Common_clownfish_curves_dnsmpl.jpg/330px-Common_clownfish_curves_dnsmpl.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/70/Common_clownfish_curves_dnsmpl.jpg/440px-Common_clownfish_curves_dnsmpl.jpg 2x" data-file-width="800" data-file-height="600" /></a><figcaption><a href="/wiki/Ocellaris_clownfish" title="Ocellaris clownfish">Ocellaris clownfish</a> and <a href="/wiki/Heteractis_magnifica" title="Heteractis magnifica">Ritter's sea anemones</a> live in a mutual service-service symbiosis, the fish driving off <a href="/wiki/Butterflyfish" title="Butterflyfish">butterflyfish</a> and the anemone's tentacles protecting the fish from predators.</figcaption></figure> <p>Strict service-service interactions are very rare, for reasons that are far from clear.<sup id="cite_ref-Ollerton06_10-1" class="reference"><a href="#cite_note-Ollerton06-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> One example is the relationship between <a href="/wiki/Sea_anemone" title="Sea anemone">sea anemones</a> and <a href="/wiki/Anemone_fish" class="mw-redirect" title="Anemone fish">anemone fish</a> in the family <a href="/wiki/Pomacentridae" title="Pomacentridae">Pomacentridae</a>: the anemones provide the fish with protection from <a href="/wiki/Predator" class="mw-redirect" title="Predator">predators</a> (which cannot tolerate the stings of the anemone's tentacles) and the fish defend the anemones against <a href="/wiki/Butterflyfish" title="Butterflyfish">butterflyfish</a> (family <a href="/wiki/Chaetodontidae" class="mw-redirect" title="Chaetodontidae">Chaetodontidae</a>), which eat anemones. However, in common with many mutualisms, there is more than one aspect to it: in the anemonefish-anemone mutualism, waste <a href="/wiki/Ammonia" title="Ammonia">ammonia</a> from the fish feeds the <a href="/wiki/Symbiotic" class="mw-redirect" title="Symbiotic">symbiotic</a> <a href="/wiki/Algae" title="Algae">algae</a> that are found in the anemone's tentacles.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> Therefore, what appears to be a service-service mutualism in fact has a service-resource component. A second example is that of the relationship between some <a href="/wiki/Ants" class="mw-redirect" title="Ants">ants</a> in the genus <i><a href="/wiki/Pseudomyrmex" title="Pseudomyrmex">Pseudomyrmex</a></i> and trees in the <a href="/wiki/Genus" title="Genus">genus</a> <i><a href="/wiki/Acacia" title="Acacia">Acacia</a></i>, such as the <a href="/wiki/Whistling_thorn" class="mw-redirect" title="Whistling thorn">whistling thorn</a> and <a href="/wiki/Bullhorn_acacia" class="mw-redirect" title="Bullhorn acacia">bullhorn acacia</a>. The <a href="/wiki/Ants" class="mw-redirect" title="Ants">ants</a> nest inside the plant's thorns. In exchange for shelter, the ants protect acacias from attack by <a href="/wiki/Herbivores" class="mw-redirect" title="Herbivores">herbivores</a> (which they frequently eat when those are small enough, introducing a resource component to this service-service relationship) and competition from other plants by trimming back vegetation that would shade the acacia. In addition, another service-resource component is present, as the ants regularly feed on <a href="/wiki/Lipid" title="Lipid">lipid</a>-rich food-bodies called <a href="/wiki/Beltian_bodies" class="mw-redirect" title="Beltian bodies">Beltian bodies</a> that are on the <i>Acacia</i> plant.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p><p>In the <a href="/wiki/Neotropics" class="mw-redirect" title="Neotropics">neotropics</a>, the ant <i><a href="/wiki/Myrmelachista_schumanni" title="Myrmelachista schumanni">Myrmelachista schumanni</a></i> makes its nest in special cavities in <i><a href="/wiki/Duroia_hirsuta" title="Duroia hirsuta">Duroia hirsute</a></i>. Plants in the vicinity that belong to other species are killed with <a href="/wiki/Formic_acid" title="Formic acid">formic acid</a>. This selective gardening can be so aggressive that small areas of the rainforest are dominated by <i>Duroia hirsute</i>. These peculiar patches are known by local people as "<a href="/wiki/Devil%27s_garden" title="Devil's garden">devil's gardens</a>".<sup id="cite_ref-Piper07_22-0" class="reference"><a href="#cite_note-Piper07-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>In some of these relationships, the cost of the ant's protection can be quite expensive. <i><a href="/wiki/Cordia" title="Cordia">Cordia</a></i> sp. trees in the <a href="/wiki/Amazon_Rainforest" class="mw-redirect" title="Amazon Rainforest">Amazonian rainforest</a> have a kind of partnership with <i><a href="/wiki/Allomerus" title="Allomerus">Allomerus</a></i> sp. ants, which make their nests in modified leaves. To increase the amount of living space available, the ants will destroy the tree's flower buds. The flowers die and leaves develop instead, providing the ants with more dwellings. Another type of <i>Allomerus</i> sp. ant lives with the <i><a href="/wiki/Hirtella" title="Hirtella">Hirtella</a></i> sp. tree in the same forests, but in this relationship, the tree has turned the tables on the ants. When the tree is ready to produce flowers, the ant abodes on certain branches begin to wither and shrink, forcing the occupants to flee, leaving the tree's flowers to develop free from ant attack.<sup id="cite_ref-Piper07_22-1" class="reference"><a href="#cite_note-Piper07-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>The term "species group" can be used to describe the manner in which individual organisms group together. In this non-taxonomic context one can refer to "same-species groups" and "mixed-species groups." While same-species groups are the norm, examples of mixed-species groups abound. For example, zebra (<i><a href="/wiki/Equus_burchelli" class="mw-redirect" title="Equus burchelli">Equus burchelli</a></i>) and wildebeest (<i><a href="/wiki/Connochaetes_taurinus" class="mw-redirect" title="Connochaetes taurinus">Connochaetes taurinus</a></i>) can remain in association during periods of long distance <a href="https://en.wiktionary.org/wiki/migration" class="extiw" title="wikt:migration">migration</a> across the <a href="/wiki/Serengeti" title="Serengeti">Serengeti</a> as a strategy for thwarting predators. <i><a href="/wiki/Cercopithecus_mitis" class="mw-redirect" title="Cercopithecus mitis">Cercopithecus mitis</a></i> and <i><a href="/wiki/Cercopithecus_ascanius" class="mw-redirect" title="Cercopithecus ascanius">Cercopithecus ascanius</a></i>, species of monkey in the <a href="/wiki/Kakamega_Forest" title="Kakamega Forest">Kakamega Forest</a> of <a href="/wiki/Kenya" title="Kenya">Kenya</a>, can stay in close proximity and travel along exactly the same routes through the forest for periods of up to 12 hours. These mixed-species groups cannot be explained by the coincidence of sharing the same habitat. Rather, they are created by the active behavioural choice of at least one of the species in question.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Mathematical_modeling">Mathematical modeling</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Mutualism_(biology)&action=edit&section=5" title="Edit section: Mathematical modeling"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Mathematical treatments of mutualisms, like the study of mutualisms in general, have lagged behind those for <a href="/wiki/Predation" title="Predation">predation</a>, or predator-prey, consumer-resource, interactions. In models of mutualisms, the terms "type I" and "type II" <a href="/wiki/Functional_response" title="Functional response">functional responses</a> refer to the linear and saturating relationships, respectively, between the <i>benefit</i> provided to an individual of species 1 (<a href="/wiki/Dependent_and_independent_variables" title="Dependent and independent variables">dependent variable</a>) and the <i>density</i> of species 2 (independent variable).<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (June 2024)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="Type_I_functional_response">Type I functional response</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Mutualism_(biology)&action=edit&section=6" title="Edit section: Type I functional response"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>One of the simplest frameworks for modeling species interactions is the <a href="/wiki/Lotka%E2%80%93Volterra_equation" class="mw-redirect" title="Lotka–Volterra equation">Lotka–Volterra equations</a>.<sup id="cite_ref-May1981_24-0" class="reference"><a href="#cite_note-May1981-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> In this model, the changes in population densities of the two mutualists are quantified as: </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle {\begin{aligned}{\frac {dN_{1}}{dt}}&=r_{1}N_{1}-\alpha _{11}N_{1}^{2}+\beta _{12}N_{1}N_{2}\\[8pt]{\frac {dN_{2}}{dt}}&=r_{2}N_{2}-\alpha _{22}N_{2}^{2}+\beta _{21}N_{1}N_{2}\end{aligned}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mtable columnalign="right left right left right left right left right left right left" rowspacing="1.1em 0.3em" columnspacing="0em 2em 0em 2em 0em 2em 0em 2em 0em 2em 0em" displaystyle="true"> <mtr> <mtd> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <mi>d</mi> <msub> <mi>N</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>1</mn> </mrow> </msub> </mrow> <mrow> <mi>d</mi> <mi>t</mi> </mrow> </mfrac> </mrow> </mtd> <mtd> <mi></mi> <mo>=</mo> <msub> <mi>r</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>1</mn> </mrow> </msub> <msub> <mi>N</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>1</mn> </mrow> </msub> <mo>−</mo> <msub> <mi>α</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>11</mn> </mrow> </msub> <msubsup> <mi>N</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>1</mn> </mrow> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> </mrow> </msubsup> <mo>+</mo> <msub> <mi>β</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>12</mn> </mrow> </msub> <msub> <mi>N</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>1</mn> </mrow> </msub> <msub> <mi>N</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> </mrow> </msub> </mtd> </mtr> <mtr> <mtd> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <mi>d</mi> <msub> <mi>N</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> </mrow> </msub> </mrow> <mrow> <mi>d</mi> <mi>t</mi> </mrow> </mfrac> </mrow> </mtd> <mtd> <mi></mi> <mo>=</mo> <msub> <mi>r</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> </mrow> </msub> <msub> <mi>N</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> </mrow> </msub> <mo>−</mo> <msub> <mi>α</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>22</mn> </mrow> </msub> <msubsup> <mi>N</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> </mrow> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> </mrow> </msubsup> <mo>+</mo> <msub> <mi>β</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>21</mn> </mrow> </msub> <msub> <mi>N</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>1</mn> </mrow> </msub> <msub> <mi>N</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> </mrow> </msub> </mtd> </mtr> </mtable> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle {\begin{aligned}{\frac {dN_{1}}{dt}}&=r_{1}N_{1}-\alpha _{11}N_{1}^{2}+\beta _{12}N_{1}N_{2}\\[8pt]{\frac {dN_{2}}{dt}}&=r_{2}N_{2}-\alpha _{22}N_{2}^{2}+\beta _{21}N_{1}N_{2}\end{aligned}}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/909bd8d58436177fb520a348b4684d3dad491975" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -5.838ex; width:35.1ex; height:12.843ex;" alt="{\displaystyle {\begin{aligned}{\frac {dN_{1}}{dt}}&=r_{1}N_{1}-\alpha _{11}N_{1}^{2}+\beta _{12}N_{1}N_{2}\\[8pt]{\frac {dN_{2}}{dt}}&=r_{2}N_{2}-\alpha _{22}N_{2}^{2}+\beta _{21}N_{1}N_{2}\end{aligned}}}"></span></dd></dl> <p>where </p> <ul><li><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle N_{i}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msub> <mi>N</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle N_{i}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/fef58cebf23adff9199f17325aefb5515fdca99d" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.671ex; width:2.666ex; height:2.509ex;" alt="{\displaystyle N_{i}}"></span> = the population density of species <i>i</i>.</li> <li><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle r_{i}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msub> <mi>r</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle r_{i}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/a0b6d651eaf432dbf1f106021c8bb499ae83fd1f" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.671ex; width:1.848ex; height:2.009ex;" alt="{\displaystyle r_{i}}"></span> = the intrinsic growth rate of the population of species <i>i</i>.</li> <li><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \alpha _{ii}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msub> <mi>α</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> <mi>i</mi> </mrow> </msub> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \alpha _{ii}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/5b5e3f22cc4f0a4e5804694ca8f0ec31526f3261" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.671ex; width:2.855ex; height:2.009ex;" alt="{\displaystyle \alpha _{ii}}"></span> = the negative effect of within-species crowding on species <i>i</i>.</li> <li><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \beta _{ij}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msub> <mi>β</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> <mi>j</mi> </mrow> </msub> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \beta _{ij}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/e761e324c0bf9a0d964805376c1c129641cf7d2f" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -1.005ex; width:2.793ex; height:2.843ex;" alt="{\displaystyle \beta _{ij}}"></span> = the beneficial effect of the density of species <i>j</i> on species <i>i</i>.</li></ul> <p>Mutualism is in essence the <a href="/wiki/Logistic_function" title="Logistic function">logistic growth equation</a> modified for mutualistic interaction. The mutualistic interaction term represents the increase in population growth of one species as a result of the presence of greater numbers of another species. As the mutualistic interactive term β is always positive, this simple model may lead to unrealistic unbounded growth.<sup id="cite_ref-G-Alg_25-0" class="reference"><a href="#cite_note-G-Alg-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> So it may be more realistic to include a further term in the formula, representing a saturation mechanism, to avoid this occurring. </p> <div class="mw-heading mw-heading3"><h3 id="Type_II_functional_response">Type II functional response</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Mutualism_(biology)&action=edit&section=7" title="Edit section: Type II functional response"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1989, David Hamilton Wright modified the above Lotka–Volterra equations by adding a new term, <i>βM</i>/<i>K</i>, to represent a mutualistic relationship.<sup id="cite_ref-Wright_26-0" class="reference"><a href="#cite_note-Wright-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Wright also considered the concept of saturation, which means that with higher densities, there is a decrease in the benefits of further increases of the mutualist population. Without saturation, depending on the size of parameter α, species densities would increase indefinitely. Because that is not possible due to environmental constraints and carrying capacity, a model that includes saturation would be more accurate. Wright's mathematical theory is based on the premise of a simple two-species mutualism model in which the benefits of mutualism become saturated due to limits posed by handling time. Wright defines handling time as the time needed to process a food item, from the initial interaction to the start of a search for new food items and assumes that processing of food and searching for food are mutually exclusive. Mutualists that display foraging behavior are exposed to the restrictions on handling time. Mutualism can be associated with symbiosis.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (June 2024)">citation needed</span></a></i>]</sup> </p> <dl><dt>Handling time interactions</dt></dl> <p>In 1959, <a href="/wiki/C._S._Holling" title="C. S. Holling">C. S. Holling</a> performed his classic disc experiment that assumed that </p> <ol><li>the number of food items captured is proportional to the allotted <a href="/wiki/Search_time" class="mw-redirect" title="Search time">searching time</a>; and</li> <li>that there is a <a href="/wiki/Handling_time" class="mw-redirect" title="Handling time">handling time</a> variable that exists separately from the notion of search time. He then developed an equation for the Type II <a href="/wiki/Functional_response" title="Functional response">functional response</a>, which showed that the feeding rate is equivalent to</li></ol> <dl><dd><dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle {\cfrac {ax}{1+axT_{H}}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <mpadded width="0" height="8.6pt" depth="3pt"> <mrow /> </mpadded> <mstyle displaystyle="false" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mi>a</mi> <mi>x</mi> </mrow> </mstyle> </mrow> <mrow> <mpadded width="0" height="8.6pt" depth="3pt"> <mrow /> </mpadded> <mstyle displaystyle="false" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mn>1</mn> <mo>+</mo> <mi>a</mi> <mi>x</mi> <msub> <mi>T</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>H</mi> </mrow> </msub> </mrow> </mstyle> </mrow> </mfrac> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle {\cfrac {ax}{1+axT_{H}}}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/24d9e3b44adc2ae4951c9be055f4d16ef80deee1" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -3.005ex; width:10.447ex; height:7.176ex;" alt="{\displaystyle {\cfrac {ax}{1+axT_{H}}}}"></span></dd></dl></dd></dl> <p>where </p> <ul><li><i>a</i> = the instantaneous discovery rate</li> <li><i>x</i> = food item density</li> <li><i>T</i><sub><i>H</i></sub> = handling time</li></ul> <p>The equation that incorporates Type II functional response and mutualism is: </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle {\frac {dN}{dt}}=N\left[r(1-cN)+{\cfrac {baM}{1+aT_{H}M}}\right]}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <mi>d</mi> <mi>N</mi> </mrow> <mrow> <mi>d</mi> <mi>t</mi> </mrow> </mfrac> </mrow> <mo>=</mo> <mi>N</mi> <mrow> <mo>[</mo> <mrow> <mi>r</mi> <mo stretchy="false">(</mo> <mn>1</mn> <mo>−</mo> <mi>c</mi> <mi>N</mi> <mo stretchy="false">)</mo> <mo>+</mo> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <mpadded width="0" height="8.6pt" depth="3pt"> <mrow /> </mpadded> <mstyle displaystyle="false" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mi>b</mi> <mi>a</mi> <mi>M</mi> </mrow> </mstyle> </mrow> <mrow> <mpadded width="0" height="8.6pt" depth="3pt"> <mrow /> </mpadded> <mstyle displaystyle="false" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mn>1</mn> <mo>+</mo> <mi>a</mi> <msub> <mi>T</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>H</mi> </mrow> </msub> <mi>M</mi> </mrow> </mstyle> </mrow> </mfrac> </mrow> </mrow> <mo>]</mo> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle {\frac {dN}{dt}}=N\left[r(1-cN)+{\cfrac {baM}{1+aT_{H}M}}\right]}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/18d5e1d77574628a6fea8238dda57cbfc20d1898" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -3.171ex; width:36.707ex; height:7.509ex;" alt="{\displaystyle {\frac {dN}{dt}}=N\left[r(1-cN)+{\cfrac {baM}{1+aT_{H}M}}\right]}"></span></dd></dl> <p>where </p> <ul><li><i>N</i> and <i>M</i> = densities of the two mutualists</li> <li><i>r</i> = intrinsic rate of increase of <i>N</i></li> <li><i>c</i> = coefficient measuring negative intraspecific interaction. This is equivalent to inverse of the <a href="/wiki/Carrying_capacity" title="Carrying capacity">carrying capacity</a>, 1/<i>K</i>, of <i>N</i>, in the <a href="/wiki/Logistic_function#Logistic_differential_equation" title="Logistic function">logistic equation</a>.</li> <li><i>a</i> = instantaneous discovery rate</li> <li><i>b</i> = coefficient converting encounters with <i>M</i> to new units of <i>N</i></li></ul> <p>or, equivalently, </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle {\frac {dN}{dt}}=N[r(1-cN)+\beta M/(X+M)]}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <mi>d</mi> <mi>N</mi> </mrow> <mrow> <mi>d</mi> <mi>t</mi> </mrow> </mfrac> </mrow> <mo>=</mo> <mi>N</mi> <mo stretchy="false">[</mo> <mi>r</mi> <mo stretchy="false">(</mo> <mn>1</mn> <mo>−</mo> <mi>c</mi> <mi>N</mi> <mo stretchy="false">)</mo> <mo>+</mo> <mi>β</mi> <mi>M</mi> <mrow class="MJX-TeXAtom-ORD"> <mo>/</mo> </mrow> <mo stretchy="false">(</mo> <mi>X</mi> <mo>+</mo> <mi>M</mi> <mo stretchy="false">)</mo> <mo stretchy="false">]</mo> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle {\frac {dN}{dt}}=N[r(1-cN)+\beta M/(X+M)]}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/c37e1855df6b3b221d64349af8299eb47ddc8535" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -2.005ex; width:37.352ex; height:5.509ex;" alt="{\displaystyle {\frac {dN}{dt}}=N[r(1-cN)+\beta M/(X+M)]}"></span></dd></dl> <p>where </p> <ul><li><i>X</i> = 1/<i>aT</i><sub>H</sub></li> <li><i>β</i> = <i>b</i>/<i>T</i><sub>H</sub></li></ul> <p>This model is most effectively applied to free-living species that encounter a number of individuals of the mutualist part in the course of their existences. Wright notes that models of biological mutualism tend to be similar qualitatively, in that the featured <a href="/wiki/Isocline" title="Isocline">isoclines</a> generally have a positive decreasing slope, and by and large similar isocline diagrams. Mutualistic interactions are best visualized as positively sloped isoclines, which can be explained by the fact that the saturation of benefits accorded to mutualism or restrictions posed by outside factors contribute to a decreasing slope. </p><p>The type II functional response is visualized as the graph of <span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle {\cfrac {baM}{1+aT_{H}M}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <mpadded width="0" height="8.6pt" depth="3pt"> <mrow /> </mpadded> <mstyle displaystyle="false" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mi>b</mi> <mi>a</mi> <mi>M</mi> </mrow> </mstyle> </mrow> <mrow> <mpadded width="0" height="8.6pt" depth="3pt"> <mrow /> </mpadded> <mstyle displaystyle="false" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mn>1</mn> <mo>+</mo> <mi>a</mi> <msub> <mi>T</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>H</mi> </mrow> </msub> <mi>M</mi> </mrow> </mstyle> </mrow> </mfrac> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle {\cfrac {baM}{1+aT_{H}M}}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/d538b33eca162d021120af54d6a07d28cf73a6ce" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -3.005ex; width:11.56ex; height:7.176ex;" alt="{\displaystyle {\cfrac {baM}{1+aT_{H}M}}}"></span> <i>vs.</i> <i>M</i>. </p> <div class="mw-heading mw-heading2"><h2 id="Structure_of_networks">Structure of networks</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Mutualism_(biology)&action=edit&section=8" title="Edit section: Structure of networks"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Mutualistic networks made up out of the interaction between plants and pollinators were found to have a similar structure in very different ecosystems on different continents, consisting of entirely different species.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> The structure of these mutualistic networks may have large consequences for the way in which pollinator communities respond to increasingly harsh conditions and on the community carrying capacity.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>Mathematical models that examine the consequences of this network structure for the stability of pollinator communities suggest that the specific way in which plant-pollinator networks are organized minimizes competition between pollinators,<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> reduce the spread of indirect effects and thus enhance ecosystem stability<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> and may even lead to strong indirect facilitation between pollinators when conditions are harsh.<sup id="cite_ref-Lever_et_al_31-0" class="reference"><a href="#cite_note-Lever_et_al-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> This means that pollinator species together can survive under harsh conditions. But it also means that pollinator species collapse simultaneously when conditions pass a critical point.<sup id="cite_ref-Garcia_et_al_32-0" class="reference"><a href="#cite_note-Garcia_et_al-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> This simultaneous collapse occurs, because pollinator species depend on each other when surviving under difficult conditions.<sup id="cite_ref-Lever_et_al_31-1" class="reference"><a href="#cite_note-Lever_et_al-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p><p>Such a community-wide collapse, involving many pollinator species, can occur suddenly when increasingly harsh conditions pass a critical point and recovery from such a collapse might not be easy. The improvement in conditions needed for pollinators to recover could be substantially larger than the improvement needed to return to conditions at which the pollinator community collapsed.<sup id="cite_ref-Lever_et_al_31-2" class="reference"><a href="#cite_note-Lever_et_al-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Humans">Humans</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Mutualism_(biology)&action=edit&section=9" title="Edit section: Humans"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Backing_sheep_at_sheepdog_competition.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/00/Backing_sheep_at_sheepdog_competition.jpg/170px-Backing_sheep_at_sheepdog_competition.jpg" decoding="async" width="170" height="227" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/00/Backing_sheep_at_sheepdog_competition.jpg/255px-Backing_sheep_at_sheepdog_competition.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/00/Backing_sheep_at_sheepdog_competition.jpg/340px-Backing_sheep_at_sheepdog_competition.jpg 2x" data-file-width="2112" data-file-height="2816" /></a><figcaption><a href="/wiki/Dog" title="Dog">Dogs</a> and <a href="/wiki/Sheep" title="Sheep">sheep</a> were among the first animals to be <a href="/wiki/Domestication" title="Domestication">domesticated</a>.</figcaption></figure> <p>Humans are involved in mutualisms with other species: their <a href="/wiki/Gut_flora" class="mw-redirect" title="Gut flora">gut flora</a> is essential for efficient <a href="/wiki/Digestion" title="Digestion">digestion</a>.<sup id="cite_ref-Sears_33-0" class="reference"><a href="#cite_note-Sears-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> Infestations of <a href="/wiki/Head_lice" class="mw-redirect" title="Head lice">head lice</a> <i>might</i> have been beneficial for humans by fostering an <a href="/wiki/Immune" class="mw-redirect" title="Immune">immune</a> response that helps to reduce the threat of <a href="/wiki/Body_louse" title="Body louse">body louse</a> borne lethal diseases.<sup id="cite_ref-mutualism_34-0" class="reference"><a href="#cite_note-mutualism-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p><p>Some relationships between humans and <a href="/wiki/Domestication" title="Domestication">domesticated</a> animals and plants are to different degrees mutualistic. For example, agricultural varieties of <a href="/wiki/Maize" title="Maize">maize</a> provide food for humans and are unable to reproduce without human intervention because the leafy sheath does not fall open, and the seedhead (the "corn on the cob") does not <a href="/wiki/Shattering_(agriculture)" title="Shattering (agriculture)">shatter</a> to scatter the seeds naturally.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (August 2022)">citation needed</span></a></i>]</sup> </p><p>In <a href="/wiki/Traditional_agriculture" class="mw-redirect" title="Traditional agriculture">traditional agriculture</a>, some plants have mutualist as <a href="/wiki/Companion_planting" title="Companion planting">companion plants</a>, providing each other with shelter, soil fertility and/or natural <a href="/wiki/Pest_control" title="Pest control">pest control</a>. For example, <a href="/wiki/Bean" title="Bean">beans</a> may grow up <a href="/wiki/Maize" title="Maize">cornstalks</a> as a trellis, while fixing nitrogen in the soil for the corn, a phenomenon that is used in <a href="/wiki/Three_Sisters_(agriculture)" title="Three Sisters (agriculture)">Three Sisters farming</a>.<sup id="cite_ref-Mount_Pleasant_2006_35-0" class="reference"><a href="#cite_note-Mount_Pleasant_2006-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p><p>One researcher has proposed that the key advantage <i><a href="/wiki/Homo_sapiens" class="mw-redirect" title="Homo sapiens">Homo sapiens</a></i> had over <a href="/wiki/Neanderthals" class="mw-redirect" title="Neanderthals">Neanderthals</a> in competing over similar habitats was the former's mutualism with dogs.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Intestinal_microbiota">Intestinal microbiota</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Mutualism_(biology)&action=edit&section=10" title="Edit section: Intestinal microbiota"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Gut_microbiota" title="Gut microbiota">microbiota in the human intestine</a> coevolved with the human species, and this relationship is considered to be a mutualism that is beneficial both to the human host and the bacteria in the gut population.<sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Mucous_membrane" title="Mucous membrane">mucous layer</a> of the intestine contains <a href="/wiki/Commensalism" title="Commensalism">commensal</a> bacteria that produce <a href="/wiki/Bacteriocin" title="Bacteriocin">bacteriocins</a>, modify the pH of the intestinal contents, and compete for nutrition to inhibit colonization by pathogens.<sup id="cite_ref-Genua2021_38-0" class="reference"><a href="#cite_note-Genua2021-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> The gut microbiota, containing trillions of <a href="/wiki/Microorganism" title="Microorganism">microorganisms</a>, possesses the metabolic capacity to produce and regulate multiple compounds that reach the circulation and act to influence the function of distal organs and systems.<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> Breakdown of the protective mucosal barrier of the gut can contribute to the development of <a href="/wiki/Colorectal_cancer" title="Colorectal cancer">colon cancer</a>.<sup id="cite_ref-Genua2021_38-1" class="reference"><a href="#cite_note-Genua2021-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Evolution_of_mutualism">Evolution of mutualism</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Mutualism_(biology)&action=edit&section=11" title="Edit section: Evolution of mutualism"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Evolution_by_type">Evolution by type</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Mutualism_(biology)&action=edit&section=12" title="Edit section: Evolution by type"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Every generation of every organism needs nutrients – and similar nutrients – more than they need particular defensive characteristics, as the fitness benefit of these vary heavily especially by environment. This may be the reason that hosts are more likely to evolve to become dependent on vertically transmitted bacterial mutualists which provide nutrients than those providing defensive benefits. This pattern is generalized beyond bacteria by Yamada et al. 2015's demonstration that undernourished <i><a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila</a></i> are heavily dependent on their fungal symbiont <i><a href="/wiki/Issatchenkia" title="Issatchenkia">Issatchenkia</a> <a href="/w/index.php?title=Issatchenkia_orientalis&action=edit&redlink=1" class="new" title="Issatchenkia orientalis (page does not exist)">orientalis</a></i> for amino acids.<sup id="cite_ref-Biedermann-Vega-2020_40-0" class="reference"><a href="#cite_note-Biedermann-Vega-2020-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Mutualism_breakdown">Mutualism breakdown</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Mutualism_(biology)&action=edit&section=13" title="Edit section: Mutualism breakdown"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Mutualisms are not static, and can be lost by evolution.<sup id="cite_ref-SachsSimms2006_41-0" class="reference"><a href="#cite_note-SachsSimms2006-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> Sachs and Simms (2006) suggest that this can occur via four main pathways: </p> <ol><li>One mutualist shifts to parasitism, and no longer benefits its partner,<sup id="cite_ref-SachsSimms2006_41-1" class="reference"><a href="#cite_note-SachsSimms2006-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> such as headlice<sup id="cite_ref-RÓZSA-2012_42-0" class="reference"><a href="#cite_note-RÓZSA-2012-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup></li> <li>One partner abandons the mutualism and lives autonomously<sup id="cite_ref-SachsSimms2006_41-2" class="reference"><a href="#cite_note-SachsSimms2006-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup></li> <li>One partner may go extinct<sup id="cite_ref-SachsSimms2006_41-3" class="reference"><a href="#cite_note-SachsSimms2006-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup></li> <li>A partner may be switched to another species<sup id="cite_ref-Werner-et-al-2018_43-0" class="reference"><a href="#cite_note-Werner-et-al-2018-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup></li></ol> <p>There are many examples of mutualism breakdown. For example, plant lineages inhabiting nutrient-rich environments have evolutionarily abandoned mycorrhizal mutualisms many times independently.<sup id="cite_ref-Wang-Qiu-2006_44-0" class="reference"><a href="#cite_note-Wang-Qiu-2006-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> Evolutionarily, headlice may have been mutualistic as they allow for early immunity to various body-louse borne disease; however, as these diseases became eradicated, the relationship has become less mutualistic and more parasitic.<sup id="cite_ref-RÓZSA-2012_42-1" class="reference"><a href="#cite_note-RÓZSA-2012-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Measuring_and_defining_mutualism">Measuring and defining mutualism</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Mutualism_(biology)&action=edit&section=14" title="Edit section: Measuring and defining mutualism"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Measuring the exact <a href="/wiki/Fitness_(biology)" title="Fitness (biology)">fitness</a> benefit to the individuals in a mutualistic relationship is not always straightforward, particularly when the individuals can receive benefits from a variety of species, for example most plant-<a href="/wiki/Pollinator" title="Pollinator">pollinator</a> mutualisms. It is therefore common to categorise mutualisms according to the closeness of the association, using terms such as <a href="/wiki/Obligate_mutualism" title="Obligate mutualism">obligate</a> and <a href="https://en.wiktionary.org/wiki/facultative" class="extiw" title="wikt:facultative">facultative</a>. Defining "closeness", however, is also problematic. It can refer to mutual dependency (the species cannot live without one another) or the biological intimacy of the relationship in relation to physical closeness (<i>e.g.</i>, one species living within the tissues of the other species).<sup id="cite_ref-Ollerton06_10-2" class="reference"><a href="#cite_note-Ollerton06-10"><span class="cite-bracket">[</span>10<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=Mutualism_(biology)&action=edit&section=15" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Arbuscular_mycorrhiza" title="Arbuscular mycorrhiza">Arbuscular mycorrhiza</a></li> <li><a href="/wiki/Co-adaptation" title="Co-adaptation">Co-adaptation</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a></li> <li><a href="/wiki/Ecological_facilitation" title="Ecological facilitation">Ecological facilitation</a></li> <li><a href="/wiki/Frugivore" title="Frugivore">Frugivore</a></li> <li><a href="/wiki/Greater_honeyguide" title="Greater honeyguide">Greater honeyguide</a> – has a mutualism with humans</li> <li><a href="/wiki/Interspecies_communication" title="Interspecies communication">Interspecies communication</a></li> <li><a href="/wiki/M%C3%BCllerian_mimicry" title="Müllerian mimicry">Müllerian mimicry</a></li> <li><a href="/wiki/Mutualisms_and_conservation" title="Mutualisms and conservation">Mutualisms and conservation</a></li> <li><i><a href="/wiki/Mutual_Aid:_A_Factor_of_Evolution" title="Mutual Aid: A Factor of Evolution">Mutual Aid: A Factor of Evolution</a></i></li> <li><a href="/wiki/Symbiogenesis" title="Symbiogenesis">Symbiogenesis</a></li> <li><a href="/wiki/Plant%E2%80%93animal_interaction" title="Plant–animal interaction">Plant–animal interaction</a></li></ul> <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=Mutualism_(biology)&action=edit&section=16" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-Bronstein15-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Bronstein15_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Bronstein15_1-1"><sup><i><b>b</b></i></sup></a></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="CITEREFBronstein2015" class="citation book cs1">Bronstein, J.L. (2015). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=hbgVDAAAQBAJ&pg=PA3">"1. The study of mutualism"</a>. <a rel="nofollow" class="external text" href="https://books.google.com/books?id=hbgVDAAAQBAJ&pg=PR5"><i>Mutualism</i></a>. Oxford University Press. pp. 3–19. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-166319-2" title="Special:BookSources/978-0-19-166319-2"><bdi>978-0-19-166319-2</bdi></a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/913513762">913513762</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=1.+The+study+of+mutualism&rft.btitle=Mutualism&rft.pages=3-19&rft.pub=Oxford+University+Press&rft.date=2015&rft_id=info%3Aoclcnum%2F913513762&rft.isbn=978-0-19-166319-2&rft.aulast=Bronstein&rft.aufirst=J.L.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DhbgVDAAAQBAJ%26pg%3DPA3&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMutualism+%28biology%29" class="Z3988"></span></span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.ixl.com/science/grade-7/classify-symbiotic-relationships">"Classify symbiotic relationships"</a>. <i>7th grade science</i>. 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Retrieved <span class="nowrap">4 December</span> 2023</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=7th+grade+science&rft.atitle=Classify+symbiotic+relationships&rft_id=https%3A%2F%2Fwww.ixl.com%2Fscience%2Fgrade-7%2Fclassify-symbiotic-relationships&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMutualism+%28biology%29" 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 class="citation web cs1"><a rel="nofollow" class="external text" href="https://yaledailynews.com/blog/2018/02/06/yale-researchers-show-that-mutualism-can-come-from-parasitism/#:~:text=The%20viruses%20and%20bacteria%20originally,benefited%20from%20one%20another%27s%20presence">"Yale researchers show that mutualism can come from parasitism"</a>. 6 February 2018.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Yale+researchers+show+that+mutualism+can+come+from+parasitism&rft.date=2018-02-06&rft_id=https%3A%2F%2Fyaledailynews.com%2Fblog%2F2018%2F02%2F06%2Fyale-researchers-show-that-mutualism-can-come-from-parasitism%2F%23%3A~%3Atext%3DThe%2520viruses%2520and%2520bacteria%2520originally%2Cbenefited%2520from%2520one%2520another%2527s%2520presence&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMutualism+%28biology%29" 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="CITEREFVan_Beneden,_Pierre-Joseph1876" class="citation book cs1"><a href="/wiki/Pierre-Joseph_van_Beneden" title="Pierre-Joseph van Beneden">Van Beneden, Pierre-Joseph</a> (1876). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/bibliography/132633#/summary"><i>Animal Parasites and Messmates</i></a>. London: Henry S. King.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Animal+Parasites+and+Messmates&rft.place=London&rft.pub=Henry+S.+King&rft.date=1876&rft.au=Van+Beneden%2C+Pierre-Joseph&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fbibliography%2F132633%23%2Fsummary&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMutualism+%28biology%29" 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"><a href="#CITEREFBronstein2015">Bronstein 2015</a>, §1.2.2.1 Mutualism versus symbiosis. p. 6</span> </li> <li id="cite_note-6"><span class="mw-cite-backlink"><b><a href="#cite_ref-6">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDouglas2014" class="citation book cs1">Douglas, Angela E. (December 2014). <i>The Symbiotic Habit</i>. 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"Mutualism and parasitism: the yin and yang of plant symbioses". <i><a href="/wiki/Current_Opinion_in_Plant_Biology" class="mw-redirect" title="Current Opinion in Plant Biology">Current Opinion in Plant Biology</a></i>. <b>9</b> (4): 364–370. <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/2006COPB....9..364P">2006COPB....9..364P</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.pbi.2006.05.008">10.1016/j.pbi.2006.05.008</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/16713732">16713732</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Current+Opinion+in+Plant+Biology&rft.atitle=Mutualism+and+parasitism%3A+the+yin+and+yang+of+plant+symbioses&rft.volume=9&rft.issue=4&rft.pages=364-370&rft.date=2006&rft_id=info%3Apmid%2F16713732&rft_id=info%3Adoi%2F10.1016%2Fj.pbi.2006.05.008&rft_id=info%3Abibcode%2F2006COPB....9..364P&rft.aulast=Paszkowski&rft.aufirst=U&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMutualism+%28biology%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPoratChadwick-Furman2004" class="citation journal cs1">Porat, D.; Chadwick-Furman, N. E. (2004). "Effects of anemonefish on giant sea anemones:expansion behavior, growth, and survival". <i><a href="/wiki/Hydrobiologia" title="Hydrobiologia">Hydrobiologia</a></i>. <b>530</b> (1–3): 513–520. <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/2004HyBio.530..513P">2004HyBio.530..513P</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs10750-004-2688-y">10.1007/s10750-004-2688-y</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:2251533">2251533</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Hydrobiologia&rft.atitle=Effects+of+anemonefish+on+giant+sea+anemones%3Aexpansion+behavior%2C+growth%2C+and+survival&rft.volume=530&rft.issue=1%E2%80%933&rft.pages=513-520&rft.date=2004&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A2251533%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1007%2Fs10750-004-2688-y&rft_id=info%3Abibcode%2F2004HyBio.530..513P&rft.aulast=Porat&rft.aufirst=D.&rft.au=Chadwick-Furman%2C+N.+E.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMutualism+%28biology%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPoratChadwick-Furman2005" class="citation journal cs1">Porat, D.; Chadwick-Furman, N. E. (2005). "Effects of anemonefish on giant sea anemones: ammonium uptake, zooxanthella content and tissue regeneration". <i>Mar. Freshw. Behav. Phys</i>. <b>38</b> (1): 43–51. <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/2005MFBP...38...43P">2005MFBP...38...43P</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F10236240500057929">10.1080/10236240500057929</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:53051081">53051081</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Mar.+Freshw.+Behav.+Phys.&rft.atitle=Effects+of+anemonefish+on+giant+sea+anemones%3A+ammonium+uptake%2C+zooxanthella+content+and+tissue+regeneration&rft.volume=38&rft.issue=1&rft.pages=43-51&rft.date=2005&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A53051081%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1080%2F10236240500057929&rft_id=info%3Abibcode%2F2005MFBP...38...43P&rft.aulast=Porat&rft.aufirst=D.&rft.au=Chadwick-Furman%2C+N.+E.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMutualism+%28biology%29" class="Z3988"></span></li> <li>Thompson, J. N. 2005. <i>The Geographic Mosaic of Coevolution</i>. University of Chicago Press. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-226-79762-5" title="Special:BookSources/978-0-226-79762-5">978-0-226-79762-5</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWright1989" class="citation journal cs1">Wright, David Hamilton (1989). "A Simple, Stable Model of Mutualism Incorporating Handling Time". <i><a href="/wiki/The_American_Naturalist" title="The American Naturalist">The American Naturalist</a></i>. <b>134</b> (4): 664–7. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1086%2F285003">10.1086/285003</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:83502337">83502337</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+American+Naturalist&rft.atitle=A+Simple%2C+Stable+Model+of+Mutualism+Incorporating+Handling+Time&rft.volume=134&rft.issue=4&rft.pages=664-7&rft.date=1989&rft_id=info%3Adoi%2F10.1086%2F285003&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A83502337%23id-name%3DS2CID&rft.aulast=Wright&rft.aufirst=David+Hamilton&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMutualism+%28biology%29" class="Z3988"></span></li></ul> </div> <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=Mutualism_(biology)&action=edit&section=18" title="Edit section: Further reading"><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/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/45px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/59px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <span style="font-weight: bold; font-style: italic;"><a href="https://commons.wikimedia.org/wiki/Category:Mutualism_(biology)" class="extiw" title="commons:Category:Mutualism (biology)">Mutualism (biology)</a></span>.</div></div> </div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1235681985"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1237033735"><div class="side-box side-box-right plainlinks sistersitebox"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1126788409"> <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/9/99/Wiktionary-logo-en-v2.svg/40px-Wiktionary-logo-en-v2.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/60px-Wiktionary-logo-en-v2.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/80px-Wiktionary-logo-en-v2.svg.png 2x" data-file-width="512" data-file-height="512" /></span></span></div> <div class="side-box-text plainlist">Look up <i><b><a href="https://en.wiktionary.org/wiki/Special:Search/mutualism_(biology)" class="extiw" title="wiktionary:Special:Search/mutualism (biology)">mutualism (biology)</a></b></i> in Wiktionary, the free dictionary.</div></div> </div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBoucherJamesKeeler1984" class="citation journal cs1">Boucher, D. G.; James, S.; Keeler, K. (1984). "The ecology of mutualism". <i><a href="/wiki/Annual_Review_of_Ecology_and_Systematics" class="mw-redirect" title="Annual Review of Ecology and Systematics">Annual Review of Ecology and Systematics</a></i>. <b>13</b>: 315–347. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.es.13.110182.001531">10.1146/annurev.es.13.110182.001531</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annual+Review+of+Ecology+and+Systematics&rft.atitle=The+ecology+of+mutualism&rft.volume=13&rft.pages=315-347&rft.date=1984&rft_id=info%3Adoi%2F10.1146%2Fannurev.es.13.110182.001531&rft.aulast=Boucher&rft.aufirst=D.+G.&rft.au=James%2C+S.&rft.au=Keeler%2C+K.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMutualism+%28biology%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBoucher1985" class="citation book cs1">Boucher, D.H. (1985). <i>The Biology of Mutualism : Ecology and Evolution</i>. Oxford University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-7099-3238-3" title="Special:BookSources/0-7099-3238-3"><bdi>0-7099-3238-3</bdi></a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/11971241">11971241</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Biology+of+Mutualism+%3A+Ecology+and+Evolution&rft.pub=Oxford+University+Press&rft.date=1985&rft_id=info%3Aoclcnum%2F11971241&rft.isbn=0-7099-3238-3&rft.aulast=Boucher&rft.aufirst=D.H.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMutualism+%28biology%29" class="Z3988"></span></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist 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<ul><li><a href="/wiki/Biological_interaction#Amensalism" title="Biological interaction">Amensalism</a></li> <li><a href="/wiki/Commensalism" title="Commensalism">Commensalism</a></li> <li><a href="/wiki/Competition_(biology)" title="Competition (biology)">Competition</a></li> <li><a href="/wiki/Deception_in_animals" title="Deception in animals">Deception in animals</a> <ul><li><a href="/wiki/Tactical_deception_in_animals" title="Tactical deception in animals">Tactical</a></li></ul></li> <li><a href="/wiki/Inquilinism" class="mw-redirect" title="Inquilinism">Inquilinism</a></li> <li><a href="/wiki/Mimicry" title="Mimicry">Mimicry</a></li> <li><a class="mw-selflink selflink">Mutualism</a></li> <li><a href="/wiki/Biological_interaction#Neutralism" title="Biological interaction">Neutralism</a></li> <li><a href="/wiki/Biological_interaction#Competition" title="Biological interaction">Synnecrosis</a></li> <li><a href="/wiki/Parasitism" title="Parasitism">Parasitism</a> <ul><li><a href="/wiki/Behavior-altering_parasite" title="Behavior-altering parasite">Behavior-altering parasite</a></li> <li><a href="/wiki/Brood_parasitism" title="Brood parasitism">Brood parasitism</a></li> <li><a href="/wiki/Host%E2%80%93parasite_coevolution" title="Host–parasite coevolution">Host–parasite coevolution</a></li> <li><a href="/wiki/Hyperparasite" title="Hyperparasite">Hyperparasite</a></li> <li><a href="/wiki/Kleptoparasitism" title="Kleptoparasitism">Kleptoparasitism</a></li> <li><a href="/wiki/Parasitic_castration" title="Parasitic castration">Parasitic castration</a></li> <li><a href="/wiki/Parasitoid" title="Parasitoid">Parasitoid</a></li></ul></li> <li><a href="/wiki/Predation" title="Predation">Predation</a> <ul><li><span style="font-size:85%;"><a href="/wiki/Carnivore" title="Carnivore">Carnivore</a></span></li> <li><span style="font-size:85%;"><a href="/wiki/Intraguild_predation" title="Intraguild predation">Intraguild</a></span></li></ul></li> <li><a href="/wiki/Symbiosis" title="Symbiosis">Symbiosis</a> <ul><li><span style="font-size:85%;"><a href="/wiki/Cleaning_symbiosis" title="Cleaning symbiosis">Cleaning symbiosis</a></span></li> <li><span style="font-size:85%;"><a href="/wiki/Mycorrhiza" title="Mycorrhiza">Mycorrhiza</a></span></li></ul></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_ecology" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evo_ecol" title="Template:Evo ecol"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evo_ecol" title="Template talk:Evo ecol"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evo_ecol" title="Special:EditPage/Template:Evo ecol"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_ecology" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Patterns of evolution</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Coloration_evidence_for_natural_selection" title="Coloration evidence for natural selection">Coloration evidence for natural selection</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergent evolution</a> <ul><li><a href="/wiki/List_of_examples_of_convergent_evolution" title="List of examples of convergent evolution">examples</a></li></ul></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergent evolution</a></li> <li><a href="/wiki/Paradox_of_the_plankton" title="Paradox of the plankton">Paradox of the plankton</a></li> <li><a href="/wiki/Predator_satiation" title="Predator satiation">Predator satiation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Signals</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a></li> <li><a href="/wiki/Anti-predator_adaptation" title="Anti-predator adaptation">Anti-predator adaptation</a> <ul><li><a href="/wiki/Alarm_signal" title="Alarm signal">Alarm signal</a></li> <li><a href="/wiki/Aposematism" title="Aposematism">Aposematism</a></li> <li><a href="/wiki/Apparent_death" title="Apparent death">Apparent death</a></li> <li><a href="/wiki/Deimatic_behaviour" title="Deimatic behaviour">Deimatic behaviour</a></li> <li><a href="/wiki/Distraction_display" title="Distraction display">Distraction display</a></li></ul></li> <li class="mw-empty-elt"></li> <li><a href="/wiki/Crypsis" title="Crypsis">Crypsis</a></li> <li><a href="/wiki/Camouflage" title="Camouflage">Camouflage</a></li> <li><a href="/wiki/Mimicry" title="Mimicry">Mimicry</a></li> <li><a href="/wiki/Unkenreflex" title="Unkenreflex">Unkenreflex</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_biology" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evolution" title="Template:Evolution"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolution" title="Template talk:Evolution"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolution" title="Special:EditPage/Template:Evolution"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_biology" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li> <li><a href="/wiki/Outline_of_evolution" title="Outline of evolution">Outline</a></li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li> <li><a href="/wiki/History_of_life" title="History of life">History of life</a></li> <li><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution" title="Evolution">Evolution</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></li> <li><a href="/wiki/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a href="/wiki/Cheating_(biology)" title="Cheating (biology)">Cheating</a></li> <li><a href="/wiki/Reciprocal_altruism" title="Reciprocal altruism">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a> <ul><li><a class="mw-selflink selflink">Mutualism</a></li></ul></li> <li><a href="/wiki/Common_descent" title="Common descent">Common descent</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergence</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergence</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li><a href="/wiki/Evidence_of_common_descent" title="Evidence of common descent">Evidence of evolution</a></li> <li><a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">Evolutionary arms race</a></li> <li><a href="/wiki/Evolutionary_pressure" title="Evolutionary pressure">Evolutionary pressure</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Extinction" title="Extinction">Extinction</a> <ul><li><a href="/wiki/Extinction_event" title="Extinction event">Event</a></li></ul></li> <li><a href="/wiki/Homology_(biology)" title="Homology (biology)">Homology</a></li> <li><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Macroevolution" title="Macroevolution">Macroevolution</a></li> <li><a href="/wiki/Microevolution" title="Microevolution">Microevolution</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Nonadaptive_radiation" title="Nonadaptive radiation">Non-adaptive radiation</a></li> <li><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></li> <li><a href="/wiki/Panspermia" title="Panspermia">Panspermia</a></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a> <ul><li><a href="/wiki/Handicap_principle" title="Handicap principle">Handicap principle</a></li></ul></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a> <ul><li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li></ul></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Taxonomy</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centered view of evolution</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Population_genetics" title="Population genetics">Population<br />genetics</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Selective_breeding" title="Selective breeding">Artificial selection</a></li> <li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness" title="Inclusive fitness">Inclusive</a></li></ul></li> <li><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a></li> <li><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a> <ul><li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a></li> <li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a> <ul><li><a href="/wiki/Sexual_selection_in_flowering_plants" title="Sexual selection in flowering plants">Flowering plants</a></li> <li><a href="/wiki/Sexual_selection_in_fungi" title="Sexual selection in fungi">Fungi</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Variation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Development</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Inversion_(evolutionary_biology)" title="Inversion (evolutionary biology)">Inversion</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Phenotypic plasticity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Taxon" title="Taxon">taxa</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_bacteria" title="Evolution of bacteria">Bacteria</a></li> <li><a href="/wiki/Evolution_of_birds" title="Evolution of birds">Birds</a> <ul><li><a href="/wiki/Origin_of_birds" title="Origin of birds">origin</a></li></ul></li> <li><a href="/wiki/Evolution_of_brachiopods" title="Evolution of brachiopods">Brachiopods</a></li> <li><a href="/wiki/Evolution_of_molluscs" title="Evolution of molluscs">Molluscs</a> <ul><li><a href="/wiki/Evolution_of_cephalopods" title="Evolution of cephalopods">Cephalopods</a></li></ul></li> <li><a href="/wiki/Dinosaur#Evolutionary_history" title="Dinosaur">Dinosaurs</a></li> <li><a href="/wiki/Evolution_of_fish" title="Evolution of fish">Fish</a></li> <li><a href="/wiki/Evolution_of_fungi" title="Evolution of fungi">Fungi</a></li> <li><a href="/wiki/Evolution_of_insects" title="Evolution of insects">Insects</a> <ul><li><a href="/wiki/Evolution_of_butterflies" title="Evolution of butterflies">butterflies</a></li></ul></li> <li><a href="/wiki/History_of_life" title="History of life">Life</a></li> <li><a href="/wiki/Evolution_of_mammals" title="Evolution of mammals">Mammals</a> <ul><li><a href="/wiki/Cat_gap" title="Cat gap">cats</a></li> <li><a href="/wiki/Canidae#Evolution" title="Canidae">canids</a> <ul><li><a href="/wiki/Evolution_of_the_wolf" title="Evolution of the wolf">wolves</a></li> <li><a href="/wiki/Domestication_of_the_dog" title="Domestication of the dog">dogs</a></li></ul></li> <li><a href="/wiki/Hyena#Evolution" title="Hyena">hyenas</a></li> <li><a href="/wiki/Evolution_of_cetaceans" title="Evolution of cetaceans">dolphins and whales</a></li> <li><a href="/wiki/Evolution_of_the_horse" title="Evolution of the horse">horses</a></li> <li><a href="/wiki/Evolution_of_Macropodidae" title="Evolution of Macropodidae">Kangaroos</a></li> <li><a href="/wiki/Evolution_of_primates" title="Evolution of primates">primates</a> <ul><li><a href="/wiki/Human_evolution" title="Human evolution">humans</a></li> <li><a href="/wiki/Evolution_of_lemurs" title="Evolution of lemurs">lemurs</a></li></ul></li> <li><a href="/wiki/Evolution_of_sirenians" title="Evolution of sirenians">sea cows</a></li></ul></li> <li><a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">Plants</a> <ul><li><a href="/wiki/Pollinator-mediated_selection" title="Pollinator-mediated selection">pollinator-mediated</a></li></ul></li> <li><a href="/wiki/Evolution_of_reptiles" title="Evolution of reptiles">Reptiles</a></li> <li><a href="/wiki/Evolution_of_spiders" title="Evolution of spiders">Spiders</a></li> <li><a href="/wiki/Evolution_of_tetrapods" title="Evolution of tetrapods">Tetrapods</a></li> <li><a href="/wiki/Viral_evolution" title="Viral evolution">Viruses</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">organs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_cells" title="Evolution of cells">Cell</a></li> <li><a href="/wiki/Models_of_DNA_evolution" title="Models of DNA evolution">DNA</a></li> <li><a href="/wiki/Evolution_of_flagella" title="Evolution of flagella">Flagella</a></li> <li><a href="/wiki/Eukaryote#Origin_of_eukaryotes" title="Eukaryote">Eukaryotes</a> <ul><li><a href="/wiki/Symbiogenesis" title="Symbiogenesis">symbiogenesis</a></li> <li><a href="/wiki/Chromosome#Eukaryotes" title="Chromosome">chromosome</a></li> <li><a href="/wiki/Endomembrane_system#Evolution" title="Endomembrane system">endomembrane system</a></li> <li><a href="/wiki/Mitochondrion#Origin_and_evolution" title="Mitochondrion">mitochondria</a></li> <li><a href="/wiki/Cell_nucleus#Evolution" title="Cell nucleus">nucleus</a></li> <li><a href="/wiki/Plastid#Origin" title="Plastid">plastids</a></li></ul></li> <li>In animals <ul><li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">eye</a></li> <li><a href="/wiki/Hair#Evolution" title="Hair">hair</a></li> <li><a href="/wiki/Evolution_of_mammalian_auditory_ossicles" title="Evolution of mammalian auditory ossicles">auditory ossicle</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">nervous system</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">brain</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Biological_process" title="Biological process">processes</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_ageing" title="Evolution of ageing">Aging</a> <ul><li><a href="/wiki/Death#Evolution_of_aging_and_mortality" title="Death">Death</a></li> <li><a href="/wiki/Programmed_cell_death#Evolutionary_origin_of_mitochondrial_apoptosis" title="Programmed cell death">Programmed cell death</a></li></ul></li> <li><a href="/wiki/Origin_of_avian_flight" title="Origin of avian flight">Avian flight</a></li> <li><a href="/wiki/Evolution_of_biological_complexity" title="Evolution of biological complexity">Biological complexity</a></li> <li><a href="/wiki/Cooperation_(evolution)" title="Cooperation (evolution)">Cooperation</a></li> <li><a href="/wiki/Evolution_of_color_vision" title="Evolution of color vision">Color vision</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">in primates</a></li></ul></li> <li><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></li> <li><a href="/wiki/Empathy#Evolution_across_species" title="Empathy">Empathy</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Ethics</a></li> <li><a href="/wiki/Evolution_of_eusociality" title="Evolution of eusociality">Eusociality</a></li> <li><a href="/wiki/Immune_system#Evolution_and_other_mechanisms" title="Immune system">Immune system</a></li> <li><a href="/wiki/Metabolism#Evolution" title="Metabolism">Metabolism</a></li> <li><a href="/wiki/Monogamy_in_animals" title="Monogamy in animals">Monogamy</a></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a></li> <li><a href="/wiki/Mosaic_evolution" title="Mosaic evolution">Mosaic evolution</a></li> <li><a href="/wiki/Multicellular_organism#Evolutionary_history" title="Multicellular organism">Multicellularity</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Sexual reproduction</a> <ul><li><a href="/wiki/Anisogamy#Evolution" title="Anisogamy">Gamete differentiation/sexes</a></li> <li><a href="/wiki/Biological_life_cycle" title="Biological life cycle">Life cycles/nuclear phases</a></li> <li><a href="/wiki/Mating_type" title="Mating type">Mating types</a></li> <li><a href="/wiki/Origin_and_function_of_meiosis" title="Origin and function of meiosis">Meiosis</a></li> <li><a href="/wiki/Sex-determination_system#Evolution" title="Sex-determination system">Sex-determination</a></li></ul></li> <li><a href="/wiki/Evolution_of_snake_venom" title="Evolution of snake venom">Snake venom</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tempo_and_Mode_in_Evolution" title="Tempo and Mode in Evolution">Tempo and modes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Phyletic_gradualism" title="Phyletic gradualism">Gradualism</a>/<a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a>/<a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Point_mutation" title="Point mutation">Micromutation</a>/<a href="/wiki/Mutationism#Later_mutationist_theories" title="Mutationism">Macromutation</a></li> <li><a href="/wiki/Uniformitarianism" title="Uniformitarianism">Uniformitarianism</a>/<a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Speciation" title="Speciation">Speciation</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allopatric_speciation" title="Allopatric speciation">Allopatric</a></li> <li><a href="/wiki/Anagenesis" title="Anagenesis">Anagenesis</a></li> <li><a href="/wiki/Catagenesis_(biology)" title="Catagenesis (biology)">Catagenesis</a></li> <li><a href="/wiki/Cladogenesis" title="Cladogenesis">Cladogenesis</a></li> <li><a href="/wiki/Cospeciation" title="Cospeciation">Cospeciation</a></li> <li><a href="/wiki/Ecological_speciation" title="Ecological speciation">Ecological</a></li> <li><a href="/wiki/Hybrid_speciation" title="Hybrid speciation">Hybrid</a></li> <li><a href="/wiki/Nonecological_speciation" title="Nonecological speciation">Non-ecological</a></li> <li><a href="/wiki/Parapatric_speciation" title="Parapatric speciation">Parapatric</a></li> <li><a href="/wiki/Peripatric_speciation" title="Peripatric speciation">Peripatric</a></li> <li><a href="/wiki/Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a></li> <li><a href="/wiki/Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance and Enlightenment</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Transmutation of species</a></li> <li><a href="/wiki/David_Hume" title="David Hume">David Hume</a> <ul><li><i><a href="/wiki/Dialogues_Concerning_Natural_Religion" title="Dialogues Concerning Natural Religion">Dialogues Concerning Natural Religion</a></i></li></ul></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> <ul><li><i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a></i></li></ul></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">History of paleontology</a></li> <li><a href="/wiki/Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li> <li><a href="/wiki/Blending_inheritance" title="Blending inheritance">Blending inheritance</a></li> <li><a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">The eclipse of Darwinism</a></li> <li><a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">History of molecular evolution</a></li> <li><a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">Extended evolutionary synthesis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Philosophy</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Darwinism" title="Darwinism">Darwinism</a></li> <li><a href="/wiki/Alternatives_to_Darwinian_evolution" title="Alternatives to Darwinian evolution">Alternatives</a> <ul><li><a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Orthogenesis" title="Orthogenesis">Orthogenesis</a></li> <li><a href="/wiki/Mutationism" title="Mutationism">Mutationism</a></li> <li><a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Structuralism_(biology)" title="Structuralism (biology)">Structuralism</a> <ul><li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">Spandrel</a></li></ul></li> <li><a href="/wiki/Theistic_evolution" title="Theistic evolution">Theistic</a></li> <li><a href="/wiki/Vitalism" title="Vitalism">Vitalism</a></li></ul></li> <li><a href="/wiki/Teleology_in_biology" title="Teleology in biology">Teleology in biology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a> <ul><li><a href="/wiki/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li></ul></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory of evolution</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Astrobiology" title="Astrobiology">Astrobiology</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a> <ul><li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Tree</a></li></ul></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><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, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Evolutionary_biology" title="Category:Evolutionary biology">Category</a></li> <li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Symbol_portal_class.svg" class="mw-file-description" title="Portal"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/16px-Symbol_portal_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/23px-Symbol_portal_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/31px-Symbol_portal_class.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span> <a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Portal</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Swarming" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Swarming" title="Template:Swarming"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Swarming" title="Template talk:Swarming"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Swarming" title="Special:EditPage/Template:Swarming"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Swarming" style="font-size:114%;margin:0 4em"><a href="/wiki/Swarm_behaviour" title="Swarm behaviour">Swarming</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Swarm_behaviour#Biological_swarming" title="Swarm behaviour">Biological swarming</a></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/Agent-based_model_in_biology" title="Agent-based model in biology">Agent-based model in biology</a></li> <li><a href="/wiki/Collective_animal_behavior" title="Collective animal behavior">Collective animal behavior</a></li> <li><a href="/wiki/Droving" title="Droving">Droving</a></li> <li><a href="/wiki/Flock_(birds)" title="Flock (birds)">Flock</a> <ul><li><a href="/wiki/Flocking_(behavior)" class="mw-redirect" title="Flocking (behavior)">flocking</a></li> <li><a href="/wiki/Sort_sol" title="Sort sol">sort sol</a></li></ul></li> <li><a href="/wiki/Herd" title="Herd">Herd</a> <ul><li><a href="/wiki/Herd_behavior" title="Herd behavior">herd behavior</a></li></ul></li> <li><a href="/wiki/Locust" title="Locust">Locust</a></li> <li><a href="/wiki/Mixed-species_foraging_flock" title="Mixed-species foraging flock">Mixed-species foraging flock</a></li> <li><a href="/wiki/Mobbing_(animal_behavior)" title="Mobbing (animal behavior)">Mobbing behavior</a> <ul><li><a href="/wiki/Feeding_frenzy" title="Feeding frenzy">feeding frenzy</a></li></ul></li> <li><a href="/wiki/Pack_(canine)" title="Pack (canine)">Pack</a> <ul><li><a href="/wiki/Pack_hunter" title="Pack hunter">pack hunter</a></li></ul></li> <li><a href="/wiki/Patterns_of_self-organization_in_ants" title="Patterns of self-organization in ants">Patterns of self-organization in ants</a> <ul><li><a href="/wiki/Ant_mill" title="Ant mill">ant mill</a></li> <li><a href="/wiki/Symmetry_breaking_of_escaping_ants" title="Symmetry breaking of escaping ants">symmetry breaking of escaping ants</a></li></ul></li> <li><a href="/wiki/Shoaling_and_schooling" title="Shoaling and schooling">Shoaling and schooling</a> <ul><li><a href="/wiki/Bait_ball" title="Bait ball">bait ball</a></li></ul></li> <li><a href="/wiki/Swarm_behaviour" title="Swarm behaviour">Swarming behaviour</a></li> <li><a href="/wiki/Swarming_(honey_bee)" title="Swarming (honey bee)">Swarming (honey bee)</a></li> <li><a href="/wiki/Swarming_motility" title="Swarming motility">Swarming motility</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="6" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Auklet_flock_Shumagins_1986.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5e/Auklet_flock_Shumagins_1986.jpg/140px-Auklet_flock_Shumagins_1986.jpg" decoding="async" width="140" height="95" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5e/Auklet_flock_Shumagins_1986.jpg/210px-Auklet_flock_Shumagins_1986.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5e/Auklet_flock_Shumagins_1986.jpg/280px-Auklet_flock_Shumagins_1986.jpg 2x" data-file-width="3756" data-file-height="2561" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Animal_migration" title="Animal migration">Animal migration</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/Animal_migration" title="Animal migration">Animal migration</a> <ul><li><a href="/wiki/Altitudinal_migration" title="Altitudinal migration">altitudinal</a></li> <li><a href="/wiki/Animal_migration_tracking" title="Animal migration tracking">tracking</a> <ul><li><a href="/wiki/History_of_wildlife_tracking_technology" title="History of wildlife tracking technology">history</a></li></ul></li> <li><a href="/wiki/Coded_wire_tag" title="Coded wire tag">coded wire tag</a></li></ul></li> <li><a href="/wiki/Bird_migration" title="Bird migration">Bird migration</a> <ul><li><a href="/wiki/Category:Bird_migration_flyways" title="Category:Bird migration flyways">flyways</a></li> <li><a href="/wiki/Reverse_migration_(birds)" title="Reverse migration (birds)">reverse migration</a></li></ul></li> <li><a href="/wiki/Cell_migration" title="Cell migration">Cell migration</a></li> <li><a href="/wiki/Fish_migration" title="Fish migration">Fish migration</a> <ul><li><a href="/wiki/Diel_vertical_migration" title="Diel vertical migration">diel vertical</a></li> <li><a href="/wiki/Lessepsian_migration" title="Lessepsian migration">Lessepsian</a></li> <li><a href="/wiki/Salmon_run" title="Salmon run">salmon run</a></li> <li><a href="/wiki/Sardine_run" title="Sardine run">sardine run</a></li></ul></li> <li><a href="/wiki/Homing_(biology)" title="Homing (biology)">Homing</a> <ul><li><a href="/wiki/Natal_homing" title="Natal homing">natal</a></li> <li><a href="/wiki/Philopatry" title="Philopatry">philopatry</a></li></ul></li> <li><a href="/wiki/Insect_migration" title="Insect migration">Insect migration</a> <ul><li><a href="/wiki/Lepidoptera_migration" title="Lepidoptera migration">butterflies</a> <ul><li><a href="/wiki/Monarch_butterfly_migration" title="Monarch butterfly migration">monarch</a></li></ul></li></ul></li> <li><a href="/wiki/Sea_turtle_migration" title="Sea turtle migration">Sea turtle migration</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Swarm_behaviour#Models" title="Swarm behaviour">Swarm algorithms</a></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/Agent-based_model" title="Agent-based model">Agent-based models</a></li> <li><a href="/wiki/Ant_colony_optimization_algorithm" class="mw-redirect" title="Ant colony optimization algorithm">Ant colony optimization</a></li> <li><a href="/wiki/Boids" title="Boids">Boids</a></li> <li><a href="/wiki/Crowd_simulation" title="Crowd simulation">Crowd simulation</a></li> <li><a href="/wiki/Particle_swarm_optimization" title="Particle swarm optimization">Particle swarm optimization</a></li> <li><a href="/wiki/Swarm_intelligence" title="Swarm intelligence">Swarm intelligence</a></li> <li><a href="/wiki/Swarm_(simulation)" title="Swarm (simulation)">Swarm (simulation)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Collective_motion" title="Collective motion">Collective motion</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/Active_matter" title="Active matter">Active matter</a></li> <li><a href="/wiki/Collective_motion" title="Collective motion">Collective motion</a></li> <li><a href="/wiki/Self-propelled_particles" title="Self-propelled particles">Self-propelled particles</a> <ul><li><a href="/wiki/Clustering_of_self-propelled_particles" title="Clustering of self-propelled particles">clustering</a></li></ul></li> <li><a href="/wiki/Vicsek_model" title="Vicsek model">Vicsek model</a></li> <li><a href="/wiki/BIO-LGCA" title="BIO-LGCA">BIO-LGCA</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Swarm_robotics" title="Swarm robotics">Swarm robotics</a></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/Ant_robotics" title="Ant robotics">Ant robotics</a></li> <li><a href="/wiki/Microbotics" title="Microbotics">Microbotics</a></li> <li><a href="/wiki/Nanorobotics" title="Nanorobotics">Nanorobotics</a></li> <li><a href="/wiki/Swarm_robotics" title="Swarm robotics">Swarm robotics</a></li> <li><a href="/wiki/Symbrion" title="Symbrion">Symbrion</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-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allee_effect" title="Allee effect">Allee effect</a></li> <li><a href="/wiki/Animal_navigation" title="Animal navigation">Animal navigation</a></li> <li><a href="/wiki/Collective_intelligence" title="Collective intelligence">Collective intelligence</a></li> <li><a href="/wiki/Decentralised_system" title="Decentralised system">Decentralised system</a></li> <li><a href="/wiki/Eusociality" title="Eusociality">Eusociality</a></li> <li><a href="/wiki/Group_size_measures" title="Group size measures">Group size measures</a></li> <li><a href="/wiki/Microbial_intelligence" title="Microbial intelligence">Microbial intelligence</a></li> <li><a class="mw-selflink selflink">Mutualism</a></li> <li><a href="/wiki/Predator_satiation" title="Predator satiation">Predator satiation</a></li> <li><a href="/wiki/Quorum_sensing" title="Quorum sensing">Quorum sensing</a></li> <li><a href="/wiki/Spatial_organization" title="Spatial organization">Spatial organization</a></li> <li><a href="/wiki/Stigmergy" title="Stigmergy">Stigmergy</a></li> <li><a href="/wiki/Swarming_(military)" title="Swarming (military)">Military swarming</a></li> <li><a href="/wiki/Task_allocation_and_partitioning_of_social_insects" class="mw-redirect" title="Task allocation and partitioning of social insects">Task allocation and partitioning of social insects</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Ecology:_Modelling_ecosystems:_Trophic_components" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Modelling_ecosystems" title="Template:Modelling ecosystems"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Modelling_ecosystems" title="Template talk:Modelling ecosystems"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Modelling_ecosystems" title="Special:EditPage/Template:Modelling ecosystems"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Ecology:_Modelling_ecosystems:_Trophic_components" style="font-size:114%;margin:0 4em"><a href="/wiki/Ecology" title="Ecology">Ecology</a>: <a href="/wiki/Ecosystem_model" title="Ecosystem model">Modelling ecosystems</a>: <a href="/wiki/Trophic_level" title="Trophic level">Trophic</a> components</div></th></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">General</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiotic_component" title="Abiotic component">Abiotic component</a></li> <li><a href="/wiki/Abiotic_stress" title="Abiotic stress">Abiotic stress</a></li> <li><a href="/wiki/Behavioral_ecology" title="Behavioral ecology">Behaviour</a></li> <li><a href="/wiki/Biogeochemical_cycle" title="Biogeochemical cycle">Biogeochemical cycle</a></li> <li><a href="/wiki/Biomass_(ecology)" title="Biomass (ecology)">Biomass</a></li> <li><a href="/wiki/Biotic_component" class="mw-redirect" title="Biotic component">Biotic component</a></li> <li><a href="/wiki/Biotic_stress" title="Biotic stress">Biotic stress</a></li> <li><a href="/wiki/Carrying_capacity" title="Carrying capacity">Carrying capacity</a></li> <li><a href="/wiki/Competition_(biology)" title="Competition (biology)">Competition</a></li> <li><a href="/wiki/Ecosystem" title="Ecosystem">Ecosystem</a></li> <li><a href="/wiki/Ecosystem_ecology" title="Ecosystem ecology">Ecosystem ecology</a></li> <li><a href="/wiki/Ecosystem_model" title="Ecosystem model">Ecosystem model</a></li> <li><a href="/wiki/Green_world_hypothesis" title="Green world hypothesis">Green world hypothesis</a></li> <li><a href="/wiki/Keystone_species" title="Keystone species">Keystone species</a></li> <li><a href="/wiki/List_of_feeding_behaviours" title="List of feeding behaviours">List of feeding behaviours</a></li> <li><a href="/wiki/Metabolic_theory_of_ecology" title="Metabolic theory of ecology">Metabolic theory of ecology</a></li> <li><a href="/wiki/Productivity_(ecology)" title="Productivity (ecology)">Productivity</a></li> <li><a href="/wiki/Resource_(biology)" title="Resource (biology)">Resource</a></li> <li><a href="/wiki/Restoration_ecology" class="mw-redirect" title="Restoration ecology">Restoration</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Autotroph" title="Autotroph">Producers</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Autotroph" title="Autotroph">Autotrophs</a></li> <li><a href="/wiki/Chemosynthesis" title="Chemosynthesis">Chemosynthesis</a></li> <li><a href="/wiki/Chemotroph" title="Chemotroph">Chemotrophs</a></li> <li><a href="/wiki/Foundation_species" title="Foundation species">Foundation species</a></li> <li><a href="/wiki/Kinetotroph" title="Kinetotroph">Kinetotrophs</a></li> <li><a href="/wiki/Mixotroph" title="Mixotroph">Mixotrophs</a></li> <li><a href="/wiki/Myco-heterotrophy" title="Myco-heterotrophy">Myco-heterotrophy</a></li> <li><a href="/wiki/Mycotroph" title="Mycotroph">Mycotroph</a></li> <li><a href="/wiki/Organotroph" title="Organotroph">Organotrophs</a></li> <li><a href="/wiki/Photoheterotroph" title="Photoheterotroph">Photoheterotrophs</a></li> <li><a href="/wiki/Photosynthesis" title="Photosynthesis">Photosynthesis</a></li> <li><a href="/wiki/Photosynthetic_efficiency" title="Photosynthetic efficiency">Photosynthetic efficiency</a></li> <li><a href="/wiki/Phototroph" title="Phototroph">Phototrophs</a></li> <li><a href="/wiki/Primary_nutritional_groups" title="Primary nutritional groups">Primary nutritional groups</a></li> <li><a href="/wiki/Primary_production" title="Primary production">Primary production</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Consumer_(food_chain)" title="Consumer (food chain)">Consumers</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Apex_predator" title="Apex predator">Apex predator</a></li> <li><a href="/wiki/Bacterivore" title="Bacterivore">Bacterivore</a></li> <li><a href="/wiki/Carnivore" title="Carnivore">Carnivores</a></li> <li><a href="/wiki/Chemoorganotroph" class="mw-redirect" title="Chemoorganotroph">Chemoorganotroph</a></li> <li><a href="/wiki/Foraging" title="Foraging">Foraging</a></li> <li><a href="/wiki/Generalist_and_specialist_species" title="Generalist and specialist species">Generalist and specialist species</a></li> <li><a href="/wiki/Intraguild_predation" title="Intraguild predation">Intraguild predation</a></li> <li><a href="/wiki/Herbivore" title="Herbivore">Herbivores</a></li> <li><a href="/wiki/Heterotroph" title="Heterotroph">Heterotroph</a></li> <li><a href="/wiki/Heterotrophic_nutrition" title="Heterotrophic nutrition">Heterotrophic nutrition</a></li> <li><a href="/wiki/Insectivore" title="Insectivore">Insectivore</a></li> <li><a href="/wiki/Mesopredator" title="Mesopredator">Mesopredators</a></li> <li><a href="/wiki/Mesopredator_release_hypothesis" title="Mesopredator release hypothesis">Mesopredator release hypothesis</a></li> <li><a href="/wiki/Omnivore" title="Omnivore">Omnivores</a></li> <li><a href="/wiki/Optimal_foraging_theory" title="Optimal foraging theory">Optimal foraging theory</a></li> <li><a href="/wiki/Planktivore" title="Planktivore">Planktivore</a></li> <li><a href="/wiki/Predation" title="Predation">Predation</a></li> <li><a href="/wiki/Prey_switching" title="Prey switching">Prey switching</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Decomposer" title="Decomposer">Decomposers</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Chemoorganoheterotrophy" class="mw-redirect" title="Chemoorganoheterotrophy">Chemoorganoheterotrophy</a></li> <li><a href="/wiki/Decomposition" title="Decomposition">Decomposition</a></li> <li><a href="/wiki/Detritivore" title="Detritivore">Detritivores</a></li> <li><a href="/wiki/Detritus" title="Detritus">Detritus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Microorganism#Habitats_and_ecology" title="Microorganism">Microorganisms</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Bacteriophage" title="Bacteriophage">Bacteriophage</a></li> <li><a href="/wiki/Lithoautotroph" title="Lithoautotroph">Lithoautotroph</a></li> <li><a href="/wiki/Lithotroph" title="Lithotroph">Lithotrophy</a></li> <li><a href="/wiki/Marine_microorganisms" title="Marine microorganisms">Marine</a></li> <li><a href="/wiki/Microbial_cooperation" title="Microbial cooperation">Microbial cooperation</a></li> <li><a href="/wiki/Microbial_ecology" title="Microbial ecology">Microbial ecology</a></li> <li><a href="/wiki/Microbial_food_web" title="Microbial food web">Microbial food web</a></li> <li><a href="/wiki/Microbial_intelligence" title="Microbial intelligence">Microbial intelligence</a></li> <li><a href="/wiki/Microbial_loop" title="Microbial loop">Microbial loop</a></li> <li><a href="/wiki/Microbial_mat" title="Microbial mat">Microbial mat</a></li> <li><a href="/wiki/Microbial_metabolism" title="Microbial metabolism">Microbial metabolism</a></li> <li><a href="/wiki/Phage_ecology" title="Phage ecology">Phage ecology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Food_web" title="Food web">Food webs</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biomagnification" title="Biomagnification">Biomagnification</a></li> <li><a href="/wiki/Ecological_efficiency" title="Ecological efficiency">Ecological efficiency</a></li> <li><a href="/wiki/Ecological_pyramid" title="Ecological pyramid">Ecological pyramid</a></li> <li><a href="/wiki/Energy_flow_(ecology)" title="Energy flow (ecology)">Energy flow</a></li> <li><a href="/wiki/Food_chain" title="Food chain">Food chain</a></li> <li><a href="/wiki/Trophic_level" title="Trophic level">Trophic level</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Example webs</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Lake_ecosystem#Trophic_relationships" title="Lake ecosystem">Lakes</a></li> <li><a href="/wiki/River_ecosystem#Trophic_relationships" title="River ecosystem">Rivers</a></li> <li><a href="/wiki/Soil_food_web" title="Soil food web">Soil</a></li> <li><a href="/wiki/Tritrophic_interactions_in_plant_defense" title="Tritrophic interactions in plant defense">Tritrophic interactions in plant defense</a></li> <li><a href="/wiki/Marine_food_web" title="Marine food web">Marine food webs</a> <ul><li><a href="/wiki/Cold_seep" title="Cold seep">cold seeps</a></li> <li><a href="/wiki/Hydrothermal_vent#Biological_communities" title="Hydrothermal vent">hydrothermal vents</a></li> <li><a href="/wiki/Intertidal_ecology" title="Intertidal ecology">intertidal</a></li> <li><a href="/wiki/Kelp_forest#Trophic_ecology" title="Kelp forest">kelp forests</a></li> <li><a href="/wiki/North_Pacific_Gyre" title="North Pacific Gyre">North Pacific Gyre</a></li> <li><a href="/wiki/Ecology_of_the_San_Francisco_Estuary#Food_web" title="Ecology of the San Francisco Estuary">San Francisco Estuary</a></li> <li><a href="/wiki/Tide_pool" title="Tide pool">tide pool</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Processes</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ascendency" title="Ascendency">Ascendency</a></li> <li><a href="/wiki/Bioaccumulation" title="Bioaccumulation">Bioaccumulation</a></li> <li><a href="/wiki/Cascade_effect_(ecology)" title="Cascade effect (ecology)">Cascade effect</a></li> <li><a href="/wiki/Climax_community" title="Climax community">Climax community</a></li> <li><a href="/wiki/Competitive_exclusion_principle" title="Competitive exclusion principle">Competitive exclusion principle</a></li> <li><a href="/wiki/Consumer%E2%80%93resource_interactions" title="Consumer–resource interactions">Consumer–resource interactions</a></li> <li><a href="/wiki/Copiotroph" title="Copiotroph">Copiotrophs</a></li> <li><a href="/wiki/Dominance_(ecology)" title="Dominance (ecology)">Dominance</a></li> <li><a href="/wiki/Ecological_network" title="Ecological network">Ecological network</a></li> <li><a href="/wiki/Ecological_succession" title="Ecological succession">Ecological succession</a></li> <li><a href="/wiki/Energy_quality" title="Energy quality">Energy quality</a></li> <li><a href="/wiki/Energy_systems_language" title="Energy systems language">Energy systems language</a></li> <li><a href="/wiki/F-ratio_(oceanography)" title="F-ratio (oceanography)">f-ratio</a></li> <li><a href="/wiki/Feed_conversion_ratio" title="Feed conversion ratio">Feed conversion ratio</a></li> <li><a href="/wiki/Feeding_frenzy" title="Feeding frenzy">Feeding frenzy</a></li> <li><a href="/wiki/Mesotrophic_soil" title="Mesotrophic soil">Mesotrophic soil</a></li> <li><a href="/wiki/Nutrient_cycle" title="Nutrient cycle">Nutrient cycle</a></li> <li><a href="/wiki/Oligotroph" title="Oligotroph">Oligotroph</a></li> <li><a href="/wiki/Paradox_of_the_plankton" title="Paradox of the plankton">Paradox of the plankton</a></li> <li><a href="/wiki/Trophic_cascade" title="Trophic cascade">Trophic cascade</a></li> <li><a href="/wiki/Trophic_mutualism" title="Trophic mutualism">Trophic mutualism</a></li> <li><a href="/wiki/Trophic_state_index" title="Trophic state index">Trophic state index</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Defense,<br />counter</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Animal_coloration" title="Animal coloration">Animal coloration</a></li> <li><a href="/wiki/Anti-predator_adaptation" title="Anti-predator adaptation">Anti-predator adaptations</a></li> <li><a href="/wiki/Camouflage" title="Camouflage">Camouflage</a></li> <li><a href="/wiki/Deimatic_behaviour" title="Deimatic behaviour">Deimatic behaviour</a></li> <li><a href="/wiki/Herbivore_adaptations_to_plant_defense" title="Herbivore adaptations to plant defense">Herbivore adaptations to plant defense</a></li> <li><a href="/wiki/Mimicry" title="Mimicry">Mimicry</a></li> <li><a href="/wiki/Plant_defense_against_herbivory" title="Plant defense against herbivory">Plant defense against herbivory</a></li> <li><a href="/wiki/Shoaling_and_schooling#Predator_avoidance" title="Shoaling and schooling">Predator avoidance in schooling fish</a></li></ul> </div></td></tr></tbody></table></div><div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Ecology:_Modelling_ecosystems:_Other_components" style="padding:3px"><table class="nowraplinks hlist mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Modelling_ecosystems" title="Template:Modelling ecosystems"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Modelling_ecosystems" title="Template talk:Modelling ecosystems"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Modelling_ecosystems" title="Special:EditPage/Template:Modelling ecosystems"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Ecology:_Modelling_ecosystems:_Other_components" style="font-size:114%;margin:0 4em"><a href="/wiki/Ecology" title="Ecology">Ecology</a>: <a href="/wiki/Ecosystem_model" title="Ecosystem model">Modelling ecosystems</a>: Other components</div></th></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Population_ecology" title="Population ecology">Population<br />ecology</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abundance_(ecology)" title="Abundance (ecology)">Abundance</a></li> <li><a href="/wiki/Allee_effect" title="Allee effect">Allee effect</a></li> <li><a href="/wiki/Consumer-resource_model" title="Consumer-resource model">Consumer-resource model</a></li> <li><a href="/wiki/Depensation" title="Depensation">Depensation</a></li> <li><a href="/wiki/Ecological_yield" title="Ecological yield">Ecological yield</a></li> <li><a href="/wiki/Effective_population_size" title="Effective population size">Effective population size</a></li> <li><a href="/wiki/Intraspecific_competition" title="Intraspecific competition">Intraspecific competition</a></li> <li><a href="/wiki/Logistic_function" title="Logistic function">Logistic function</a></li> <li><a href="/wiki/Malthusian_growth_model" title="Malthusian growth model">Malthusian growth model</a></li> <li><a href="/wiki/Maximum_sustainable_yield" title="Maximum sustainable yield">Maximum sustainable yield</a></li> <li><a href="/wiki/Overpopulation" title="Overpopulation">Overpopulation</a></li> <li><a href="/wiki/Overexploitation" title="Overexploitation">Overexploitation</a></li> <li><a href="/wiki/Population_cycle" title="Population cycle">Population cycle</a></li> <li><a href="/wiki/Population_dynamics" title="Population dynamics">Population dynamics</a></li> <li><a href="/wiki/Population_model" title="Population model">Population modeling</a></li> <li><a href="/wiki/Population_size" title="Population size">Population size</a></li> <li><a href="/wiki/Lotka%E2%80%93Volterra_equations" title="Lotka–Volterra equations">Predator–prey (Lotka–Volterra) equations</a></li> <li><a href="/wiki/Recruitment_(biology)" title="Recruitment (biology)">Recruitment</a></li> <li><a href="/wiki/Small_population_size" title="Small population size">Small population size</a></li> <li><a href="/wiki/Ecological_stability" title="Ecological stability">Stability</a> <ul><li><a href="/wiki/Ecological_resilience" title="Ecological resilience">Resilience</a></li> <li><a href="/wiki/Resistance_(ecology)" title="Resistance (ecology)">Resistance</a></li></ul></li> <li><a href="/wiki/Random_generalized_Lotka%E2%80%93Volterra_model" title="Random generalized Lotka–Volterra model">Random generalized Lotka–Volterra model</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Species</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Density_dependence" title="Density dependence">Density-dependent inhibition</a></li> <li><a href="/wiki/Ecological_effects_of_biodiversity" title="Ecological effects of biodiversity">Ecological effects of biodiversity</a></li> <li><a href="/wiki/Ecological_extinction" title="Ecological extinction">Ecological extinction</a></li> <li><a href="/wiki/Endemism" title="Endemism">Endemic species</a></li> <li><a href="/wiki/Flagship_species" title="Flagship species">Flagship species</a></li> <li><a href="/wiki/Ordination_(statistics)" title="Ordination (statistics)">Gradient analysis</a></li> <li><a href="/wiki/Bioindicator" title="Bioindicator">Indicator species</a></li> <li><a href="/wiki/Introduced_species" title="Introduced species">Introduced species</a></li> <li><a href="/wiki/Invasive_species" title="Invasive species">Invasive species</a> / <a href="/wiki/Native_species" title="Native species">Native species</a></li> <li><a href="/wiki/Latitudinal_gradients_in_species_diversity" title="Latitudinal gradients in species diversity">Latitudinal gradients in species diversity</a></li> <li><a href="/wiki/Minimum_viable_population" title="Minimum viable population">Minimum viable population</a></li> <li><a href="/wiki/Unified_neutral_theory_of_biodiversity" title="Unified neutral theory of biodiversity">Neutral theory</a></li> <li><a href="/wiki/Occupancy%E2%80%93abundance_relationship" title="Occupancy–abundance relationship">Occupancy–abundance relationship</a></li> <li><a href="/wiki/Population_viability_analysis" title="Population viability analysis">Population viability analysis</a></li> <li><a href="/wiki/Priority_effect" title="Priority effect">Priority effect</a></li> <li><a href="/wiki/Rapoport%27s_rule" title="Rapoport's rule">Rapoport's rule</a></li> <li><a href="/wiki/Relative_abundance_distribution" title="Relative abundance distribution">Relative abundance distribution</a></li> <li><a href="/wiki/Relative_species_abundance" title="Relative species abundance">Relative species abundance</a></li> <li><a href="/wiki/Species_diversity" title="Species diversity">Species diversity</a></li> <li><a href="/wiki/Species_homogeneity" title="Species homogeneity">Species homogeneity</a></li> <li><a href="/wiki/Species_richness" title="Species richness">Species richness</a></li> <li><a href="/wiki/Species_distribution" title="Species distribution">Species distribution</a></li> <li><a href="/wiki/Species%E2%80%93area_relationship" title="Species–area relationship">Species–area curve</a></li> <li><a href="/wiki/Umbrella_species" title="Umbrella species">Umbrella species</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Species<br />interaction</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Antibiosis" title="Antibiosis">Antibiosis</a></li> <li><a href="/wiki/Biological_interaction" title="Biological interaction">Biological interaction</a></li> <li><a href="/wiki/Commensalism" title="Commensalism">Commensalism</a></li> <li><a href="/wiki/Community_(ecology)" title="Community (ecology)">Community ecology</a></li> <li><a href="/wiki/Ecological_facilitation" title="Ecological facilitation">Ecological facilitation</a></li> <li><a href="/wiki/Interspecific_competition" title="Interspecific competition">Interspecific competition</a></li> <li><a class="mw-selflink selflink">Mutualism</a></li> <li><a href="/wiki/Parasitism" title="Parasitism">Parasitism</a></li> <li><a href="/wiki/Storage_effect" title="Storage effect">Storage effect</a></li> <li><a href="/wiki/Symbiosis" title="Symbiosis">Symbiosis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Spatial_ecology" title="Spatial ecology">Spatial<br />ecology</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Cross-boundary_subsidy" title="Cross-boundary subsidy">Cross-boundary subsidy</a></li> <li><a href="/wiki/Cline_(biology)" title="Cline (biology)">Ecocline</a></li> <li><a href="/wiki/Ecotone" title="Ecotone">Ecotone</a></li> <li><a href="/wiki/Ecotype" title="Ecotype">Ecotype</a></li> <li><a href="/wiki/Disturbance_(ecology)" title="Disturbance (ecology)">Disturbance</a></li> <li><a href="/wiki/Edge_effects" title="Edge effects">Edge effects</a></li> <li><a href="/wiki/Foster%27s_rule" title="Foster's rule">Foster's rule</a></li> <li><a href="/wiki/Habitat_fragmentation" title="Habitat fragmentation">Habitat fragmentation</a></li> <li><a href="/wiki/Ideal_free_distribution" title="Ideal free distribution">Ideal free distribution</a></li> <li><a href="/wiki/Intermediate_disturbance_hypothesis" title="Intermediate disturbance hypothesis">Intermediate disturbance hypothesis</a></li> <li><a href="/wiki/Insular_biogeography" title="Insular biogeography">Insular biogeography</a></li> <li><a href="/wiki/Land_change_modeling" title="Land change modeling">Land change modeling</a></li> <li><a href="/wiki/Landscape_ecology" title="Landscape ecology">Landscape ecology</a></li> <li><a href="/wiki/Landscape_epidemiology" title="Landscape epidemiology">Landscape epidemiology</a></li> <li><a href="/wiki/Landscape_limnology" title="Landscape limnology">Landscape limnology</a></li> <li><a href="/wiki/Metapopulation" title="Metapopulation">Metapopulation</a></li> <li><a href="/wiki/Patch_dynamics" title="Patch dynamics">Patch dynamics</a></li> <li><a href="/wiki/R/K_selection_theory" title="R/K selection theory"><i>r</i>/<i>K</i> selection theory</a></li> <li><a href="/wiki/Resource_selection_function" title="Resource selection function">Resource selection function</a></li> <li><a href="/wiki/Source%E2%80%93sink_dynamics" title="Source–sink dynamics">Source–sink dynamics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Ecological_niche" title="Ecological niche">Niche</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ecological_niche" title="Ecological niche">Ecological niche</a></li> <li><a href="/wiki/Ecological_trap" title="Ecological trap">Ecological trap</a></li> <li><a href="/wiki/Ecosystem_engineer" title="Ecosystem engineer">Ecosystem engineer</a></li> <li><a href="/wiki/Species_distribution_modelling" title="Species distribution modelling">Environmental niche modelling</a></li> <li><a href="/wiki/Guild_(ecology)" title="Guild (ecology)">Guild</a></li> <li><a href="/wiki/Habitat" title="Habitat">Habitat</a> <ul><li><a href="/wiki/Marine_habitat" title="Marine habitat">marine habitats</a></li></ul></li> <li><a href="/wiki/Limiting_similarity" title="Limiting similarity">Limiting similarity</a></li> <li><a href="/wiki/Niche_apportionment_models" title="Niche apportionment models">Niche apportionment models</a></li> <li><a href="/wiki/Niche_construction" title="Niche construction">Niche construction</a></li> <li><a href="/wiki/Niche_differentiation" class="mw-redirect" title="Niche differentiation">Niche differentiation</a></li> <li><a href="/wiki/Ontogenetic_niche_shift" title="Ontogenetic niche shift">Ontogenetic niche shift</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Non-trophic_networks" title="Non-trophic networks">Other<br />networks</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Assembly_rules" title="Assembly rules">Assembly rules</a></li> <li><a href="/wiki/Bateman%27s_principle" title="Bateman's principle">Bateman's principle</a></li> <li><a href="/wiki/Bioluminescence" title="Bioluminescence">Bioluminescence</a></li> <li><a href="/wiki/Ecological_collapse" class="mw-redirect" title="Ecological collapse">Ecological collapse</a></li> <li><a href="/wiki/Ecological_debt" title="Ecological debt">Ecological debt</a></li> <li><a href="/wiki/Ecological_debt" title="Ecological debt">Ecological deficit</a></li> <li><a href="/wiki/Energy_flow_(ecology)" title="Energy flow (ecology)">Ecological energetics</a></li> <li><a href="/wiki/Ecological_indicator" title="Ecological indicator">Ecological indicator</a></li> <li><a href="/wiki/Ecological_threshold" title="Ecological threshold">Ecological threshold</a></li> <li><a href="/wiki/Ecosystem_diversity" title="Ecosystem diversity">Ecosystem diversity</a></li> <li><a href="/wiki/Emergence" title="Emergence">Emergence</a></li> <li><a href="/wiki/Extinction_debt" title="Extinction debt">Extinction debt</a></li> <li><a href="/wiki/Kleiber%27s_law" title="Kleiber's law">Kleiber's law</a></li> <li><a href="/wiki/Liebig%27s_law_of_the_minimum" title="Liebig's law of the minimum">Liebig's law of the minimum</a></li> <li><a href="/wiki/Marginal_value_theorem" title="Marginal value theorem">Marginal value theorem</a></li> <li><a href="/wiki/Thorson%27s_rule" title="Thorson's rule">Thorson's rule</a></li> <li><a href="/wiki/Xerosere" title="Xerosere">Xerosere</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allometry" title="Allometry">Allometry</a></li> <li><a href="/wiki/Alternative_stable_state" title="Alternative stable state">Alternative stable state</a></li> <li><a href="/wiki/Balance_of_nature" title="Balance of nature">Balance of nature</a></li> <li><a href="/wiki/Biological_data_visualization" title="Biological data visualization">Biological data visualization</a></li> <li><a href="/wiki/Ecological_economics" title="Ecological economics">Ecological economics</a></li> <li><a href="/wiki/Ecological_footprint" title="Ecological footprint">Ecological footprint</a></li> <li><a href="/wiki/Ecological_forecasting" title="Ecological forecasting">Ecological forecasting</a></li> <li><a href="/wiki/Environmental_humanities" title="Environmental humanities">Ecological humanities</a></li> <li><a href="/wiki/Ecological_stoichiometry" title="Ecological stoichiometry">Ecological stoichiometry</a></li> <li><a href="/wiki/Ecopath" title="Ecopath">Ecopath</a></li> <li><a href="/wiki/Ecosystem_based_fisheries" class="mw-redirect" title="Ecosystem based fisheries">Ecosystem based fisheries</a></li> <li><a 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ecology</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div><a href="/wiki/Outline_of_ecology" title="Outline of ecology">Outline of ecology</a></div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style></div><div role="navigation" class="navbox authority-control" aria-label="Navbox" style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Help:Authority_control" title="Help:Authority control">Authority control databases</a>: National <span class="mw-valign-text-top noprint" typeof="mw:File/Frameless"><a 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