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class="vector-toc-numb">2</span> <span>Social interaction</span> </div> </a> <ul id="toc-Social_interaction-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Foraging_advantages" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Foraging_advantages"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Foraging advantages</span> </div> </a> <ul id="toc-Foraging_advantages-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Reproductive_advantages" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Reproductive_advantages"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Reproductive advantages</span> </div> </a> <ul id="toc-Reproductive_advantages-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Hydrodynamic_efficiency" class="vector-toc-list-item vector-toc-level-1 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class="vector-toc-list"> </ul> </li> <li id="toc-How_fish_school" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#How_fish_school"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>How fish school</span> </div> </a> <ul id="toc-How_fish_school-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Describing_shoal_structure" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Describing_shoal_structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Describing shoal structure</span> </div> </a> <ul id="toc-Describing_shoal_structure-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Modelling_school_behaviour" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Modelling_school_behaviour"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>Modelling school behaviour</span> </div> </a> <button aria-controls="toc-Modelling_school_behaviour-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Modelling school behaviour subsection</span> </button> <ul id="toc-Modelling_school_behaviour-sublist" class="vector-toc-list"> <li id="toc-Mathematical_models" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mathematical_models"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.1</span> <span>Mathematical models</span> </div> </a> <ul id="toc-Mathematical_models-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolutionary_models" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Evolutionary_models"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.2</span> <span>Evolutionary models</span> </div> </a> <ul id="toc-Evolutionary_models-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Mapping_the_formation_of_schools" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Mapping_the_formation_of_schools"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>Mapping the formation of schools</span> </div> </a> <ul id="toc-Mapping_the_formation_of_schools-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Leadership_and_decision-making" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Leadership_and_decision-making"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>Leadership and decision-making</span> </div> </a> <ul id="toc-Leadership_and_decision-making-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Shoal_choice" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Shoal_choice"> <div class="vector-toc-text"> <span class="vector-toc-numb">13</span> <span>Shoal choice</span> </div> </a> <ul id="toc-Shoal_choice-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Commercial_fishing" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Commercial_fishing"> <div class="vector-toc-text"> <span class="vector-toc-numb">14</span> <span>Commercial fishing</span> </div> </a> <ul id="toc-Commercial_fishing-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_examples" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_examples"> <div class="vector-toc-text"> <span class="vector-toc-numb">15</span> <span>Further examples</span> </div> </a> <ul id="toc-Further_examples-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">16</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Notes" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Notes"> <div class="vector-toc-text"> <span class="vector-toc-numb">17</span> <span>Notes</span> </div> </a> <ul id="toc-Notes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">18</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">19</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-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">20</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" 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href="https://bg.wikipedia.org/wiki/%D0%A0%D0%B8%D0%B1%D0%B5%D0%BD_%D0%BF%D0%B0%D1%81%D0%B0%D0%B6" title="Рибен пасаж – Bulgarian" lang="bg" hreflang="bg" data-title="Рибен пасаж" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Mola_de_peix" title="Mola de peix – Catalan" lang="ca" hreflang="ca" data-title="Mola de peix" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Schwarmfisch" title="Schwarmfisch – German" lang="de" hreflang="de" data-title="Schwarmfisch" 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/Kalaparv" title="Kalaparv – Estonian" lang="et" hreflang="et" data-title="Kalaparv" 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/Cardumen" title="Cardumen – Spanish" lang="es" hreflang="es" data-title="Cardumen" 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/Fi%C5%9Daro" title="Fiŝaro – Esperanto" lang="eo" hreflang="eo" data-title="Fiŝaro" 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/Sarda" title="Sarda – Basque" lang="eu" hreflang="eu" data-title="Sarda" 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%AF%D8%B3%D8%AA%D9%87_%D9%85%D8%A7%D9%87%DB%8C_%D9%88_%D8%AA%D9%88%D8%AF%D9%87_%D9%85%D8%A7%D9%87%DB%8C" title="دسته ماهی و توده ماهی – Persian" lang="fa" hreflang="fa" data-title="دسته ماهی و توده ماهی" 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/Banc_(poisson)" title="Banc (poisson) – French" lang="fr" hreflang="fr" data-title="Banc (poisson)" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Cardume" title="Cardume – Galician" lang="gl" hreflang="gl" data-title="Cardume" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Kawanan_dan_rombongan" title="Kawanan dan rombongan – Indonesian" lang="id" hreflang="id" data-title="Kawanan dan rombongan" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Fiskitorfa" title="Fiskitorfa – Icelandic" lang="is" hreflang="is" data-title="Fiskitorfa" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target"><span>Íslenska</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Banco_(pesci)" title="Banco (pesci) – Italian" lang="it" hreflang="it" data-title="Banco (pesci)" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-csb mw-list-item"><a href="https://csb.wikipedia.org/wiki/Wa%C5%82a_rib" title="Wała rib – Kashubian" lang="csb" hreflang="csb" data-title="Wała rib" data-language-autonym="Kaszëbsczi" data-language-local-name="Kashubian" class="interlanguage-link-target"><span>Kaszëbsczi</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/School_(vissen)" title="School (vissen) – Dutch" lang="nl" hreflang="nl" data-title="School (vissen)" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Stim" title="Stim – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Stim" 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/Stimfisk" title="Stimfisk – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Stimfisk" 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/%C5%81awica_ryb" title="Ławica ryb – Polish" lang="pl" hreflang="pl" data-title="Ławica ryb" 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/Cardume" title="Cardume – Portuguese" lang="pt" hreflang="pt" data-title="Cardume" 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%A1%D1%82%D0%B0%D0%B8_%D0%B8_%D0%BA%D0%BE%D1%81%D1%8F%D0%BA%D0%B8_%D1%80%D1%8B%D0%B1" title="Стаи и косяки рыб – Russian" lang="ru" hreflang="ru" data-title="Стаи и косяки рыб" 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/Shoaling_and_schooling" title="Shoaling and schooling – Simple English" lang="en-simple" hreflang="en-simple" data-title="Shoaling and schooling" 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-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Kalojen_parvik%C3%A4ytt%C3%A4ytyminen" title="Kalojen parvikäyttäytyminen – Finnish" lang="fi" hreflang="fi" data-title="Kalojen parvikäyttäytyminen" 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/Fiskstim" title="Fiskstim – Swedish" lang="sv" hreflang="sv" data-title="Fiskstim" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%AE%E0%AE%BE%E0%AE%AA%E0%AF%8D%E0%AE%AA%E0%AF%81_%E0%AE%AE%E0%AF%80%E0%AE%A9%E0%AF%8D%E0%AE%95%E0%AE%B3%E0%AF%8D_%E0%AE%AE%E0%AE%B1%E0%AF%8D%E0%AE%B1%E0%AF%81%E0%AE%AE%E0%AF%8D_%E0%AE%95%E0%AF%82%E0%AE%9F%E0%AF%8D%E0%AE%9F_%E0%AE%AE%E0%AF%80%E0%AE%A9%E0%AF%8D%E0%AE%95%E0%AE%B3%E0%AF%8D" title="மாப்பு மீன்கள் மற்றும் கூட்ட மீன்கள் – Tamil" lang="ta" hreflang="ta" data-title="மாப்பு மீன்கள் மற்றும் கூட்ட மீன்கள்" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E9%AD%9A%E7%BE%A3" title="魚羣 – Cantonese" lang="yue" hreflang="yue" data-title="魚羣" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target"><span>粵語</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a 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<div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">In biology, any group of fish that stay together for social reasons</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">This article is about shoaling and schooling fish. For other uses, see <a href="/wiki/Shoal_(disambiguation)" class="mw-disambig" title="Shoal (disambiguation)">Shoal (disambiguation)</a> and <a href="/wiki/School_(disambiguation)" class="mw-disambig" title="School (disambiguation)">School (disambiguation)</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">"School of fish" redirects here. For the alternative rock band, see <a href="/wiki/School_of_Fish" title="School of Fish">School of Fish</a>.</div> <p class="mw-empty-elt"> </p> <style data-mw-deduplicate="TemplateStyles:r1237032888/mw-parser-output/.tmulti">.mw-parser-output .tmulti .multiimageinner{display:flex;flex-direction:column}.mw-parser-output .tmulti .trow{display:flex;flex-direction:row;clear:left;flex-wrap:wrap;width:100%;box-sizing:border-box}.mw-parser-output .tmulti .tsingle{margin:1px;float:left}.mw-parser-output .tmulti .theader{clear:both;font-weight:bold;text-align:center;align-self:center;background-color:transparent;width:100%}.mw-parser-output .tmulti .thumbcaption{background-color:transparent}.mw-parser-output .tmulti .text-align-left{text-align:left}.mw-parser-output .tmulti .text-align-right{text-align:right}.mw-parser-output .tmulti .text-align-center{text-align:center}@media all and (max-width:720px){.mw-parser-output .tmulti .thumbinner{width:100%!important;box-sizing:border-box;max-width:none!important;align-items:center}.mw-parser-output .tmulti .trow{justify-content:center}.mw-parser-output .tmulti .tsingle{float:none!important;max-width:100%!important;box-sizing:border-box;text-align:center}.mw-parser-output .tmulti .tsingle .thumbcaption{text-align:left}.mw-parser-output .tmulti .trow>.thumbcaption{text-align:center}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .tmulti .multiimageinner img{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .tmulti .multiimageinner img{background-color:white}}</style><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:304px;max-width:304px"><div class="trow"><div class="theader" style="text-align:center">Shoaling and schooling</div></div><div class="trow"><div class="tsingle" style="width:302px;max-width:302px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Maldives_Surgeonfish,_Acanthurus_leucosternon.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/5/50/Maldives_Surgeonfish%2C_Acanthurus_leucosternon.jpg/300px-Maldives_Surgeonfish%2C_Acanthurus_leucosternon.jpg" decoding="async" width="300" height="225" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/50/Maldives_Surgeonfish%2C_Acanthurus_leucosternon.jpg/450px-Maldives_Surgeonfish%2C_Acanthurus_leucosternon.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/50/Maldives_Surgeonfish%2C_Acanthurus_leucosternon.jpg/600px-Maldives_Surgeonfish%2C_Acanthurus_leucosternon.jpg 2x" data-file-width="3072" data-file-height="2304" /></a></span></div><div class="thumbcaption">These <a href="/wiki/Acanthurus_leucosternon" title="Acanthurus leucosternon">powder blue tangs</a> are <b>shoaling</b>. They are swimming somewhat independently, but in such a way that they stay connected, forming a social group.</div></div></div><div class="trow"><div class="tsingle" style="width:302px;max-width:302px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Lutjanus_kasmira_school.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6b/Lutjanus_kasmira_school.jpg/300px-Lutjanus_kasmira_school.jpg" decoding="async" width="300" height="225" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6b/Lutjanus_kasmira_school.jpg/450px-Lutjanus_kasmira_school.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6b/Lutjanus_kasmira_school.jpg/600px-Lutjanus_kasmira_school.jpg 2x" data-file-width="2048" data-file-height="1536" /></a></span></div><div class="thumbcaption">These <a href="/wiki/Bluestripe_snapper" class="mw-redirect" title="Bluestripe snapper">bluestripe snapper</a> are <b>schooling</b>. They are all swimming in the same direction in a coordinated way.</div></div></div></div></div> <p>In biology, any group of <a href="/wiki/Fish" title="Fish">fish</a> that stay together for social reasons are <b>shoaling</b>, and if the group is swimming in the same direction in a coordinated manner, they are <b>schooling</b>.<sup id="cite_ref-Pitcher_365_1-0" class="reference"><a href="#cite_note-Pitcher_365-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> In common usage, the terms are sometimes used rather loosely.<sup id="cite_ref-Pitcher_365_1-1" class="reference"><a href="#cite_note-Pitcher_365-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> About one quarter of fish species shoal all their lives, and about one half shoal for part of their lives.<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> </p><p>Fish derive many benefits from shoaling behaviour including defence against predators (through better predator detection and by diluting the chance of individual capture), enhanced <a href="/wiki/Foraging" title="Foraging">foraging</a> success, and higher success in finding a mate. It is also likely that fish benefit from shoal membership through increased <a href="/wiki/Hydrodynamic" class="mw-redirect" title="Hydrodynamic">hydrodynamic</a> efficiency. </p><p>Fish use many traits to choose shoalmates. Generally they prefer larger shoals, shoalmates of their own species, shoalmates similar in size and appearance to themselves, healthy fish, and kin (when recognized). </p><p>The <b>oddity effect</b> posits that any shoal member that stands out in appearance will be preferentially targeted by predators. This may explain why fish prefer to shoal with individuals that resemble themselves. The oddity effect thus tends to homogenize shoals. </p> <meta property="mw:PageProp/toc" /> <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=Shoaling_and_schooling&action=edit&section=1" title="Edit section: Overview"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1251242444">.mw-parser-output .ambox{border:1px solid #a2a9b1;border-left:10px solid #36c;background-color:#fbfbfb;box-sizing:border-box}.mw-parser-output .ambox+link+.ambox,.mw-parser-output .ambox+link+style+.ambox,.mw-parser-output .ambox+link+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+style+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+link+.ambox{margin-top:-1px}html body.mediawiki .mw-parser-output .ambox.mbox-small-left{margin:4px 1em 4px 0;overflow:hidden;width:238px;border-collapse:collapse;font-size:88%;line-height:1.25em}.mw-parser-output .ambox-speedy{border-left:10px solid #b32424;background-color:#fee7e6}.mw-parser-output .ambox-delete{border-left:10px solid #b32424}.mw-parser-output .ambox-content{border-left:10px solid #f28500}.mw-parser-output .ambox-style{border-left:10px solid #fc3}.mw-parser-output .ambox-move{border-left:10px solid #9932cc}.mw-parser-output .ambox-protection{border-left:10px solid #a2a9b1}.mw-parser-output .ambox .mbox-text{border:none;padding:0.25em 0.5em;width:100%}.mw-parser-output .ambox .mbox-image{border:none;padding:2px 0 2px 0.5em;text-align:center}.mw-parser-output .ambox .mbox-imageright{border:none;padding:2px 0.5em 2px 0;text-align:center}.mw-parser-output .ambox .mbox-empty-cell{border:none;padding:0;width:1px}.mw-parser-output .ambox .mbox-image-div{width:52px}@media(min-width:720px){.mw-parser-output .ambox{margin:0 10%}}@media print{body.ns-0 .mw-parser-output .ambox{display:none!important}}</style><table class="box-More_citations_needed_section plainlinks metadata ambox ambox-content ambox-Refimprove" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Question_book-new.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>needs additional citations for <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">verification</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Shoaling_and_schooling" title="Special:EditPage/Shoaling and schooling">improve this article</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a> in this section. Unsourced material may be challenged and removed.</span> <span class="date-container"><i>(<span class="date">January 2021</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Heringsschwarm.gif" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4d/Heringsschwarm.gif/220px-Heringsschwarm.gif" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/4/4d/Heringsschwarm.gif 1.5x" data-file-width="320" data-file-height="240" /></a><figcaption>Underwater video loop of a school of <a href="/wiki/Herring" title="Herring">herrings</a> migrating at high speed to their spawning grounds in the <a href="/wiki/Baltic_Sea" title="Baltic Sea">Baltic Sea</a></figcaption></figure> <p>An <b>aggregation</b> of fish is the general term for any collection of fish that have gathered together in some locality. Fish aggregations can be structured or unstructured. An unstructured aggregation might be a group of mixed species and sizes that have gathered randomly near some local resource, such as food or nesting sites. </p><p>If, in addition, the aggregation comes together in an interactive, social way, they may be said to be <b>shoaling</b>.<sup id="cite_ref-Pitcher_365_1-2" class="reference"><a href="#cite_note-Pitcher_365-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>a<span class="cite-bracket">]</span></a></sup> Although shoaling fish can relate to each other in a loose way, with each fish swimming and foraging somewhat independently, they are nonetheless aware of the other members of the group as shown by the way they adjust behaviour such as swimming, so as to remain close to the other fish in the group. Shoaling groups can include fish of disparate sizes and can include mixed-species subgroups. </p><p>If the shoal becomes more tightly organised, with the fish synchronising their swimming so they all move at the same speed and in the same direction, then the fish may be said to be <b>schooling</b>.<sup id="cite_ref-Pitcher_365_1-3" class="reference"><a href="#cite_note-Pitcher_365-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>b<span class="cite-bracket">]</span></a></sup> Schooling fish are usually of the same species and the same age/size. Fish schools move with the individual members precisely spaced from each other. The schools undertake complicated manoeuvres, as though the schools have minds of their own.<sup id="cite_ref-Moyle_and_Cech_6-0" class="reference"><a href="#cite_note-Moyle_and_Cech-6"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><p>The intricacies of schooling are far from fully understood, especially the swimming and feeding energetics. Many hypotheses to explain the function of schooling have been suggested, such as better orientation, <a href="/wiki/Synchronization" title="Synchronization">synchronized</a> hunting, predator confusion and reduced risk of being found. Schooling also has disadvantages, such as excretion buildup in the breathing media and oxygen and food depletion. The way the fish array in the school probably gives energy saving advantages, though this is controversial.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Great_Barracuda_off_the_Netherland_Antilles.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/81/Great_Barracuda_off_the_Netherland_Antilles.jpg/220px-Great_Barracuda_off_the_Netherland_Antilles.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/81/Great_Barracuda_off_the_Netherland_Antilles.jpg/330px-Great_Barracuda_off_the_Netherland_Antilles.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/81/Great_Barracuda_off_the_Netherland_Antilles.jpg/440px-Great_Barracuda_off_the_Netherland_Antilles.jpg 2x" data-file-width="600" data-file-height="450" /></a><figcaption>Schools of <a href="/wiki/Forage_fish" title="Forage fish">forage fish</a> often accompany large predator fish. Here a school of <a href="/wiki/Bar_jack" title="Bar jack">jacks</a> accompany a <a href="/wiki/Great_barracuda" title="Great barracuda">great barracuda</a>.</figcaption></figure> <p>Fish can be <a href="/wiki/Obligate" title="Obligate">obligate</a> or facultative (optional) shoalers.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Obligate shoalers, such as <a href="/wiki/Tuna" title="Tuna">tunas</a>, <a href="/wiki/Herring" title="Herring">herrings</a> and <a href="/wiki/Anchovy" title="Anchovy">anchovy</a>, spend all of their time shoaling or schooling, and become agitated if separated from the group. Facultative shoalers, such as <a href="/wiki/Atlantic_cod" title="Atlantic cod">Atlantic cod</a>, <a href="/wiki/Pollachius_virens" title="Pollachius virens">saiths</a> and some <a href="/wiki/Carangid" class="mw-redirect" title="Carangid">carangids</a>, shoal only some of the time, perhaps for reproductive purposes.<sup id="cite_ref-Partridge1980_9-0" class="reference"><a href="#cite_note-Partridge1980-9"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p>Shoaling fish can shift into a disciplined and coordinated school, then shift back to an amorphous shoal within seconds. Such shifts are triggered by changes of activity from feeding, resting, travelling or avoiding predators.<sup id="cite_ref-Moyle_and_Cech_6-1" class="reference"><a href="#cite_note-Moyle_and_Cech-6"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><p>When schooling fish stop to feed, they break ranks and become shoals. Shoals are more vulnerable to predator attack. The shape a shoal or school takes depends on the type of fish and what the fish are doing. Schools that are travelling can form long thin lines, or squares or ovals or amoeboid shapes. Fast moving schools usually form a wedge shape, while shoals that are feeding tend to become circular.<sup id="cite_ref-Moyle_and_Cech_6-2" class="reference"><a href="#cite_note-Moyle_and_Cech-6"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Forage_fish" title="Forage fish">Forage fish</a> are small fish which are preyed on by larger predators for food. Predators include other larger fish, <a href="/wiki/Seabird" title="Seabird">seabirds</a> and <a href="/wiki/Marine_mammal" title="Marine mammal">marine mammals</a>. Typical ocean forage fish are small, <a href="/wiki/Filter_feeder" title="Filter feeder">filter-feeding</a> fish such as <a href="/wiki/Herring" title="Herring">herring</a>, <a href="/wiki/Anchovies" class="mw-redirect" title="Anchovies">anchovies</a> and <a href="/wiki/Menhaden" title="Menhaden">menhaden</a>. Forage fish compensate for their small size by forming schools. Some swim in synchronised grids with their mouths open so they can efficiently filter feed on <a href="/wiki/Plankton" title="Plankton">plankton</a>.<sup id="cite_ref-Kils,_1992_10-0" class="reference"><a href="#cite_note-Kils,_1992-10"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> These schools can become huge, moving along coastlines and <a href="/wiki/Fish_migration" title="Fish migration">migrating</a> across open oceans. The shoals are concentrated food resources for the great marine predators. </p> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:School_jacks_klein.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/School_jacks_klein.JPG/220px-School_jacks_klein.JPG" decoding="async" width="220" height="146" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/School_jacks_klein.JPG/330px-School_jacks_klein.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b1/School_jacks_klein.JPG/440px-School_jacks_klein.JPG 2x" data-file-width="800" data-file-height="531" /></a><figcaption>A school of fish has many eyes that can scan for food or threats</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><span><video id="mwe_player_0" poster="//upload.wikimedia.org/wikipedia/commons/thumb/d/d1/Sardines_reacting_to_tuna_-_MBA.webm/220px--Sardines_reacting_to_tuna_-_MBA.webm.jpg" controls="" preload="none" data-mw-tmh="" class="mw-file-element" width="220" height="124" data-durationhint="30" data-mwtitle="Sardines_reacting_to_tuna_-_MBA.webm" data-mwprovider="wikimediacommons" resource="/wiki/File:Sardines_reacting_to_tuna_-_MBA.webm"><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/d/d1/Sardines_reacting_to_tuna_-_MBA.webm/Sardines_reacting_to_tuna_-_MBA.webm.480p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="480p.vp9.webm" data-width="854" data-height="480" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/d/d1/Sardines_reacting_to_tuna_-_MBA.webm/Sardines_reacting_to_tuna_-_MBA.webm.720p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="720p.vp9.webm" data-width="1280" data-height="720" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/d/d1/Sardines_reacting_to_tuna_-_MBA.webm/Sardines_reacting_to_tuna_-_MBA.webm.1080p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="1080p.vp9.webm" data-width="1920" data-height="1080" /><source src="//upload.wikimedia.org/wikipedia/commons/d/d1/Sardines_reacting_to_tuna_-_MBA.webm" type="video/webm; codecs="vp9, vorbis"" data-width="1920" data-height="1080" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/d/d1/Sardines_reacting_to_tuna_-_MBA.webm/Sardines_reacting_to_tuna_-_MBA.webm.240p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="240p.vp9.webm" data-width="426" data-height="240" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/d/d1/Sardines_reacting_to_tuna_-_MBA.webm/Sardines_reacting_to_tuna_-_MBA.webm.360p.webm" type="video/webm; codecs="vp8, vorbis"" data-transcodekey="360p.webm" data-width="640" data-height="360" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/d/d1/Sardines_reacting_to_tuna_-_MBA.webm/Sardines_reacting_to_tuna_-_MBA.webm.360p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="360p.vp9.webm" data-width="640" data-height="360" /></video></span><figcaption><a href="/wiki/Pacific_sardine" class="mw-redirect" title="Pacific sardine">Pacific sardine</a> school reacting to attention from <a href="/wiki/Yellowfin_tuna" title="Yellowfin tuna">yellowfin tuna</a></figcaption></figure> <p>These sometimes immense gatherings fuel the <a href="/wiki/Ocean_food_web" class="mw-redirect" title="Ocean food web">ocean food web</a>. Most forage fish are <a href="/wiki/Pelagic_fish" title="Pelagic fish">pelagic fish</a>, which means they form their schools in open water, and not on or near the bottom (<a href="/wiki/Demersal_fish" title="Demersal fish">demersal fish</a>). Forage fish are short-lived, and go mostly unnoticed by humans. The predators are keenly focused on the shoals, acutely aware of their numbers and whereabouts, and make <a href="/wiki/Fish_migration" title="Fish migration">migrations</a> themselves, often in schools of their own, that can span thousands of miles to connect with, or stay connected with them.<sup id="cite_ref-NCMC_11-0" class="reference"><a href="#cite_note-NCMC-11"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Herring" title="Herring">Herring</a> are among the more spectacular schooling fish. They aggregate together in huge numbers. The largest schools are often formed during migrations by merging with smaller schools. "Chains" of schools one hundred kilometres (60 miles) long have been observed of <a href="/wiki/Mugil" title="Mugil">mullet</a> migrating in the <a href="/wiki/Caspian_Sea" title="Caspian Sea">Caspian Sea</a>. Radakov estimated herring schools in the North Atlantic can occupy up to 4.8 cubic kilometres (1.2 cubic miles) with fish densities between 0.5 and 1.0 fish/cubic metre (<style data-mw-deduplicate="TemplateStyles:r1154941027">.mw-parser-output .frac{white-space:nowrap}.mw-parser-output .frac .num,.mw-parser-output .frac .den{font-size:80%;line-height:0;vertical-align:super}.mw-parser-output .frac .den{vertical-align:sub}.mw-parser-output .sr-only{border:0;clip:rect(0,0,0,0);clip-path:polygon(0px 0px,0px 0px,0px 0px);height:1px;margin:-1px;overflow:hidden;padding:0;position:absolute;width:1px}</style><span class="frac"><span class="num">3</span>⁄<span class="den">8</span></span> to <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1154941027"><span class="frac"><span class="num">3</span>⁄<span class="den">4</span></span> fish per cubic yard), totalling about three billion fish in a single school.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> These schools move along coastlines and traverse the open oceans. Herring schools in general have very precise arrangements which allow the school to maintain relatively constant cruising speeds. Herrings have excellent hearing, and their schools react very rapidly to a predator. The herrings keep a certain distance from a moving scuba diver or a cruising predator like a killer whale, forming a vacuole which looks like a doughnut from a spotter plane.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p>Many species of large predatory fish also school, including many <a href="/wiki/Highly_migratory_fish" class="mw-redirect" title="Highly migratory fish">highly migratory fish</a>, such as <a href="/wiki/Tuna" title="Tuna">tuna</a> and some oceangoing <a href="/wiki/Shark" title="Shark">sharks</a>. <a href="/wiki/Cetacean" class="mw-redirect" title="Cetacean">Cetaceans</a> such as dolphins, porpoises and whales, operate in organised social groups called <i>pods</i>. </p><p>"Shoaling behaviour is generally described as a trade-off between the anti-predator benefits of living in groups and the costs of increased foraging competition."<sup id="cite_ref-Hoare_et_al._2000_14-0" class="reference"><a href="#cite_note-Hoare_et_al._2000-14"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Landa (1998) argues that the cumulative advantages of shoaling, as elaborated below, are strong selective inducements for fish to join shoals.<sup id="cite_ref-Landa_15-0" class="reference"><a href="#cite_note-Landa-15"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Parrish <i>et al.</i> (2002) argue similarly that schooling is a classic example of <a href="/wiki/Emergence" title="Emergence">emergence</a>, where there are properties that are possessed by the school but not by the individual fish. Emergent properties give an evolutionary advantage to members of the school which non members do not receive.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Social_interaction">Social interaction</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=2" title="Edit section: Social interaction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Support for the social and genetic function of aggregations, especially those formed by fish, can be seen in several aspects of their behaviour. For instance, experiments have shown that individual fish removed from a school will have a higher respiratory rate than those found in the school.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> This effect has been attributed to stress, and the effect of being with conspecifics therefore appears to be a calming one and a powerful social motivation for remaining in an aggregation.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Herring, for instance, will become very agitated if they are isolated from conspecifics.<sup id="cite_ref-Partridge1980_9-1" class="reference"><a href="#cite_note-Partridge1980-9"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Because of their adaptation to schooling behaviour they are rarely displayed in <a href="/wiki/Aquarium" title="Aquarium">aquaria</a>. Even with the best facilities aquaria can offer they become fragile and sluggish compared to their quivering energy in wild schools.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Foraging_advantages">Foraging advantages</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=3" title="Edit section: Foraging advantages"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Upwelling_image1.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/Upwelling_image1.jpg/300px-Upwelling_image1.jpg" decoding="async" width="300" height="197" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/Upwelling_image1.jpg/450px-Upwelling_image1.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/f/fd/Upwelling_image1.jpg 2x" data-file-width="514" data-file-height="337" /></a><figcaption>Some coastal <a href="/wiki/Upwelling" title="Upwelling">upwellings</a> (red) provide plankton-rich feeding grounds for shoals of <a href="/wiki/Forage_fish" title="Forage fish">forage fish</a>, which in turn attract larger predator fish.</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Forage_fish#Hunting_copepods" title="Forage fish">Forage fish § Hunting copepods</a></div> <p>It has also been proposed that swimming in groups enhances foraging success. This ability was demonstrated by Pitcher and others in their study of foraging behaviour in shoaling <a href="/wiki/Cyprinid" class="mw-redirect" title="Cyprinid">cyprinids</a>.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> In this study, the time it took for groups of <a href="/wiki/Common_minnow" title="Common minnow">minnows</a> and goldfish to find a patch of food was quantified. The number of fishes in the groups was varied, and a statistically significant decrease in the amount of time necessary for larger groups to find food was established. Further support for an enhanced foraging capability of schools is seen in the structure of schools of predatory fish. Partridge and others analysed the school structure of Atlantic bluefin tuna from aerial photographs and found that the school assumed a parabolic shape, a fact that was suggestive of cooperative hunting in this species.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p><p>"The reason for this is the presence of many eyes searching for the food. Fish in shoals "share" information by monitoring each other's behaviour closely. Feeding behaviour in one fish quickly stimulates food-searching behaviour in others.<sup id="cite_ref-Pitcher_1993_21-0" class="reference"><a href="#cite_note-Pitcher_1993-21"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p><p>Fertile feeding grounds for forage fish are provided by ocean upwellings. <a href="/wiki/Oceanic_gyre" class="mw-redirect" title="Oceanic gyre">Oceanic gyres</a> are large-scale <a href="/wiki/Ocean_currents" class="mw-redirect" title="Ocean currents">ocean currents</a> caused by the <a href="/wiki/Coriolis_effect" class="mw-redirect" title="Coriolis effect">Coriolis effect</a>. Wind-driven surface currents interact with these gyres and the underwater topography, such as <a href="/wiki/Seamount" title="Seamount">seamounts</a>, <a href="/wiki/Fishing_bank" class="mw-redirect" title="Fishing bank">fishing banks</a>, and the edge of <a href="/wiki/Continental_shelf" title="Continental shelf">continental shelves</a>, to produce downwellings and <a href="/wiki/Upwelling" title="Upwelling">upwellings</a>.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> These can transport nutrients which plankton thrive on. The result can be rich feeding grounds attractive to the plankton feeding forage fish. In turn, the forage fish themselves become a feeding ground for larger predator fish. Most upwellings are coastal, and many of them support some of the most productive fisheries in the world. Regions of notable upwelling include coastal <a href="/wiki/Peru" title="Peru">Peru</a>, <a href="/wiki/Chile" title="Chile">Chile</a>, <a href="/wiki/Arabian_Sea" title="Arabian Sea">Arabian Sea</a>, western <a href="/wiki/South_Africa" title="South Africa">South Africa</a>, eastern <a href="/wiki/New_Zealand" title="New Zealand">New Zealand</a> and the <a href="/wiki/California" title="California">California</a> coast.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p><p><a href="/wiki/Copepod" title="Copepod">Copepods</a>, the primary <a href="/wiki/Zooplankton" title="Zooplankton">zooplankton</a>, are a major item on the forage fish menu. They are a group of small <a href="/wiki/Crustacean" title="Crustacean">crustaceans</a> found in ocean and <a href="/wiki/Fresh_water" title="Fresh water">freshwater</a> <a href="/wiki/Habitat_(ecology)" class="mw-redirect" title="Habitat (ecology)">habitats</a>. Copepods are typically one millimetre (0.04 in) to two millimetres (0.08 in) long, with a teardrop shaped body. Some scientists say they form the largest animal <a href="/wiki/Biomass" title="Biomass">biomass</a> on the planet.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> Copepods are very alert and evasive. They have large <a href="/wiki/Antenna_(biology)" title="Antenna (biology)">antennae</a> (see photo below left). When they spread their antennae they can sense the pressure wave from an approaching fish and jump with great speed over a few centimeters. If copepod concentrations reach high levels, schooling herrings adopt a method called <a href="/wiki/Ram_feeding" class="mw-redirect" title="Ram feeding">ram feeding</a>. In the photo below, herring ram feed on a school of <a href="/wiki/Copepod" title="Copepod">copepods</a>. They swim with their mouth wide open and their <a href="/wiki/Operculum_(fish)" title="Operculum (fish)">opercula</a> fully expanded.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p> <ul class="gallery mw-gallery-traditional"> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Copepodkils.jpg" class="mw-file-description" title="This copepod has its antenna spread (click to enlarge). The antenna detects the pressure wave of an approaching fish."><img alt="This copepod has its antenna spread (click to enlarge). The antenna detects the pressure wave of an approaching fish." src="//upload.wikimedia.org/wikipedia/commons/thumb/2/28/Copepodkils.jpg/160px-Copepodkils.jpg" decoding="async" width="160" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/28/Copepodkils.jpg/240px-Copepodkils.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/28/Copepodkils.jpg/320px-Copepodkils.jpg 2x" data-file-width="2000" data-file-height="1500" /></a></span></div> <div class="gallerytext">This <a href="/wiki/Copepod" title="Copepod">copepod</a> has its antenna spread (click to enlarge). The antenna detects the pressure wave of an approaching fish.</div> </li> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Herring2.jpg" class="mw-file-description" title="Copepods are a major food source for forage fish like this Atlantic herring."><img alt="Copepods are a major food source for forage fish like this Atlantic herring." src="//upload.wikimedia.org/wikipedia/commons/thumb/2/25/Herring2.jpg/160px-Herring2.jpg" decoding="async" width="160" height="61" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/2/25/Herring2.jpg 1.5x" data-file-width="211" data-file-height="81" /></a></span></div> <div class="gallerytext">Copepods are a major food source for forage fish like this <a href="/wiki/Atlantic_herring" title="Atlantic herring">Atlantic herring</a>.</div> </li> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Herringramkils.jpg" class="mw-file-description" title="School of herrings ram-feeding on a school of copepods, with opercula expanded so their red gills are visible"><img alt="School of herrings ram-feeding on a school of copepods, with opercula expanded so their red gills are visible" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/98/Herringramkils.jpg/160px-Herringramkils.jpg" decoding="async" width="160" height="101" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/98/Herringramkils.jpg/240px-Herringramkils.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/98/Herringramkils.jpg/320px-Herringramkils.jpg 2x" data-file-width="793" data-file-height="500" /></a></span></div> <div class="gallerytext">School of herrings ram-feeding on a school of copepods, with <a href="/wiki/Operculum_(fish)" title="Operculum (fish)">opercula</a> expanded so their red gills are visible</div> </li> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Synchropredation.gif" class="mw-file-description" title="Animation showing how herrings hunting in a synchronised way can capture the very alert and evasive copepod"><img alt="Animation showing how herrings hunting in a synchronised way can capture the very alert and evasive copepod" src="//upload.wikimedia.org/wikipedia/commons/thumb/5/59/Synchropredation.gif/160px-Synchropredation.gif" decoding="async" width="160" height="100" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/59/Synchropredation.gif/240px-Synchropredation.gif 1.5x, //upload.wikimedia.org/wikipedia/commons/5/59/Synchropredation.gif 2x" data-file-width="320" data-file-height="200" /></a></span></div> <div class="gallerytext">Animation showing how herrings hunting in a synchronised way can capture the very alert and evasive copepod</div> </li> </ul> <p>The fish swim in a grid where the distance between them is the same as the jump length of their prey, as indicated in the animation above right. In the animation, juvenile herring hunt the <a href="/wiki/Copepod" title="Copepod">copepods</a> in this synchronised way. The copepods sense with their <a href="/wiki/Antenna_(biology)" title="Antenna (biology)">antennae</a> the pressure-wave of an approaching herring and react with a fast escape jump. The length of the jump is fairly constant. The fish align themselves in a grid with this characteristic jump length. A copepod can dart about 80 times before it tires. After a jump, it takes it 60 milliseconds to spread its antennae again, and this time delay becomes its undoing, as the almost endless stream of herrings allows a herring to eventually snap the copepod. A single juvenile herring could never catch a large copepod.<sup id="cite_ref-Kils,_1992_10-1" class="reference"><a href="#cite_note-Kils,_1992-10"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <div style="clear:both;" class=""></div> <div class="mw-heading mw-heading2"><h2 id="Reproductive_advantages">Reproductive advantages</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=4" title="Edit section: Reproductive advantages"><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/Fish_migration" title="Fish migration">Fish migration</a></div> <p>A third proposed benefit of fish groups is that they serve a reproductive function. They provide increased access to potential mates, since finding a mate in a shoal does not take much energy. And for migrating fish that navigate long distances to spawn, it is likely that the navigation of the shoal, with an input from all the shoal members, will be better than that taken by an individual fish.<sup id="cite_ref-Moyle_and_Cech_6-3" class="reference"><a href="#cite_note-Moyle_and_Cech-6"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources"><span title="This citation requires a reference to the specific page or range of pages in which the material appears. (March 2024)">page needed</span></a></i>]</sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Capelin-iceland.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Capelin-iceland.svg/220px-Capelin-iceland.svg.png" decoding="async" width="220" height="217" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Capelin-iceland.svg/330px-Capelin-iceland.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Capelin-iceland.svg/440px-Capelin-iceland.svg.png 2x" data-file-width="509" data-file-height="503" /></a><figcaption>Migration of Icelandic capelin</figcaption></figure> <p>Forage fish often make great <a href="/wiki/Fish_migration" title="Fish migration">migrations</a> between their spawning, feeding and nursery grounds. Schools of a particular stock usually travel in a triangle between these grounds. For example, one stock of herrings have their spawning ground in southern <a href="/wiki/Norway" title="Norway">Norway</a>, their feeding ground in <a href="/wiki/Iceland" title="Iceland">Iceland</a>, and their nursery ground in northern Norway. Wide triangular journeys such as these may be important because forage fish, when feeding, cannot distinguish their own offspring.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p><p><a href="/wiki/Capelin" title="Capelin">Capelin</a> are a forage fish of the <a href="/wiki/Smelt_(fish)" title="Smelt (fish)">smelt</a> family found in the <a href="/wiki/Atlantic_Ocean" title="Atlantic Ocean">Atlantic</a> and <a href="/wiki/Arctic_Ocean" title="Arctic Ocean">Arctic</a> oceans. In summer, they graze on dense swarms of <a href="/wiki/Plankton" title="Plankton">plankton</a> at the edge of the ice shelf. Larger capelin also eat <a href="/wiki/Krill" title="Krill">krill</a> and other <a href="/wiki/Crustacean" title="Crustacean">crustaceans</a>. The capelin move inshore in large schools to spawn and migrate in spring and summer to feed in plankton rich areas between <a href="/wiki/Iceland" title="Iceland">Iceland</a>, <a href="/wiki/Greenland" title="Greenland">Greenland</a>, and <a href="/wiki/Jan_Mayen" title="Jan Mayen">Jan Mayen</a>. The migration is affected by <a href="/wiki/Ocean_current" title="Ocean current">ocean currents</a>. Around Iceland maturing capelin make large northward feeding migrations in spring and summer. The return migration takes place in September to November. The spawning migration starts north of Iceland in December or January.<sup id="cite_ref-Gjøsæter_1998_24-0" class="reference"><a href="#cite_note-Gjøsæter_1998-24"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Hydrodynamic_efficiency">Hydrodynamic efficiency</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=5" title="Edit section: Hydrodynamic efficiency"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>This theory states that groups of fish may save energy when swimming together, much in the way that bicyclists may <a href="/wiki/Drafting_(racing)" class="mw-redirect" title="Drafting (racing)">draft</a> one another in a <a href="/wiki/Peloton" title="Peloton">peloton</a>. Geese flying in a Vee formation are also thought to save energy by flying in the updraft of the <a href="/wiki/Wingtip_vortices" title="Wingtip vortices">wingtip vortex</a> generated by the previous animal in the formation.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> Increased efficiencies in swimming in groups have been proposed for schools of fish and <a href="/wiki/Antarctic_krill" title="Antarctic krill">Antarctic krill</a>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p><p>It would seem reasonable to think that the regular spacing and size uniformity of fish in schools would result in hydrodynamic efficiencies.<sup id="cite_ref-Hoare_et_al._2000_14-1" class="reference"><a href="#cite_note-Hoare_et_al._2000-14"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> While early laboratory-based experiments failed to detect hydrodynamic benefits created by the neighbours of a fish in a school,<sup id="cite_ref-Pitcher_1993_21-1" class="reference"><a href="#cite_note-Pitcher_1993-21"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> it is thought that efficiency gains do occur in the wild. More recent experiments with groups of fish swimming in flumes support this, with fish reducing their swimming costs by as much as 20% as compared to when the same fish are swimming in isolation.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Landa (1998) argued that the leader of a school constantly changes, because while being in the body of a school gives a hydrodynamic advantage, the leader will be the first to the food.<sup id="cite_ref-Landa_15-1" class="reference"><a href="#cite_note-Landa-15"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> More recent work suggests that, after individuals at the front of the school encounter and ingest more food, they then relocate further back within the school due to the locomotor constraints generated during meal digestion.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Predator_avoidance">Predator avoidance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=6" title="Edit section: Predator avoidance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Moofushi_Kandu_fish.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/32/Moofushi_Kandu_fish.jpg/300px-Moofushi_Kandu_fish.jpg" decoding="async" width="300" height="225" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/32/Moofushi_Kandu_fish.jpg/450px-Moofushi_Kandu_fish.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/32/Moofushi_Kandu_fish.jpg/600px-Moofushi_Kandu_fish.jpg 2x" data-file-width="1632" data-file-height="1224" /></a><figcaption>Schooling predator <a href="/wiki/Bluefin_trevally" title="Bluefin trevally">bluefin trevally</a> size up schooling <a href="/wiki/Anchovy" title="Anchovy">anchovies</a></figcaption></figure> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Red_Fish_at_Papah%C4%81naumoku%C4%81kea_(cropped).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Red_Fish_at_Papah%C4%81naumoku%C4%81kea_%28cropped%29.jpg/300px-Red_Fish_at_Papah%C4%81naumoku%C4%81kea_%28cropped%29.jpg" decoding="async" width="300" height="256" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Red_Fish_at_Papah%C4%81naumoku%C4%81kea_%28cropped%29.jpg/450px-Red_Fish_at_Papah%C4%81naumoku%C4%81kea_%28cropped%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Red_Fish_at_Papah%C4%81naumoku%C4%81kea_%28cropped%29.jpg/600px-Red_Fish_at_Papah%C4%81naumoku%C4%81kea_%28cropped%29.jpg 2x" data-file-width="2395" data-file-height="2043" /></a><figcaption>Many eyes provide a higher level of vigilance</figcaption></figure> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Schooling_response_time_in_face_of_predator.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3b/Schooling_response_time_in_face_of_predator.png/300px-Schooling_response_time_in_face_of_predator.png" decoding="async" width="300" height="199" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3b/Schooling_response_time_in_face_of_predator.png/450px-Schooling_response_time_in_face_of_predator.png 1.5x, //upload.wikimedia.org/wikipedia/commons/3/3b/Schooling_response_time_in_face_of_predator.png 2x" data-file-width="559" data-file-height="370" /></a><figcaption>Schooling response time in face of predator<sup id="cite_ref-Aparna2011_29-0" class="reference"><a href="#cite_note-Aparna2011-29"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>It is commonly observed that schooling fish are particularly in danger of being eaten if they are separated from the school.<sup id="cite_ref-Moyle_and_Cech_6-4" class="reference"><a href="#cite_note-Moyle_and_Cech-6"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Several <a href="/wiki/Antipredator_adaptations" class="mw-redirect" title="Antipredator adaptations">anti-predator functions</a> of fish schools have been proposed. </p><p>One potential method by which fish schools might thwart <a href="/wiki/Predator" class="mw-redirect" title="Predator">predators</a> is the "predator confusion effect" proposed and demonstrated by Milinski and Heller (1978).<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> This theory is based on the idea that it becomes difficult for predators to choose individual prey from groups because the many moving targets create a sensory overload of the predator's visual channel. Milinski and Heller's findings have been corroborated both in experiment<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> and computer simulations.<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Olson_RS,_Hintze_A,_Dyer_FC,_Knoester_DB,_Adami_C_2013_20130305_35-0" class="reference"><a href="#cite_note-Olson_RS,_Hintze_A,_Dyer_FC,_Knoester_DB,_Adami_C_2013_20130305-35"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> "Shoaling fish are the same size and silvery, so it is difficult for a visually oriented predator to pick an individual out of a mass of twisting, flashing fish and then have enough time to grab its prey before it disappears into the shoal."<sup id="cite_ref-Moyle_and_Cech_6-5" class="reference"><a href="#cite_note-Moyle_and_Cech-6"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><p>Schooling behaviour confuses the <a href="/wiki/Lateral_line" title="Lateral line">lateral line organ (LLO)</a> as well as the <a href="/wiki/Electrosensory" class="mw-redirect" title="Electrosensory">electrosensory</a> system (ESS) of predators.<sup id="cite_ref-Larsson2009_36-0" class="reference"><a href="#cite_note-Larsson2009-36"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Larsson2012_38-0" class="reference"><a href="#cite_note-Larsson2012-38"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> Fin movements of a single fish act as a point-shaped wave source, emitting a gradient by which predators might localize it. Since fields of many fish will overlap, schooling should obscure this gradient, perhaps mimicking pressure waves of a larger animal, and more likely confuse the lateral line perception.<sup id="cite_ref-Larsson2009_36-1" class="reference"><a href="#cite_note-Larsson2009-36"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> The LLO is essential in the final stages of a predator attack.<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> Electro-receptive animals may localize a field source by using spatial non-uniformities. To produce separate signals, individual prey must be about five body widths apart. If objects are too close together to be distinguished, they will form a blurred image.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> Based on this it was suggested that schooling may confuse the ESS of predators.<sup id="cite_ref-Larsson2009_36-2" class="reference"><a href="#cite_note-Larsson2009-36"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p><p>A third potential anti-predator effect of animal aggregations is the "many eyes" hypothesis. This theory states that as the size of the group increases, the task of scanning the environment for predators can be spread out over many individuals. Not only does this <a href="/wiki/Mass_Collaboration" class="mw-redirect" title="Mass Collaboration">mass collaboration</a> presumably provide a higher level of vigilance, it could also allow more time for individual feeding.<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> </p><p>A fourth hypothesis for an anti-predatory effect of fish schools is the "encounter dilution" effect. The dilution effect is an elaboration of <a href="/wiki/Safety_in_numbers" title="Safety in numbers">safety in numbers</a>, and interacts with the confusion effect.<sup id="cite_ref-Pitcher_1993_21-2" class="reference"><a href="#cite_note-Pitcher_1993-21"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> A given predator attack will eat a smaller proportion of a large shoal than a small shoal.<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> Hamilton proposed that animals aggregate because of a "selfish" avoidance of a predator and was thus a form of cover-seeking.<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> Another formulation of the theory was given by Turner and Pitcher and was viewed as a combination of detection and attack probabilities.<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> In the detection component of the theory, it was suggested that potential prey might benefit by living together since a predator is less likely to chance upon a single group than a scattered distribution. In the attack component, it was thought that an attacking predator is less likely to eat a particular fish when a greater number of fish are present. In sum, a fish has an advantage if it is in the larger of two groups, assuming that the probability of detection and attack does not increase disproportionately with the size of the group.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p><p>Schooling forage fish are subject to constant attacks by predators. An example is the attacks that take place during the African <a href="/wiki/Sardine_run" title="Sardine run">sardine run</a>. The African sardine run is a spectacular migration by millions of silvery sardines along the southern <a href="/wiki/Coastline" class="mw-redirect" title="Coastline">coastline</a> of Africa. In terms of biomass, the sardine run could rival East Africa's great <a href="/wiki/Blue_Wildebeest" class="mw-redirect" title="Blue Wildebeest">wildebeest migration</a>.<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Sardine" title="Sardine">Sardines</a> have a short life-cycle, living only two or three years. Adult sardines, about two years old, mass on the <a href="/wiki/Agulhas_Bank" title="Agulhas Bank">Agulhas Bank</a> where they spawn during spring and summer, releasing tens of thousands of eggs into the water. The adult sardines then make their way in hundreds of shoals towards the sub-tropical waters of the <a href="/wiki/Indian_Ocean" title="Indian Ocean">Indian Ocean</a>. A larger shoal might be 7 kilometres (4.3 mi) long, 1.5 kilometres (0.93 mi) wide and 30 metres (98 ft) deep. Huge numbers of sharks, dolphins, tuna, sailfish, <a href="/wiki/Cape_fur_seal" class="mw-redirect" title="Cape fur seal">Cape fur seals</a> and even killer whales congregate and follow the shoals, creating a <a href="/wiki/Feeding_frenzy" title="Feeding frenzy">feeding frenzy</a> along the coastline.<sup id="cite_ref-baitball_48-0" class="reference"><a href="#cite_note-baitball-48"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> </p> <div style="clear:both;" class=""></div> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:A_tornado_of_fish.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0d/A_tornado_of_fish.jpg/300px-A_tornado_of_fish.jpg" decoding="async" width="300" height="225" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0d/A_tornado_of_fish.jpg/450px-A_tornado_of_fish.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0d/A_tornado_of_fish.jpg/600px-A_tornado_of_fish.jpg 2x" data-file-width="1024" data-file-height="768" /></a><figcaption>A big <a href="/wiki/Bait_ball" title="Bait ball">bait ball</a> swirling around a loose kelp stipe</figcaption></figure> <p>When threatened, sardines (and other forage fish) instinctively group together and create massive <a href="/wiki/Bait_ball" title="Bait ball">bait balls</a>. Bait balls can be up to 20 metres (66 ft) in diameter. They are short lived, seldom lasting longer than 20 minutes. The fish eggs, left behind at the Agulhas Banks, drift north west with the current into waters off the west coast, where the larvae develop into juvenile fish. When they are old enough, they aggregate into dense shoals and migrate southwards, returning to the Agulhas banks to restart the cycle.<sup id="cite_ref-baitball_48-1" class="reference"><a href="#cite_note-baitball-48"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> </p><p>The development of schooling behavior was probably associated with an increased quality of perception, predatory lifestyle and size sorting mechanisms to avoid cannibalism.<sup id="cite_ref-Larsson2012_38-1" class="reference"><a href="#cite_note-Larsson2012-38"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> In filter-feeding ancestors, before vision and the <a href="/wiki/Lateral_line" title="Lateral line">octavolateralis system</a> (OLS) had developed, the risk of predation would have been limited and mainly due to invertebrate predators. Hence, at that time, safety in numbers was probably not a major incentive for gathering in shoals or schools. The development of vision and the OLS would have permitted detection of potential prey. This could have led to an increased potential for cannibalism within the shoal. On the other hand, increased quality of perception would give small individuals a chance to escape or to never join a shoal with larger fish. It has been shown that small fish avoid joining a group with larger fish, although big fish do not avoid joining small conspecifics.<sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> This sorting mechanism based on increased quality of perception could have resulted in homogeneity of size of fish in shoals, which would increase the capacity for moving in synchrony.<sup id="cite_ref-Larsson2012_38-2" class="reference"><a href="#cite_note-Larsson2012-38"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Predator_countermeasures">Predator countermeasures</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=7" title="Edit section: Predator countermeasures"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:School_of_blackfin_barracuda_(sphyraena_qenie).JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/45/School_of_blackfin_barracuda_%28sphyraena_qenie%29.JPG/280px-School_of_blackfin_barracuda_%28sphyraena_qenie%29.JPG" decoding="async" width="280" height="298" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/45/School_of_blackfin_barracuda_%28sphyraena_qenie%29.JPG/420px-School_of_blackfin_barracuda_%28sphyraena_qenie%29.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/45/School_of_blackfin_barracuda_%28sphyraena_qenie%29.JPG/560px-School_of_blackfin_barracuda_%28sphyraena_qenie%29.JPG 2x" data-file-width="2609" data-file-height="2781" /></a><figcaption> School of <a href="/wiki/Blackfin_barracuda" title="Blackfin barracuda">blackfin barracuda</a> lined up in attack mode</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Antipredator_adaptations" class="mw-redirect" title="Antipredator adaptations">Antipredator adaptations</a></div> <p>Predators have devised various countermeasures to undermine the defensive shoaling and schooling manoeuvres of forage fish. The <a href="/wiki/Sailfish" title="Sailfish">sailfish</a> raises its sail to make it appear much larger so it can herd a school of fish or squid. <a href="/wiki/Swordfish" title="Swordfish">Swordfish</a> charge at high speed through forage fish schools, slashing with their swords to kill or stun prey. They then turn and return to consume their "catch". <a href="/wiki/Thresher_shark" title="Thresher shark">Thresher sharks</a> use their long tails to stun shoaling fishes. Before striking, the sharks compact schools of prey by swimming around them and splashing the water with their tails, often in pairs or small groups. Threshers swim in circles to drive schooling prey into a compact mass, before striking them sharply with the upper lobe of its tail to stun them.<sup id="cite_ref-seitz_50-0" class="reference"><a href="#cite_note-seitz-50"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Spinner_shark" title="Spinner shark">Spinner sharks</a> charge vertically through the school, spinning on their axis with their mouths open and snapping all around. The shark's momentum at the end of these spiralling runs often carries it into the air.<sup id="cite_ref-compagno_52-0" class="reference"><a href="#cite_note-compagno-52"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-marinebio_53-0" class="reference"><a href="#cite_note-marinebio-53"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> </p> <ul class="gallery mw-gallery-traditional"> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 110px;"><span typeof="mw:File"><a href="/wiki/File:Istiophorus_platypterus.jpg" class="mw-file-description" title="Sailfish herd with their sails."><img alt="Sailfish herd with their sails." src="//upload.wikimedia.org/wikipedia/commons/thumb/5/50/Istiophorus_platypterus.jpg/149px-Istiophorus_platypterus.jpg" decoding="async" width="149" height="80" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/50/Istiophorus_platypterus.jpg/224px-Istiophorus_platypterus.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/50/Istiophorus_platypterus.jpg/298px-Istiophorus_platypterus.jpg 2x" data-file-width="1000" data-file-height="537" /></a></span></div> <div class="gallerytext">Sailfish herd with their sails.</div> </li> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 110px;"><span typeof="mw:File"><a href="/wiki/File:Xiphias_gladius1.jpg" class="mw-file-description" title="Swordfish slash with their swords."><img alt="Swordfish slash with their swords." src="//upload.wikimedia.org/wikipedia/commons/thumb/9/98/Xiphias_gladius1.jpg/160px-Xiphias_gladius1.jpg" decoding="async" width="160" height="59" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/98/Xiphias_gladius1.jpg/240px-Xiphias_gladius1.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/98/Xiphias_gladius1.jpg/320px-Xiphias_gladius1.jpg 2x" data-file-width="700" data-file-height="259" /></a></span></div> <div class="gallerytext">Swordfish slash with their swords.</div> </li> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 110px;"><span typeof="mw:File"><a href="/wiki/File:Alopias_vulpinus.jpg" class="mw-file-description" title="Thresher sharks strike with their tails."><img alt="Thresher sharks strike with their tails." src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Alopias_vulpinus.jpg/145px-Alopias_vulpinus.jpg" decoding="async" width="145" height="80" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Alopias_vulpinus.jpg/217px-Alopias_vulpinus.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Alopias_vulpinus.jpg/290px-Alopias_vulpinus.jpg 2x" data-file-width="1800" data-file-height="996" /></a></span></div> <div class="gallerytext">Thresher sharks strike with their tails.</div> </li> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 110px;"><span typeof="mw:File"><a href="/wiki/File:Carcharhinus_brevipinna.jpg" class="mw-file-description" title="Spinner sharks spin on their long axis."><img alt="Spinner sharks spin on their long axis." src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6d/Carcharhinus_brevipinna.jpg/160px-Carcharhinus_brevipinna.jpg" decoding="async" width="160" height="47" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6d/Carcharhinus_brevipinna.jpg/240px-Carcharhinus_brevipinna.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6d/Carcharhinus_brevipinna.jpg/320px-Carcharhinus_brevipinna.jpg 2x" data-file-width="577" data-file-height="168" /></a></span></div> <div class="gallerytext">Spinner sharks spin on their long axis.</div> </li> </ul> <div style="clear:both;" class=""></div> <ul class="gallery mw-gallery-packed" style="float:right;"> <li class="gallerybox" style="width: 374.66666666667px"> <div class="thumb" style="width: 372.66666666667px;"><span typeof="mw:File"><a href="/wiki/File:Barrier_feeding_in_bottlenose_dolphins.jpg" class="mw-file-description" title="↑  A team of common bottlenose dolphins cooperate to make schooling fish jump in the air. In this vulnerable position the fish are easy prey for the dolphins.[52]"><img alt="↑  A team of common bottlenose dolphins cooperate to make schooling fish jump in the air. In this vulnerable position the fish are easy prey for the dolphins.[52]" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2e/Barrier_feeding_in_bottlenose_dolphins.jpg/559px-Barrier_feeding_in_bottlenose_dolphins.jpg" decoding="async" width="373" height="450" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2e/Barrier_feeding_in_bottlenose_dolphins.jpg/839px-Barrier_feeding_in_bottlenose_dolphins.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2e/Barrier_feeding_in_bottlenose_dolphins.jpg/1119px-Barrier_feeding_in_bottlenose_dolphins.jpg 2x" data-file-width="1446" data-file-height="1745" /></a></span></div> <div class="gallerytext"><div class="center" style="width:auto; margin-left:auto; margin-right:auto;">↑  A team of <a href="/wiki/Common_bottlenose_dolphin" title="Common bottlenose dolphin">common bottlenose dolphins</a> cooperate to make schooling fish jump in the air. In this vulnerable position the fish are easy prey for the dolphins.<sup id="cite_ref-Gazda2005_54-0" class="reference"><a href="#cite_note-Gazda2005-54"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup></div></div> </li> </ul> <p>Some predators, such as dolphins, hunt in groups of their own. One technique employed by many dolphin species is <a href="/wiki/Herding" title="Herding">herding</a>, where a pod will control a school of fish while individual members take turns ploughing through and feeding on the more tightly packed school (a formation commonly known as a <a href="/wiki/Bait_ball" title="Bait ball">bait ball</a>). Corralling is a method where fish are chased to shallow water where they are more easily captured. In <a href="/wiki/South_Carolina" title="South Carolina">South Carolina</a>, the Atlantic <a href="/wiki/Bottlenose_dolphin" title="Bottlenose dolphin">bottlenose dolphin</a> takes this one step further with what has become known as strand feeding, where the fish are driven onto mud banks and retrieved from there.<sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Common_bottlenose_dolphin" title="Common bottlenose dolphin">Common bottlenose dolphins</a> have been observed using another technique. One dolphin acts as a "driver" and herds a school of fish towards several other dolphins who form a barrier. The driver dolphin slaps its fluke which makes the fish leap into the air. As the fish leap, the driver dolphin moves with the barrier dolphins and catches the fish in the air.<sup id="cite_ref-Gazda2005_54-1" class="reference"><a href="#cite_note-Gazda2005-54"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> This type of cooperative role specialization seems to be more common in <a href="/wiki/Marine_animal" class="mw-redirect" title="Marine animal">marine animals</a> than in <a href="/wiki/Terrestrial_animals" class="mw-redirect" title="Terrestrial animals">terrestrial animals</a>, perhaps because the oceans have more variability in prey diversity, <a href="/wiki/Biomass" title="Biomass">biomass</a>, and predator mobility.<sup id="cite_ref-Gazda2005_54-2" class="reference"><a href="#cite_note-Gazda2005-54"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p><p>During the <a href="/wiki/Sardine_run" title="Sardine run">sardine run</a>, as many as 18,000 dolphins, behaving like sheepdogs, herd the sardines into bait balls, or corral them in shallow water. Once the bait balls are rounded up, the dolphins and other predators take turns ploughing through them, gorging on the fish as they sweep through. Seabirds also attack them from above, flocks of <a href="/wiki/Gannet" title="Gannet">gannets</a>, <a href="/wiki/Cormorant" title="Cormorant">cormorants</a>, <a href="/wiki/Tern" title="Tern">terns</a> and <a href="/wiki/Gull" title="Gull">gulls</a>. Some of these seabirds plummet from heights of 30 metres (100 feet), plunging through the water leaving vapour-like trails, similar to that of fighter planes.<sup id="cite_ref-baitball_48-2" class="reference"><a href="#cite_note-baitball-48"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Gannet" title="Gannet">Gannets</a> plunge into the water at up to 100 kilometres per hour (60 mph). They have air sacs under their skin in their face and chest which act like <a href="/wiki/Bubble-wrap" class="mw-redirect" title="Bubble-wrap">bubble-wrap</a>, cushioning the impact with the water. </p><p>Subsets of <a href="/wiki/Bottlenose_dolphin" title="Bottlenose dolphin">bottlenose dolphin</a> populations in <a href="/wiki/Mauritania" title="Mauritania">Mauritania</a> are known to engage in interspecific cooperative fishing with human fishermen. The dolphins drive a school of fish towards the shore where humans await with their nets. In the confusion of casting nets, the dolphins catch a large number of fish as well. Intraspecific cooperative foraging techniques have also been observed, and some propose that these behaviours are transmitted through cultural means. Rendell & Whitehead have proposed a structure for the study of culture in cetaceans.<sup id="cite_ref-Rendell_2001_56-0" class="reference"><a href="#cite_note-Rendell_2001-56"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> </p><p>Some whales <a href="/wiki/Lunge_feed" class="mw-redirect" title="Lunge feed">lunge feed</a> on bait balls.<sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> Lunge feeding is an extreme feeding method, in which the whale accelerates from below a bait ball to a high velocity and then opens its mouth to a large gape angle. This generates the water pressure required to expand its mouth and engulf and filter a huge amount of water and fish. Lunge feeding by the huge <a href="/wiki/Rorqual" title="Rorqual">rorquals</a> is said to be the largest <a href="/wiki/Biomechanics" title="Biomechanics">biomechanical</a> event on Earth.<sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> </p> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox" style="width: 260px; clear: right; float:right;margin:0 0 1.5em 1.5em"><tbody><tr><th colspan="2" class="infobox-above" style="font-size:115%">External videos</th></tr><tr><td colspan="2" class="infobox-full-data" style="text-align: left"><span typeof="mw:File"><span><img alt="video icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/16px-Nuvola_apps_kaboodle.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/24px-Nuvola_apps_kaboodle.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/32px-Nuvola_apps_kaboodle.svg.png 2x" data-file-width="128" data-file-height="128" /></span></span> <a rel="nofollow" class="external text" href="http://www.lifeinthefastlane.ca/wp-content/uploads/2008/11/sharks_sardines_4sfw.jpg">Dolphins herd sardines.</a><sup id="cite_ref-baitball_48-3" class="reference"><a href="#cite_note-baitball-48"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup></td></tr><tr><td colspan="2" class="infobox-full-data" style="text-align: left"><span typeof="mw:File"><span><img alt="video icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/16px-Nuvola_apps_kaboodle.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/24px-Nuvola_apps_kaboodle.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/32px-Nuvola_apps_kaboodle.svg.png 2x" data-file-width="128" data-file-height="128" /></span></span> <a rel="nofollow" class="external text" href="http://www.lifeinthefastlane.ca/wp-content/uploads/2008/11/sharks_sardines_5sfw.jpg">Gannets "divebomb" sardines.</a><sup id="cite_ref-baitball_48-4" class="reference"><a href="#cite_note-baitball-48"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup></td></tr><tr><td colspan="2" class="infobox-full-data" style="text-align: left"><span typeof="mw:File"><span><img alt="video icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/16px-Nuvola_apps_kaboodle.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/24px-Nuvola_apps_kaboodle.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/32px-Nuvola_apps_kaboodle.svg.png 2x" data-file-width="128" data-file-height="128" /></span></span> <a rel="nofollow" class="external text" href="http://www.blackwellpublishing.com/helfman/video55.asp">Cooperative hunting by Sailfish.</a></td></tr></tbody></table> <ul class="gallery mw-gallery-packed" style="float:left;"> <li class="gallerybox" style="width: 265.33333333333px"> <div class="thumb" style="width: 263.33333333333px;"><span typeof="mw:File"><a href="/wiki/File:Humpback_lunge_feeding.jpg" class="mw-file-description" title="A pair of humpback whales, a species of rorqual, lunge feeding"><img alt="A pair of humpback whales, a species of rorqual, lunge feeding" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/84/Humpback_lunge_feeding.jpg/395px-Humpback_lunge_feeding.jpg" decoding="async" width="264" height="180" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/8/84/Humpback_lunge_feeding.jpg 1.5x" data-file-width="547" data-file-height="374" /></a></span></div> <div class="gallerytext">A pair of humpback whales, a species of rorqual, lunge feeding</div> </li> <li class="gallerybox" style="width: 199.33333333333px"> <div class="thumb" style="width: 197.33333333333px;"><span typeof="mw:File"><a href="/wiki/File:Morus_serrator.jpg" class="mw-file-description" title="Gannets "divebomb" at high speed"><img alt="Gannets "divebomb" at high speed" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Morus_serrator.jpg/296px-Morus_serrator.jpg" decoding="async" width="198" height="180" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Morus_serrator.jpg/445px-Morus_serrator.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/65/Morus_serrator.jpg/593px-Morus_serrator.jpg 2x" data-file-width="2048" data-file-height="1866" /></a></span></div> <div class="gallerytext">Gannets "divebomb" at high speed</div> </li> </ul> <div style="clear:both;" class=""></div> <div class="mw-heading mw-heading2"><h2 id="How_fish_school">How fish school</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=8" title="Edit section: How fish school"><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:Barracuda_Tornado.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/97/Barracuda_Tornado.jpg/220px-Barracuda_Tornado.jpg" decoding="async" width="220" height="138" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/97/Barracuda_Tornado.jpg/330px-Barracuda_Tornado.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/97/Barracuda_Tornado.jpg/440px-Barracuda_Tornado.jpg 2x" data-file-width="500" data-file-height="313" /></a><figcaption>A "tornado" of schooling <a href="/wiki/Barracuda" title="Barracuda">barracudas</a></figcaption></figure> <p>Fish schools swim in disciplined phalanxes, with some species, such as herrings, able to stream up and down at impressive speeds, twisting this way and that, and making startling changes in the shape of the school, without collisions. It is as if their motions are choreographed, though they are not. There must be very fast response systems to allow the fish to do this. Young fish practice schooling techniques in pairs, and then in larger groups as their techniques and senses mature. The schooling behaviour develops instinctively and is not learned from older fish. To school the way they do, fish require sensory systems which can respond with great speed to small changes in their position relative to their neighbour. Most schools lose their schooling abilities after dark, and just shoal. This indicates that vision is important to schooling. The importance of vision is also indicated by the behaviour of fish who have been temporarily blinded. Schooling species have eyes on the sides of their heads, which means they can easily see their neighbours. Also, schooling species often have "schooling marks" on their shoulders or the base of their tails, or visually prominent stripes, which provide reference marks when schooling,<sup id="cite_ref-Bone_59-0" class="reference"><a href="#cite_note-Bone-59"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> similar in function to <a href="/wiki/Motion_capture#Passive_markers" title="Motion capture">passive markers</a> in artificial motion capture. However fish without these markers will still engage in schooling behaviour,<sup id="cite_ref-bbcnews_60-0" class="reference"><a href="#cite_note-bbcnews-60"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> though perhaps not as efficiently.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p><p>Other senses are also used. Pheromones or sound may also play a part but supporting evidence has not been found so far. The <a href="/wiki/Lateral_line" title="Lateral line">lateral line</a> is a line running along each side of the fish from the gill covers to the base of the tail. In laboratory experiments the lateral lines of schooling fish have been removed. They swam closer, leading to a theory that the lateral lines provide additional stimuli input when the fish get too close.<sup id="cite_ref-Bone_59-1" class="reference"><a href="#cite_note-Bone-59"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> The lateral-line system is very sensitive to changes in water currents and vibration in the water. It uses receptors called <a href="/wiki/Neuromasts" class="mw-redirect" title="Neuromasts">neuromasts</a>, each of which is composed of a group of hair cells. The hairs are surrounded by a protruding jelly-like <a href="/wiki/Lateral_line" title="Lateral line">cupula</a>, typically 0.1 to 0.2 mm long. The hair cells in the lateral line are similar to the hair cells inside the vertebrate inner ear, indicating that the lateral line and the inner ear share a common origin.<sup id="cite_ref-Moyle_and_Cech_6-6" class="reference"><a href="#cite_note-Moyle_and_Cech-6"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Describing_shoal_structure">Describing shoal structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=9" title="Edit section: Describing shoal structure"><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/Collective_animal_behavior" title="Collective animal behavior">Collective animal behavior</a></div> <p>It is difficult to observe and describe the three dimensional structure of real world fish shoals because of the large number of fish involved. Techniques include the use of recent advances in <a href="/wiki/Fisheries_acoustics" title="Fisheries acoustics">fisheries acoustics</a>.<sup id="cite_ref-sensor_61-0" class="reference"><a href="#cite_note-sensor-61"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> </p><p>Parameters defining a fish shoal include: </p> <ul><li><a href="/wiki/Group_size_measures" title="Group size measures">Shoal size</a> – The number of fish in the shoal. A remote sensing technique has been used near the edge of the continental shelf off the east coast of North America to take images of fish shoals. The shoals – most likely made up of <a href="/wiki/Atlantic_herring" title="Atlantic herring">Atlantic herring</a>, <a href="/wiki/Scup" title="Scup">scup</a>, <a href="/wiki/Hake" title="Hake">hake</a>, and black sea bass – were said to contain "tens of millions" of fish and stretched for "many kilometers".<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup></li> <li>Density – The density of a fish shoal is the number of fish divided by the volume occupied by the shoal. Density is not necessarily a constant throughout the group. Fish in schools typically have a density of about one fish per cube of body length.<sup id="cite_ref-63" class="reference"><a href="#cite_note-63"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup></li></ul> <ul class="gallery mw-gallery-packed"> <li class="gallerybox" style="width: 175.33333333333px"> <div class="thumb" style="width: 173.33333333333px;"><span typeof="mw:File"><a href="/wiki/File:Jon_hanson_-_schooling_bannerfish_school_(by-sa).jpg" class="mw-file-description" title="Low density"><img alt="Low density" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/94/Jon_hanson_-_schooling_bannerfish_school_%28by-sa%29.jpg/260px-Jon_hanson_-_schooling_bannerfish_school_%28by-sa%29.jpg" decoding="async" width="174" height="130" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/94/Jon_hanson_-_schooling_bannerfish_school_%28by-sa%29.jpg/391px-Jon_hanson_-_schooling_bannerfish_school_%28by-sa%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/94/Jon_hanson_-_schooling_bannerfish_school_%28by-sa%29.jpg/520px-Jon_hanson_-_schooling_bannerfish_school_%28by-sa%29.jpg 2x" data-file-width="1280" data-file-height="960" /></a></span></div> <div class="gallerytext">Low density</div> </li> <li class="gallerybox" style="width: 175.33333333333px"> <div class="thumb" style="width: 173.33333333333px;"><span typeof="mw:File"><a href="/wiki/File:School_of_Pterocaesio_chrysozona_in_Papua_New_Guinea_1.jpg" class="mw-file-description" title="High density"><img alt="High density" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/School_of_Pterocaesio_chrysozona_in_Papua_New_Guinea_1.jpg/260px-School_of_Pterocaesio_chrysozona_in_Papua_New_Guinea_1.jpg" decoding="async" width="174" height="130" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/School_of_Pterocaesio_chrysozona_in_Papua_New_Guinea_1.jpg/391px-School_of_Pterocaesio_chrysozona_in_Papua_New_Guinea_1.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/86/School_of_Pterocaesio_chrysozona_in_Papua_New_Guinea_1.jpg/520px-School_of_Pterocaesio_chrysozona_in_Papua_New_Guinea_1.jpg 2x" data-file-width="1632" data-file-height="1224" /></a></span></div> <div class="gallerytext">High density</div> </li> <li class="gallerybox" style="width: 197.33333333333px"> <div class="thumb" style="width: 195.33333333333px;"><span typeof="mw:File"><a href="/wiki/File:Lutjanus_viridis.jpg" class="mw-file-description" title="Low polarity"><img alt="Low polarity" src="//upload.wikimedia.org/wikipedia/commons/thumb/7/75/Lutjanus_viridis.jpg/293px-Lutjanus_viridis.jpg" decoding="async" width="196" height="130" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/75/Lutjanus_viridis.jpg/440px-Lutjanus_viridis.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/75/Lutjanus_viridis.jpg/585px-Lutjanus_viridis.jpg 2x" data-file-width="4229" data-file-height="2819" /></a></span></div> <div class="gallerytext">Low polarity</div> </li> <li class="gallerybox" style="width: 175.33333333333px"> <div class="thumb" style="width: 173.33333333333px;"><span typeof="mw:File"><a href="/wiki/File:Sixfinger_threadfin_school.jpg" class="mw-file-description" title="High polarity"><img alt="High polarity" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Sixfinger_threadfin_school.jpg/260px-Sixfinger_threadfin_school.jpg" decoding="async" width="174" height="130" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Sixfinger_threadfin_school.jpg/391px-Sixfinger_threadfin_school.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/86/Sixfinger_threadfin_school.jpg/520px-Sixfinger_threadfin_school.jpg 2x" data-file-width="720" data-file-height="540" /></a></span></div> <div class="gallerytext">High polarity</div> </li> </ul> <ul><li>Polarity – The group polarity describes the extent to which the fish are all pointing in the same direction. In order to determine this parameter, the average orientation of all animals in the group is determined. For each animal, the angular difference between its orientation and the group orientation is then found. The group polarity is the average of these differences.<sup id="cite_ref-64" class="reference"><a href="#cite_note-64"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup></li> <li>Nearest neighbour distance – The nearest neighbour distance (NND) describes the distance between the centroid of one fish (the focal fish) and the centroid of the fish nearest to the focal fish. This parameter can be found for each fish in an aggregation and then averaged. Care must be taken to account for the fish located at the edge of a fish aggregation, since these fish have no neighbour in one direction. The NND is also related to the packing density. For schooling fish the NND is usually between one-half and one body length.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup></li> <li>Nearest neighbour position – In a polar coordinate system, the nearest neighbour position describes the angle and distance of the nearest neighbour to a focal fish.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup></li> <li><a href="/wiki/Packing_density" title="Packing density">Packing fraction</a> – The packing fraction is a parameter borrowed from physics to define the organization (or state i.e. solid, liquid, or gas) of 3D fish groups. It is an alternative measure to density. In this parameter, the aggregation is idealized as an ensemble of solid spheres, with each fish at the center of a sphere. The packing fraction is defined as the ratio of the total volume occupied by all individual spheres divided by the global volume of the aggregation. Values range from zero to one, where a small packing fraction represents a dilute system like a gas.<sup id="cite_ref-cavagna_65-0" class="reference"><a href="#cite_note-cavagna-65"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup></li> <li>Integrated conditional density – This parameter measures the density at various length scales and therefore describes the homogeneity of density throughout an animal group.<sup id="cite_ref-cavagna_65-1" class="reference"><a href="#cite_note-cavagna-65"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Pair_distribution_function" title="Pair distribution function">Pair distribution function</a> – This parameter is usually used in physics to characterize the degree of spatial order in a system of particles. It also describes the density, but this measure describes the density at a distance away from a given point. Cavagna et al. found that flocks of starlings exhibited more structure than a gas but less than a liquid.<sup id="cite_ref-cavagna_65-2" class="reference"><a href="#cite_note-cavagna-65"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup></li></ul> <div class="mw-heading mw-heading2"><h2 id="Modelling_school_behaviour">Modelling school behaviour</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=10" title="Edit section: Modelling school behaviour"><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/Collective_animal_behaviour" class="mw-redirect" title="Collective animal behaviour">Collective animal behaviour</a> and <a href="/wiki/Swarm_intelligence" title="Swarm intelligence">Swarm intelligence</a></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1237032888/mw-parser-output/.tmulti"><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:404px;max-width:404px"><div class="trow"><div class="tsingle" style="width:402px;max-width:402px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Metric_vs_topological_distance_for_animal_aggregations.png" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/b/bf/Metric_vs_topological_distance_for_animal_aggregations.png/400px-Metric_vs_topological_distance_for_animal_aggregations.png" decoding="async" width="400" height="244" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/bf/Metric_vs_topological_distance_for_animal_aggregations.png/600px-Metric_vs_topological_distance_for_animal_aggregations.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/bf/Metric_vs_topological_distance_for_animal_aggregations.png/800px-Metric_vs_topological_distance_for_animal_aggregations.png 2x" data-file-width="859" data-file-height="524" /></a></span></div><div class="thumbcaption"><div class="center" style="width:auto; margin-left:auto; margin-right:auto;">↑ Diagram illustrating the difference between <i>metric distance</i> and <i>topological distance</i> in reference to fish schools</div></div></div></div><div class="trow"><div class="tsingle" style="width:402px;max-width:402px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Large_fish_school.png" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/ea/Large_fish_school.png/400px-Large_fish_school.png" decoding="async" width="400" height="247" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/e/ea/Large_fish_school.png 1.5x" data-file-width="508" data-file-height="314" /></a></span></div><div class="thumbcaption"><div class="center" style="width:auto; margin-left:auto; margin-right:auto;">↑ Large schools, like this one, still follow positional evaluation and are regulated by the same density and volume characteristics as smaller schools<sup id="cite_ref-Aparna2011_29-1" class="reference"><a href="#cite_note-Aparna2011-29"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup></div> <hr /> <p><span typeof="mw:File"><a href="/wiki/File:Nuvola_apps_kaboodle.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/16px-Nuvola_apps_kaboodle.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/24px-Nuvola_apps_kaboodle.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/32px-Nuvola_apps_kaboodle.svg.png 2x" data-file-width="128" data-file-height="128" /></a></span> <a rel="nofollow" class="external text" href="http://www.red3d.com/cwr/boids/">Boids simulation</a> – needs Java </p> <span typeof="mw:File"><a href="/wiki/File:Nuvola_apps_kaboodle.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/16px-Nuvola_apps_kaboodle.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/24px-Nuvola_apps_kaboodle.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nuvola_apps_kaboodle.svg/32px-Nuvola_apps_kaboodle.svg.png 2x" data-file-width="128" data-file-height="128" /></a></span> <a rel="nofollow" class="external text" href="http://phet.colorado.edu/sims/self-driven-particle-model/self-driven-particle-model_en.jar">Interactive simulation of self-propelled particles</a><sup id="cite_ref-66" class="reference"><a href="#cite_note-66"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> – needs Java</div></div></div></div></div> <div class="mw-heading mw-heading3"><h3 id="Mathematical_models">Mathematical models</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=11" title="Edit section: Mathematical models"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1251242444"><table class="box-More_citations_needed_section plainlinks metadata ambox ambox-content ambox-Refimprove" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Question_book-new.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>needs additional citations for <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">verification</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Shoaling_and_schooling" title="Special:EditPage/Shoaling and schooling">improve this article</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a> in this section. Unsourced material may be challenged and removed.</span> <span class="date-container"><i>(<span class="date">January 2021</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p>The observational approach is complemented by the mathematical modelling of schools. The most common mathematical models of schools instruct the individual animals to follow three rules: </p> <ol><li>Move in the same direction as your neighbour</li> <li>Remain close to your neighbours</li> <li>Avoid collisions with your neighbours</li></ol> <p>An example of such a simulation is the <a href="/wiki/Boids" title="Boids">boids</a> program created by <a href="/wiki/Craig_Reynolds_(computer_graphics)" title="Craig Reynolds (computer graphics)">Craig Reynolds</a> in 1986.<sup id="cite_ref-67" class="reference"><a href="#cite_note-67"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> Another is the <a href="/wiki/Self-propelled_particle" class="mw-redirect" title="Self-propelled particle">self-propelled particle</a> model introduced by <a href="/wiki/Tam%C3%A1s_Vicsek" title="Tamás Vicsek">Vicsek</a> <i>et al.</i> in 1995<sup id="cite_ref-Vicsek1995_68-0" class="reference"><a href="#cite_note-Vicsek1995-68"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> Many current models use variations on these rules. For instance, many models implement these three rules through layered zones around each fish. </p> <ol><li>In the zone of repulsion very close to the fish, the focal fish will seek to distance itself from its neighbours in order to avoid a collision.</li> <li>In the slightly further away zone of alignment, a focal fish will seek to align its direction of motion with its neighbours.</li> <li>In the outmost zone of attraction, which extends as far away from the focal fish as it is able to sense, the focal fish will seek to move towards a neighbour.</li></ol> <p>The shape of these zones will necessarily be affected by the sensory capabilities of the fish. Fish rely on both vision and on hydrodynamic signals relayed through its <a href="/wiki/Lateral_line" title="Lateral line">lateral line</a>. Antarctic krill rely on vision and on hydrodynamic signals relayed through its <a href="/wiki/Antenna_(biology)" title="Antenna (biology)">antennae</a>. </p><p>In a masters thesis published in 2008, Moshi Charnell produced schooling behaviour without using the alignment matching component of an individual's behaviour.<sup id="cite_ref-69" class="reference"><a href="#cite_note-69"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> His model reduces the three basic rules to the following two rules: </p> <ol><li>Remain close to your neighbours</li> <li>Avoid collisions with your neighbours</li></ol> <p>In a paper published in 2009, researchers from Iceland recount their application of an interacting particle model to the capelin stock around Iceland, successfully predicting the spawning migration route for 2008.<sup id="cite_ref-70" class="reference"><a href="#cite_note-70"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Evolutionary_models">Evolutionary models</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=12" title="Edit section: Evolutionary models"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In order to gain insight into <i>why</i> animals evolve <a href="/wiki/Swarming_behaviour" class="mw-redirect" title="Swarming behaviour">swarming behaviour</a>, scientists have turned to evolutionary models that simulate populations of evolving animals. Typically these studies use a <a href="/wiki/Genetic_algorithm" title="Genetic algorithm">genetic algorithm</a> to simulate <a href="/wiki/Evolution" title="Evolution">evolution</a> over many generations in the model. These studies have investigated a number of hypotheses explaining why animals evolve swarming behaviour, such as the <a href="/wiki/Selfish_herd_theory" title="Selfish herd theory">selfish herd theory</a>,<sup id="cite_ref-71" class="reference"><a href="#cite_note-71"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-72" class="reference"><a href="#cite_note-72"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-73" class="reference"><a href="#cite_note-73"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> the predator confusion effect,<sup id="cite_ref-Olson_RS,_Hintze_A,_Dyer_FC,_Knoester_DB,_Adami_C_2013_20130305_35-1" class="reference"><a href="#cite_note-Olson_RS,_Hintze_A,_Dyer_FC,_Knoester_DB,_Adami_C_2013_20130305-35"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-75" class="reference"><a href="#cite_note-75"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> the dilution effect,<sup id="cite_ref-76" class="reference"><a href="#cite_note-76"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-77" class="reference"><a href="#cite_note-77"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> and the many eyes theory.<sup id="cite_ref-78" class="reference"><a href="#cite_note-78"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Mapping_the_formation_of_schools">Mapping the formation of schools</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=13" title="Edit section: Mapping the formation of schools"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 2009, building on recent advances in <a href="/wiki/Acoustics" title="Acoustics">acoustic</a> imaging,<sup id="cite_ref-sensor_61-1" class="reference"><a href="#cite_note-sensor-61"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-79" class="reference"><a href="#cite_note-79"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> a group of <a href="/wiki/MIT" class="mw-redirect" title="MIT">MIT</a> researchers observed for "the first time the formation and subsequent migration of a huge shoal of fish."<sup id="cite_ref-Makris_80-0" class="reference"><a href="#cite_note-Makris-80"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> The results provide the first field confirmation of general theories about how large groups behave, from <a href="/wiki/Locust_swarm" class="mw-redirect" title="Locust swarm">locust swarms</a> to <a href="/wiki/Flocking_(behavior)" class="mw-redirect" title="Flocking (behavior)">bird flocks</a>.<sup id="cite_ref-PhysOrg_81-0" class="reference"><a href="#cite_note-PhysOrg-81"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> </p><p>The researchers imaged <a href="/wiki/Spawn_(biology)" class="mw-redirect" title="Spawn (biology)">spawning</a> <a href="/wiki/Atlantic_herring" title="Atlantic herring">Atlantic herring</a> off <a href="/wiki/Georges_Bank" title="Georges Bank">Georges Bank</a>. They found that the fish come together from deeper water in the evening, shoaling in a disordered way. A chain reaction triggers when the population density reaches a critical value, like an <a href="/wiki/Audience_wave" class="mw-redirect" title="Audience wave">audience wave</a> travelling around a sport stadium. A rapid transition then occurs, and the fish become highly polarised and synchronized in the manner of schooling fish. After the transition, the schools start migrating, extending up to 40 kilometres (25 mi) across the ocean, to shallow parts of the bank. There they spawn during the night. In the morning, the fish school back to deeper water again and then disband. Small groups of leaders were also discovered that significantly influenced much larger groups.<sup id="cite_ref-PhysOrg_81-1" class="reference"><a href="#cite_note-PhysOrg-81"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Leadership_and_decision-making">Leadership and decision-making</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=14" title="Edit section: Leadership and decision-making"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Fish schools are faced with decisions they must make if they are to remain together. For example, a decision might be which direction to swim when confronted by a predator, which areas to stop and forage, or when and where to migrate.<sup id="cite_ref-82" class="reference"><a href="#cite_note-82"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Quorum_sensing" title="Quorum sensing">Quorum sensing</a> can function as a collective decision-making process in any <a href="/wiki/Decentralised_system" title="Decentralised system">decentralised system</a>. A quorum response has been defined as "a steep increase in the probability of group members performing a given behaviour once a threshold minimum number of their group mates already performing that behaviour is exceeded".<sup id="cite_ref-Ward2012_83-0" class="reference"><a href="#cite_note-Ward2012-83"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup> A recent investigation showed that small groups of fish used consensus decision-making when deciding which fish model to follow. The fish did this by a simple quorum rule such that individuals watched the decisions of others before making their own decisions. This technique generally resulted in the 'correct' decision but occasionally cascaded into the 'incorrect' decision. In addition, as the group size increased, the fish made more accurate decisions in following the more attractive fish model.<sup id="cite_ref-84" class="reference"><a href="#cite_note-84"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> Consensus decision-making, a form of <a href="/wiki/Collective_intelligence" title="Collective intelligence">collective intelligence</a>, thus effectively uses information from multiple sources to generally reach the correct conclusion. Such behaviour has also been demonstrated in the shoaling behaviour of <a href="/wiki/Threespine_stickleback" class="mw-redirect" title="Threespine stickleback">threespine sticklebacks</a>.<sup id="cite_ref-Ward2012_83-1" class="reference"><a href="#cite_note-Ward2012-83"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup> </p><p>Other open questions of shoaling behaviour include identifying which individuals are responsible for the direction of shoal movement. In the case of <a href="/wiki/Fish_migration" title="Fish migration">migratory</a> movement, most members of a shoal seem to know where they are going. Observations on the foraging behaviour of captive <a href="/wiki/Golden_shiner" title="Golden shiner">golden shiner</a> (a kind of <a href="/wiki/Minnow" title="Minnow">minnow</a>) found they formed shoals which were led by a small number of experienced individuals who knew when and where food was available.<sup id="cite_ref-85" class="reference"><a href="#cite_note-85"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> If all golden shiners in a shoal have similar knowledge of food availability, there are a few individuals that still emerge as natural leaders (being at the front more often) and behavioural tests suggest they are naturally bolder.<sup id="cite_ref-86" class="reference"><a href="#cite_note-86"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> Smaller golden shiners appear more willing than larger ones to be near the front of the shoal, perhaps because they are hungrier.<sup id="cite_ref-87" class="reference"><a href="#cite_note-87"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> Observations on the <a href="/wiki/Common_roach" title="Common roach">common roach</a> have shown that food-deprived individuals tend to be at the front of a shoal, where they obtain more food<sup id="cite_ref-88" class="reference"><a href="#cite_note-88"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-89" class="reference"><a href="#cite_note-89"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> but where they may also be more vulnerable to ambush predators.<sup id="cite_ref-90" class="reference"><a href="#cite_note-90"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup> Individuals that are wary of predation tend to seek more central positions within shoals.<sup id="cite_ref-91" class="reference"><a href="#cite_note-91"><span class="cite-bracket">[</span>89<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Shoal_choice">Shoal choice</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=15" title="Edit section: Shoal choice"><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:Glass_fish.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/67/Glass_fish.jpg/220px-Glass_fish.jpg" decoding="async" width="220" height="186" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/67/Glass_fish.jpg/330px-Glass_fish.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/67/Glass_fish.jpg/440px-Glass_fish.jpg 2x" data-file-width="2764" data-file-height="2336" /></a><figcaption> Fish, such as these <a href="/wiki/Sweeper" title="Sweeper">sweepers</a>, usually prefer to join larger schools which contain members of their own species matching their own size</figcaption></figure> <p>Experimental studies of shoal preference are relatively easy to perform. An aquarium containing a choosing fish is sandwiched between two aquaria containing different shoals, and the choosing fish is assumed to spend more time next to the shoal it prefers. Studies of this kind have identified several factors important for shoal preference.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p><p>Fish generally prefer larger shoals.<sup id="cite_ref-92" class="reference"><a href="#cite_note-92"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-93" class="reference"><a href="#cite_note-93"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup> This makes sense, as larger shoal usually provide better protection against predators. Indeed, the preference for larger shoals seems stronger when predators are nearby,<sup id="cite_ref-94" class="reference"><a href="#cite_note-94"><span class="cite-bracket">[</span>92<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-95" class="reference"><a href="#cite_note-95"><span class="cite-bracket">[</span>93<span class="cite-bracket">]</span></a></sup> or in species that rely more on shoaling than body armour against predation.<sup id="cite_ref-96" class="reference"><a href="#cite_note-96"><span class="cite-bracket">[</span>94<span class="cite-bracket">]</span></a></sup> Larger shoals may also find food faster, though that food would have to be shared amongst more individuals. Competition may mean that hungry individuals might prefer smaller shoals or exhibit a lesser preference for very large shoals, as shown in sticklebacks.<sup id="cite_ref-97" class="reference"><a href="#cite_note-97"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-98" class="reference"><a href="#cite_note-98"><span class="cite-bracket">[</span>96<span class="cite-bracket">]</span></a></sup> </p><p>Fish prefer to shoal with their own species. Sometimes, several species may become mingled in one shoal, but when a predator is presented to such shoals, the fish reorganize themselves so that each individual ends up being closer to members of its own species.<sup id="cite_ref-99" class="reference"><a href="#cite_note-99"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> </p><p>Fish tend to prefer shoals made up of individuals that match their own size.<sup id="cite_ref-100" class="reference"><a href="#cite_note-100"><span class="cite-bracket">[</span>98<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-101" class="reference"><a href="#cite_note-101"><span class="cite-bracket">[</span>99<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-102" class="reference"><a href="#cite_note-102"><span class="cite-bracket">[</span>100<span class="cite-bracket">]</span></a></sup> This makes sense as predators have an easier time catching individuals that stand out in a shoal. Some fish may even prefer shoals of another species if this means a better match in current body size.<sup id="cite_ref-103" class="reference"><a href="#cite_note-103"><span class="cite-bracket">[</span>101<span class="cite-bracket">]</span></a></sup> As for shoal size however, hunger can affect the preference for similarly sized fish; large fish, for example, might prefer to associate with smaller ones because of the competitive advantage they will gain over these shoalmates. In <a href="/wiki/Golden_shiner" title="Golden shiner">golden shiner</a>, large satiated fish prefer to associate with other large individuals, but hungry ones prefer smaller shoalmates.<sup id="cite_ref-104" class="reference"><a href="#cite_note-104"><span class="cite-bracket">[</span>102<span class="cite-bracket">]</span></a></sup> </p><p>Fish prefer to shoal with individuals with which the choosing fish is already familiar. This has been demonstrated in <a href="/wiki/Guppy" title="Guppy">guppies</a>,<sup id="cite_ref-105" class="reference"><a href="#cite_note-105"><span class="cite-bracket">[</span>103<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-106" class="reference"><a href="#cite_note-106"><span class="cite-bracket">[</span>104<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Threespine_stickleback" class="mw-redirect" title="Threespine stickleback">threespine stickleback</a>,<sup id="cite_ref-107" class="reference"><a href="#cite_note-107"><span class="cite-bracket">[</span>105<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Banded_killifish" title="Banded killifish">banded killifish</a>,<sup id="cite_ref-108" class="reference"><a href="#cite_note-108"><span class="cite-bracket">[</span>106<span class="cite-bracket">]</span></a></sup> the surfperch <i><a href="/wiki/Embiotoca" title="Embiotoca">Embiotoca</a> jacksoni</i>,<sup id="cite_ref-109" class="reference"><a href="#cite_note-109"><span class="cite-bracket">[</span>107<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Mexican_tetra" title="Mexican tetra">Mexican tetra</a>,<sup id="cite_ref-110" class="reference"><a href="#cite_note-110"><span class="cite-bracket">[</span>108<span class="cite-bracket">]</span></a></sup> and various minnows.<sup id="cite_ref-111" class="reference"><a href="#cite_note-111"><span class="cite-bracket">[</span>109<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-112" class="reference"><a href="#cite_note-112"><span class="cite-bracket">[</span>110<span class="cite-bracket">]</span></a></sup> A study with the <a href="/wiki/White_Cloud_Mountain_minnow" title="White Cloud Mountain minnow">White Cloud Mountain minnow</a> has also found that choosing fish prefer to shoal with individuals that have consumed the same diet as themselves.<sup id="cite_ref-113" class="reference"><a href="#cite_note-113"><span class="cite-bracket">[</span>111<span class="cite-bracket">]</span></a></sup> </p><p>Sticklebacks and killifish have been shown to prefer shoals made up of healthy individuals over parasitized ones, on the basis of visual signs of parasitism and abnormal behaviour by the parasitized fish.<sup id="cite_ref-114" class="reference"><a href="#cite_note-114"><span class="cite-bracket">[</span>112<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-115" class="reference"><a href="#cite_note-115"><span class="cite-bracket">[</span>113<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-116" class="reference"><a href="#cite_note-116"><span class="cite-bracket">[</span>114<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-117" class="reference"><a href="#cite_note-117"><span class="cite-bracket">[</span>115<span class="cite-bracket">]</span></a></sup> Zebrafish prefer shoals that consist of well-fed (greater stomach width) fish over food-deprived ones.<sup id="cite_ref-118" class="reference"><a href="#cite_note-118"><span class="cite-bracket">[</span>116<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Threespine_stickleback" class="mw-redirect" title="Threespine stickleback">Threespine stickleback</a> prefer to join a shoal made up of bold individuals rather than shy ones.<sup id="cite_ref-119" class="reference"><a href="#cite_note-119"><span class="cite-bracket">[</span>117<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Pterophyllum" title="Pterophyllum">Angelfish</a> prefer shoals made up of subordinate rather than dominant individuals.<sup id="cite_ref-120" class="reference"><a href="#cite_note-120"><span class="cite-bracket">[</span>118<span class="cite-bracket">]</span></a></sup> <a href="/wiki/European_minnow" class="mw-redirect" title="European minnow">European minnow</a> can discriminate between shoals composed of good versus poor competitors, even in the absence of obvious cues such as differences in aggressiveness, size, or feeding rate; they prefer to associate with the poor competitors.<sup id="cite_ref-121" class="reference"><a href="#cite_note-121"><span class="cite-bracket">[</span>119<span class="cite-bracket">]</span></a></sup> All of this suggests a strategy to obtain food, as bolder individuals should be more likely to find food, while subordinates would offer less competition for the discovered food.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p><p>Fish prefer to join shoals that are actively feeding.<sup id="cite_ref-122" class="reference"><a href="#cite_note-122"><span class="cite-bracket">[</span>120<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-123" class="reference"><a href="#cite_note-123"><span class="cite-bracket">[</span>121<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Golden_shiner" title="Golden shiner">Golden shiner</a> can also detect the anticipatory activity of shoals that expect to be fed soon, and preferentially join such shoals.<sup id="cite_ref-124" class="reference"><a href="#cite_note-124"><span class="cite-bracket">[</span>122<span class="cite-bracket">]</span></a></sup> Zebrafish also choose shoals that are more active.<sup id="cite_ref-125" class="reference"><a href="#cite_note-125"><span class="cite-bracket">[</span>123<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Commercial_fishing">Commercial fishing</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=16" title="Edit section: Commercial fishing"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The schooling behaviour of fish is exploited on an industrial scale by the <a href="/wiki/Commercial_fishing" title="Commercial fishing">commercial fishing</a> industry. Huge <a href="/wiki/Purse_seiner" class="mw-redirect" title="Purse seiner">purse seiner</a> vessels use spotter planes to locate schooling fish, such as tuna, cod, mackerel and forage fish. They can capture huge schools by rapidly encircling them with <a href="/wiki/Purse_seine" class="mw-redirect" title="Purse seine">purse seine nets</a> with the help of fast auxiliary boats and sophisticated sonar, which can track the shape of the shoal.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Further_examples">Further examples</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=17" title="Edit section: Further examples"><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:Blacksmithfish_300.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Blacksmithfish_300.jpg/220px-Blacksmithfish_300.jpg" decoding="async" width="220" height="148" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Blacksmithfish_300.jpg/330px-Blacksmithfish_300.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Blacksmithfish_300.jpg/440px-Blacksmithfish_300.jpg 2x" data-file-width="1239" data-file-height="831" /></a><figcaption>School of <a href="/wiki/Chromis_punctipinnis" title="Chromis punctipinnis">blacksmiths</a> being cleaned by parasite eating fish</figcaption></figure> <p><a href="/wiki/Chromis_punctipinnis" title="Chromis punctipinnis">Blacksmith</a> fish live in loose shoals. They have a symbiotic relationship with the parasite eating senorita fish. When they encounter a shoal of senorita fish, they stop and form a tight ball and hang upside down (pictured), each fish waiting its turn to be cleaned. The senorita fish pick dead tissues and external parasites, like parasitic copecods and isocods, from the skin of other fishes.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2016)">citation needed</span></a></i>]</sup> </p><p>Some shoals engage in <a href="/wiki/Mobbing_behaviour" class="mw-redirect" title="Mobbing behaviour">mobbing behaviour</a>. For example, <a href="/wiki/Bluegill" title="Bluegill">bluegills</a> form large nesting colonies and sometimes attack <a href="/wiki/Common_snapping_turtle" title="Common snapping turtle">snapping turtles</a>. This may function to advertise their presence, drive the predator from the area, or aid in cultural transmission of predator recognition.<sup id="cite_ref-Dominey_126-0" class="reference"><a href="#cite_note-Dominey-126"><span class="cite-bracket">[</span>124<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Piranha" title="Piranha">Piranha</a> have a reputation as fearless fish that hunt in ferocious packs. However, recent research, which "started off with the premise that they school as a means of cooperative hunting", discovered that they were in fact rather fearful fish, like other fish, which schooled for protection from their predators, such as cormorants, caimans and dolphins. Piranhas are "basically like regular fish with large teeth".<sup id="cite_ref-127" class="reference"><a href="#cite_note-127"><span class="cite-bracket">[</span>125<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Humboldt_squid" title="Humboldt squid">Humboldt squid</a> are large carnivorous marine invertebrates that move in schools of up to 1,200 individuals. They swim at speeds of up to 24 kilometres per hour (15 mph or 13 kn) propelled by water ejected through a <a href="/wiki/Hyponome" class="mw-redirect" title="Hyponome">siphon</a> and by two triangular fins. Their <a href="/wiki/Tentacle" title="Tentacle">tentacles</a> bear suckers lined with sharp teeth with which they grasp prey and drag it towards a large, sharp beak. During the day the Humboldt squid behave similar to <a href="/wiki/Mesopelagic_fish" class="mw-redirect" title="Mesopelagic fish">mesopelagic fish</a>, living at depths of 200 to 700 m (660 to 2,300 ft). Electronic tagging has shown that they also undergo <a href="/wiki/Diel_vertical_migration" title="Diel vertical migration">diel vertical migrations</a> which bring them closer to the surface from dusk to dawn.<sup id="cite_ref-Gilly_128-0" class="reference"><a href="#cite_note-Gilly-128"><span class="cite-bracket">[</span>126<span class="cite-bracket">]</span></a></sup> They hunt near the surface at night, taking advantage of the dark to use their keen vision to feed on more plentiful prey. The squid feed primarily on small <a href="/wiki/Fish" title="Fish">fish</a>, <a href="/wiki/Crustaceans" class="mw-redirect" title="Crustaceans">crustaceans</a>, <a href="/wiki/Cephalopods" class="mw-redirect" title="Cephalopods">cephalopods</a>, and <a href="/wiki/Copepod" title="Copepod">copepod</a>, and hunt for their prey in a cooperative fashion, the first observation of such behaviour in invertebrates.<sup id="cite_ref-zimmerman_129-0" class="reference"><a href="#cite_note-zimmerman-129"><span class="cite-bracket">[</span>127<span class="cite-bracket">]</span></a></sup> The Humboldt squid is also known to quickly devour larger prey when cooperatively hunting in groups. Humboldt squid are known for their speed in feasting on hooked fish, sharks, and squid, even from their own species and shoal,<sup id="cite_ref-130" class="reference"><a href="#cite_note-130"><span class="cite-bracket">[</span>128<span class="cite-bracket">]</span></a></sup> and have been known to attack fishermen and divers.<sup id="cite_ref-131" class="reference"><a href="#cite_note-131"><span class="cite-bracket">[</span>129<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=Shoaling_and_schooling&action=edit&section=18" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><span><video id="mwe_player_1" poster="//upload.wikimedia.org/wikipedia/commons/thumb/b/b2/Bigeye_trevally.ogv/220px--Bigeye_trevally.ogv.jpg" controls="" preload="none" data-mw-tmh="" class="mw-file-element" width="220" height="165" data-durationhint="10" data-mwtitle="Bigeye_trevally.ogv" data-mwprovider="wikimediacommons" resource="/wiki/File:Bigeye_trevally.ogv"><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/b/b2/Bigeye_trevally.ogv/Bigeye_trevally.ogv.240p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="240p.vp9.webm" data-width="320" data-height="240" /><source src="//upload.wikimedia.org/wikipedia/commons/b/b2/Bigeye_trevally.ogv" type="video/ogg; codecs="theora, vorbis"" data-width="320" data-height="240" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/b/b2/Bigeye_trevally.ogv/Bigeye_trevally.ogv.360p.webm" type="video/webm; codecs="vp8, vorbis"" data-transcodekey="360p.webm" data-width="320" data-height="240" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/b/b2/Bigeye_trevally.ogv/Bigeye_trevally.ogv.144p.mjpeg.mov" type="video/quicktime" data-transcodekey="144p.mjpeg.mov" data-width="192" data-height="144" /></video></span><figcaption>Schooling <a href="/wiki/Bigeye_trevally" title="Bigeye trevally">bigeye trevally</a></figcaption></figure> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 18em;"> <ul><li><a href="/wiki/Allee_effect" title="Allee effect">Allee effect</a></li> <li><a href="/wiki/Antipredator_adaptation" class="mw-redirect" title="Antipredator adaptation">Antipredator adaptation</a></li> <li><a href="/wiki/Cellular_automaton" title="Cellular automaton">Cellular automaton</a></li> <li><a href="/wiki/Krill#Swarming" title="Krill">Krill#Swarming</a></li> <li><a href="/wiki/Lek_(mating_arena)" class="mw-redirect" title="Lek (mating arena)">Lek (mating arena)</a></li> <li><a href="/wiki/Mobile_Bay_jubilee" title="Mobile Bay jubilee">Mobile Bay jubilee</a></li> <li><a href="/wiki/Optimal_foraging_theory" title="Optimal foraging theory">Optimal foraging theory</a></li> <li><a href="/wiki/Predator_satiation" title="Predator satiation">Predator satiation</a></li> <li><a href="/wiki/Schreckstoff" title="Schreckstoff">Schreckstoff</a></li> <li><a href="/wiki/The_Blue_Planet" title="The Blue Planet">The Blue Planet</a></li> <li><i><a href="/wiki/The_Shoals_of_Herring" title="The Shoals of Herring">The Shoals of Herring</a></i></li></ul> </div><div style="clear:right;" class=""></div> <div class="mw-heading mw-heading2"><h2 id="Notes">Notes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=19" title="Edit section: Notes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-lower-alpha"> <div class="mw-references-wrap"><ol class="references"> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text">Other collective nouns used for fish include a <i>draught</i> of fish, a <i>drift</i> of fish, or a <i>scale</i> of fish. Collective nouns used for specific fish or marine animal species groups include a <i>grind</i> of blackfish, a <i>troubling</i> of goldfish, <i>glean</i> of herrings, <i>bind</i> or <i>run</i> of salmon, <i>shiver</i> of sharks, <i>fever</i> of stingrays, <i>taint</i> of tilapia, <i>hover</i> of trouts and <i>pod</i> of whales.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2016)">citation needed</span></a></i>]</sup></span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text">Shoaling is a special case of aggregating, and schooling is a special case of shoaling. While schooling and shoaling mean different things within biology, they are often treated as <a href="/wiki/Synonym" title="Synonym">synonyms</a> by non-specialists, with speakers of <a href="/wiki/British_English" title="British English">British English</a> tending to use "shoaling" to describe any grouping of fish, while speakers of <a href="/wiki/American_English" title="American English">American English</a> tend to use "schooling" just as loosely.<sup id="cite_ref-Pitcher_365_1-4" class="reference"><a href="#cite_note-Pitcher_365-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=20" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239543626"><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-Pitcher_365-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Pitcher_365_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Pitcher_365_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Pitcher_365_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Pitcher_365_1-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Pitcher_365_1-4"><sup><i><b>e</b></i></sup></a></span> <span class="reference-text">Pitcher and Parish 1993, page 365.</span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFShaw1978" class="citation journal cs1">Shaw, E (1978). "Schooling fishes". <i><a href="/wiki/American_Scientist" title="American Scientist">American Scientist</a></i>. <b>66</b> (2): 166–175. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1978AmSci..66..166S">1978AmSci..66..166S</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=American+Scientist&rft.atitle=Schooling+fishes&rft.volume=66&rft.issue=2&rft.pages=166-175&rft.date=1978&rft_id=info%3Abibcode%2F1978AmSci..66..166S&rft.aulast=Shaw&rft.aufirst=E&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text">Helfman G., Collette B., & Facey D.: The Diversity of Fishes, Blackwell Publishing, p 375, 1997, <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-86542-256-7" title="Special:BookSources/0-86542-256-7">0-86542-256-7</a></span> </li> <li id="cite_note-Moyle_and_Cech-6"><span class="mw-cite-backlink">^ <a href="#cite_ref-Moyle_and_Cech_6-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Moyle_and_Cech_6-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Moyle_and_Cech_6-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Moyle_and_Cech_6-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Moyle_and_Cech_6-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Moyle_and_Cech_6-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Moyle_and_Cech_6-6"><sup><i><b>g</b></i></sup></a></span> <span class="reference-text">Moyle, PB and Cech, JJ (2003) <i>Fishes, An Introduction to Ichthyology.</i> 5th Ed, Benjamin Cummings. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-13-100847-2" title="Special:BookSources/978-0-13-100847-2">978-0-13-100847-2</a></span> </li> <li id="cite_note-7"><span class="mw-cite-backlink"><b><a href="#cite_ref-7">^</a></b></span> <span class="reference-text">Pitcher, TJ and <a href="/wiki/Julia_Parrish" title="Julia Parrish">Parrish, JK</a> (1993) Behaviour of Teleost Fishes, Chp 12: <a rel="nofollow" class="external text" href="https://books.google.com/books?id=10lElCT7v5wC&pg=PA363"><i>Functions of shoaling behaviour in teleosts</i></a> Springer. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-412-42930-9" title="Special:BookSources/978-0-412-42930-9">978-0-412-42930-9</a></span> </li> <li id="cite_note-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-8">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBreder1967" class="citation journal cs1">Breder, C. M. Jr. (1967). "On the survival value of fish schools". <i>Zoologica</i>. <b>52</b>: 25–40.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Zoologica&rft.atitle=On+the+survival+value+of+fish+schools&rft.volume=52&rft.pages=25-40&rft.date=1967&rft.aulast=Breder&rft.aufirst=C.+M.+Jr.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></span> </li> <li id="cite_note-Partridge1980-9"><span class="mw-cite-backlink">^ <a href="#cite_ref-Partridge1980_9-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Partridge1980_9-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPartridgePitcherCullenWilson1980" class="citation journal cs1">Partridge, B.; Pitcher, T.; Cullen, M.; Wilson, J. (1980). "The three-dimensional structure of fish schools". <i>Behav Ecol Sociobiol</i>. <b>6</b> (4): 277–288. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2FBF00292770">10.1007/BF00292770</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:8032766">8032766</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Behav+Ecol+Sociobiol&rft.atitle=The+three-dimensional+structure+of+fish+schools&rft.volume=6&rft.issue=4&rft.pages=277-288&rft.date=1980&rft_id=info%3Adoi%2F10.1007%2FBF00292770&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A8032766%23id-name%3DS2CID&rft.aulast=Partridge&rft.aufirst=B.&rft.au=Pitcher%2C+T.&rft.au=Cullen%2C+M.&rft.au=Wilson%2C+J.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></span> </li> <li id="cite_note-Kils,_1992-10"><span class="mw-cite-backlink">^ <a href="#cite_ref-Kils,_1992_10-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Kils,_1992_10-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKils1992" class="citation journal cs1">Kils, U. (1992). "The ecoSCOPE and dynIMAGE: Microscale tools for in situ studies of predator-prey interactions". <i>Arch Hydrobiol Beih</i>. <b>36</b>: 83–96.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Arch+Hydrobiol+Beih&rft.atitle=The+ecoSCOPE+and+dynIMAGE%3A+Microscale+tools+for+in+situ+studies+of+predator-prey+interactions&rft.volume=36&rft.pages=83-96&rft.date=1992&rft.aulast=Kils&rft.aufirst=U.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></span> </li> <li id="cite_note-NCMC-11"><span class="mw-cite-backlink"><b><a href="#cite_ref-NCMC_11-0">^</a></b></span> <span class="reference-text"><a href="/wiki/National_Coalition_for_Marine_Conservation" class="mw-redirect" title="National Coalition for Marine Conservation">National Coalition for Marine Conservation</a>: <a rel="nofollow" class="external text" href="http://www.savethefish.org/PDF_files/fish_file_10_forage_121906.pdf">Forage fish</a></span> </li> <li id="cite_note-12"><span class="mw-cite-backlink"><b><a href="#cite_ref-12">^</a></b></span> <span class="reference-text">Radakov DV (1973) <i>Schooling in the ecology of fish.</i> Israel Program for Scientific Translation, translated by Mill H. Halsted Press, New York. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-7065-1351-6" title="Special:BookSources/978-0-7065-1351-6">978-0-7065-1351-6</a></span> </li> <li id="cite_note-13"><span class="mw-cite-backlink"><b><a href="#cite_ref-13">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNøttestadAxelsen1999" class="citation journal cs1">Nøttestad, L.; Axelsen, B. E. (1999). <a rel="nofollow" class="external text" href="https://wayback.archive-it.org/all/20081217092609/http://www.imr.no/__data/page/3939/Killer%20whale%20paper_1%20C%20J%20Zoo%2077,%201540-46.pdf">"Herring schooling manoeuvres in response to killer whale attacks"</a> <span class="cs1-format">(PDF)</span>. <i>Canadian Journal of Zoology</i>. <b>77</b> (10): 1540–1546. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1139%2Fz99-124">10.1139/z99-124</a>. 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(2006). <a rel="nofollow" class="external text" href="http://ic.ucsc.edu/~kudela/OS130/Readings/Gilly,2006.pdf">"Vertical and horizontal migrations by the jumbo squid <i>Dosidicus gigas</i> revealed by electronic tagging"</a> <span class="cs1-format">(PDF)</span>. <i>Marine Ecology Progress Series</i>. <b>324</b>: 1–17. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2006MEPS..324....1G">2006MEPS..324....1G</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.3354%2Fmeps324001">10.3354/meps324001</a></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Marine+Ecology+Progress+Series&rft.atitle=Vertical+and+horizontal+migrations+by+the+jumbo+squid+Dosidicus+gigas+revealed+by+electronic+tagging&rft.volume=324&rft.pages=1-17&rft.date=2006&rft_id=info%3Adoi%2F10.3354%2Fmeps324001&rft_id=info%3Abibcode%2F2006MEPS..324....1G&rft.aulast=Gilly&rft.aufirst=W.F.&rft.au=Markaida%2C+U.&rft.au=Baxter%2C+C.H.&rft.au=Block%2C+B.A.&rft.au=Boustany%2C+A.&rft.au=Zeidberg%2C+L.&rft.au=Reisenbichler%2C+K.&rft.au=Robison%2C+B.&rft.au=Bazzino%2C+G.&rft.au=Salinas%2C+C.&rft_id=http%3A%2F%2Fic.ucsc.edu%2F~kudela%2FOS130%2FReadings%2FGilly%2C2006.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></span> </li> <li id="cite_note-zimmerman-129"><span class="mw-cite-backlink"><b><a href="#cite_ref-zimmerman_129-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFZimmermann2006" class="citation web cs1">Zimmermann, Tim (July 2006). <a rel="nofollow" class="external text" href="http://outside.away.com/outside/features/200607/sea-of-cortez-humboldt-squid-1.html">"Behold the Humboldt squid"</a>. <i>Outside Online</i><span class="reference-accessdate">. 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Retrieved <span class="nowrap">13 September</span> 2014</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=The+Curious+Case+of+the+Cannibal+Squid+%E2%80%93+National+Wildlife+Federation&rft_id=http%3A%2F%2Fwww.nwf.org%2FNews-and-Magazines%2FNational-Wildlife%2FAnimals%2FArchives%2F2005%2FThe-Curious-Case-of-the-Cannibal-Squid.aspx&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></span> </li> <li id="cite_note-131"><span class="mw-cite-backlink"><b><a href="#cite_ref-131">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFThomas2007" class="citation news cs1">Thomas, Pete (26 March 2007). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20121024120038/http://articles.latimes.com/2007/mar/26/sports/sp-squid26/2">"Warning lights of the sea"</a>. <i>Los Angeles Times</i>. Archived from <a rel="nofollow" class="external text" href="http://articles.latimes.com/2007/mar/26/sports/sp-squid26/2">the original</a> on 24 October 2012.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Los+Angeles+Times&rft.atitle=Warning+lights+of+the+sea&rft.date=2007-03-26&rft.aulast=Thomas&rft.aufirst=Pete&rft_id=http%3A%2F%2Farticles.latimes.com%2F2007%2Fmar%2F26%2Fsports%2Fsp-squid26%2F2&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=21" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin refbegin-columns references-column-width" style="column-width: 30em"> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBonabeauDagorn1995" class="citation journal cs1">Bonabeau, E; Dagorn, L (1995). <a rel="nofollow" class="external text" href="http://horizon.documentation.ird.fr/exl-doc/pleins_textes/pleins_textes_7/b_fdi_57-58/010024411.pdf">"Possible universality in the size distribution of fish schools"</a> <span class="cs1-format">(PDF)</span>. <i>Physical Review</i>. <b>51</b> (6): R5220–R5223. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1995PhRvE..51.5220B">1995PhRvE..51.5220B</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1103%2Fphysreve.51.r5220">10.1103/physreve.51.r5220</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/9963400">9963400</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Physical+Review&rft.atitle=Possible+universality+in+the+size+distribution+of+fish+schools&rft.volume=51&rft.issue=6&rft.pages=R5220-R5223&rft.date=1995&rft_id=info%3Apmid%2F9963400&rft_id=info%3Adoi%2F10.1103%2Fphysreve.51.r5220&rft_id=info%3Abibcode%2F1995PhRvE..51.5220B&rft.aulast=Bonabeau&rft.aufirst=E&rft.au=Dagorn%2C+L&rft_id=http%3A%2F%2Fhorizon.documentation.ird.fr%2Fexl-doc%2Fpleins_textes%2Fpleins_textes_7%2Fb_fdi_57-58%2F010024411.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></li> <li>Boinski S and Garber PA (2000) <a rel="nofollow" class="external text" href="https://books.google.com/books?id=KvffRQh5-noC&pg=PA574"><i>On the Move: How and why Animals Travel in Groups</i></a> University of Chicago Press. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-226-06339-3" title="Special:BookSources/978-0-226-06339-3">978-0-226-06339-3</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBreder1954" class="citation journal cs1">Breder, CM (1954). "Equations Descriptive of Fish Schools and Other Animal Aggregations". <i>Ecology</i>. <b>35</b> (3): 361–370. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1954Ecol...35..361B">1954Ecol...35..361B</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.2307%2F1930099">10.2307/1930099</a>. <a href="/wiki/JSTOR_(identifier)" class="mw-redirect" title="JSTOR (identifier)">JSTOR</a> <a rel="nofollow" class="external text" href="https://www.jstor.org/stable/1930099">1930099</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Ecology&rft.atitle=Equations+Descriptive+of+Fish+Schools+and+Other+Animal+Aggregations&rft.volume=35&rft.issue=3&rft.pages=361-370&rft.date=1954&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F1930099%23id-name%3DJSTOR&rft_id=info%3Adoi%2F10.2307%2F1930099&rft_id=info%3Abibcode%2F1954Ecol...35..361B&rft.aulast=Breder&rft.aufirst=CM&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></li> <li>Childress S (1981) <a rel="nofollow" class="external text" href="https://books.google.com/books?id=YZ2zOzsvTLoC&pg=PA155"><i>Mechanics of Swimming and Flying</i></a> Cambridge University Press. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-521-28071-6" title="Special:BookSources/978-0-521-28071-6">978-0-521-28071-6</a></li> <li>Camazine S, Deneubourg JL, <a href="/wiki/Nigel_R._Franks" title="Nigel R. Franks">Franks NR</a>, Sneyd J, Theraulaz G and Bonabeau E (2003) <i>Self-Organization in Biological Systems.</i> Princeton University Press. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-691-11624-2" title="Special:BookSources/978-0-691-11624-2">978-0-691-11624-2</a> – especially <a rel="nofollow" class="external text" href="https://books.google.com/books?id=zMgyNN6Ufj0C&pg=PA7">Chapter 11</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEvansFinnieaManica2007" class="citation journal cs1">Evans, SR; Finniea, M; Manica, A (2007). "Shoaling preferences in decapod crustacea". <i>Animal Behaviour</i>. <b>74</b> (6): 1691–1696. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.anbehav.2007.03.017">10.1016/j.anbehav.2007.03.017</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:53150496">53150496</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Animal+Behaviour&rft.atitle=Shoaling+preferences+in+decapod+crustacea&rft.volume=74&rft.issue=6&rft.pages=1691-1696&rft.date=2007&rft_id=info%3Adoi%2F10.1016%2Fj.anbehav.2007.03.017&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A53150496%23id-name%3DS2CID&rft.aulast=Evans&rft.aufirst=SR&rft.au=Finniea%2C+M&rft.au=Manica%2C+A&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDelcourtPoncin2012" class="citation journal cs1">Delcourt, J; Poncin, P (2012). "Shoals and schools: back to the heuristic definitions and quantitative references". <i>Reviews in Fish Biology and Fisheries</i>. <b>22</b> (3): 595–619. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2012RFBF...22..595D">2012RFBF...22..595D</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs11160-012-9260-z">10.1007/s11160-012-9260-z</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:18306602">18306602</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Reviews+in+Fish+Biology+and+Fisheries&rft.atitle=Shoals+and+schools%3A+back+to+the+heuristic+definitions+and+quantitative+references&rft.volume=22&rft.issue=3&rft.pages=595-619&rft.date=2012&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A18306602%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1007%2Fs11160-012-9260-z&rft_id=info%3Abibcode%2F2012RFBF...22..595D&rft.aulast=Delcourt&rft.aufirst=J&rft.au=Poncin%2C+P&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></li> <li>Gautrais, J., Jost, C. & Theraulaz, G. (2008) <a rel="nofollow" class="external text" href="http://www.sekj.org/PDF/anz45-free/anz45-415.pdf">Key behavioural factors in a self-organised fish school model.</a> Annales Zoologici Fennici 45: 415–428.</li> <li>Godin, JJ (1997) <a rel="nofollow" class="external text" href="https://books.google.com/books?id=cF2Uw8HPA60C"><i>Behavioural Ecology of Teleost Fishes</i></a> Oxford University Press. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-850503-7" title="Special:BookSources/978-0-19-850503-7">978-0-19-850503-7</a></li> <li>Ghosh S and Ramamoorthy CV (2004) <a rel="nofollow" class="external text" href="https://archive.org/details/springer_10.1007-b97343/page/n66">Design for Networked Information Technology Systems</a> Springer. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-387-95544-5" title="Special:BookSources/978-0-387-95544-5">978-0-387-95544-5</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHagerHelfman1991" class="citation journal cs1">Hager, MC; Helfman, GS (1991). 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Archived from <a rel="nofollow" class="external text" href="https://137.146.28.140/biology/BI358j/Readings/Partridge%201982.pdf">the original</a> <span class="cs1-format">(PDF)</span> on 3 July 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Scientific+American&rft.atitle=The+structure+and+function+of+fish+schools&rft.volume=246&rft.issue=6&rft.pages=114-123&rft.date=1982&rft_id=info%3Apmid%2F7201674&rft_id=info%3Adoi%2F10.1038%2Fscientificamerican0682-114&rft_id=info%3Abibcode%2F1982SciAm.246f.114P&rft.aulast=Partridge&rft.aufirst=BL&rft_id=https%3A%2F%2F137.146.28.140%2Fbiology%2FBI358j%2FReadings%2FPartridge%25201982.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPitcher1983" class="citation journal cs1">Pitcher, TJ (1983). 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"Fish in larger shoals find food faster". <i>Behavioral Ecology and Sociobiology</i>. <b>10</b> (2): 149–151. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2FBF00300175">10.1007/BF00300175</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:6340986">6340986</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Behavioral+Ecology+and+Sociobiology&rft.atitle=Fish+in+larger+shoals+find+food+faster&rft.volume=10&rft.issue=2&rft.pages=149-151&rft.date=1982&rft_id=info%3Adoi%2F10.1007%2FBF00300175&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A6340986%23id-name%3DS2CID&rft.aulast=Pitcher&rft.aufirst=TJ&rft.au=Magurran%2C+AE&rft.au=Winfield%2C+IJ&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></li> <li>Pitcher TJ (2010) <a rel="nofollow" class="external text" href="https://books.google.com/books?id=kkRKJCofvXMC">"Fish schooling"</a> In: Steele JH, Thorpe SA and Turekian KK (Eds.) <i>Marine Biology,</i> Academic Press, pages 337–349. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-08-096480-5" title="Special:BookSources/978-0-08-096480-5">978-0-08-096480-5</a>.</li> <li>Pryor K and Norris KS (1998) <a rel="nofollow" class="external text" href="https://books.google.com/books?id=yPlKzPGFUvwC&pg=PA57"><i>Dolphin Societies: Discoveries and Puzzles</i></a> University of California Press. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-520-21656-3" title="Special:BookSources/0-520-21656-3">0-520-21656-3</a>, <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-520-21656-3" title="Special:BookSources/978-0-520-21656-3">978-0-520-21656-3</a></li> <li>Ross DA (2000) <a rel="nofollow" class="external text" href="https://archive.org/details/fishermansocean0000ross/page/169"><i>The Fisherman's Ocean</i></a> Stackpole Books. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-8117-2771-6" title="Special:BookSources/978-0-8117-2771-6">978-0-8117-2771-6</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFScalabrinMassé1993" class="citation journal cs1">Scalabrin, C; Massé, J (1993). <a rel="nofollow" class="external text" href="https://doi.org/10.1051%2Falr%3A1993027">"Acoustic detection of the spatial and temporal distribution of fish shoals in the Bay of Biscay"</a>. <i>Aquatic Living Resources</i>. <b>6</b> (3): 269–283. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1993AqLR....6..269S">1993AqLR....6..269S</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1051%2Falr%3A1993027">10.1051/alr:1993027</a></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Aquatic+Living+Resources&rft.atitle=Acoustic+detection+of+the+spatial+and+temporal+distribution+of+fish+shoals+in+the+Bay+of+Biscay&rft.volume=6&rft.issue=3&rft.pages=269-283&rft.date=1993&rft_id=info%3Adoi%2F10.1051%2Falr%3A1993027&rft_id=info%3Abibcode%2F1993AqLR....6..269S&rft.aulast=Scalabrin&rft.aufirst=C&rft.au=Mass%C3%A9%2C+J&rft_id=https%3A%2F%2Fdoi.org%2F10.1051%252Falr%253A1993027&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSenoNakai1995" class="citation journal cs1">Seno, H; Nakai, K (1995). 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"Collective motion". <i>Physics Reports</i>. <b>517</b> (3–4): 71–140. <a href="/wiki/ArXiv_(identifier)" class="mw-redirect" title="ArXiv (identifier)">arXiv</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://arxiv.org/abs/1010.5017">1010.5017</a></span>. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2012PhR...517...71V">2012PhR...517...71V</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.physrep.2012.03.004">10.1016/j.physrep.2012.03.004</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:119109873">119109873</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Physics+Reports&rft.atitle=Collective+motion&rft.volume=517&rft.issue=3%E2%80%934&rft.pages=71-140&rft.date=2012&rft_id=info%3Aarxiv%2F1010.5017&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A119109873%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1016%2Fj.physrep.2012.03.004&rft_id=info%3Abibcode%2F2012PhR...517...71V&rft.aulast=Vicsek&rft.aufirst=A&rft.au=Zafeiris%2C+A&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></li> <li>White TI (2007) <a rel="nofollow" class="external text" href="https://books.google.com/books?id=X53qu2C0xA8C&pg=PA128"><i>In Defense of Dolphins</i></a> Blackwell Publishing. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4051-5779-7" title="Special:BookSources/978-1-4051-5779-7">978-1-4051-5779-7</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWolf1985" class="citation journal cs1">Wolf, NG (1985). "Odd fish abandon mixed-species groups when threatened". <i>Behavioral Ecology and Sociobiology</i>. <b>17</b> (1): 47–52. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fbf00299428">10.1007/bf00299428</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:11935938">11935938</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Behavioral+Ecology+and+Sociobiology&rft.atitle=Odd+fish+abandon+mixed-species+groups+when+threatened&rft.volume=17&rft.issue=1&rft.pages=47-52&rft.date=1985&rft_id=info%3Adoi%2F10.1007%2Fbf00299428&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A11935938%23id-name%3DS2CID&rft.aulast=Wolf&rft.aufirst=NG&rfr_id=info%3Asid%2Fen.wikipedia.org%3AShoaling+and+schooling" class="Z3988"></span></li> <li>Wootton, RJ (1998) <a rel="nofollow" class="external text" href="https://books.google.com/books?id=uYOOfHPJSgEC&pg=PA188"><i>Ecology of Teleost Fishes</i></a> Springer. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-412-64200-5" title="Special:BookSources/978-0-412-64200-5">978-0-412-64200-5</a></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Shoaling_and_schooling&action=edit&section=22" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/45px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/59px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <span style="font-weight: bold; font-style: italic;"><a href="https://commons.wikimedia.org/wiki/Category:Schools_of_fish" class="extiw" title="commons:Category:Schools of fish">Schools of fish</a></span>.</div></div> </div> <ul><li><a rel="nofollow" class="external text" href="http://www.collective-behavior.com/index.html">Collective Animal Behavior website organized around David Sumpter's book (2008) by the same name</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20090722004518/http://angel.elte.hu/starling/">STARFLAG project: Description of starling flocking project</a></li> <li><a rel="nofollow" class="external text" href="http://www.cbid.gatech.edu/">Center for Biologically Inspired Design at Georgia Tech</a></li> <li><a rel="nofollow" class="external text" href="http://www.math.uu.se/~david/web/index.html">David Sumpter's research website</a></li> <li><a rel="nofollow" class="external text" href="http://icouzin.princeton.edu/">Iain Couzin's research website</a></li> <li><a rel="nofollow" class="external text" href="http://www.fish.washington.edu/people/parrish/">Website of Julia Parrish, an animal aggregation researcher</a></li> <li>Pelagic Fisheries Research Program (2002) <a rel="nofollow" class="external text" href="https://web.archive.org/web/20090918183551/http://www2.hawaii.edu/~dagorn/workshop.htm"><i>Current status and new directions for studying schooling and aggregation behavior of pelagic fish</i></a></li> <li>Clover, Charles (2008) <a rel="nofollow" class="external text" href="https://web.archive.org/web/20080227035904/http://www.telegraph.co.uk/earth/main.jhtml?xml=/earth/2008/02/26/eafish126.xml"><i>Fish can count to four – but no higher</i></a> Telegraph Media Group.</li> <li><a rel="nofollow" class="external text" href="http://www.pac.dfo-mpo.gc.ca/sci/herring/hertags/pages/default4_e.htm">Herring Migratory Behaviour</a></li> <li><a rel="nofollow" class="external text" href="http://jgaffuri.github.io/Vectag/sims/fish.html">Example of schooling simulation</a></li> <li>Bhaduri, Aparna (2010) <a rel="nofollow" class="external text" href="http://cnx.org/contents/1137e6e6-b7bd-4b03-83e9-addeb2183873@3/Schooling_in_Fish">Schooling in Fish</a> <i><a href="/wiki/OpenStax_College" class="mw-redirect" title="OpenStax College">OpenStax College</a></i>. 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style="width:1%"><a href="/wiki/Fish_anatomy" title="Fish anatomy">Anatomy</a><br /> and<br /><a href="/wiki/Fish_physiology" title="Fish physiology">physiology</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/Age_determination_in_fish" title="Age determination in fish">Age determination</a></li> <li><a href="/wiki/Anguilliformity" title="Anguilliformity">Anguilliformity</a></li> <li><a href="/wiki/Fish_bone" title="Fish bone">Bone</a> <ul><li><a href="/wiki/Dermal_bone" title="Dermal bone">dermal</a></li> <li><a href="/wiki/Intramembranous_ossification" title="Intramembranous ossification">intramembranous ossification</a></li></ul></li> <li><a href="/wiki/Cleithrum" title="Cleithrum">Cleithrum</a></li> <li><a href="/wiki/Chromatophore" title="Chromatophore">Chromatophore</a></li> <li><a href="/wiki/Fish_fin" title="Fish fin">Fins</a> <ul><li><a href="/wiki/Dorsal_fin" title="Dorsal fin">dorsal fin</a></li></ul></li> <li><a href="/wiki/Fish_gill" title="Fish gill">Gill</a> <ul><li><a href="/wiki/Branchial_arch" title="Branchial arch">branchial arch</a></li> <li><a href="/wiki/Gill_raker" title="Gill raker">gill raker</a></li> <li><a href="/wiki/Gill_slit" title="Gill slit">gill slit</a></li> <li><a href="/wiki/Pharyngeal_arch" title="Pharyngeal arch">pharyngeal arch</a></li> <li><a href="/wiki/Pharyngeal_slit" title="Pharyngeal slit">pharyngeal slit</a></li> <li><a href="/wiki/Pseudobranch" title="Pseudobranch">pseudobranch</a></li></ul></li> <li><a href="/wiki/Glossohyal" title="Glossohyal">Glossohyal</a></li> <li><a href="/wiki/Fish_jaw" title="Fish jaw">Jaw</a> <ul><li><a href="/wiki/Hyomandibula" title="Hyomandibula">hyomandibula</a></li> <li><a href="/wiki/Pharyngeal_jaw" title="Pharyngeal jaw">pharyngeal jaw</a></li></ul></li> <li><a href="/wiki/Leydig%27s_organ" title="Leydig's organ">Leydig's organ</a></li> <li><a href="/wiki/Mauthner_cell" title="Mauthner cell">Mauthner cell</a></li> <li><a href="/wiki/Meristics" title="Meristics">Meristics</a></li> <li><a href="/wiki/Operculum_(fish)" title="Operculum (fish)">Operculum</a> <ul><li><a href="/wiki/Operculum_papillare" title="Operculum papillare">papillare</a></li></ul></li> <li><a href="/wiki/Papilla_(fish_anatomy)" title="Papilla (fish anatomy)">Papilla</a></li> <li><a href="/wiki/Photophore" title="Photophore">Photophore</a></li> <li><a href="/wiki/Root_effect" title="Root effect">Root effect</a></li> <li><a href="/wiki/Shark_cartilage" title="Shark cartilage">Shark cartilage</a></li> <li><a href="/wiki/Fish_scale" title="Fish scale">Scales</a> <ul><li><a href="/wiki/Ganoine" title="Ganoine">ganoine</a></li></ul></li> <li><a href="/wiki/Spiral_valve" title="Spiral valve">Spiral valve</a></li> <li><a href="/wiki/Suckermouth" title="Suckermouth">Suckermouth</a></li> <li><a href="/wiki/Swim_bladder" title="Swim bladder">Swim bladder</a> <ul><li><a href="/wiki/Physoclisti" title="Physoclisti">physoclisti</a></li> <li><a href="/wiki/Physostome" title="Physostome">physostome</a></li></ul></li> <li><a href="/wiki/Teeth" class="mw-redirect" title="Teeth">Teeth</a> <ul><li><a href="/wiki/Pharyngeal_teeth" title="Pharyngeal teeth">pharyngeal</a></li> <li><a href="/wiki/Shark_tooth" title="Shark tooth">shark</a></li></ul></li> <li><a href="/wiki/Teleost_leptins" title="Teleost leptins">Teleost leptins</a></li> <li><a href="/wiki/Digital_Fish_Library" title="Digital Fish Library">Digital Library</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sensory_systems_in_fish" title="Sensory systems in fish">Sensory<br />systems</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/Ampullae_of_Lorenzini" title="Ampullae of Lorenzini">Ampullae of Lorenzini</a></li> <li><a href="/wiki/Barbel_(anatomy)" title="Barbel (anatomy)">Barbel</a></li> <li><a href="/wiki/Hydrodynamic_reception" title="Hydrodynamic reception">Hydrodynamic reception</a></li> <li><a href="/wiki/Electroreception_and_electrogenesis" title="Electroreception and electrogenesis">Electroreception and electrogenesis</a></li> <li><a href="/wiki/Jamming_avoidance_response" title="Jamming avoidance response">Jamming avoidance response</a></li> <li><a href="/wiki/Lateral_line" title="Lateral line">Lateral line</a></li> <li><a href="/wiki/Otolith" title="Otolith">Otolith</a></li> <li><a href="/wiki/Pain_in_fish" title="Pain in fish">Capacity for pain</a></li> <li><a href="/wiki/Schreckstoff" title="Schreckstoff">Schreckstoff</a></li> <li><a href="/wiki/Surface_wave_detection_by_animals" title="Surface wave detection by animals">Surface wave detection</a></li> <li><a href="/wiki/Vision_in_fishes" class="mw-redirect" title="Vision in fishes">Vision</a></li> <li><a href="/wiki/Weberian_apparatus" title="Weberian apparatus">Weberian apparatus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Fish_reproduction" title="Fish reproduction">Reproduction</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/Bubble_nest" title="Bubble nest">Bubble nest</a></li> <li><a href="/wiki/Clasper" title="Clasper">Clasper</a></li> <li><a href="/wiki/Egg_case_(Chondrichthyes)" title="Egg case (Chondrichthyes)">Egg case</a></li> <li><a href="/wiki/Fish_development" title="Fish development">Development</a></li> <li><a href="/wiki/Ichthyoplankton" title="Ichthyoplankton">Ichthyoplankton</a></li> <li><a href="/wiki/Juvenile_fish" title="Juvenile fish">Juvenile</a></li> <li><a href="/wiki/Life_history_theory" title="Life history theory">Life history theory</a></li> <li><a href="/wiki/Milt" title="Milt">Milt</a></li> <li><a href="/wiki/Mouthbrooder" title="Mouthbrooder">Mouthbrooder</a></li> <li><a href="/wiki/Polyandry_in_fish" title="Polyandry in fish">Polyandry</a></li> <li><a href="/wiki/Pregnancy_in_fish" title="Pregnancy in fish">Pregnancy</a></li> <li><a href="/wiki/Roe" title="Roe">Roe</a></li> <li><a href="/wiki/Sequential_hermaphroditism" title="Sequential hermaphroditism">Sequential hermaphroditism</a></li> <li><a href="/wiki/Spawn_(biology)" class="mw-redirect" title="Spawn (biology)">Spawning</a> <ul><li><a href="/wiki/Spawning_trigger" title="Spawning trigger">triggers</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Fish_locomotion" title="Fish locomotion">Locomotion</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/Fin_and_flipper_locomotion" title="Fin and flipper locomotion">Fin and flipper locomotion</a></li> <li><a href="/wiki/Amphibious_fish" title="Amphibious fish">Amphibious</a></li> <li><a href="/wiki/Walking_fish" title="Walking fish">Walking</a></li> <li><a href="/wiki/Flying_fish" title="Flying fish">Flying</a></li> <li><a href="/wiki/Undulatory_locomotion" title="Undulatory locomotion">Undulatory locomotion</a></li> <li><a href="/wiki/Tradeoffs_for_locomotion_in_air_and_water" title="Tradeoffs for locomotion in air and water">Tradeoffs for locomotion in air and water</a></li> <li><a href="/wiki/RoboTuna" title="RoboTuna">RoboTuna</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other<br />behaviour</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/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/Bait_ball" title="Bait ball">Bait ball</a></li> <li><a href="/wiki/Bottom_feeder" title="Bottom feeder">Bottom feeders</a></li> <li><a href="/wiki/Cleaner_fish" title="Cleaner fish">Cleaner fish</a></li> <li><a href="/wiki/Corallivore" title="Corallivore">Corallivory</a></li> <li><a href="/wiki/Diel_vertical_migration" title="Diel vertical migration">Diel vertical migration</a></li> <li><a href="/wiki/Electric_fish" title="Electric fish">Electric fish</a></li> <li><a href="/wiki/Filter_feeder" title="Filter feeder">Filter feeders</a></li> <li><a href="/wiki/Forage_fish" title="Forage fish">Forage fish</a></li> <li><a href="/wiki/Fish_migration" title="Fish migration">Migratory</a></li> <li><a href="/wiki/Paedophagy" title="Paedophagy">Paedophagy</a></li> <li><a href="/wiki/Predatory_fish" title="Predatory fish">Predatory</a></li> <li><a href="/wiki/Salmon_run" title="Salmon run">Salmon run</a></li> <li><a href="/wiki/Sardine_run" title="Sardine run">Sardine run</a></li> <li><a href="/wiki/Lepidophagy" title="Lepidophagy">Scale eaters</a></li> <li><a class="mw-selflink selflink">Schooling fish</a></li> <li><a href="/wiki/Sleep_in_fish" title="Sleep in fish">Sleep</a></li> <li><a href="/wiki/Venomous_fish" title="Venomous fish">Venomous</a></li> <li><a href="/wiki/Fish_intelligence" title="Fish intelligence">Intelligence</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">By <a href="/wiki/Marine_habitat" title="Marine habitat">habitat</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/Cavefish" title="Cavefish">Cave</a></li> <li><a href="/wiki/Coastal_fish" title="Coastal fish">Coastal</a></li> <li><a href="/wiki/Coldwater_fish" title="Coldwater fish">Coldwater</a></li> <li><a href="/wiki/Coral_reef_fish" title="Coral reef fish">Coral reef</a></li> <li><a href="/wiki/Deep-sea_fish" title="Deep-sea fish">Deep-sea</a></li> <li><a href="/wiki/Demersal_fish" title="Demersal fish">Demersal</a></li> <li><a href="/wiki/Euryhaline" title="Euryhaline">Euryhaline</a></li> <li><a href="/wiki/Freshwater_fish" title="Freshwater fish">Freshwater</a></li> <li><a href="/wiki/Groundfish" class="mw-redirect" title="Groundfish">Groundfish</a></li> <li><a href="/wiki/Pelagic_fish" title="Pelagic fish">Pelagic</a></li> <li><a href="/wiki/Tropical_fish" title="Tropical fish">Tropical</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other types</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/Bait_fish" title="Bait fish">Bait</a></li> <li><a href="/wiki/Coarse_fishing" title="Coarse fishing">Coarse</a></li> <li><a href="/wiki/Diversity_of_fish" title="Diversity of fish">Diversity</a></li> <li><a href="/wiki/Game_fish" title="Game fish">Game</a></li> <li><a href="/wiki/Genetically_modified_fish" title="Genetically modified fish">Genetically modified</a></li> <li><a href="/wiki/Hallucinogenic_fish" title="Hallucinogenic fish">Hallucinogenic</a></li> <li><a href="/wiki/Oily_fish" title="Oily fish">Oily</a></li> <li><a href="/wiki/Poisonous_fish" title="Poisonous fish">Poisonous</a></li> <li><a href="/wiki/Rough_fish" title="Rough fish">Rough</a></li> <li><a href="/wiki/Whitefish_(fisheries_term)" title="Whitefish (fisheries term)">Whitefish</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/List_of_commercially_important_fish_species" title="List of commercially important fish species">Commerce</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Fish_farming" title="Fish farming">Farming</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/Carp#Aquaculture" title="Carp">Carp</a></li> <li><a href="/wiki/Aquaculture_of_catfish" title="Aquaculture of catfish">Catfish</a></li> <li><a href="/wiki/Octopus_aquaculture" title="Octopus aquaculture">Octopus</a></li> <li><a href="/wiki/Aquaculture_of_salmonids" title="Aquaculture of salmonids">Salmonids</a></li> <li><a href="/wiki/Aquaculture_of_tilapia" title="Aquaculture of tilapia">Tilapia</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Wild_fisheries" title="Wild fisheries">Wild fisheries</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/Predatory_fish" title="Predatory fish">Predatory</a> <ul><li><a href="/wiki/Billfish" title="Billfish">billfish</a></li> <li><a href="/wiki/Mackerel" title="Mackerel">mackerel</a></li> <li><a href="/wiki/Salmon" title="Salmon">salmon</a></li> <li><a href="/wiki/Tuna" title="Tuna">tuna</a></li></ul></li> <li><a href="/wiki/Forage_fish" title="Forage fish">Forage</a> <ul><li><a href="/wiki/Anchovy" title="Anchovy">anchovy</a></li> <li><a href="/wiki/Herring" title="Herring">herring</a></li> <li><a href="/wiki/Sardine" title="Sardine">sardine</a></li> <li><a href="/wiki/Sprat" title="Sprat">sprats</a></li></ul></li> <li><a href="/wiki/Demersal_fish" title="Demersal fish">Demersal</a> <ul><li><a href="/wiki/Cod" title="Cod">cod</a></li> <li><a href="/wiki/Flatfish" title="Flatfish">flatfish</a></li> <li><a href="/wiki/Pollock" title="Pollock">pollock</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Major groups</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/Agnatha" title="Agnatha">Jawless</a> <ul><li><a href="/wiki/Hagfish" title="Hagfish">hagfish</a></li> <li><a href="/wiki/Hyperoartia" class="mw-redirect" title="Hyperoartia">lampreys</a></li></ul></li> <li><a href="/wiki/Chondrichthyes" title="Chondrichthyes">Cartilaginous</a> <ul><li><a href="/wiki/Holocephali" title="Holocephali">chimaeras</a></li> <li><a href="/wiki/Shark" title="Shark">sharks</a></li> <li><a href="/wiki/Batoidea" class="mw-redirect" title="Batoidea">rays</a></li></ul></li> <li><a href="/wiki/Osteichthyes" title="Osteichthyes">Bony</a> <ul><li><a href="/wiki/Actinopterygii" title="Actinopterygii">spiny-finned</a></li> <li><a href="/wiki/Sarcopterygii" title="Sarcopterygii">fleshy-finned</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Lists</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/Lists_of_aquarium_life" title="Lists of aquarium life">Aquarium life</a></li> <li><a href="/wiki/Blind_fish" title="Blind fish">Blind</a></li> <li><a href="/wiki/List_of_fish_common_names" title="List of fish common names">Fish common names</a></li> <li><a href="/wiki/List_of_fish_families" title="List of fish families">Fish families</a></li> <li><a href="/wiki/Glossary_of_ichthyology" title="Glossary of ichthyology">Glossary of ichthyology</a></li> <li><a href="/wiki/List_of_largest_fish" title="List of largest fish">Largest</a></li> <li><a href="/wiki/List_of_smallest_fish" title="List of smallest fish">Smallest</a></li> <li>Threatened <ul><li><a href="/wiki/List_of_threatened_rays" title="List of threatened rays">rays</a></li> <li><a href="/wiki/List_of_threatened_sharks" title="List of threatened sharks">sharks</a></li></ul></li> <li><a href="/wiki/Lists_of_prehistoric_fish" title="Lists of prehistoric fish">Prehistoric</a></li> <li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Lists_of_fishes" title="Category:Lists of fishes">more lists...</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="3"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <b><a href="/wiki/Category:Fish" title="Category:Fish">Category</a></b></li> <li><span class="noviewer" typeof="mw:File"><span title="WikiProject"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/16px-People_icon.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/24px-People_icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/32px-People_icon.svg.png 2x" data-file-width="100" data-file-height="100" /></span></span> <b><a href="/wiki/Wikipedia:WikiProject_Fishes" title="Wikipedia:WikiProject Fishes">WikiProject</a></b></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Swarming" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Swarming" title="Template:Swarming"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Swarming" title="Template talk:Swarming"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Swarming" title="Special:EditPage/Template:Swarming"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Swarming" style="font-size:114%;margin:0 4em"><a href="/wiki/Swarm_behaviour" title="Swarm behaviour">Swarming</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Swarm_behaviour#Biological_swarming" title="Swarm behaviour">Biological swarming</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Agent-based_model_in_biology" title="Agent-based model in biology">Agent-based model in biology</a></li> <li><a href="/wiki/Collective_animal_behavior" title="Collective animal behavior">Collective animal behavior</a></li> <li><a href="/wiki/Droving" title="Droving">Droving</a></li> <li><a href="/wiki/Flock_(birds)" title="Flock (birds)">Flock</a> <ul><li><a href="/wiki/Flocking_(behavior)" class="mw-redirect" title="Flocking (behavior)">flocking</a></li> <li><a href="/wiki/Sort_sol" title="Sort sol">sort sol</a></li></ul></li> <li><a href="/wiki/Herd" title="Herd">Herd</a> <ul><li><a href="/wiki/Herd_behavior" title="Herd behavior">herd behavior</a></li></ul></li> <li><a href="/wiki/Locust" title="Locust">Locust</a></li> <li><a href="/wiki/Mixed-species_foraging_flock" title="Mixed-species foraging flock">Mixed-species foraging flock</a></li> <li><a href="/wiki/Mobbing_(animal_behavior)" title="Mobbing (animal behavior)">Mobbing behavior</a> <ul><li><a href="/wiki/Feeding_frenzy" title="Feeding frenzy">feeding frenzy</a></li></ul></li> <li><a href="/wiki/Pack_(canine)" title="Pack (canine)">Pack</a> <ul><li><a href="/wiki/Pack_hunter" title="Pack hunter">pack hunter</a></li></ul></li> <li><a href="/wiki/Patterns_of_self-organization_in_ants" title="Patterns of self-organization in ants">Patterns of self-organization in ants</a> <ul><li><a href="/wiki/Ant_mill" title="Ant mill">ant mill</a></li> <li><a href="/wiki/Symmetry_breaking_of_escaping_ants" title="Symmetry breaking of escaping ants">symmetry breaking of escaping ants</a></li></ul></li> <li><a class="mw-selflink selflink">Shoaling and schooling</a> <ul><li><a href="/wiki/Bait_ball" title="Bait ball">bait ball</a></li></ul></li> <li><a href="/wiki/Swarm_behaviour" title="Swarm behaviour">Swarming behaviour</a></li> <li><a href="/wiki/Swarming_(honey_bee)" title="Swarming (honey bee)">Swarming (honey bee)</a></li> <li><a href="/wiki/Swarming_motility" title="Swarming motility">Swarming motility</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="6" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Auklet_flock_Shumagins_1986.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5e/Auklet_flock_Shumagins_1986.jpg/140px-Auklet_flock_Shumagins_1986.jpg" decoding="async" width="140" height="95" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5e/Auklet_flock_Shumagins_1986.jpg/210px-Auklet_flock_Shumagins_1986.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5e/Auklet_flock_Shumagins_1986.jpg/280px-Auklet_flock_Shumagins_1986.jpg 2x" data-file-width="3756" data-file-height="2561" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Animal_migration" title="Animal migration">Animal migration</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Animal_migration" title="Animal migration">Animal migration</a> <ul><li><a href="/wiki/Altitudinal_migration" title="Altitudinal migration">altitudinal</a></li> <li><a href="/wiki/Animal_migration_tracking" title="Animal migration tracking">tracking</a> <ul><li><a href="/wiki/History_of_wildlife_tracking_technology" title="History of wildlife tracking technology">history</a></li></ul></li> <li><a href="/wiki/Coded_wire_tag" title="Coded wire tag">coded wire tag</a></li></ul></li> <li><a href="/wiki/Bird_migration" title="Bird migration">Bird migration</a> <ul><li><a href="/wiki/Category:Bird_migration_flyways" title="Category:Bird migration flyways">flyways</a></li> <li><a href="/wiki/Reverse_migration_(birds)" title="Reverse migration (birds)">reverse migration</a></li></ul></li> <li><a href="/wiki/Cell_migration" title="Cell migration">Cell migration</a></li> <li><a href="/wiki/Fish_migration" title="Fish migration">Fish migration</a> <ul><li><a href="/wiki/Diel_vertical_migration" title="Diel vertical migration">diel vertical</a></li> <li><a href="/wiki/Lessepsian_migration" title="Lessepsian migration">Lessepsian</a></li> <li><a href="/wiki/Salmon_run" title="Salmon run">salmon run</a></li> <li><a href="/wiki/Sardine_run" title="Sardine run">sardine run</a></li></ul></li> <li><a href="/wiki/Homing_(biology)" title="Homing (biology)">Homing</a> <ul><li><a href="/wiki/Natal_homing" title="Natal homing">natal</a></li> <li><a href="/wiki/Philopatry" title="Philopatry">philopatry</a></li></ul></li> <li><a href="/wiki/Insect_migration" title="Insect migration">Insect migration</a> <ul><li><a href="/wiki/Lepidoptera_migration" title="Lepidoptera migration">butterflies</a> <ul><li><a href="/wiki/Monarch_butterfly_migration" title="Monarch butterfly migration">monarch</a></li></ul></li></ul></li> <li><a href="/wiki/Sea_turtle_migration" title="Sea turtle migration">Sea turtle migration</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Swarm_behaviour#Models" title="Swarm behaviour">Swarm algorithms</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Agent-based_model" title="Agent-based model">Agent-based models</a></li> <li><a href="/wiki/Ant_colony_optimization_algorithm" class="mw-redirect" title="Ant colony optimization algorithm">Ant colony optimization</a></li> <li><a href="/wiki/Boids" title="Boids">Boids</a></li> <li><a href="/wiki/Crowd_simulation" title="Crowd simulation">Crowd simulation</a></li> <li><a href="/wiki/Particle_swarm_optimization" title="Particle swarm optimization">Particle swarm optimization</a></li> <li><a href="/wiki/Swarm_intelligence" title="Swarm intelligence">Swarm intelligence</a></li> <li><a href="/wiki/Swarm_(simulation)" title="Swarm (simulation)">Swarm (simulation)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Collective_motion" title="Collective motion">Collective motion</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Active_matter" title="Active matter">Active matter</a></li> <li><a href="/wiki/Collective_motion" title="Collective motion">Collective motion</a></li> <li><a href="/wiki/Self-propelled_particles" title="Self-propelled particles">Self-propelled particles</a> <ul><li><a href="/wiki/Clustering_of_self-propelled_particles" title="Clustering of self-propelled particles">clustering</a></li></ul></li> <li><a href="/wiki/Vicsek_model" title="Vicsek model">Vicsek model</a></li> <li><a href="/wiki/BIO-LGCA" title="BIO-LGCA">BIO-LGCA</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Swarm_robotics" title="Swarm robotics">Swarm robotics</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ant_robotics" title="Ant robotics">Ant robotics</a></li> <li><a href="/wiki/Microbotics" title="Microbotics">Microbotics</a></li> <li><a href="/wiki/Nanorobotics" title="Nanorobotics">Nanorobotics</a></li> <li><a href="/wiki/Swarm_robotics" title="Swarm robotics">Swarm robotics</a></li> <li><a href="/wiki/Symbrion" title="Symbrion">Symbrion</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related topics</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allee_effect" title="Allee effect">Allee effect</a></li> <li><a href="/wiki/Animal_navigation" title="Animal navigation">Animal navigation</a></li> <li><a href="/wiki/Collective_intelligence" title="Collective intelligence">Collective intelligence</a></li> <li><a href="/wiki/Decentralised_system" title="Decentralised system">Decentralised system</a></li> <li><a href="/wiki/Eusociality" title="Eusociality">Eusociality</a></li> <li><a href="/wiki/Group_size_measures" title="Group size measures">Group size measures</a></li> <li><a href="/wiki/Microbial_intelligence" title="Microbial intelligence">Microbial intelligence</a></li> <li><a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li> <li><a href="/wiki/Predator_satiation" title="Predator satiation">Predator satiation</a></li> <li><a href="/wiki/Quorum_sensing" title="Quorum sensing">Quorum sensing</a></li> <li><a href="/wiki/Spatial_organization" title="Spatial organization">Spatial organization</a></li> <li><a href="/wiki/Stigmergy" title="Stigmergy">Stigmergy</a></li> <li><a href="/wiki/Swarming_(military)" title="Swarming (military)">Military swarming</a></li> <li><a href="/wiki/Task_allocation_and_partitioning_of_social_insects" class="mw-redirect" title="Task allocation and partitioning of social insects">Task allocation and partitioning of social insects</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Fishery_science_topics" style="padding:3px"><table class="nowraplinks 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:Fishery_science_topics" title="Template:Fishery science topics"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Fishery_science_topics" title="Template talk:Fishery science topics"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Fishery_science_topics" title="Special:EditPage/Template:Fishery science topics"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Fishery_science_topics" style="font-size:114%;margin:0 4em"><a href="/wiki/Fishery" title="Fishery">Fishery</a> science topics</div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd hlist" 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="Fisheries_science_and_wild_fisheries" style="font-size:114%;margin:0 4em"><a href="/wiki/Fisheries_science" title="Fisheries science">Fisheries science</a> and <a href="/wiki/Wild_fisheries" title="Wild fisheries">wild fisheries</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:7em"><a href="/wiki/Fisheries_science" title="Fisheries science">Fisheries<br />science</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/Population_dynamics_of_fisheries" title="Population dynamics of fisheries">Population dynamics of fisheries</a></li> <li><a href="/wiki/Shifting_baseline" title="Shifting baseline">Shifting baseline</a></li> <li><a href="/wiki/Fish_stocks" title="Fish stocks">Fish stock</a></li> <li><a href="/wiki/Fish_mortality" title="Fish mortality">Fish mortality</a></li> <li><a href="/wiki/Condition_index_in_fish" title="Condition index in fish">Condition index</a></li> <li><a href="/wiki/Stock_assessment" title="Stock assessment">Stock assessment</a></li> <li><a href="/wiki/Fish_measurement" title="Fish measurement">Fish measurement</a></li> <li><a href="/wiki/Standard_weight_in_fish" title="Standard weight in fish">Standard weight in fish</a></li> <li><a href="/wiki/Fish_counter" title="Fish counter">Fish counter</a></li> <li><a href="/wiki/Data_storage_tag" title="Data storage tag">Data storage tag</a></li> <li><a href="/wiki/Catch_per_unit_effort" title="Catch per unit effort">Catch per unit effort</a></li> <li><a href="/wiki/Otolith_microchemical_analysis" title="Otolith microchemical analysis">Otolith microchemical analysis</a></li> <li><a href="/wiki/Biomass_(ecology)" title="Biomass (ecology)">Biomass</a></li> <li><a href="/wiki/Fisheries_acoustics" title="Fisheries acoustics">Fisheries acoustics</a></li> <li><a href="/wiki/Acoustic_tag" title="Acoustic tag">Acoustic tag</a></li> <li><a href="/wiki/Coded_wire_tag" title="Coded wire tag">Coded wire tag</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/Pop-up_satellite_archival_tag" title="Pop-up satellite archival tag">Pop-up satellite archival tag</a></li> <li><a href="/wiki/EcoSCOPE" title="EcoSCOPE">EcoSCOPE</a></li> <li><a href="/wiki/Age_class_structure" title="Age class structure">Age class structure</a></li> <li><a href="/wiki/Trophic_level" title="Trophic level">Trophic level</a></li> <li><a href="/wiki/Trophic_cascade" title="Trophic cascade">Trophic cascades</a></li> <li><a href="/wiki/Hydrographic_containment" title="Hydrographic containment">Hydrographic containment</a></li> <li><a href="/wiki/Match/mismatch" title="Match/mismatch">Match/mismatch hypothesis</a></li> <li><a href="/wiki/Climate_change_and_fisheries" title="Climate change and fisheries">Climate change and fisheries</a></li> <li><a href="/wiki/Marine_biology" title="Marine biology">Marine biology</a></li> <li><a href="/wiki/Aquatic_ecosystem" title="Aquatic ecosystem">Aquatic ecosystems</a></li> <li><a href="/wiki/Bioeconomics_(fisheries)" title="Bioeconomics (fisheries)">Bioeconomics</a></li> <li><a href="/wiki/EconMult" title="EconMult">EconMult</a></li> <li><a href="/wiki/Ecopath" title="Ecopath">Ecopath</a></li> <li><a href="/wiki/FishBase" title="FishBase">FishBase</a></li> <li><a href="/wiki/Census_of_Marine_Life" title="Census of Marine Life">Census of Marine Life</a></li> <li><a href="/wiki/OSTM/Jason-2" title="OSTM/Jason-2">OSTM/Jason-2</a></li> <li><a href="/wiki/Category:Fisheries_databases" title="Category:Fisheries databases">Fisheries databases</a></li> <li><a href="/wiki/Category:Fisheries_and_aquaculture_research_institutes" title="Category:Fisheries and aquaculture research institutes">Institutes</a></li> <li><a href="/wiki/Fisheries_science#Notable_contributors" title="Fisheries science">Fisheries scientists</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="2" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Great_Barracuda_off_the_Netherland_Antilles.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/81/Great_Barracuda_off_the_Netherland_Antilles.jpg/140px-Great_Barracuda_off_the_Netherland_Antilles.jpg" decoding="async" width="140" height="105" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/81/Great_Barracuda_off_the_Netherland_Antilles.jpg/210px-Great_Barracuda_off_the_Netherland_Antilles.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/81/Great_Barracuda_off_the_Netherland_Antilles.jpg/280px-Great_Barracuda_off_the_Netherland_Antilles.jpg 2x" data-file-width="600" data-file-height="450" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em"><a href="/wiki/Wild_fisheries" title="Wild fisheries">Wild<br />fisheries</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/Ocean_fisheries" title="Ocean fisheries">Ocean fisheries</a></li> <li><a href="/wiki/Diversity_of_fish" title="Diversity of fish">Diversity of fish</a></li> <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/Demersal_fish" title="Demersal fish">Demersal fish</a></li> <li><a href="/wiki/Forage_fish" title="Forage fish">Forage fish</a></li> <li><a href="/wiki/Pelagic_fish" title="Pelagic fish">Pelagic fish</a></li> <li><a href="/wiki/Cod_fisheries" title="Cod fisheries">Cod fisheries</a></li> <li><a href="/wiki/Crab_fisheries" title="Crab fisheries">Crab fisheries</a></li> <li><a href="/wiki/Eel_life_history" title="Eel life history">Eel fisheries</a></li> <li><a href="/wiki/Krill_fishery" title="Krill fishery">Krill fisheries</a></li> <li><a href="/wiki/Kelp#Kelp_in_history_and_culture" title="Kelp">Kelp fisheries</a></li> <li><a href="/wiki/Lobster_fishing" title="Lobster fishing">Lobster fisheries</a></li> <li><a href="/wiki/Shrimp_fishery" title="Shrimp fishery">Shrimp fishery</a></li> <li><a href="/wiki/Eel_ladder" class="mw-redirect" title="Eel ladder">Eel ladder</a></li> <li><a href="/wiki/Fish_ladder" title="Fish ladder">Fish ladder</a></li> <li><a href="/wiki/Fish_screen" title="Fish screen">Fish screen</a></li> <li><a href="/wiki/Fish_migration" title="Fish migration">Migration</a></li> <li><a href="/wiki/Salmon_run" title="Salmon run">Salmon run</a></li> <li><a href="/wiki/Sardine_run" title="Sardine run">Sardine run</a></li> <li><a class="mw-selflink selflink">Shoaling and schooling</a></li> <li><a href="/wiki/Marine_habitat" title="Marine habitat">Marine habitat</a></li> <li><a href="/wiki/Marine_snow" title="Marine snow">Marine snow</a></li> <li><a href="/wiki/Water_column" title="Water column">Water column</a></li> <li><a href="/wiki/Upwelling" title="Upwelling">Upwelling</a></li> <li><a href="/wiki/Humboldt_Current" title="Humboldt Current">Humboldt current</a></li> <li><a href="/wiki/Algal_bloom" title="Algal bloom">Algal blooms</a></li> <li><a href="/wiki/Dead_zone_(ecology)" title="Dead zone (ecology)">Dead zones</a></li> <li><a href="/wiki/Fish_kill" title="Fish kill">Fish kill</a></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 hlist" 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="Fisheries_law,_management,_sustainability_and_conservation" style="font-size:114%;margin:0 4em"><a href="/wiki/Fisheries_law" title="Fisheries law">Fisheries law</a>, <a href="/wiki/Fisheries_management" title="Fisheries management">management</a>, <a href="/wiki/Sustainable_fishery" title="Sustainable fishery">sustainability</a> and <a href="/wiki/Marine_conservation" title="Marine conservation">conservation</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:7em">Law</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/Fisheries_law" title="Fisheries law">Fisheries law</a></li> <li><a href="/wiki/Exclusive_economic_zone" title="Exclusive economic zone">Exclusive economic zone</a></li> <li><a href="/wiki/Illegal,_unreported_and_unregulated_fishing" title="Illegal, unreported and unregulated fishing">Illegal, unreported and unregulated fishing</a></li> <li><a href="/wiki/United_Nations_Convention_on_the_Law_of_the_Sea" title="United Nations Convention on the Law of the Sea">United Nations Convention on the Law of the Sea</a> <ul><li><a href="/wiki/United_Nations_Fish_Stocks_Agreement" title="United Nations Fish Stocks Agreement">United Nations Fish Stocks Agreement</a></li></ul></li> <li><a href="/wiki/Fisheries_Convention" title="Fisheries Convention">Fisheries Convention</a></li> <li><a href="/wiki/Magnuson%E2%80%93Stevens_Fishery_Conservation_and_Management_Act" title="Magnuson–Stevens Fishery Conservation and Management Act">Magnuson–Stevens Act</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="6" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Yellowfin_tuna_nurp.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Yellowfin_tuna_nurp.jpg/140px-Yellowfin_tuna_nurp.jpg" decoding="async" width="140" height="99" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Yellowfin_tuna_nurp.jpg/210px-Yellowfin_tuna_nurp.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Yellowfin_tuna_nurp.jpg/280px-Yellowfin_tuna_nurp.jpg 2x" data-file-width="1558" data-file-height="1106" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em">Management</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/Fisheries_management" title="Fisheries management">Fisheries management</a></li> <li><a href="/wiki/Fisheries_monitoring_control_and_surveillance" title="Fisheries monitoring control and surveillance">Monitoring control and surveillance</a></li> <li><a href="/wiki/Vessel_monitoring_system" title="Vessel monitoring system">Vessel monitoring system</a></li> <li><a href="/wiki/Fishery_Resources_Monitoring_System" title="Fishery Resources Monitoring System">Fishery Resources Monitoring System</a></li> <li><a href="/wiki/Catch_reporting" title="Catch reporting">Catch reporting</a></li> <li><a href="/wiki/Fisheries_observer" title="Fisheries observer">Fisheries observer</a></li> <li><a href="/wiki/Catch_share" title="Catch share">Catch share</a></li> <li><a href="/wiki/Individual_fishing_quota" title="Individual fishing quota">Individual fishing quota</a></li> <li><a href="/wiki/Minimum_landing_size" title="Minimum landing size">Minimum landing size</a></li> <li><a href="/wiki/Pulse_fishing" title="Pulse fishing">Pulse fishing</a></li> <li><a href="/wiki/Slot_limit" title="Slot limit">Slot limit</a></li> <li><a href="/wiki/Common_Fisheries_Policy" title="Common Fisheries Policy">EU Common Fisheries Policy</a></li> <li><a href="/wiki/European_Fishery_MLS" title="European Fishery MLS">EU MLS</a></li> <li><a href="/wiki/European_Fisheries_Control_Agency" title="European Fisheries Control Agency">European Fisheries Control Agency</a></li> <li><a href="/wiki/National_Marine_Fisheries_Service" title="National Marine Fisheries Service">National Marine Fisheries Service</a></li> <li><a href="/wiki/Regional_fishery_body" title="Regional fishery body">Regional fishery body</a> <ul><li><a href="/wiki/Regional_fishery_management_organisation" class="mw-redirect" title="Regional fishery management organisation">Regional fishery management organisation</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em">Sustainability</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/Sustainable_fishery" title="Sustainable fishery">Sustainable fishery</a></li> <li><a href="/wiki/Maximum_sustainable_yield" title="Maximum sustainable yield">Maximum sustainable yield</a></li> <li><a href="/wiki/List_of_commercially_important_fish_species" title="List of commercially important fish species">List of commercially important fish species</a></li> <li><a href="/wiki/Sustainable_seafood" title="Sustainable seafood">Sustainable seafood</a></li> <li><a href="/wiki/Overfishing" title="Overfishing">Overfishing</a></li> <li><a href="/wiki/Environmental_impact_of_fishing" title="Environmental impact of fishing">Environmental impact of fishing</a></li> <li><a href="/wiki/Fishing_down_the_food_web" title="Fishing down the food web">Fishing down the food web</a></li> <li><a href="/wiki/Destructive_fishing_practices" title="Destructive fishing practices">Destructive fishing practices</a></li> <li><a href="/wiki/Future_of_Marine_Animal_Populations" title="Future of Marine Animal Populations">Future of Marine Animal Populations</a></li> <li><i><a href="/wiki/The_Sunken_Billions" title="The Sunken Billions">The Sunken Billions</a></i></li> <li><i><a href="/wiki/The_End_of_the_Line_(book)" title="The End of the Line (book)">The End of the Line</a></i></li> <li><a href="/wiki/Bycatch" title="Bycatch">Bycatch</a></li> <li><a href="/wiki/Discards" title="Discards">Discards</a></li> <li><a href="/wiki/Incidental_catch" title="Incidental catch">Incidental catch</a></li> <li><a href="/wiki/Cetacean_bycatch" title="Cetacean bycatch">Cetacean bycatch</a></li> <li><a href="/wiki/Turtle_excluder_device" title="Turtle excluder device">Turtle excluder device</a></li> <li><a href="/wiki/Shrimp-Turtle_Case" title="Shrimp-Turtle Case"><i>Shrimp-Turtle</i> case</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em">Conservation</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/Marine_protected_area" title="Marine protected area">Marine protected area</a></li> <li><a href="/wiki/Marine_reserve" title="Marine reserve">Marine reserve</a></li> <li><a href="/wiki/Marine_conservation" title="Marine conservation">Marine conservation</a></li> <li><a href="/wiki/Marine_conservation_activism" title="Marine conservation activism">Marine conservation activism</a></li> <li><a href="/wiki/Salmon_conservation" title="Salmon conservation">Salmon conservation</a></li> <li><a href="/wiki/Grey_nurse_shark_conservation" title="Grey nurse shark conservation">Grey nurse shark conservation</a></li> <li><a href="/wiki/Shark_sanctuary" title="Shark sanctuary">Shark sanctuary</a></li> <li><a href="/wiki/European_Environmental_Agency" class="mw-redirect" title="European Environmental Agency">European Environmental Agency</a></li> <li><a href="/wiki/United_States_Fish_and_Wildlife_Service" title="United States Fish and Wildlife Service">U.S. Fish and Wildlife Service</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em">Advocacy</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/FishAct" title="FishAct">FishAct</a></li> <li><a href="/wiki/Marine_Stewardship_Council" title="Marine Stewardship Council">Marine Stewardship Council</a></li> <li><a href="/wiki/Aquaculture_Stewardship_Council" title="Aquaculture Stewardship Council">Aquaculture Stewardship Council</a></li> <li><a href="/wiki/Friend_of_the_Sea" title="Friend of the Sea">Friend of the Sea</a></li> <li><a href="/wiki/SeaChoice" title="SeaChoice">SeaChoice</a></li> <li><a href="/wiki/Seafood_Watch" title="Seafood Watch">Seafood Watch</a></li> <li><a href="/wiki/Ocean_Conservancy" title="Ocean Conservancy">Ocean Conservancy</a></li> <li><a href="/wiki/Ocean_Outcomes" title="Ocean Outcomes">Ocean Outcomes</a></li> <li><a href="/wiki/Oceana_(non-profit_group)" title="Oceana (non-profit group)">Oceana</a></li> <li><a href="/wiki/Save_Our_Seas_Foundation" title="Save Our Seas Foundation">Save Our Seas Foundation</a></li> <li><a href="/wiki/Sea_Around_Us_(organization)" title="Sea Around Us (organization)">Sea Around Us</a></li> <li><a href="/wiki/WorldFish" title="WorldFish">WorldFish</a></li> <li><a href="/wiki/Defying_Ocean%27s_End" title="Defying Ocean's End">Defying Ocean's End</a></li> <li><a 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