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<span>Overview</span> </div> </a> <ul id="toc-Overview-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Biomass_transfer_efficiency" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Biomass_transfer_efficiency"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Biomass transfer efficiency</span> </div> </a> <ul id="toc-Biomass_transfer_efficiency-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolution" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Evolution</span> </div> </a> <ul id="toc-Evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Fractional_trophic_levels" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Fractional_trophic_levels"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Fractional trophic levels</span> </div> </a> <ul id="toc-Fractional_trophic_levels-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mean_trophic_level" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Mean_trophic_level"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Mean trophic level</span> </div> </a> <ul id="toc-Mean_trophic_level-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-FiB_index" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#FiB_index"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>FiB index</span> </div> </a> <ul id="toc-FiB_index-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Tritrophic_and_other_interactions" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Tritrophic_and_other_interactions"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Tritrophic and other interactions</span> </div> </a> <ul id="toc-Tritrophic_and_other_interactions-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">9</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">10</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button 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Available in 33 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-33" 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">33 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/Trofiese_vlak" title="Trofiese vlak – Afrikaans" lang="af" hreflang="af" data-title="Trofiese vlak" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target"><span>Afrikaans</span></a></li><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%85%D8%B3%D8%AA%D9%88%D9%89_%D8%BA%D8%B0%D8%A7%D8%A6%D9%8A" title="مستوى غذائي – Arabic" lang="ar" hreflang="ar" data-title="مستوى غذائي" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/Nivel_tr%C3%B3ficu" title="Nivel tróficu – Asturian" lang="ast" hreflang="ast" data-title="Nivel tróficu" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target"><span>Asturianu</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Nivell_tr%C3%B2fic" title="Nivell tròfic – Catalan" lang="ca" hreflang="ca" data-title="Nivell tròfic" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Trofisk_niveau" title="Trofisk niveau – Danish" lang="da" hreflang="da" data-title="Trofisk niveau" 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/Trophieniveau" title="Trophieniveau – German" lang="de" hreflang="de" data-title="Trophieniveau" 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/Troofiline_tase" title="Troofiline tase – Estonian" lang="et" hreflang="et" data-title="Troofiline tase" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Nivel_tr%C3%B3fico" title="Nivel trófico – Spanish" lang="es" hreflang="es" data-title="Nivel trófico" 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/Trofa_nivelo" title="Trofa nivelo – Esperanto" lang="eo" hreflang="eo" data-title="Trofa nivelo" 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/Maila_trofiko" title="Maila trofiko – Basque" lang="eu" hreflang="eu" data-title="Maila trofiko" 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/%D8%B3%D8%B7%D8%AD_%D9%BE%D8%B1%D9%88%D8%B1%D8%AF%DA%AF%DB%8C" title="سطح پروردگی – Persian" lang="fa" hreflang="fa" data-title="سطح پروردگی" 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/Niveau_trophique" title="Niveau trophique – French" lang="fr" hreflang="fr" data-title="Niveau trophique" 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/Leibh%C3%A9al_tr%C3%B3fach" title="Leibhéal trófach – Irish" lang="ga" hreflang="ga" data-title="Leibhéal trófach" 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/Nivel_tr%C3%B3fico" title="Nivel trófico – Galician" lang="gl" hreflang="gl" data-title="Nivel trófico" 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%98%81%EC%96%91_%EB%8B%A8%EA%B3%84" title="영양 단계 – Korean" lang="ko" hreflang="ko" data-title="영양 단계" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Livello_trofico" title="Livello trofico – Italian" lang="it" hreflang="it" data-title="Livello trofico" 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%A8%D7%9E%D7%AA_%D7%94%D7%96%D7%A0%D7%94" title="רמת הזנה – Hebrew" lang="he" hreflang="he" data-title="רמת הזנה" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Trofisk_niv%C3%A5" title="Trofisk nivå – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Trofisk nivå" 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/Trofisk_niv%C3%A5" title="Trofisk nivå – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Trofisk nivå" 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/Poziom_troficzny" title="Poziom troficzny – Polish" lang="pl" hreflang="pl" data-title="Poziom troficzny" 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/N%C3%ADvel_tr%C3%B3fico" title="Nível trófico – Portuguese" lang="pt" hreflang="pt" data-title="Nível trófico" 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%A2%D1%80%D0%BE%D1%84%D0%B8%D1%87%D0%B5%D1%81%D0%BA%D0%B8%D0%B9_%D1%83%D1%80%D0%BE%D0%B2%D0%B5%D0%BD%D1%8C" title="Трофический уровень – Russian" lang="ru" hreflang="ru" data-title="Трофический уровень" 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/Trophic_level" title="Trophic level – Simple English" lang="en-simple" hreflang="en-simple" data-title="Trophic level" 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/Trofi%C4%8Dna_struktura_ekosistema" title="Trofična struktura ekosistema – Slovenian" lang="sl" hreflang="sl" data-title="Trofična struktura ekosistema" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%A2%D1%80%D0%BE%D1%84%D0%B8%D1%87%D0%BA%D0%B8_%D0%BD%D0%B8%D0%B2%D0%BE" title="Трофички ниво – Serbian" lang="sr" hreflang="sr" data-title="Трофички ниво" 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/Trofi%C4%8Dki_nivo" title="Trofički nivo – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Trofički nivo" 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/Trofiataso" title="Trofiataso – Finnish" lang="fi" hreflang="fi" data-title="Trofiataso" 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/Trofisk_struktur" title="Trofisk struktur – Swedish" lang="sv" hreflang="sv" data-title="Trofisk struktur" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%A2%D1%80%D0%BE%D1%84%D1%96%D1%87%D0%BD%D0%B8%D0%B9_%D1%80%D1%96%D0%B2%D0%B5%D0%BD%D1%8C" title="Трофічний рівень – Ukrainian" lang="uk" hreflang="uk" data-title="Трофічний рівень" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/B%E1%BA%ADc_dinh_d%C6%B0%E1%BB%A1ng" title="Bậc dinh dưỡng – Vietnamese" lang="vi" hreflang="vi" data-title="Bậc dinh dưỡng" 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-wuu mw-list-item"><a href="https://wuu.wikipedia.org/wiki/%E8%90%A5%E5%85%BB%E7%BA%A7" title="营养级 – Wu" lang="wuu" hreflang="wuu" data-title="营养级" data-language-autonym="吴语" data-language-local-name="Wu" class="interlanguage-link-target"><span>吴语</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E7%87%9F%E9%A4%8A%E7%B4%9A" title="營養級 – Cantonese" lang="yue" hreflang="yue" data-title="營養級" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target"><span>粵語</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E7%87%9F%E9%A4%8A%E7%B4%9A" title="營養級 – Chinese" lang="zh" hreflang="zh" data-title="營養級" 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class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Position of an organism in a food chain</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Not to be confused with <a href="/wiki/Trophic_level_index" title="Trophic level index">Trophic level index</a>.</div> <p class="mw-empty-elt"> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Far_Pastures_-_geograph.org.uk_-_52967.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/38/Far_Pastures_-_geograph.org.uk_-_52967.jpg/260px-Far_Pastures_-_geograph.org.uk_-_52967.jpg" decoding="async" width="260" height="195" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/38/Far_Pastures_-_geograph.org.uk_-_52967.jpg/390px-Far_Pastures_-_geograph.org.uk_-_52967.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/38/Far_Pastures_-_geograph.org.uk_-_52967.jpg/520px-Far_Pastures_-_geograph.org.uk_-_52967.jpg 2x" data-file-width="640" data-file-height="480" /></a><figcaption><b>First trophic level</b>. The plants in this image, and the <a href="/wiki/Algae" title="Algae">algae</a> and <a href="/wiki/Phytoplankton" title="Phytoplankton">phytoplankton</a> in the lake, are <a href="/wiki/Primary_producer" class="mw-redirect" title="Primary producer">primary producers</a>. They take nutrients from the soil or the water, and manufacture their own food by <a href="/wiki/Photosynthesis" title="Photosynthesis">photosynthesis</a>, using energy from the sun.</figcaption></figure> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/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/trophic" class="extiw" title="wiktionary:trophic"> trophic</a></b></i> in Wiktionary, the free dictionary.</div></div> </div> <p>The <b>trophic level</b> of an <a href="/wiki/Organism" title="Organism">organism</a> is the position it occupies in a <a href="/wiki/Food_web" title="Food web">food web</a>. Within a food web, a <a href="/wiki/Food_chain" title="Food chain">food chain</a> is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it is from the start of the chain. A food web starts at trophic level 1 with <a href="/wiki/Primary_producer" class="mw-redirect" title="Primary producer">primary producers</a> such as plants, can move to <a href="/wiki/Herbivore" title="Herbivore">herbivores</a> at level 2, <a href="/wiki/Carnivore" title="Carnivore">carnivores</a> at level 3 or higher, and typically finish with <a href="/wiki/Apex_predator" title="Apex predator">apex predators</a> at level 4 or 5. The path along the chain can form either a one-way flow or a part of a wider food "web". Ecological communities with higher <a href="/wiki/Biodiversity" title="Biodiversity">biodiversity</a> form more complex trophic paths. </p><p>The word <i>trophic</i> derives from the <a href="/wiki/Ancient_Greek" title="Ancient Greek">Greek</a> <a href="https://en.wiktionary.org/wiki/%CF%84%CF%81%CE%BF%CF%86%CE%AE" class="extiw" title="wikt:τροφή">τροφή</a> (trophē) referring to food or nourishment.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Trophic_level&action=edit&section=1" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The concept of trophic level was developed by <a href="/wiki/Raymond_Lindeman" title="Raymond Lindeman">Raymond Lindeman</a> (1942), based on the terminology of <a href="/wiki/August_Thienemann" title="August Thienemann">August Thienemann</a> (1926): "producers", "consumers", and "reducers" (modified to "decomposers" by Lindeman).<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><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> <div class="mw-heading mw-heading2"><h2 id="Overview">Overview</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Trophic_level&action=edit&section=2" title="Edit section: Overview"><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:ConsumerWikiPDiag.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cb/ConsumerWikiPDiag.svg/290px-ConsumerWikiPDiag.svg.png" decoding="async" width="290" height="214" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cb/ConsumerWikiPDiag.svg/435px-ConsumerWikiPDiag.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cb/ConsumerWikiPDiag.svg/580px-ConsumerWikiPDiag.svg.png 2x" data-file-width="998" data-file-height="736" /></a><figcaption><a href="/wiki/Consumer-resource_systems" class="mw-redirect" title="Consumer-resource systems">Consumer categories</a> based on material eaten (plant: green shades are live, brown shades are dead; animal: red shades are live, purple shades are dead; or particulate: grey shades) and feeding strategy (gatherer: lighter shade of each color; miner: darker shade of each color)</figcaption></figure> <p>The three basic ways in which organisms get food are as producers, consumers, and decomposers. </p> <ul><li>Producers (<a href="/wiki/Autotroph" title="Autotroph">autotrophs</a>) are typically <a href="/wiki/Plant" title="Plant">plants</a> or <a href="/wiki/Algae" title="Algae">algae</a>. Plants and algae do not usually eat other organisms, but pull nutrients from the soil or the ocean and manufacture their own food using <a href="/wiki/Photosynthesis" title="Photosynthesis">photosynthesis</a>. For this reason, they are called <a href="/wiki/Primary_producer" class="mw-redirect" title="Primary producer">primary producers</a>. In this way, it is energy from the sun that usually powers the base of the food chain.<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> An exception occurs in deep-sea <a href="/wiki/Hydrothermal_vents" class="mw-redirect" title="Hydrothermal vents">hydrothermal ecosystems</a>, where there is no sunlight. Here primary producers manufacture food through a process called <a href="/wiki/Chemosynthesis" title="Chemosynthesis">chemosynthesis</a>.<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></li> <li><a href="/wiki/Consumers_(food_chain)" class="mw-redirect" title="Consumers (food chain)">Consumers</a> (<a href="/wiki/Heterotroph" title="Heterotroph">heterotrophs</a>) are <a href="/wiki/Species" title="Species">species</a> that cannot manufacture their own food and need to consume other organisms. Animals that eat primary producers (like plants) are called <a href="/wiki/Herbivore" title="Herbivore">herbivores</a>. Animals that eat other animals are called <a href="/wiki/Carnivore" title="Carnivore">carnivores</a>, and animals that eat both plants and other animals are called <a href="/wiki/Omnivore" title="Omnivore">omnivores</a>.</li> <li><a href="/wiki/Decomposer" title="Decomposer">Decomposers</a> (<a href="/wiki/Detritivores" class="mw-redirect" title="Detritivores">detritivores</a>) break down dead plant and animal material and wastes and release it again as energy and nutrients into the ecosystem for recycling. Decomposers, such as <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> and <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a> (mushrooms), feed on waste and dead matter, converting it into inorganic chemicals that can be recycled as mineral nutrients for plants to use again.</li></ul> <p>Trophic levels can be represented by numbers, starting at level 1 with plants. Further trophic levels are numbered subsequently according to how far the organism is along the food chain. </p> <dl><dt>Level 1</dt> <dd>Plants and algae make their own food and are called producers.</dd> <dt>Level 2</dt> <dd>Herbivores eat plants and are called primary consumers.</dd> <dt>Level 3</dt> <dd>Carnivores that eat herbivores are called secondary consumers.</dd> <dt>Level 4</dt> <dd>Carnivores that eat other carnivores are called tertiary consumers.</dd> <dt><a href="/wiki/Apex_predator" title="Apex predator">Apex predator</a></dt> <dd>By definition, healthy adult <i>apex predators</i> have no predators (with members of their own species a possible exception) and are at the highest numbered level of their food web.</dd></dl> <ul class="gallery mw-gallery-traditional center"> <li class="gallerybox" style="width: 215px"> <div class="thumb" style="width: 210px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Sylvilagus_floridanus.jpg" class="mw-file-description" title="Second trophic level Rabbits eat plants at the first trophic level, so they are primary consumers."><img alt="Second trophic level Rabbits eat plants at the first trophic level, so they are primary consumers." src="//upload.wikimedia.org/wikipedia/commons/thumb/7/70/Sylvilagus_floridanus.jpg/120px-Sylvilagus_floridanus.jpg" decoding="async" width="120" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/70/Sylvilagus_floridanus.jpg/180px-Sylvilagus_floridanus.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/70/Sylvilagus_floridanus.jpg/240px-Sylvilagus_floridanus.jpg 2x" data-file-width="1300" data-file-height="1300" /></a></span></div> <div class="gallerytext"><b>Second trophic level</b><br /><a href="/wiki/Rabbit" title="Rabbit">Rabbits</a> eat plants at the first trophic level, so they are primary consumers.</div> </li> <li class="gallerybox" style="width: 215px"> <div class="thumb" style="width: 210px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Vulpes_vulpes_with_prey.jpg" class="mw-file-description" title="Third trophic level Foxes eat rabbits at the second trophic level, so they are secondary consumers."><img alt="Third trophic level Foxes eat rabbits at the second trophic level, so they are secondary consumers." src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/Vulpes_vulpes_with_prey.jpg/180px-Vulpes_vulpes_with_prey.jpg" decoding="async" width="180" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/Vulpes_vulpes_with_prey.jpg/270px-Vulpes_vulpes_with_prey.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8a/Vulpes_vulpes_with_prey.jpg/360px-Vulpes_vulpes_with_prey.jpg 2x" data-file-width="1897" data-file-height="1262" /></a></span></div> <div class="gallerytext"><b>Third trophic level</b><br /><a href="/wiki/Fox" title="Fox">Foxes</a> eat rabbits at the second trophic level, so they are secondary consumers.</div> </li> <li class="gallerybox" style="width: 215px"> <div class="thumb" style="width: 210px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Aquila_chrysaetos_1_(Bohu%C5%A1_%C4%8C%C3%AD%C4%8Del).jpg" class="mw-file-description" title="Fourth trophic level Golden eagles eat foxes at the third trophic level, so they are tertiary consumers."><img alt="Fourth trophic level Golden eagles eat foxes at the third trophic level, so they are tertiary consumers." src="//upload.wikimedia.org/wikipedia/commons/thumb/8/89/Aquila_chrysaetos_1_%28Bohu%C5%A1_%C4%8C%C3%AD%C4%8Del%29.jpg/180px-Aquila_chrysaetos_1_%28Bohu%C5%A1_%C4%8C%C3%AD%C4%8Del%29.jpg" decoding="async" width="180" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/89/Aquila_chrysaetos_1_%28Bohu%C5%A1_%C4%8C%C3%AD%C4%8Del%29.jpg/270px-Aquila_chrysaetos_1_%28Bohu%C5%A1_%C4%8C%C3%AD%C4%8Del%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/89/Aquila_chrysaetos_1_%28Bohu%C5%A1_%C4%8C%C3%AD%C4%8Del%29.jpg/360px-Aquila_chrysaetos_1_%28Bohu%C5%A1_%C4%8C%C3%AD%C4%8Del%29.jpg 2x" data-file-width="1024" data-file-height="682" /></a></span></div> <div class="gallerytext"><b>Fourth trophic level</b><br /><a href="/wiki/Golden_eagle" title="Golden eagle">Golden eagles</a> eat foxes at the third trophic level, so they are tertiary consumers.</div> </li> <li class="gallerybox" style="width: 215px"> <div class="thumb" style="width: 210px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Fungi_in_Borneo.jpg" class="mw-file-description" title="Decomposers The fungi on this tree feed on dead matter, converting it back to nutrients that primary producers can use."><img alt="Decomposers The fungi on this tree feed on dead matter, converting it back to nutrients that primary producers can use." src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d4/Fungi_in_Borneo.jpg/90px-Fungi_in_Borneo.jpg" decoding="async" width="90" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d4/Fungi_in_Borneo.jpg/135px-Fungi_in_Borneo.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d4/Fungi_in_Borneo.jpg/179px-Fungi_in_Borneo.jpg 2x" data-file-width="449" data-file-height="600" /></a></span></div> <div class="gallerytext"><b><a href="/wiki/Decomposer" title="Decomposer">Decomposers</a></b><br />The fungi on this tree feed on dead matter, converting it back to nutrients that <a href="/wiki/Primary_producer" class="mw-redirect" title="Primary producer">primary producers</a> can use.</div> </li> </ul> <p>In real-world <a href="/wiki/Ecosystem" title="Ecosystem">ecosystems</a>, there is more than one food chain for most organisms, since most organisms eat more than one kind of food or are eaten by more than one type of predator. A diagram that sets out the intricate network of intersecting and overlapping food chains for an ecosystem is called its <a href="/wiki/Food_web" title="Food web">food web</a>.<sup id="cite_ref-Lisowski-etal_6-0" class="reference"><a href="#cite_note-Lisowski-etal-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Decomposers are often left off food webs, but if included, they mark the end of a food chain.<sup id="cite_ref-Lisowski-etal_6-1" class="reference"><a href="#cite_note-Lisowski-etal-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Thus food chains start with primary producers and end with decay and decomposers. Since decomposers recycle nutrients, leaving them so they can be reused by primary producers, they are sometimes regarded as occupying their own trophic level.<sup id="cite_ref-AHSD_7-0" class="reference"><a href="#cite_note-AHSD-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Pauly&Palomares_8-0" class="reference"><a href="#cite_note-Pauly&Palomares-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p>The trophic level of a species may vary if it has a choice of diet. Virtually all plants and phytoplankton are purely phototrophic and are at exactly level 1.0. Many worms are at around 2.1; insects 2.2; jellyfish 3.0; birds 3.6.<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> A 2013 study estimates the average trophic level of human beings at 2.21, similar to pigs or anchovies.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> This is only an average, and plainly both modern and ancient human eating habits are complex and vary greatly. For example, a traditional Inuit living on a diet consisting primarily of seals would have a trophic level of nearly 5.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Biomass_transfer_efficiency">Biomass transfer efficiency</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Trophic_level&action=edit&section=3" title="Edit section: Biomass transfer efficiency"><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:Ecological_Pyramid.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3a/Ecological_Pyramid.svg/340px-Ecological_Pyramid.svg.png" decoding="async" width="340" height="244" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3a/Ecological_Pyramid.svg/510px-Ecological_Pyramid.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3a/Ecological_Pyramid.svg/680px-Ecological_Pyramid.svg.png 2x" data-file-width="512" data-file-height="368" /></a><figcaption>An <a href="/wiki/Energy_pyramid" class="mw-redirect" title="Energy pyramid">energy pyramid</a> illustrates how much energy is needed as it flows upward to support the next trophic level. Only about 10% of the energy transferred between each trophic level is converted to <a href="/wiki/Biomass_(ecology)" title="Biomass (ecology)">biomass</a>.</figcaption></figure> <p>In general, each trophic level relates to the one below it by absorbing some of the energy it consumes, and in this way can be regarded as resting on, or supported by, the next lower trophic level. Food chains can be diagrammed to illustrate the amount of energy that moves from one feeding level to the next in a food chain. This is called an <a href="/wiki/Energy_pyramid" class="mw-redirect" title="Energy pyramid">energy pyramid</a>. The energy transferred between levels can also be thought of as approximating to a transfer in <a href="/wiki/Biomass_(ecology)" title="Biomass (ecology)">biomass</a>, so energy pyramids can also be viewed as biomass pyramids, picturing the amount of biomass that results at higher levels from biomass consumed at lower levels. However, when primary producers grow rapidly and are consumed rapidly, the biomass at any one moment may be low; for example, phytoplankton (producer) biomass can be low compared to the zooplankton (consumer) biomass in the same area of ocean.<sup id="cite_ref-Behrenfeld_2014_pp._880–887_12-0" class="reference"><a href="#cite_note-Behrenfeld_2014_pp._880–887-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p><p>The efficiency with which energy or biomass is transferred from one trophic level to the next is called the <a href="/wiki/Ecological_efficiency" title="Ecological efficiency">ecological efficiency</a>. Consumers at each level convert on average only about 10% of the chemical energy in their food to their own organic tissue (the <a href="/wiki/Ten_percent_law" class="mw-redirect" title="Ten percent law">ten-per cent law</a>). For this reason, food chains rarely extend for more than 5 or 6 levels. At the lowest trophic level (the bottom of the food chain), plants convert about 1% of the sunlight they receive into chemical energy. It follows from this that the total energy originally present in the incident sunlight that is finally embodied in a tertiary consumer is about 0.001%<sup id="cite_ref-AHSD_7-1" class="reference"><a href="#cite_note-AHSD-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Evolution">Evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Trophic_level&action=edit&section=4" title="Edit section: Evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Both the number of trophic levels and the complexity of relationships between them evolve as life diversifies through time, the exception being intermittent mass extinction events.<sup id="cite_ref-SahneyBenton2008_13-0" class="reference"><a href="#cite_note-SahneyBenton2008-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Fractional_trophic_levels">Fractional trophic levels</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Trophic_level&action=edit&section=5" title="Edit section: Fractional trophic levels"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Orca_porpoising.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/38/Orca_porpoising.jpg/220px-Orca_porpoising.jpg" decoding="async" width="220" height="148" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/38/Orca_porpoising.jpg/330px-Orca_porpoising.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/38/Orca_porpoising.jpg/440px-Orca_porpoising.jpg 2x" data-file-width="700" data-file-height="471" /></a><figcaption>Killer whales (<a href="/wiki/Orca" title="Orca">orca</a>) are <a href="/wiki/Apex_predator" title="Apex predator">apex predators</a> but they are divided into separate populations that hunt specific prey, such as tuna, small sharks, and seals.</figcaption></figure> <p>Food webs largely define ecosystems, and the trophic levels define the position of organisms within the webs. But these trophic levels are not always simple integers, because organisms often feed at more than one trophic level.<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> For example, some carnivores also eat plants, and some plants are carnivores. A large carnivore may eat both smaller carnivores and herbivores; the bobcat eats rabbits, but the mountain lion eats both bobcats and rabbits. Animals can also eat each other; the <a href="/wiki/American_bullfrog" title="American bullfrog">bullfrog</a> eats <a href="/wiki/Crayfish" title="Crayfish">crayfish</a> and crayfish eat young bullfrogs. The feeding habits of a juvenile animal, and, as a consequence, its trophic level, can change as it grows up. </p><p>The fisheries scientist <a href="/wiki/Daniel_Pauly" title="Daniel Pauly">Daniel Pauly</a> sets the values of trophic levels to one in plants and detritus, two in herbivores and detritivores (primary consumers), three in secondary consumers, and so on. The definition of the trophic level, TL, for any consumer species is:<sup id="cite_ref-Pauly&Palomares_8-1" class="reference"><a href="#cite_note-Pauly&Palomares-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <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 TL_{i}=1+\sum _{j}(TL_{j}\cdot DC_{ij})\!}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>T</mi> <msub> <mi>L</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> <mo>=</mo> <mn>1</mn> <mo>+</mo> <munder> <mo>∑</mo> <mrow class="MJX-TeXAtom-ORD"> <mi>j</mi> </mrow> </munder> <mo stretchy="false">(</mo> <mi>T</mi> <msub> <mi>L</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>j</mi> </mrow> </msub> <mo>⋅</mo> <mi>D</mi> <msub> <mi>C</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> <mi>j</mi> </mrow> </msub> <mo stretchy="false">)</mo> <mspace width="negativethinmathspace" /> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle TL_{i}=1+\sum _{j}(TL_{j}\cdot DC_{ij})\!}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/3c8fbf55e3325ca7fb0239a63ddd419d7ba2f88c" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -3.338ex; margin-right: -0.166ex; width:26.935ex; height:5.843ex;" alt="{\displaystyle TL_{i}=1+\sum _{j}(TL_{j}\cdot DC_{ij})\!}"></span></dd></dl></dd></dl> <p>where <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 TL_{j}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>T</mi> <msub> <mi>L</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>j</mi> </mrow> </msub> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle TL_{j}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/54749d24c562f963e31fc6cb492aaf5196a17cc2" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -1.005ex; width:4.129ex; height:2.843ex;" alt="{\displaystyle TL_{j}}"></span> is the fractional trophic level of the prey <i>j</i>, and <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 DC_{ij}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>D</mi> <msub> <mi>C</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> <mi>j</mi> </mrow> </msub> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle DC_{ij}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/64f12feb826cbfbe172e998d78874c6cd9d9a675" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -1.005ex; width:5.063ex; height:2.843ex;" alt="{\displaystyle DC_{ij}}"></span> represents the fraction of <i>j</i> in the diet of <i>i</i>. That is, the consumer trophic level is one plus the <a href="/wiki/Weighted_average" class="mw-redirect" title="Weighted average">weighted average</a> of how much different trophic levels contribute to its food. </p><p>In the case of marine ecosystems, the trophic level of most fish and other marine consumers takes a value between 2.0 and 5.0. The upper value, 5.0, is unusual, even for large fish,<sup id="cite_ref-r1_16-0" class="reference"><a href="#cite_note-r1-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> though it occurs in apex predators of marine mammals, such as polar bears and orcas.<sup id="cite_ref-Pauly1998b_17-0" class="reference"><a href="#cite_note-Pauly1998b-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p><p>In addition to observational studies of animal behavior, and quantification of animal stomach contents, trophic level can be quantified through <a href="/wiki/Stable_isotope" class="mw-redirect" title="Stable isotope">stable isotope</a> analysis of animal tissues such as <a href="/wiki/Muscle" title="Muscle">muscle</a>, <a href="/wiki/Skin" title="Skin">skin</a>, <a href="/wiki/Hair" title="Hair">hair</a>, bone <a href="/wiki/Collagen" title="Collagen">collagen</a>. This is because there is a consistent increase in the nitrogen isotopic composition at each trophic level caused by fractionations that occur with the synthesis of biomolecules; the magnitude of this increase in nitrogen isotopic composition is approximately 3–4‰.<sup id="cite_ref-r2_18-0" class="reference"><a href="#cite_note-r2-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><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> </p> <div class="mw-heading mw-heading2"><h2 id="Mean_trophic_level">Mean trophic level</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Trophic_level&action=edit&section=6" title="Edit section: Mean trophic level"><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:Bluefin-big.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/18/Bluefin-big.jpg/220px-Bluefin-big.jpg" decoding="async" width="220" height="145" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/18/Bluefin-big.jpg/330px-Bluefin-big.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/18/Bluefin-big.jpg/440px-Bluefin-big.jpg 2x" data-file-width="1728" data-file-height="1140" /></a><figcaption>The mean trophic level of the world fisheries catch has steadily declined because many high trophic level fish, such as this <a href="/wiki/Tuna" title="Tuna">tuna</a>, have been <a href="/wiki/Overfished" class="mw-redirect" title="Overfished">overfished</a>.</figcaption></figure> <p>In fisheries, the mean trophic level for the fisheries catch across an entire area or ecosystem is calculated for year <span class="texhtml mvar" style="font-style:italic;">y</span> as: </p> <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 TL_{y}={\frac {\sum _{i}(TL_{i}\cdot Y_{iy})}{\sum _{i}Y_{iy}}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>T</mi> <msub> <mi>L</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>y</mi> </mrow> </msub> <mo>=</mo> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <munder> <mo>∑</mo> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </munder> <mo stretchy="false">(</mo> <mi>T</mi> <msub> <mi>L</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> <mo>⋅</mo> <msub> <mi>Y</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> <mi>y</mi> </mrow> </msub> <mo stretchy="false">)</mo> </mrow> <mrow> <munder> <mo>∑</mo> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </munder> <msub> <mi>Y</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> <mi>y</mi> </mrow> </msub> </mrow> </mfrac> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle TL_{y}={\frac {\sum _{i}(TL_{i}\cdot Y_{iy})}{\sum _{i}Y_{iy}}}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/979a5ed588bc3803e3505d0c774c49a763c1a4a5" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -2.671ex; width:21.931ex; height:6.509ex;" alt="{\displaystyle TL_{y}={\frac {\sum _{i}(TL_{i}\cdot Y_{iy})}{\sum _{i}Y_{iy}}}}"></span></dd></dl></dd></dl> <p>where <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 Y_{iy}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msub> <mi>Y</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> <mi>y</mi> </mrow> </msub> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle Y_{iy}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/4979c162a2b4e11f55a5b8e62c7c37c052abd317" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -1.005ex; width:2.967ex; height:2.843ex;" alt="{\displaystyle Y_{iy}}"></span> is the annual catch of the species or group <span class="texhtml mvar" style="font-style:italic;">i</span> in year <span class="texhtml mvar" style="font-style:italic;">y</span>, and <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 \ TL_{i}\ }"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mtext> </mtext> <mi>T</mi> <msub> <mi>L</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> <mtext> </mtext> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \ TL_{i}\ }</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/10ba36ab9191a61c4e0bebee456e62b34a336f40" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.671ex; width:5.18ex; height:2.509ex;" alt="{\displaystyle \ TL_{i}\ }"></span> is the trophic level for species <span class="texhtml mvar" style="font-style:italic;">i</span> as defined above.<sup id="cite_ref-Pauly&Palomares_8-2" class="reference"><a href="#cite_note-Pauly&Palomares-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p>Fish at higher trophic levels usually have a higher economic value, which can result in <a href="/wiki/Overfishing" title="Overfishing">overfishing</a> at the higher trophic levels. Earlier reports found precipitous declines in mean trophic level of <a href="/wiki/Fisheries" class="mw-redirect" title="Fisheries">fisheries</a> catch, in a process known as <a href="/wiki/Fishing_down_the_food_web" title="Fishing down the food web">fishing down the food web</a>.<sup id="cite_ref-r3_20-0" class="reference"><a href="#cite_note-r3-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> However, more recent work finds no relation between economic value and trophic level;<sup id="cite_ref-r4_21-0" class="reference"><a href="#cite_note-r4-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> and that mean trophic levels in catches, surveys and stock assessments have not in fact declined, suggesting that fishing down the food web is not a global phenomenon.<sup id="cite_ref-r5_22-0" class="reference"><a href="#cite_note-r5-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> However Pauly <i>et al</i>. note that trophic levels peaked at 3.4 in 1970 in the northwest and west-central Atlantic, followed by a subsequent decline to 2.9 in 1994. They report a shift away from long-lived, piscivorous, high-trophic-level bottom fishes, such as cod and haddock, to short-lived, planktivorous, low-trophic-level invertebrates (e.g., shrimp) and small, pelagic fish (e.g., herring). This shift from high-trophic-level fishes to low-trophic-level invertebrates and fishes is a response to changes in the relative abundance of the preferred catch. They consider that this is part of the global fishery collapse,<sup id="cite_ref-Pauly1998b_17-1" class="reference"><a href="#cite_note-Pauly1998b-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Pauly1998a_23-0" class="reference"><a href="#cite_note-Pauly1998a-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> which finds an echo in the overfished Mediterranean Sea.<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>Humans have a mean trophic level of about 2.21, about the same as a pig or an anchovy.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="FiB_index">FiB index</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Trophic_level&action=edit&section=7" title="Edit section: FiB index"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Since biomass <a href="/wiki/Ecological_efficiency" title="Ecological efficiency">transfer efficiencies</a> are only about 10%, it follows that the rate of biological production is much greater at lower trophic levels than it is at higher levels. Fisheries catch, at least, to begin with, will tend to increase as the trophic level declines. At this point the fisheries will target species lower in the food web.<sup id="cite_ref-Pauly1998a_23-1" class="reference"><a href="#cite_note-Pauly1998a-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> In 2000, this led Pauly and others to construct a "Fisheries in Balance" index, usually called the FiB index.<sup id="cite_ref-Pauly2000_27-0" class="reference"><a href="#cite_note-Pauly2000-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> The FiB index is defined, for any year <i>y</i>, by<sup id="cite_ref-Pauly&Palomares_8-3" class="reference"><a href="#cite_note-Pauly&Palomares-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <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 FiB_{y}=\log {\frac {Y_{y}/(TE)^{TL_{y}}}{Y_{0}/(TE)^{TL_{0}}}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>F</mi> <mi>i</mi> <msub> <mi>B</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>y</mi> </mrow> </msub> <mo>=</mo> <mi>log</mi> <mo>⁡</mo> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <msub> <mi>Y</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>y</mi> </mrow> </msub> <mrow class="MJX-TeXAtom-ORD"> <mo>/</mo> </mrow> <mo stretchy="false">(</mo> <mi>T</mi> <mi>E</mi> <msup> <mo stretchy="false">)</mo> <mrow class="MJX-TeXAtom-ORD"> <mi>T</mi> <msub> <mi>L</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>y</mi> </mrow> </msub> </mrow> </msup> </mrow> <mrow> <msub> <mi>Y</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>0</mn> </mrow> </msub> <mrow class="MJX-TeXAtom-ORD"> <mo>/</mo> </mrow> <mo stretchy="false">(</mo> <mi>T</mi> <mi>E</mi> <msup> <mo stretchy="false">)</mo> <mrow class="MJX-TeXAtom-ORD"> <mi>T</mi> <msub> <mi>L</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>0</mn> </mrow> </msub> </mrow> </msup> </mrow> </mfrac> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle FiB_{y}=\log {\frac {Y_{y}/(TE)^{TL_{y}}}{Y_{0}/(TE)^{TL_{0}}}}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/723b788c1add3e5b535b706b789ff761983786b3" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -2.671ex; width:24.779ex; height:6.843ex;" alt="{\displaystyle FiB_{y}=\log {\frac {Y_{y}/(TE)^{TL_{y}}}{Y_{0}/(TE)^{TL_{0}}}}}"></span></dd></dl></dd></dl> <p>where <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 Y_{y}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msub> <mi>Y</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>y</mi> </mrow> </msub> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle Y_{y}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/7fd0f3653c219cdf1f338410797fa3715211367e" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -1.005ex; width:2.4ex; height:2.843ex;" alt="{\displaystyle Y_{y}}"></span> is the catch at year <i>y</i>, <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 TL_{y}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>T</mi> <msub> <mi>L</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>y</mi> </mrow> </msub> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle TL_{y}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/280ca70d94d7a65934f054d0fb0c678ccf58c84e" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -1.005ex; width:4.268ex; height:2.843ex;" alt="{\displaystyle TL_{y}}"></span> is the mean trophic level of the catch at year <i>y</i>, <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 Y_{0}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msub> <mi>Y</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>0</mn> </mrow> </msub> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle Y_{0}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/d4305e31e20568c004e0e4c8540bbfe2730a42cb" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.671ex; width:2.405ex; height:2.509ex;" alt="{\displaystyle Y_{0}}"></span> is the catch, <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 TL_{0}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>T</mi> <msub> <mi>L</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>0</mn> </mrow> </msub> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle TL_{0}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/eeba29429ef80ddcab8eb1e5acc5536985f29158" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.671ex; width:4.273ex; height:2.509ex;" alt="{\displaystyle TL_{0}}"></span> the mean trophic level of the catch at the start of the series being analyzed, and <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 TE}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>T</mi> <mi>E</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle TE}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/4269b5099d7b300c074d573991af09dc3d2a1c1c" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.338ex; width:3.412ex; height:2.176ex;" alt="{\displaystyle TE}"></span> is the transfer efficiency of biomass or energy between trophic levels. </p><p>The FiB index is stable (zero) over periods of time when changes in trophic levels are matched by appropriate changes in the catch in the opposite direction. The index increases if catches increase for any reason, e.g. higher fish biomass, or geographic expansion.<sup id="cite_ref-Pauly&Palomares_8-4" class="reference"><a href="#cite_note-Pauly&Palomares-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Such decreases explain the "backward-bending" plots of trophic level versus catch originally observed by Pauly and others in 1998.<sup id="cite_ref-Pauly1998a_23-2" class="reference"><a href="#cite_note-Pauly1998a-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Tritrophic_and_other_interactions">Tritrophic and other interactions</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Trophic_level&action=edit&section=8" title="Edit section: Tritrophic and other interactions"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Tritrophic_interactions_in_plant_defense" title="Tritrophic interactions in plant defense">Tritrophic interactions in plant defense</a></div> <p>One aspect of trophic levels is called tritrophic interaction. Ecologists often restrict their research to two trophic levels as a way of simplifying the analysis; however, this can be misleading if tritrophic interactions (such as plant–herbivore–predator) are not easily understood by simply adding pairwise interactions (plant-herbivore plus herbivore–predator, for example). Significant interactions can occur between the first trophic level (plant) and the third trophic level (a predator) in determining herbivore population growth, for example. Simple genetic changes may yield morphological variants in plants that then differ in their resistance to herbivores because of the effects of the plant architecture on enemies of the herbivore.<sup id="cite_ref-Kareiva&Sahakain_28-0" class="reference"><a href="#cite_note-Kareiva&Sahakain-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Plants can also <a href="/wiki/Plant_defense_against_herbivory" title="Plant defense against herbivory">develop defenses against herbivores</a> such as chemical defenses.<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> </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=Trophic_level&action=edit&section=9" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Cascade_effect_(ecology)" title="Cascade effect (ecology)">Cascade effect</a></li> <li><a href="/wiki/Energy_flow_(ecology)" title="Energy flow (ecology)">Energy flow (ecology)</a></li> <li><a href="/wiki/Marine_trophic_level" class="mw-redirect" title="Marine trophic level">Marine trophic level</a></li> <li><a href="/wiki/Mesopredator_release_hypothesis" title="Mesopredator release hypothesis">Mesopredator release hypothesis</a></li> <li><a href="/wiki/Trophic_cascade" title="Trophic cascade">Trophic cascade</a></li> <li><a href="/wiki/Trophic_dynamics" class="mw-redirect" title="Trophic dynamics">Trophic dynamics</a> – <i>Food web</i></li> <li><a href="/wiki/Trophic_state_index" title="Trophic state index">Trophic state index</a> – applied to lakes</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=Trophic_level&action=edit&section=10" 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 reflist-columns references-column-width" style="column-width: 25em;"> <ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite class="citation encyclopaedia cs1"><a rel="nofollow" class="external text" href="https://www.merriam-webster.com/dictionary/trophic">"Trophic"</a>. <i>Merriam-Webster Dictionary</i><span class="reference-accessdate">. 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href="/wiki/Ecosystem_model" title="Ecosystem model">Modelling ecosystems</a>: <a class="mw-selflink selflink">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 class="mw-selflink selflink">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 href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">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 href="/wiki/Endolith" title="Endolith">Endolith</a></li> <li><a href="/wiki/Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></li> <li><a href="/wiki/Functional_ecology" title="Functional ecology">Functional ecology</a></li> <li><a href="/wiki/Industrial_ecology" title="Industrial ecology">Industrial ecology</a></li> <li><a href="/wiki/Macroecology" title="Macroecology">Macroecology</a></li> <li><a href="/wiki/Microecosystem" title="Microecosystem">Microecosystem</a></li> <li><a href="/wiki/Natural_environment" title="Natural environment">Natural environment</a></li> <li><a href="/wiki/Regime_shift" title="Regime shift">Regime shift</a></li> <li><a href="/wiki/Sexecology" title="Sexecology">Sexecology</a></li> <li><a href="/wiki/Systems_ecology" title="Systems ecology">Systems ecology</a></li> <li><a href="/wiki/Urban_ecology" title="Urban ecology">Urban ecology</a></li> <li><a href="/wiki/Theoretical_ecology" title="Theoretical ecology">Theoretical 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"></div><div role="navigation" class="navbox" aria-labelledby="Aquatic_ecosystems" 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:Aquatic_ecosystems" title="Template:Aquatic ecosystems"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Aquatic_ecosystems" title="Template talk:Aquatic ecosystems"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Aquatic_ecosystems" title="Special:EditPage/Template:Aquatic ecosystems"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Aquatic_ecosystems" style="font-size:114%;margin:0 4em"><a href="/wiki/Aquatic_ecosystem" title="Aquatic ecosystem">Aquatic ecosystems</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="General_components_and_freshwater_ecosystems" style="font-size:114%;margin:0 4em">General components and <a href="/wiki/Freshwater_ecosystem" title="Freshwater ecosystem">freshwater ecosystems</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">General</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/Acoustic_ecology" title="Acoustic ecology">Acoustic ecology</a></li> <li><a href="/wiki/Algal_bloom" title="Algal bloom">Algal bloom</a></li> <li><a href="/wiki/Anoxic_waters" title="Anoxic waters">Anoxic waters</a></li> <li><a href="/wiki/Aquatic_adaptation" class="mw-redirect" title="Aquatic adaptation">Aquatic adaptation</a></li> <li><a href="/wiki/Aquatic_animal" title="Aquatic animal">Aquatic animal</a> <ul><li><a href="/wiki/Aquatic_insect" title="Aquatic insect">Insect</a></li> <li><a href="/wiki/Aquatic_mammal" title="Aquatic mammal">Mammal</a></li> <li><a href="/wiki/Water_bird" title="Water bird">Water bird</a></li></ul></li> <li><a href="/wiki/Aquatic_biomonitoring" title="Aquatic biomonitoring">Aquatic biomonitoring</a></li> <li><a href="/wiki/Aquatic_plant" title="Aquatic plant">Aquatic plant</a></li> <li><a href="/wiki/Agent-based_model_in_biology#Aquatic_population_dynamics" title="Agent-based model in biology">Aquatic population dynamics</a></li> <li><a href="/wiki/Aquatic_predation" class="mw-redirect" title="Aquatic predation">Aquatic predation</a></li> <li><a href="/wiki/Aquatic_respiration" title="Aquatic respiration">Aquatic respiration</a></li> <li><a href="/wiki/Aquatic_science" title="Aquatic science">Aquatic science</a></li> <li><a href="/wiki/Aquatic_toxicology" title="Aquatic toxicology">Aquatic toxicology</a></li> <li><a href="/wiki/Benthos" title="Benthos">Benthos</a></li> <li><a href="/wiki/Bioluminescence" title="Bioluminescence">Bioluminescence</a></li> <li><a href="/wiki/Biomass_(ecology)" title="Biomass (ecology)">Biomass</a></li> <li><a href="/wiki/Cascade_effect_(ecology)" title="Cascade effect (ecology)">Cascade effect</a></li> <li><a href="/wiki/Colored_dissolved_organic_matter" title="Colored dissolved organic matter">Colored dissolved organic matter</a></li> <li><a href="/wiki/Dead_zone_(ecology)" title="Dead zone (ecology)">Dead zone</a></li> <li><a href="/wiki/Ecohydrology" title="Ecohydrology">Ecohydrology</a></li> <li><a href="/wiki/Eutrophication" title="Eutrophication">Eutrophication</a></li> <li><a href="/wiki/Fisheries_science" title="Fisheries science">Fisheries science</a></li> <li><a href="/wiki/Food_chain" title="Food chain">Food chain</a></li> <li><a href="/wiki/Food_web" title="Food web">Food web</a></li> <li><a href="/wiki/GIS_and_aquatic_science" title="GIS and aquatic science">GIS and aquatic science</a></li> <li><a href="/wiki/Hydrobiology" title="Hydrobiology">Hydrobiology</a></li> <li><a href="/wiki/Hypoxia_(environmental)" title="Hypoxia (environmental)">Hypoxia</a></li> <li><a href="/wiki/Macrobenthos" title="Macrobenthos">Macrobenthos</a></li> <li><a href="/wiki/Meiobenthos" title="Meiobenthos">Meiobenthos</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_loop" title="Microbial loop">Microbial loop</a></li> <li><a href="/wiki/Nekton" title="Nekton">Nekton</a></li> <li><a href="/wiki/Neuston" title="Neuston">Neuston</a></li> <li><a href="/wiki/Particle_(ecology)" title="Particle (ecology)">Particle</a></li> <li><a href="/wiki/Pelagic_zone" title="Pelagic zone">Pelagic zone</a></li> <li><a href="/wiki/Photic_zone" title="Photic zone">Photic zone</a></li> <li><a href="/wiki/Phytoplankton" title="Phytoplankton">Phytoplankton</a></li> <li><a href="/wiki/Plankton" title="Plankton">Plankton</a></li> <li><a href="/wiki/Pleuston" class="mw-redirect" title="Pleuston">Pleuston</a></li> <li><a href="/wiki/Productivity_(ecology)" title="Productivity (ecology)">Productivity</a></li> <li><a href="/wiki/Ramsar_Convention" title="Ramsar Convention">Ramsar Convention</a></li> <li><a href="/wiki/Sediment_trap" title="Sediment trap">Sediment trap</a></li> <li><a href="/wiki/Shoaling_and_schooling" title="Shoaling and schooling">Shoaling and schooling</a></li> <li><a href="/wiki/Siltation" title="Siltation">Siltation</a></li> <li><a href="/wiki/Spawn_(biology)" class="mw-redirect" title="Spawn (biology)">Spawn</a></li> <li><a href="/wiki/Isotope_analysis#Stable_isotope_analysis_in_aquatic_ecosystems" title="Isotope analysis">Stable isotope analysis in aquatic ecosystems</a></li> <li><a href="/wiki/Substrate_(marine_biology)" class="mw-redirect" title="Substrate (marine biology)">Substrate</a></li> <li><a href="/wiki/Thermal_pollution" title="Thermal pollution">Thermal pollution</a></li> <li><a class="mw-selflink selflink">Trophic level</a></li> <li><a href="/wiki/Underwater_camouflage" title="Underwater camouflage">Underwater camouflage</a></li> <li><a href="/wiki/Water_column" title="Water column">Water column</a></li> <li><a href="/wiki/Zooplankton" title="Zooplankton">Zooplankton</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="3" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Aerial_view_of_Slapton_Sands_and_northern_part_of_Ley_-_geograph.org.uk_-_1434412.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Aerial_view_of_Slapton_Sands_and_northern_part_of_Ley_-_geograph.org.uk_-_1434412.jpg/140px-Aerial_view_of_Slapton_Sands_and_northern_part_of_Ley_-_geograph.org.uk_-_1434412.jpg" decoding="async" width="140" height="202" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Aerial_view_of_Slapton_Sands_and_northern_part_of_Ley_-_geograph.org.uk_-_1434412.jpg/210px-Aerial_view_of_Slapton_Sands_and_northern_part_of_Ley_-_geograph.org.uk_-_1434412.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Aerial_view_of_Slapton_Sands_and_northern_part_of_Ley_-_geograph.org.uk_-_1434412.jpg/280px-Aerial_view_of_Slapton_Sands_and_northern_part_of_Ley_-_geograph.org.uk_-_1434412.jpg 2x" data-file-width="382" data-file-height="551" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Freshwater</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/Freshwater_biology" title="Freshwater biology">Freshwater biology</a></li> <li><a href="/wiki/Freshwater_biome" class="mw-redirect" title="Freshwater biome">Freshwater biome</a></li> <li><a href="/wiki/Freshwater_environmental_quality_parameters" title="Freshwater environmental quality parameters">Freshwater environmental quality parameters</a></li> <li><a href="/wiki/Freshwater_fish" title="Freshwater fish">Freshwater fish</a></li> <li><a href="/wiki/Hyporheic_zone" title="Hyporheic zone">Hyporheic zone</a></li> <li><a href="/wiki/Limnology" title="Limnology">Limnology</a></li> <li><a href="/wiki/Lake_ecosystem" title="Lake ecosystem">Lake ecosystem</a></li> <li><a href="/wiki/Lake_stratification" title="Lake stratification">Lake stratification</a></li> <li><a href="/wiki/Macrophyte" class="mw-redirect" title="Macrophyte">Macrophyte</a></li> <li><a href="/wiki/Pond" title="Pond">Pond</a> <ul><li><a href="/wiki/Fish_pond" title="Fish pond">Fish pond</a></li></ul></li> <li><a href="/wiki/Rheotaxis" title="Rheotaxis">Rheotaxis</a></li> <li><a href="/wiki/River_ecosystem" title="River ecosystem">River ecosystem</a></li> <li><a href="/wiki/Stream_bed" title="Stream bed">Stream bed</a></li> <li><a href="/wiki/Stream_pool" title="Stream pool">Stream pool</a></li> <li><a href="/wiki/Trophic_state_index" title="Trophic state index">Trophic state index</a></li> <li><a href="/wiki/Upland_and_lowland" title="Upland and lowland">Upland and lowland</a></li> <li><a href="/wiki/Water_garden" title="Water garden">Water garden</a></li> <li><a href="/wiki/Wetland" title="Wetland">Wetland</a> <ul><li><a href="/wiki/Bog" title="Bog">Bog</a></li> <li><a href="/wiki/Brackish_marsh" title="Brackish marsh">Brackish marsh</a></li> <li><a href="/wiki/Fen" title="Fen">Fen</a></li> <li><a href="/wiki/Freshwater_marsh" title="Freshwater marsh">Freshwater marsh</a></li> <li><a href="/wiki/Freshwater_swamp_forest" title="Freshwater swamp forest">Freshwater swamp forest</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Ecoregions</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/List_of_freshwater_ecoregions_(WWF)" title="List of freshwater ecoregions (WWF)">List of freshwater ecoregions (WWF)</a> <ul><li><a href="/wiki/List_of_freshwater_ecoregions_in_Africa_and_Madagascar" title="List of freshwater ecoregions in Africa and Madagascar">Africa and Madagascar</a></li> <li><a href="/wiki/List_of_freshwater_ecoregions_of_Latin_America_and_the_Caribbean" title="List of freshwater ecoregions of Latin America and the Caribbean">Latin America and the Caribbean</a></li></ul></li> <li><a href="/wiki/List_of_marine_ecoregions" title="List of marine ecoregions">List of marine ecoregions</a></li> <li>Specific examples <ul><li><a href="/wiki/Geography_and_ecology_of_the_Everglades" title="Geography and ecology of the Everglades">Everglades</a></li> <li><a href="/wiki/Freshwater_ecology_of_Maharashtra" title="Freshwater ecology of Maharashtra">Maharashtra</a></li> <li><a href="/wiki/Ecosystem_of_the_North_Pacific_Subtropical_Gyre" title="Ecosystem of the North Pacific Subtropical Gyre">North Pacific Subtropical Gyre</a></li> <li><a href="/wiki/Ecology_of_the_San_Francisco_Estuary" title="Ecology of the San Francisco Estuary">San Francisco Estuary</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Marine_ecosystems" style="font-size:114%;margin:0 4em"><a href="/wiki/Marine_ecosystem" title="Marine ecosystem">Marine ecosystems</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">General</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/Deep_scattering_layer" title="Deep scattering layer">Deep scattering layer</a></li> <li><a href="/wiki/Diel_vertical_migration" title="Diel vertical migration">Diel vertical migration</a></li> <li><a href="/wiki/F-ratio_(oceanography)" title="F-ratio (oceanography)">f-ratio</a></li> <li><a href="/wiki/Iron_fertilization" title="Iron fertilization">Iron fertilization</a></li> <li><a href="/wiki/Large_marine_ecosystem" class="mw-redirect" title="Large marine ecosystem">Large marine ecosystem</a></li> <li><a href="/wiki/Marine_biology" title="Marine biology">Marine biology</a></li> <li><a href="/wiki/Marine_chemistry" title="Marine chemistry">Marine chemistry</a></li> <li><a href="/wiki/Marine_food_web" title="Marine food web">Marine food web</a></li> <li><a href="/wiki/Marine_primary_production" title="Marine primary production">Marine primary production</a></li> <li><a href="/wiki/Marine_snow" title="Marine snow">Marine snow</a></li> <li><a href="/wiki/Ocean_fertilization" title="Ocean fertilization">Ocean fertilization</a></li> <li><a href="/wiki/Oceanic_physical-biological_process" title="Oceanic physical-biological process">Oceanic physical-biological process</a></li> <li><a href="/wiki/Ocean_turbidity" title="Ocean turbidity">Ocean turbidity</a></li> <li><a href="/wiki/Photophore" title="Photophore">Photophore</a></li> <li><a href="/wiki/Thorson%27s_rule" title="Thorson's rule">Thorson's rule</a></li> <li><a href="/wiki/Upwelling" title="Upwelling">Upwelling</a></li> <li><a href="/wiki/Viral_shunt" title="Viral shunt">Viral shunt</a></li> <li><a href="/wiki/Whale_fall" title="Whale fall">Whale fall</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="4" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Maldivesfish2.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/35/Maldivesfish2.jpg/140px-Maldivesfish2.jpg" decoding="async" width="140" height="95" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/35/Maldivesfish2.jpg/210px-Maldivesfish2.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/35/Maldivesfish2.jpg/280px-Maldivesfish2.jpg 2x" data-file-width="1818" data-file-height="1228" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Marine_life" title="Marine life">Marine life</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Census_of_Marine_Life" title="Census of Marine Life">Census of Marine Life</a></li> <li><a href="/wiki/Deep-sea_community" title="Deep-sea community">Deep-sea community</a></li> <li><a href="/wiki/Deep-water_coral" title="Deep-water coral">Deep-water coral</a></li> <li><a href="/wiki/Marine_fungi" title="Marine fungi">Marine fungi</a></li> <li><a href="/wiki/Marine_invertebrates" title="Marine invertebrates">Marine invertebrates</a></li> <li><a href="/wiki/Marine_larval_ecology" title="Marine larval ecology">Marine larval ecology</a></li> <li><a href="/wiki/Seagrass" title="Seagrass">Seagrass</a></li> <li><a href="/wiki/Seashore_wildlife" title="Seashore wildlife">Seashore wildlife</a></li> <li><a href="/wiki/Wild_fisheries" title="Wild fisheries">Wild fisheries</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Marine_microorganisms" title="Marine microorganisms">Microorganisms</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/Marine_bacteriophage" class="mw-redirect" title="Marine bacteriophage">Marine bacteriophage</a></li> <li><a href="/wiki/Marine_prokaryotes" title="Marine prokaryotes">Marine prokaryotes</a></li> <li><a href="/wiki/Marine_protists" title="Marine protists">Marine protists</a></li> <li><a href="/wiki/Marine_viruses" title="Marine viruses">Marine viruses</a></li> <li><a href="/wiki/Paradox_of_the_plankton" title="Paradox of the plankton">Paradox of the plankton</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Marine_vertebrate" title="Marine vertebrate">Vertebrates</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/Marine_mammal" title="Marine mammal">Marine mammal</a></li> <li><a href="/wiki/Marine_reptile" title="Marine reptile">Marine reptile</a></li> <li><a href="/wiki/Saltwater_fish" title="Saltwater fish">Saltwater fish</a> <ul><li><a href="/wiki/Coastal_fish" title="Coastal fish">Coastal fish</a></li> <li><a href="/wiki/Coral_reef_fish" title="Coral reef fish">Coral reef fish</a></li> <li><a href="/wiki/Deep-sea_fish" title="Deep-sea fish">Deep-sea fish</a></li> <li><a href="/wiki/Demersal_fish" title="Demersal fish">Demersal fish</a></li> <li><a href="/wiki/Pelagic_fish" title="Pelagic fish">Pelagic fish</a></li></ul></li> <li><a href="/wiki/Seabird" title="Seabird">Seabird</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Marine_habitat" title="Marine habitat">Marine habitats</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/Bay_mud" title="Bay mud">Bay mud</a></li> <li><a href="/wiki/Marine_coastal_ecosystem" title="Marine coastal ecosystem">Marine coastal ecosystem</a></li> <li><a href="/wiki/Coastal_biogeomorphology" title="Coastal biogeomorphology">Coastal biogeomorphology</a></li> <li><a href="/wiki/Cold_seep" title="Cold seep">Cold seep</a></li> <li><a href="/wiki/Coral_reef" title="Coral reef">Coral reef</a></li> <li><a href="/wiki/Davidson_Seamount#Ecology" title="Davidson Seamount">Davidson Seamount § Ecology</a></li> <li><a href="/wiki/Estuary" title="Estuary">Estuary</a></li> <li><a href="/wiki/Intertidal_ecology" title="Intertidal ecology">Intertidal ecology</a></li> <li><a href="/wiki/Intertidal_wetland" title="Intertidal wetland">Intertidal wetland</a></li> <li><a href="/wiki/Kelp_forest" title="Kelp forest">Kelp forest</a></li> <li><a href="/wiki/Hydrothermal_vent" title="Hydrothermal vent">Hydrothermal vent</a></li> <li><a href="/wiki/Lagoon" title="Lagoon">Lagoon</a></li> <li><a href="/wiki/Mangrove" title="Mangrove">Mangrove</a></li> <li><a href="/wiki/Marine_biomes" class="mw-redirect" title="Marine biomes">Marine biomes</a></li> <li><a href="/wiki/Mudflat" title="Mudflat">Mudflat</a></li> <li><a href="/wiki/Oyster_reef" title="Oyster reef">Oyster reef</a></li> <li><a href="/wiki/Rocky_shore" title="Rocky shore">Rocky shore</a></li> <li><a href="/wiki/Salt_marsh" title="Salt marsh">Salt marsh</a></li> <li><a href="/wiki/Salt_pannes_and_pools" title="Salt pannes and pools">Salt pannes and pools</a></li> <li><a href="/wiki/Seagrass_meadow" title="Seagrass meadow">Seagrass meadow</a></li> <li><a href="/wiki/Sponge_ground" title="Sponge 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