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<span>History</span> </div> </a> <button aria-controls="toc-History-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 History subsection</span> </button> <ul id="toc-History-sublist" class="vector-toc-list"> <li id="toc-The_beginnings_of_ethology" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#The_beginnings_of_ethology"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>The beginnings of ethology</span> </div> </a> <ul id="toc-The_beginnings_of_ethology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Growth_of_the_field" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Growth_of_the_field"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Growth of the field</span> </div> </a> <ul id="toc-Growth_of_the_field-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Social_ethology" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Social_ethology"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Social ethology</span> </div> </a> <ul id="toc-Social_ethology-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Determinants_of_behaviour" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Determinants_of_behaviour"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Determinants of behaviour</span> </div> </a> <button aria-controls="toc-Determinants_of_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 Determinants of behaviour subsection</span> </button> <ul id="toc-Determinants_of_behaviour-sublist" class="vector-toc-list"> <li id="toc-Instinct" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Instinct"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Instinct</span> </div> </a> <ul id="toc-Instinct-sublist" class="vector-toc-list"> <li id="toc-Fixed_action_patterns" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Fixed_action_patterns"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1.1</span> <span>Fixed action patterns</span> </div> </a> <ul id="toc-Fixed_action_patterns-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Learning" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Learning"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Learning</span> </div> </a> <ul id="toc-Learning-sublist" class="vector-toc-list"> <li id="toc-Habituation" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Habituation"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.1</span> <span>Habituation</span> </div> </a> <ul id="toc-Habituation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Associative_learning" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Associative_learning"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.2</span> <span>Associative learning</span> </div> </a> <ul id="toc-Associative_learning-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Imprinting" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Imprinting"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.3</span> <span>Imprinting</span> </div> </a> <ul id="toc-Imprinting-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cultural_learning" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Cultural_learning"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.4</span> <span>Cultural learning</span> </div> </a> <ul id="toc-Cultural_learning-sublist" class="vector-toc-list"> <li id="toc-Observational_learning" class="vector-toc-list-item vector-toc-level-4"> <a class="vector-toc-link" href="#Observational_learning"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.4.1</span> <span>Observational learning</span> </div> </a> <ul id="toc-Observational_learning-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Imitation" class="vector-toc-list-item vector-toc-level-4"> <a class="vector-toc-link" href="#Imitation"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.4.2</span> <span>Imitation</span> </div> </a> <ul id="toc-Imitation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Stimulus_and_local_enhancement" class="vector-toc-list-item vector-toc-level-4"> <a class="vector-toc-link" href="#Stimulus_and_local_enhancement"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.4.3</span> <span>Stimulus and local enhancement</span> </div> </a> <ul id="toc-Stimulus_and_local_enhancement-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Social_transmission" class="vector-toc-list-item vector-toc-level-4"> <a class="vector-toc-link" href="#Social_transmission"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.4.4</span> <span>Social transmission</span> </div> </a> <ul id="toc-Social_transmission-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Teaching" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Teaching"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.5</span> <span>Teaching</span> </div> </a> <ul id="toc-Teaching-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Mating_and_the_fight_for_supremacy" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Mating_and_the_fight_for_supremacy"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Mating and the fight for supremacy</span> </div> </a> <ul id="toc-Mating_and_the_fight_for_supremacy-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Social_behaviour" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Social_behaviour"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Social behaviour</span> </div> </a> <button aria-controls="toc-Social_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 Social behaviour subsection</span> </button> <ul id="toc-Social_behaviour-sublist" class="vector-toc-list"> <li id="toc-Benefits_and_costs_of_group_living" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Benefits_and_costs_of_group_living"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Benefits and costs of group living</span> </div> </a> <ul id="toc-Benefits_and_costs_of_group_living-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Group_size" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Group_size"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Group size</span> </div> </a> <ul id="toc-Group_size-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Tinbergen's_four_questions_for_ethologists" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Tinbergen's_four_questions_for_ethologists"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Tinbergen's four questions for ethologists</span> </div> </a> <ul id="toc-Tinbergen's_four_questions_for_ethologists-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <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">10</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" title="Table of Contents" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Ethology</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 81 languages" > <label id="p-lang-btn-label" for="p-lang-btn-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--action-progressive mw-portlet-lang-heading-81" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">81 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/Etologie" title="Etologie – Afrikaans" lang="af" hreflang="af" data-title="Etologie" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target"><span>Afrikaans</span></a></li><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%B9%D9%84%D9%85_%D8%B3%D9%84%D9%88%D9%83_%D8%A7%D9%84%D8%AD%D9%8A%D9%88%D8%A7%D9%86" title="علم سلوك الحيوان – Arabic" lang="ar" hreflang="ar" data-title="علم سلوك الحيوان" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-as mw-list-item"><a href="https://as.wikipedia.org/wiki/%E0%A6%87%E0%A6%A5%E2%80%99%E0%A6%B2%E0%A6%9C%E0%A7%80" title="ইথ’লজী – Assamese" lang="as" hreflang="as" data-title="ইথ’লজী" data-language-autonym="অসমীয়া" data-language-local-name="Assamese" class="interlanguage-link-target"><span>অসমীয়া</span></a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/Etolox%C3%ADa" title="Etoloxía – Asturian" lang="ast" hreflang="ast" data-title="Etoloxía" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target"><span>Asturianu</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Etologiya" title="Etologiya – Azerbaijani" lang="az" hreflang="az" data-title="Etologiya" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/H%C3%AAng-%C3%BBi-ha%CC%8Dk" title="Hêng-ûi-ha̍k – Minnan" lang="nan" hreflang="nan" data-title="Hêng-ûi-ha̍k" data-language-autonym="閩南語 / Bân-lâm-gú" data-language-local-name="Minnan" class="interlanguage-link-target"><span>閩南語 / Bân-lâm-gú</span></a></li><li class="interlanguage-link interwiki-ba mw-list-item"><a href="https://ba.wikipedia.org/wiki/%D0%AD%D1%82%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F" title="Этология – Bashkir" lang="ba" hreflang="ba" data-title="Этология" data-language-autonym="Башҡортса" data-language-local-name="Bashkir" class="interlanguage-link-target"><span>Башҡортса</span></a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%AD%D1%82%D0%B0%D0%BB%D0%BE%D0%B3%D1%96%D1%8F" title="Эталогія – Belarusian" lang="be" hreflang="be" data-title="Эталогія" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target"><span>Беларуская</span></a></li><li class="interlanguage-link interwiki-be-x-old mw-list-item"><a href="https://be-tarask.wikipedia.org/wiki/%D0%AD%D1%82%D0%B0%D0%BB%D1%91%D0%B3%D1%96%D1%8F" title="Эталёгія – Belarusian (Taraškievica orthography)" lang="be-tarask" hreflang="be-tarask" data-title="Эталёгія" data-language-autonym="Беларуская (тарашкевіца)" data-language-local-name="Belarusian (Taraškievica orthography)" class="interlanguage-link-target"><span>Беларуская (тарашкевіца)</span></a></li><li class="interlanguage-link interwiki-bcl mw-list-item"><a href="https://bcl.wikipedia.org/wiki/Etolohiya" title="Etolohiya – Central Bikol" lang="bcl" hreflang="bcl" data-title="Etolohiya" data-language-autonym="Bikol Central" data-language-local-name="Central Bikol" class="interlanguage-link-target"><span>Bikol Central</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%95%D1%82%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F" title="Етология – Bulgarian" lang="bg" hreflang="bg" data-title="Етология" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Etologija" title="Etologija – Bosnian" lang="bs" hreflang="bs" data-title="Etologija" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Etologia" title="Etologia – Catalan" lang="ca" hreflang="ca" data-title="Etologia" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Etologie" title="Etologie – Czech" lang="cs" hreflang="cs" data-title="Etologie" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Etologi" title="Etologi – Danish" lang="da" hreflang="da" data-title="Etologi" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Ethologie" title="Ethologie – German" lang="de" hreflang="de" data-title="Ethologie" 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/Etoloogia" title="Etoloogia – Estonian" lang="et" hreflang="et" data-title="Etoloogia" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%97%CE%B8%CE%BF%CE%BB%CE%BF%CE%B3%CE%AF%CE%B1" title="Ηθολογία – Greek" lang="el" hreflang="el" data-title="Ηθολογία" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Etolog%C3%ADa" title="Etología – Spanish" lang="es" hreflang="es" data-title="Etología" 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/Etologio" title="Etologio – Esperanto" lang="eo" hreflang="eo" data-title="Etologio" 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/Etologia" title="Etologia – Basque" lang="eu" hreflang="eu" data-title="Etologia" 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%B1%D9%81%D8%AA%D8%A7%D8%B1%D8%B4%D9%86%D8%A7%D8%B3%DB%8C_%D8%AC%D8%A7%D9%86%D9%88%D8%B1%D8%A7%D9%86" title="رفتارشناسی جانوران – Persian" lang="fa" hreflang="fa" data-title="رفتارشناسی جانوران" 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/%C3%89thologie" title="Éthologie – French" lang="fr" hreflang="fr" data-title="Éthologie" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-fy mw-list-item"><a href="https://fy.wikipedia.org/wiki/Gedrachsbiology" title="Gedrachsbiology – Western Frisian" lang="fy" hreflang="fy" data-title="Gedrachsbiology" data-language-autonym="Frysk" data-language-local-name="Western Frisian" class="interlanguage-link-target"><span>Frysk</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Eiteola%C3%ADocht" title="Eiteolaíocht – Irish" lang="ga" hreflang="ga" data-title="Eiteolaíocht" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Etolox%C3%ADa" title="Etoloxía – Galician" lang="gl" hreflang="gl" data-title="Etoloxía" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EB%8F%99%EB%AC%BC%ED%96%89%EB%8F%99%ED%95%99" title="동물행동학 – Korean" lang="ko" hreflang="ko" data-title="동물행동학" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D4%B7%D5%A9%D5%B8%D5%AC%D5%B8%D5%A3%D5%AB%D5%A1" title="Էթոլոգիա – Armenian" lang="hy" hreflang="hy" data-title="Էթոլոգիա" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Etologija" title="Etologija – Croatian" lang="hr" hreflang="hr" data-title="Etologija" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-io mw-list-item"><a href="https://io.wikipedia.org/wiki/Etologio" title="Etologio – Ido" lang="io" hreflang="io" data-title="Etologio" data-language-autonym="Ido" data-language-local-name="Ido" class="interlanguage-link-target"><span>Ido</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Etologi" title="Etologi – Indonesian" lang="id" hreflang="id" data-title="Etologi" 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-ia mw-list-item"><a href="https://ia.wikipedia.org/wiki/Ethologia" title="Ethologia – Interlingua" lang="ia" hreflang="ia" data-title="Ethologia" data-language-autonym="Interlingua" data-language-local-name="Interlingua" class="interlanguage-link-target"><span>Interlingua</span></a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Atferlisfr%C3%A6%C3%B0i" title="Atferlisfræði – Icelandic" lang="is" hreflang="is" data-title="Atferlisfræði" 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/Etologia" title="Etologia – Italian" lang="it" hreflang="it" data-title="Etologia" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%90%D7%AA%D7%95%D7%9C%D7%95%D7%92%D7%99%D7%94" title="אתולוגיה – Hebrew" lang="he" hreflang="he" data-title="אתולוגיה" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%94%E1%83%97%E1%83%9D%E1%83%9A%E1%83%9D%E1%83%92%E1%83%98%E1%83%90" title="ეთოლოგია – Georgian" lang="ka" hreflang="ka" data-title="ეთოლოგია" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%AD%D1%82%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F" title="Этология – Kazakh" lang="kk" hreflang="kk" data-title="Этология" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ht mw-list-item"><a href="https://ht.wikipedia.org/wiki/Etoloji" title="Etoloji – Haitian Creole" lang="ht" hreflang="ht" data-title="Etoloji" data-language-autonym="Kreyòl ayisyen" data-language-local-name="Haitian Creole" class="interlanguage-link-target"><span>Kreyòl ayisyen</span></a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%AD%D1%82%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F" title="Этология – Kyrgyz" lang="ky" hreflang="ky" data-title="Этология" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target"><span>Кыргызча</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Ethologia" title="Ethologia – Latin" lang="la" hreflang="la" data-title="Ethologia" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Etolo%C4%A3ija" title="Etoloģija – Latvian" lang="lv" hreflang="lv" data-title="Etoloģija" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target"><span>Latviešu</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Etologija" title="Etologija – Lithuanian" lang="lt" hreflang="lt" data-title="Etologija" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-li mw-list-item"><a href="https://li.wikipedia.org/wiki/Ethologie" title="Ethologie – Limburgish" lang="li" hreflang="li" data-title="Ethologie" data-language-autonym="Limburgs" data-language-local-name="Limburgish" class="interlanguage-link-target"><span>Limburgs</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Etol%C3%B3gia" title="Etológia – Hungarian" lang="hu" hreflang="hu" data-title="Etológia" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%8E%E0%B4%A4%E0%B5%8D%E0%B4%A4%E0%B5%8B%E0%B4%B3%E0%B4%9C%E0%B4%BF" title="എത്തോളജി – Malayalam" lang="ml" hreflang="ml" data-title="എത്തോളജി" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-xmf mw-list-item"><a href="https://xmf.wikipedia.org/wiki/%E1%83%94%E1%83%97%E1%83%9D%E1%83%9A%E1%83%9D%E1%83%92%E1%83%98%E1%83%90" title="ეთოლოგია – Mingrelian" lang="xmf" hreflang="xmf" data-title="ეთოლოგია" data-language-autonym="მარგალური" data-language-local-name="Mingrelian" class="interlanguage-link-target"><span>მარგალური</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Etologi" title="Etologi – Malay" lang="ms" hreflang="ms" data-title="Etologi" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Ethologie" title="Ethologie – Dutch" lang="nl" hreflang="nl" data-title="Ethologie" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E5%8B%95%E7%89%A9%E8%A1%8C%E5%8B%95%E5%AD%A6" title="動物行動学 – Japanese" lang="ja" hreflang="ja" data-title="動物行動学" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Atferdsbiologi" title="Atferdsbiologi – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Atferdsbiologi" 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/%C3%85tferdsbiologi" title="Åtferdsbiologi – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Åtferdsbiologi" 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-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Etologia" title="Etologia – Occitan" lang="oc" hreflang="oc" data-title="Etologia" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Etologiya" title="Etologiya – Uzbek" lang="uz" hreflang="uz" data-title="Etologiya" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target"><span>Oʻzbekcha / ўзбекча</span></a></li><li class="interlanguage-link interwiki-pms mw-list-item"><a href="https://pms.wikipedia.org/wiki/Etolog%C3%ACa" title="Etologìa – Piedmontese" lang="pms" hreflang="pms" data-title="Etologìa" data-language-autonym="Piemontèis" data-language-local-name="Piedmontese" class="interlanguage-link-target"><span>Piemontèis</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Etologia" title="Etologia – Polish" lang="pl" hreflang="pl" data-title="Etologia" 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/Etologia" title="Etologia – Portuguese" lang="pt" hreflang="pt" data-title="Etologia" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Etologie" title="Etologie – Romanian" lang="ro" hreflang="ro" data-title="Etologie" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%AD%D1%82%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F" title="Этология – Russian" lang="ru" hreflang="ru" data-title="Этология" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/Etologjia" title="Etologjia – Albanian" lang="sq" hreflang="sq" data-title="Etologjia" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target"><span>Shqip</span></a></li><li class="interlanguage-link interwiki-scn mw-list-item"><a href="https://scn.wikipedia.org/wiki/Etulugg%C3%ACa" title="Etuluggìa – Sicilian" lang="scn" hreflang="scn" data-title="Etuluggìa" data-language-autonym="Sicilianu" data-language-local-name="Sicilian" class="interlanguage-link-target"><span>Sicilianu</span></a></li><li class="interlanguage-link interwiki-si mw-list-item"><a href="https://si.wikipedia.org/wiki/%E0%B6%A0%E0%B6%BB%E0%B7%8A%E0%B6%BA%E0%B7%8F_%E0%B7%80%E0%B7%92%E0%B6%AF%E0%B7%8A%E2%80%8D%E0%B6%BA%E0%B7%8F%E0%B7%80" title="චර්යා විද්‍යාව – Sinhala" lang="si" hreflang="si" data-title="චර්යා විද්‍යාව" data-language-autonym="සිංහල" data-language-local-name="Sinhala" class="interlanguage-link-target"><span>සිංහල</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Ethology" title="Ethology – Simple English" lang="en-simple" hreflang="en-simple" data-title="Ethology" 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-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Etol%C3%B3gia" title="Etológia – Slovak" lang="sk" hreflang="sk" data-title="Etológia" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Etologija" title="Etologija – Slovenian" lang="sl" hreflang="sl" data-title="Etologija" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/Etologija" title="Etologija – Serbian" lang="sr" hreflang="sr" data-title="Etologija" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Etologija" title="Etologija – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Etologija" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Etologia" title="Etologia – Finnish" lang="fi" hreflang="fi" data-title="Etologia" 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/Etologi" title="Etologi – Swedish" lang="sv" hreflang="sv" data-title="Etologi" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/Etolohiya" title="Etolohiya – Tagalog" lang="tl" hreflang="tl" data-title="Etolohiya" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target"><span>Tagalog</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%B5%E0%AE%BF%E0%AE%B2%E0%AE%99%E0%AF%8D%E0%AE%95%E0%AE%BF%E0%AE%A9_%E0%AE%A8%E0%AE%9F%E0%AE%A4%E0%AF%8D%E0%AE%A4%E0%AF%88%E0%AE%AF%E0%AE%BF%E0%AE%AF%E0%AE%B2%E0%AF%8D" title="விலங்கின நடத்தையியல் – Tamil" lang="ta" hreflang="ta" data-title="விலங்கின நடத்தையியல்" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-tt mw-list-item"><a href="https://tt.wikipedia.org/wiki/%D0%AD%D1%82%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F" title="Этология – Tatar" lang="tt" hreflang="tt" data-title="Этология" data-language-autonym="Татарча / tatarça" data-language-local-name="Tatar" class="interlanguage-link-target"><span>Татарча / tatarça</span></a></li><li class="interlanguage-link interwiki-tet mw-list-item"><a href="https://tet.wikipedia.org/wiki/Etolojia" title="Etolojia – Tetum" lang="tet" hreflang="tet" data-title="Etolojia" data-language-autonym="Tetun" data-language-local-name="Tetum" class="interlanguage-link-target"><span>Tetun</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%9E%E0%B8%A4%E0%B8%95%E0%B8%B4%E0%B8%81%E0%B8%A3%E0%B8%A3%E0%B8%A1%E0%B8%A7%E0%B8%B4%E0%B8%97%E0%B8%A2%E0%B8%B2" title="พฤติกรรมวิทยา – Thai" lang="th" hreflang="th" data-title="พฤติกรรมวิทยา" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Etoloji" title="Etoloji – Turkish" lang="tr" hreflang="tr" data-title="Etoloji" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%95%D1%82%D0%BE%D0%BB%D0%BE%D0%B3%D1%96%D1%8F" title="Етологія – Ukrainian" lang="uk" hreflang="uk" data-title="Етологія" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D8%AE%D9%84%D9%82%DB%8C%D8%A7%D8%AA" title="خلقیات – Urdu" lang="ur" hreflang="ur" data-title="خلقیات" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target"><span>اردو</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/T%E1%BA%ADp_t%C3%ADnh_h%E1%BB%8Dc" title="Tập tính học – Vietnamese" lang="vi" hreflang="vi" data-title="Tập tính học" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-war mw-list-item"><a href="https://war.wikipedia.org/wiki/Etolohiya" title="Etolohiya – Waray" lang="war" hreflang="war" data-title="Etolohiya" data-language-autonym="Winaray" data-language-local-name="Waray" class="interlanguage-link-target"><span>Winaray</span></a></li><li class="interlanguage-link interwiki-wuu mw-list-item"><a href="https://wuu.wikipedia.org/wiki/%E5%8A%A8%E7%89%A9%E8%A1%8C%E4%B8%BA%E5%AD%A6" title="动物行为学 – Wu" lang="wuu" hreflang="wuu" data-title="动物行为学" data-language-autonym="吴语" data-language-local-name="Wu" class="interlanguage-link-target"><span>吴语</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E5%8B%95%E7%89%A9%E8%A1%8C%E7%82%BA%E5%AD%B8" title="動物行為學 – Cantonese" lang="yue" 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href="/wiki/Ethnology" title="Ethnology">ethnology</a> or <a href="/wiki/Ecology" title="Ecology">ecology</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">"Animal behaviour" redirects here. For other uses, see <a href="/wiki/Animal_behaviour_(disambiguation)" class="mw-disambig" title="Animal behaviour (disambiguation)">Animal behaviour (disambiguation)</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">For the journal, see <a href="/wiki/Ethology_(journal)" title="Ethology (journal)"><i>Ethology</i> (journal)</a>.</div> <div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Scientific objective study of non-human animal behaviour</div> <p class="mw-empty-elt"> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Waggle_dance_photo.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/eb/Waggle_dance_photo.png/220px-Waggle_dance_photo.png" decoding="async" width="220" height="216" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/eb/Waggle_dance_photo.png/330px-Waggle_dance_photo.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/eb/Waggle_dance_photo.png/440px-Waggle_dance_photo.png 2x" data-file-width="645" data-file-height="632" /></a><figcaption><a href="/wiki/Honey_bee" title="Honey bee">Honeybee</a> workers perform the <a href="/wiki/Waggle_dance" title="Waggle dance">waggle dance</a> to indicate the range and direction of food.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Grebe_Courtship_(detail).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/07/Grebe_Courtship_%28detail%29.jpg/220px-Grebe_Courtship_%28detail%29.jpg" decoding="async" width="220" height="215" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/07/Grebe_Courtship_%28detail%29.jpg/330px-Grebe_Courtship_%28detail%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/07/Grebe_Courtship_%28detail%29.jpg/440px-Grebe_Courtship_%28detail%29.jpg 2x" data-file-width="1143" data-file-height="1116" /></a><figcaption><a href="/wiki/Great_crested_grebe" title="Great crested grebe">Great crested grebes</a> perform a complex synchronised <a href="/wiki/Courtship_display" title="Courtship display">courtship display</a>.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Fighting_impalas_edit2.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0b/Fighting_impalas_edit2.jpg/220px-Fighting_impalas_edit2.jpg" decoding="async" width="220" height="117" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0b/Fighting_impalas_edit2.jpg/330px-Fighting_impalas_edit2.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0b/Fighting_impalas_edit2.jpg/440px-Fighting_impalas_edit2.jpg 2x" data-file-width="2210" data-file-height="1173" /></a><figcaption>Male <a href="/wiki/Impala" title="Impala">impalas</a> fighting during the <a href="/wiki/Rut_(mammalian_reproduction)" title="Rut (mammalian reproduction)">rut</a> </figcaption></figure> <p><b>Ethology</b> is a branch of <a href="/wiki/Zoology" title="Zoology">zoology</a> that studies the <a href="/wiki/Behavior" title="Behavior">behaviour</a> of non-human <a href="/wiki/Animal" title="Animal">animals</a>. It has its scientific roots in the work of <a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> and of American and German <a href="/wiki/Ornithology" title="Ornithology">ornithologists</a> of the late 19th and early 20th century, including <a href="/wiki/Charles_Otis_Whitman" title="Charles Otis Whitman">Charles O. Whitman</a>, <a href="/wiki/Oskar_Heinroth" title="Oskar Heinroth">Oskar Heinroth</a>, and <a href="/wiki/Wallace_Craig" title="Wallace Craig">Wallace Craig</a>. The modern discipline of ethology is generally considered to have begun during the 1930s with the work of the Dutch biologist <a href="/wiki/Nikolaas_Tinbergen" title="Nikolaas Tinbergen">Nikolaas Tinbergen</a> and the Austrian biologists <a href="/wiki/Konrad_Lorenz" title="Konrad Lorenz">Konrad Lorenz</a> and <a href="/wiki/Karl_von_Frisch" title="Karl von Frisch">Karl von Frisch</a>, the three winners of the 1973 <a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a>. Ethology combines laboratory and field science, with a strong relation to <a href="/wiki/Neuroanatomy" title="Neuroanatomy">neuroanatomy</a>, <a href="/wiki/Ecology" title="Ecology">ecology</a>, and <a href="/wiki/Evolutionary_biology" title="Evolutionary biology">evolutionary biology</a>. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Etymology">Etymology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=1" title="Edit section: Etymology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The modern term <i>ethology</i> derives from the <a href="/wiki/Greek_language" title="Greek language">Greek language</a>: <a href="https://en.wiktionary.org/wiki/%E1%BC%A6%CE%B8%CE%BF%CF%82" class="extiw" title="wikt:ἦθος">ἦθος</a>, <i><a href="/wiki/Ethos" title="Ethos">ethos</a></i> meaning "character" and <span title="Ancient Greek (to 1453)-language text"><span lang="grc"><a href="https://en.wiktionary.org/wiki/-%CE%BB%CE%BF%CE%B3%CE%AF%CE%B1" class="extiw" title="wikt:-λογία">-λογία</a></span></span>, <i><a href="https://en.wiktionary.org/wiki/-logia" class="extiw" title="wikt:-logia">-logia</a></i> meaning "the study of". The term was first popularized by the American <a href="/wiki/Entomologist" class="mw-redirect" title="Entomologist">entomologist</a> <a href="/wiki/William_Morton_Wheeler" title="William Morton Wheeler">William Morton Wheeler</a> in 1902.<sup id="cite_ref-insectbehavior_1-0" class="reference"><a href="#cite_note-insectbehavior-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=2" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="The_beginnings_of_ethology">The beginnings of ethology</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=3" title="Edit section: The beginnings of ethology"><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:Charles_Darwin_1880.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/45/Charles_Darwin_1880.jpg/170px-Charles_Darwin_1880.jpg" decoding="async" width="170" height="233" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/45/Charles_Darwin_1880.jpg/255px-Charles_Darwin_1880.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/45/Charles_Darwin_1880.jpg/340px-Charles_Darwin_1880.jpg 2x" data-file-width="2986" data-file-height="4096" /></a><figcaption><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> (1809–1882) explored the expression of emotions in animals.</figcaption></figure> <p>Ethologists have been concerned particularly with the <a href="/wiki/Evolution" title="Evolution">evolution</a> of behaviour and its understanding in terms of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>. In one sense, the first modern ethologist was <a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a>, whose 1872 book <i><a href="/wiki/The_Expression_of_the_Emotions_in_Man_and_Animals" title="The Expression of the Emotions in Man and Animals">The Expression of the Emotions in Man and Animals</a></i> influenced many ethologists. He pursued his interest in behaviour by encouraging his protégé <a href="/wiki/George_Romanes" title="George Romanes">George Romanes</a>, who investigated animal learning and intelligence using an <a href="/wiki/Anthropomorphism" title="Anthropomorphism">anthropomorphic</a> method, <a href="/wiki/Anecdotal_cognitivism" title="Anecdotal cognitivism">anecdotal cognitivism</a>, that did not gain scientific support.<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>Other early ethologists, such as <a href="/wiki/Eug%C3%A8ne_Marais" title="Eugène Marais">Eugène Marais</a>, <a href="/wiki/Charles_Otis_Whitman" title="Charles Otis Whitman">Charles O. Whitman</a>, <a href="/wiki/Oskar_Heinroth" title="Oskar Heinroth">Oskar Heinroth</a>, <a href="/wiki/Wallace_Craig" title="Wallace Craig">Wallace Craig</a> and <a href="/wiki/Julian_Huxley" title="Julian Huxley">Julian Huxley</a>, instead concentrated on behaviours that can be called <a href="/wiki/Instinct" title="Instinct">instinctive</a> in that they occur in all members of a species under specified circumstances.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-insectbehavior_1-1" class="reference"><a href="#cite_note-insectbehavior-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Their starting point for studying the behaviour of a new species was to construct an <a href="/wiki/Ethogram" title="Ethogram">ethogram</a>, a description of the main types of behaviour with their frequencies of occurrence. This provided an objective, cumulative database of behaviour.<sup id="cite_ref-insectbehavior_1-2" class="reference"><a href="#cite_note-insectbehavior-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Growth_of_the_field">Growth of the field</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=4" title="Edit section: Growth of the field"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Due to the work of <a href="/wiki/Konrad_Lorenz" title="Konrad Lorenz">Konrad Lorenz</a> and <a href="/wiki/Niko_Tinbergen" class="mw-redirect" title="Niko Tinbergen">Niko Tinbergen</a>, ethology developed strongly in continental Europe during the years prior to <a href="/wiki/World_War_II" title="World War II">World War II</a>.<sup id="cite_ref-insectbehavior_1-3" class="reference"><a href="#cite_note-insectbehavior-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> After the war, Tinbergen moved to the <a href="/wiki/University_of_Oxford" title="University of Oxford">University of Oxford</a>, and ethology became stronger in the <a href="/wiki/United_Kingdom" title="United Kingdom">UK</a>, with the additional influence of <a href="/wiki/William_Homan_Thorpe" title="William Homan Thorpe">William Thorpe</a>, <a href="/wiki/Robert_Hinde" title="Robert Hinde">Robert Hinde</a>, and <a href="/wiki/Patrick_Bateson" title="Patrick Bateson">Patrick Bateson</a> at the <a href="/wiki/University_of_Cambridge" title="University of Cambridge">University of Cambridge</a>.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>Lorenz, Tinbergen, and von Frisch were jointly awarded the <a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a> in 1973 for their work of developing ethology.<sup id="cite_ref-Nobelprize_6-0" class="reference"><a href="#cite_note-Nobelprize-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>Ethology is now a well-recognized scientific discipline, with its own journals such as <i><a href="/wiki/Animal_Behaviour_(journal)" title="Animal Behaviour (journal)">Animal Behaviour</a></i>, <i><a href="/wiki/Applied_Animal_Behaviour_Science" class="mw-redirect" title="Applied Animal Behaviour Science">Applied Animal Behaviour Science</a></i>, <i><a href="/wiki/Animal_Cognition" title="Animal Cognition">Animal Cognition</a></i>, <i><a href="/wiki/Behaviour_(journal)" title="Behaviour (journal)">Behaviour</a></i>, <i><a href="/wiki/Behavioral_Ecology_(journal)" title="Behavioral Ecology (journal)">Behavioral Ecology</a></i> and <i><a href="/wiki/Ethology_(journal)" title="Ethology (journal)">Ethology</a></i>. In 1972, the <a href="/wiki/International_Society_for_Human_Ethology" title="International Society for Human Ethology">International Society for Human Ethology</a> was founded along with its journal, <i>Human Ethology</i>.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Social_ethology">Social ethology</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=5" title="Edit section: Social ethology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1972, the English ethologist John H. Crook distinguished comparative ethology from social ethology, and argued that much of the ethology that had existed so far was really comparative ethology—examining animals as individuals—whereas, in the future, ethologists would need to concentrate on the behaviour of social groups of animals and the social structure within them.<sup id="cite_ref-Crook_Goss-Custard_1972_pp._277–312_8-0" class="reference"><a href="#cite_note-Crook_Goss-Custard_1972_pp._277–312-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/E._O._Wilson" title="E. O. Wilson">E. O. Wilson</a>'s book <i><a href="/wiki/Sociobiology:_The_New_Synthesis" title="Sociobiology: The New Synthesis">Sociobiology: The New Synthesis</a></i> appeared in 1975,<sup id="cite_ref-WilsonSociobiology_9-0" class="reference"><a href="#cite_note-WilsonSociobiology-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> and since that time, the study of behaviour has been much more concerned with social aspects. It has been driven by the Darwinism associated with Wilson, <a href="/wiki/Robert_Trivers" title="Robert Trivers">Robert Trivers</a>, and <a href="/wiki/W._D._Hamilton" title="W. D. Hamilton">W. D. Hamilton</a>. The related development of <a href="/wiki/Behavioural_ecology" class="mw-redirect" title="Behavioural ecology">behavioural ecology</a> has helped transform ethology.<sup id="cite_ref-Davies2012_10-0" class="reference"><a href="#cite_note-Davies2012-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Furthermore, a substantial rapprochement with <a href="/wiki/Comparative_psychology" title="Comparative psychology">comparative psychology</a> has occurred, so the modern scientific study of behaviour offers a spectrum of approaches. In 2020, Tobias Starzak and Albert Newen from the Institute of Philosophy II at the <a href="/wiki/Ruhr_University_Bochum" title="Ruhr University Bochum">Ruhr University Bochum</a> postulated that animals may have beliefs.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Determinants_of_behaviour">Determinants of behaviour</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=6" title="Edit section: Determinants of behaviour"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Behaviour is determined by three major factors, namely inborn <b>instincts</b>, <b>learning</b>, and <b>environmental factors</b>. The latter include abiotic and biotic factors. Abiotic factors such as <a href="/wiki/Temperature" title="Temperature">temperature</a> or <a href="/wiki/Light" title="Light">light</a> conditions have dramatic effects on animals, especially if they are <a href="/wiki/Ectotherm" title="Ectotherm">ectothermic</a> or <a href="/wiki/Nocturnality" title="Nocturnality">nocturnal</a>. Biotic factors include members of the same species (e.g. sexual behavior), <a href="/wiki/Predation" title="Predation">predators</a> (fight or flight), or <a href="/wiki/Parasitism" title="Parasitism">parasites</a> and <a href="/wiki/Disease" title="Disease">diseases</a>.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Instinct">Instinct</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=7" title="Edit section: Instinct"><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">Main article: <a href="/wiki/Instinct" title="Instinct">Instinct</a></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Larus_Dominicanus_with_young.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/88/Larus_Dominicanus_with_young.jpg/220px-Larus_Dominicanus_with_young.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/88/Larus_Dominicanus_with_young.jpg/330px-Larus_Dominicanus_with_young.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/88/Larus_Dominicanus_with_young.jpg/440px-Larus_Dominicanus_with_young.jpg 2x" data-file-width="3072" data-file-height="2048" /></a><figcaption><a href="/wiki/Kelp_gull" title="Kelp gull">Kelp gull</a> chicks peck at red spot on mother's beak to stimulate regurgitating reflex</figcaption></figure> <p><i><a href="/wiki/Webster%27s_Dictionary" title="Webster's Dictionary">Webster's Dictionary</a></i> defines <a href="/wiki/Instinct" title="Instinct">instinct</a> as "A largely inheritable and unalterable tendency of an organism to make a complex and specific response to environmental stimuli without involving reason".<sup id="cite_ref-Merriam_13-0" class="reference"><a href="#cite_note-Merriam-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> This covers fixed action patterns like beak movements of bird chicks,<sup id="cite_ref-Campbell_1996_14-0" class="reference"><a href="#cite_note-Campbell_1996-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> and the waggle dance of honeybees.<sup id="cite_ref-Buchmann_2006_15-0" class="reference"><a href="#cite_note-Buchmann_2006-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Fixed_action_patterns">Fixed action patterns</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=8" title="Edit section: Fixed action patterns"><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">Main article: <a href="/wiki/Fixed_action_pattern" title="Fixed action pattern">Fixed action pattern</a></div> <p>An important development, associated with the name of Konrad Lorenz though probably due more to his teacher, <a href="/wiki/Oskar_Heinroth" title="Oskar Heinroth">Oskar Heinroth</a>, was the identification of <a href="/wiki/Fixed_action_pattern" title="Fixed action pattern">fixed action patterns</a>. Lorenz popularized these as instinctive responses that would occur reliably in the presence of identifiable stimuli called sign stimuli or "releasing stimuli". Fixed action patterns are now considered to be instinctive behavioural sequences that are relatively invariant within the species and that almost inevitably run to completion.<sup id="cite_ref-Campbell_1996_14-1" class="reference"><a href="#cite_note-Campbell_1996-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p><p>One example of a releaser is the <a href="/wiki/Beak" title="Beak">beak</a> movements of many bird species performed by newly hatched chicks, which stimulates the mother to regurgitate food for her offspring.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Other examples are the classic studies by Tinbergen on the <a href="/wiki/Fixed_action_pattern#Greylag_goose_egg-retrieval_behavior" title="Fixed action pattern">egg-retrieval behaviour</a> and the effects of a "<a href="/wiki/Supernormal_stimulus" title="Supernormal stimulus">supernormal stimulus</a>" on the behaviour of <a href="/wiki/Graylag_goose" class="mw-redirect" title="Graylag goose">graylag geese</a>.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p><p>One investigation of this kind was the study of the <a href="/wiki/Waggle_dance" title="Waggle dance">waggle dance</a> ("dance language") in <a href="/wiki/Bee_learning_and_communication" title="Bee learning and communication">bee communication</a> by <a href="/wiki/Karl_von_Frisch" title="Karl von Frisch">Karl von Frisch</a>.<sup id="cite_ref-Buchmann_2006_15-1" class="reference"><a href="#cite_note-Buchmann_2006-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Learning">Learning</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=9" title="Edit section: Learning"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading4"><h4 id="Habituation">Habituation</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=10" title="Edit section: Habituation"><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">Main article: <a href="/wiki/Habituation" title="Habituation">Habituation</a></div> <p>Habituation is a simple form of learning and occurs in many animal taxa. It is the process whereby an animal ceases responding to a stimulus. Often, the response is an innate behavior. Essentially, the animal learns not to respond to irrelevant stimuli. For example, <a href="/wiki/Prairie_dog" title="Prairie dog">prairie dogs</a> (<i>Cynomys ludovicianus</i>) give alarm calls when predators approach, causing all individuals in the group to quickly scramble down burrows. When prairie dog towns are located near trails used by humans, giving alarm calls every time a person walks by is expensive in terms of time and energy. Habituation to humans is therefore an important behavior in this context.<sup id="cite_ref-Breed_19-0" class="reference"><a href="#cite_note-Breed-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Associative_learning">Associative learning</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=11" title="Edit section: Associative learning"><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">Main article: <a href="/wiki/Association_(psychology)" title="Association (psychology)">Association (psychology)</a></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/Classical_conditioning" title="Classical conditioning">Classical conditioning</a> and <a href="/wiki/Operant_conditioning" title="Operant conditioning">Operant conditioning</a></div> <p>Associative learning in animal behaviour is any learning process in which a new response becomes associated with a particular stimulus.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> The first studies of associative learning were made by the Russian <a href="/wiki/Physiologist" class="mw-redirect" title="Physiologist">physiologist</a> <a href="/wiki/Ivan_Pavlov" title="Ivan Pavlov">Ivan Pavlov</a>, who observed that dogs trained to associate food with the ringing of a bell would salivate on hearing the bell.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Imprinting">Imprinting</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=12" title="Edit section: Imprinting"><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">Main article: <a href="/wiki/Imprinting_(psychology)" title="Imprinting (psychology)">Imprinting (psychology)</a></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Moose-Imprinting-sr81-15.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8d/Moose-Imprinting-sr81-15.jpg/220px-Moose-Imprinting-sr81-15.jpg" decoding="async" width="220" height="146" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8d/Moose-Imprinting-sr81-15.jpg/330px-Moose-Imprinting-sr81-15.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8d/Moose-Imprinting-sr81-15.jpg/440px-Moose-Imprinting-sr81-15.jpg 2x" data-file-width="1024" data-file-height="681" /></a><figcaption><a href="/wiki/Imprinting_(psychology)" title="Imprinting (psychology)">Imprinting</a> in a <a href="/wiki/Moose" title="Moose">moose</a>.</figcaption></figure> <p><a href="/wiki/Imprinting_(psychology)" title="Imprinting (psychology)">Imprinting</a> enables the young to discriminate the members of their own species, vital for reproductive success. This important type of learning only takes place in a very limited period of time. <a href="/wiki/Konrad_Lorenz" title="Konrad Lorenz">Konrad Lorenz</a> observed that the young of birds such as <a href="/wiki/Goose" title="Goose">geese</a> and <a href="/wiki/Chicken" title="Chicken">chickens</a> followed their mothers spontaneously from almost the first day after they were hatched, and he discovered that this response could be imitated by an arbitrary stimulus if the eggs were incubated artificially and the stimulus were presented during a critical period that continued for a few days after hatching.<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Cultural_learning">Cultural learning</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=13" title="Edit section: Cultural learning"><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">Main article: <a href="/wiki/Cultural_transmission_in_animals" class="mw-redirect" title="Cultural transmission in animals">Cultural transmission in animals</a></div> <div class="mw-heading mw-heading5"><h5 id="Observational_learning">Observational learning</h5><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=14" title="Edit section: Observational learning"><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">Main article: <a href="/wiki/Observational_learning" title="Observational learning">Observational learning</a></div> <div class="mw-heading mw-heading5"><h5 id="Imitation">Imitation</h5><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=15" title="Edit section: Imitation"><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">Main article: <a href="/wiki/Imitation" title="Imitation">Imitation</a></div> <p><a href="/wiki/Imitation" title="Imitation">Imitation</a> is an advanced behavior whereby an animal observes and exactly replicates the behavior of another. The National Institutes of Health reported that capuchin monkeys preferred the company of researchers who imitated them to that of researchers who did not. The monkeys not only spent more time with their imitators but also preferred to engage in a simple task with them even when provided with the option of performing the same task with a non-imitator.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Imitation has been observed in recent research on chimpanzees; not only did these chimps copy the actions of another individual, when given a choice, the chimps preferred to imitate the actions of the higher-ranking elder chimpanzee as opposed to the lower-ranking young chimpanzee.<sup id="cite_ref-FerrariHorner2010_26-0" class="reference"><a href="#cite_note-FerrariHorner2010-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading5"><h5 id="Stimulus_and_local_enhancement">Stimulus and local enhancement</h5><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=16" title="Edit section: Stimulus and local enhancement"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Animals can learn using observational learning but without the process of imitation. One way is <i>stimulus enhancement</i> in which individuals become interested in an object as the result of observing others interacting with the object.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Increased interest in an object can result in object manipulation which allows for new object-related behaviours by trial-and-error learning. Haggerty (1909) devised an experiment in which a monkey climbed up the side of a cage, placed its arm into a wooden chute, and pulled a rope in the chute to release food. Another monkey was provided an opportunity to obtain the food after watching a monkey go through this process on four occasions. The monkey performed a different method and finally succeeded after trial-and-error.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> In local enhancement, a demonstrator attracts an observer's attention to a particular location.<sup id="cite_ref-mechanisms_29-0" class="reference"><a href="#cite_note-mechanisms-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> Local enhancement has been observed to transmit foraging information among birds, rats and pigs.<sup id="cite_ref-foraging_30-0" class="reference"><a href="#cite_note-foraging-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> The stingless bee (<i><a href="/wiki/Trigona_corvina" title="Trigona corvina">Trigona corvina</a></i>) uses local enhancement to locate other members of their colony and food resources.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading5"><h5 id="Social_transmission">Social transmission</h5><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=17" title="Edit section: Social transmission"><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">See also: <a href="/wiki/Cultural_transmission_in_animals" class="mw-redirect" title="Cultural transmission in animals">Cultural transmission in animals</a></div> <p>A well-documented example of social transmission of a behaviour occurred in a group of <a href="/wiki/Macaque" title="Macaque">macaques</a> on <a href="/wiki/Hachijojima" class="mw-redirect" title="Hachijojima">Hachijojima</a> Island, Japan. The macaques lived in the inland forest until the 1960s, when a group of researchers started giving them potatoes on the beach: soon, they started venturing onto the beach, picking the potatoes from the sand, and cleaning and eating them.<sup id="cite_ref-WilsonSociobiology_9-1" class="reference"><a href="#cite_note-WilsonSociobiology-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> About one year later, an individual was observed bringing a potato to the sea, putting it into the water with one hand, and cleaning it with the other. This behaviour was soon expressed by the individuals living in contact with her; when they gave birth, this behaviour was also expressed by their young—a form of social transmission.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Teaching">Teaching</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=18" title="Edit section: Teaching"><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">See also: <a href="/wiki/Animal_culture#Teaching" title="Animal culture">Animal culture § Teaching</a></div> <p>Teaching is a highly specialized aspect of learning in which the "teacher" (demonstrator) adjusts their behaviour to increase the probability of the "pupil" (observer) achieving the desired end-result of the behaviour. For example, <a href="/wiki/Orca" title="Orca">orcas</a> are known to intentionally beach themselves to catch <a href="/wiki/Pinniped" title="Pinniped">pinniped</a> prey.<sup id="cite_ref-rendell_33-0" class="reference"><a href="#cite_note-rendell-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> Mother orcas teach their young to catch pinnipeds by pushing them onto the shore and encouraging them to attack the prey. Because the mother orca is altering her behaviour to help her offspring learn to catch prey, this is evidence of teaching.<sup id="cite_ref-rendell_33-1" class="reference"><a href="#cite_note-rendell-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> Teaching is not limited to mammals. Many insects, for example, have been observed demonstrating various forms of teaching to obtain food. <a href="/wiki/Ant" title="Ant">Ants</a>, for example, will guide each other to food sources through a process called "<a href="/wiki/Tandem_running" title="Tandem running">tandem running</a>," in which an ant will guide a companion ant to a source of food.<sup id="cite_ref-hoppit_34-0" class="reference"><a href="#cite_note-hoppit-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> It has been suggested that the pupil ant is able to learn this route to obtain food in the future or teach the route to other ants. This behaviour of teaching is also exemplified by crows, specifically <a href="/wiki/New_Caledonian_crow" title="New Caledonian crow">New Caledonian crows</a>. The adults (whether individual or in families) teach their young adolescent offspring how to construct and utilize tools. For example, <i><a href="/wiki/Pandanus" title="Pandanus">Pandanus</a></i> branches are used to extract insects and other larvae from holes within trees.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Mating_and_the_fight_for_supremacy">Mating and the fight for supremacy</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=19" title="Edit section: Mating and the fight for supremacy"><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:Courtship_of_sarus_cranes.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/75/Courtship_of_sarus_cranes.jpg/220px-Courtship_of_sarus_cranes.jpg" decoding="async" width="220" height="180" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/75/Courtship_of_sarus_cranes.jpg/330px-Courtship_of_sarus_cranes.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/75/Courtship_of_sarus_cranes.jpg/440px-Courtship_of_sarus_cranes.jpg 2x" data-file-width="697" data-file-height="570" /></a><figcaption><a href="/wiki/Courtship_display" title="Courtship display">Courtship display</a> of a <a href="/wiki/Sarus_crane" title="Sarus crane">sarus crane</a> </figcaption></figure> <p>Individual <a href="/wiki/Reproduction" title="Reproduction">reproduction</a> is the most important phase in the proliferation of individuals or genes within a species: for this reason, there exist complex <a href="/wiki/Mating" title="Mating">mating</a> <a href="/wiki/Ritual" title="Ritual">rituals</a>, which can be very complex even if they are often regarded as fixed action patterns. The <a href="/wiki/Stickleback" title="Stickleback">stickleback</a>'s complex mating ritual, studied by Tinbergen, is regarded as a notable example.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p><p>Often in <a href="/wiki/Social_relation" title="Social relation">social life</a>, animals fight for the right to reproduce, as well as social supremacy. A common example of fighting for social and sexual supremacy is the so-called <a href="/wiki/Pecking_order" class="mw-redirect" title="Pecking order">pecking order</a> among <a href="/wiki/Poultry" title="Poultry">poultry</a>. Every time a group of poultry cohabitate for a certain time length, they establish a pecking order. In these groups, one chicken dominates the others and can peck without being pecked. A second chicken can peck all the others except the first, and so on. Chickens higher in the pecking order may at times be distinguished by their healthier appearance when compared to lower level chickens.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (June 2020)">citation needed</span></a></i>]</sup> While the pecking order is establishing, frequent and violent fights can happen, but once established, it is broken only when other individuals enter the group, in which case the pecking order re-establishes from scratch.<sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Social_behaviour">Social behaviour</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=20" title="Edit section: Social behaviour"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Several animal species, including humans, tend to live in groups. <a href="/wiki/Group_size_measures" title="Group size measures">Group size</a> is a major aspect of their social environment. <a href="/wiki/Social_relation" title="Social relation">Social life</a> is probably a complex and effective survival strategy. It may be regarded as a sort of <a href="/wiki/Symbiosis" title="Symbiosis">symbiosis</a> among individuals of the same species: a <a href="/wiki/Society" title="Society">society</a> is composed of a group of individuals belonging to the same species living within well-defined rules on <a href="/wiki/Food" title="Food">food</a> management, role assignments and reciprocal dependence.<sup id="cite_ref-Mandal_2015_38-0" class="reference"><a href="#cite_note-Mandal_2015-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p><p>When <a href="/wiki/Biologist" title="Biologist">biologists</a> interested in <a href="/wiki/Evolution_theory" class="mw-redirect" title="Evolution theory">evolution theory</a> first started examining social behaviour, some apparently unanswerable questions arose, such as how the birth of <a href="/wiki/Reproduce" class="mw-redirect" title="Reproduce">sterile</a> <a href="/wiki/Caste" title="Caste">castes</a>, like in <a href="/wiki/Bee" title="Bee">bees</a>, could be explained through an evolving mechanism that emphasizes the reproductive success of as many individuals as possible, or why, amongst animals living in small groups like <a href="/wiki/Squirrel" title="Squirrel">squirrels</a>, an individual would risk its own life to save the rest of the group. These behaviours may be examples of <a href="/wiki/Altruism_(biology)" title="Altruism (biology)">altruism</a>.<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> Not all behaviours are altruistic, as indicated by the table below. For example, revengeful behaviour was at one point claimed to have been observed exclusively in <i><a href="/wiki/Homo_sapiens" class="mw-redirect" title="Homo sapiens">Homo sapiens</a></i>. However, other species have been reported to be vengeful including chimpanzees,<sup id="cite_ref-McCullough2008_40-0" class="reference"><a href="#cite_note-McCullough2008-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> as well as anecdotal reports of vengeful camels.<sup id="cite_ref-Waal2001_41-0" class="reference"><a href="#cite_note-Waal2001-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> </p> <table class="wikitable" style="margin: 1em auto;"> <caption>Classification of social behaviours<sup id="cite_ref-Mandal_2015_38-1" class="reference"><a href="#cite_note-Mandal_2015-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </caption> <tbody><tr> <th>Type of behaviour </th> <th>Effect on the donor </th> <th>Effect on the receiver </th></tr> <tr> <td><a href="/wiki/Egotism" title="Egotism">Egoistic</a> </td> <td>Neutral to Increases fitness </td> <td>Decreases fitness </td></tr> <tr> <td><a href="/wiki/Cooperation" title="Cooperation">Cooperative</a> </td> <td>Neutral to Increases fitness </td> <td>Neutral to Increases fitness </td></tr> <tr> <td><a href="/wiki/Altruism" title="Altruism">Altruistic</a> </td> <td>Decreases fitness </td> <td>Neutral to Increases fitness </td></tr> <tr> <td><a href="/wiki/Revenge" title="Revenge">Revengeful</a> </td> <td>Decreases fitness </td> <td>Decreases fitness </td></tr></tbody></table> <p><a href="/wiki/Altruistic" class="mw-redirect" title="Altruistic">Altruistic</a> behaviour has been explained by the <a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">gene-centred view of evolution</a>.<sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Benefits_and_costs_of_group_living">Benefits and costs of group living</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=21" title="Edit section: Benefits and costs of group living"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>One advantage of group living is decreased predation. If the number of predator attacks stays the same despite increasing prey group size, each prey has a reduced risk of predator attacks through the dilution effect.<sup id="cite_ref-Davies2012_10-1" class="reference"><a href="#cite_note-Davies2012-10"><span class="cite-bracket">[</span>10<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. (December 2016)">page needed</span></a></i>]</sup> Further, according to the <a href="/wiki/Selfish_herd_theory" title="Selfish herd theory">selfish herd theory</a>, the fitness benefits associated with group living vary depending on the location of an individual within the group. The theory suggests that conspecifics positioned at the centre of a group will reduce the likelihood predations while those at the periphery will become more vulnerable to attack.<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> In groups, prey can also actively reduce their predation risk through more effective defence tactics, or through earlier detection of predators through increased vigilance.<sup id="cite_ref-Davies2012_10-2" class="reference"><a href="#cite_note-Davies2012-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p><p>Another advantage of group living is an increased ability to forage for food. Group members may exchange information about food sources, facilitating the process of <a href="/wiki/Foraging#Group_foraging" title="Foraging">resource location</a>.<sup id="cite_ref-Davies2012_10-3" class="reference"><a href="#cite_note-Davies2012-10"><span class="cite-bracket">[</span>10<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. (December 2016)">page needed</span></a></i>]</sup> Honeybees are a notable example of this, using the <a href="/wiki/Waggle_dance" title="Waggle dance">waggle dance</a> to communicate the location of flowers to the rest of their hive.<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> Predators also receive benefits from <a href="/wiki/Cooperative_hunting" class="mw-redirect" title="Cooperative hunting">hunting in groups</a>, through using better strategies and being able to take down larger prey.<sup id="cite_ref-Davies2012_10-4" class="reference"><a href="#cite_note-Davies2012-10"><span class="cite-bracket">[</span>10<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. (December 2016)">page needed</span></a></i>]</sup> </p><p>Some disadvantages accompany living in groups. Living in close proximity to other animals can facilitate the transmission of parasites and disease, and groups that are too large may also experience greater competition for resources and mates.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Group_size">Group size</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=22" title="Edit section: Group size"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Theoretically, social animals should have optimal group sizes that maximize the benefits and minimize the costs of group living. However, in nature, most groups are stable at slightly larger than optimal sizes.<sup id="cite_ref-Davies2012_10-5" class="reference"><a href="#cite_note-Davies2012-10"><span class="cite-bracket">[</span>10<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. (December 2016)">page needed</span></a></i>]</sup> Because it generally benefits an individual to join an optimally-sized group, despite slightly decreasing the advantage for all members, groups may continue to increase in size until it is more advantageous to remain alone than to join an overly full group.<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> </p> <div style="clear:both;" class=""></div> <div class="mw-heading mw-heading2"><h2 id="Tinbergen's_four_questions_for_ethologists"><span id="Tinbergen.27s_four_questions_for_ethologists"></span>Tinbergen's four questions for ethologists</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ethology&action=edit&section=23" title="Edit section: Tinbergen's four questions for ethologists"><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">Main article: <a href="/wiki/Tinbergen%27s_four_questions" title="Tinbergen's four questions">Tinbergen's four questions</a></div> <p>Tinbergen argued that ethology needed to include four kinds of explanation in any instance of behaviour:<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> </p> <ul><li>Function – How does the behaviour affect the animal's chances of survival and reproduction? Why does the animal respond that way instead of some other way?</li> <li>Causation – What are the stimuli that elicit the response, and how has it been modified by recent learning?</li> <li>Development – How does the behaviour change with age, and what early experiences are necessary for the animal to display the behaviour?</li> <li>Evolutionary history – How does the behaviour compare with similar behaviour in related species, and how might it have begun through the process of <a href="/wiki/Phylogeny" class="mw-redirect" title="Phylogeny">phylogeny</a>?</li></ul> <p>These explanations are complementary rather than mutually exclusive—all instances of behaviour require an explanation at each of these four levels. For example, the function of eating is to acquire nutrients (which ultimately aids survival and reproduction), but the immediate cause of eating is hunger (causation). Hunger and eating are evolutionarily ancient and are found in many species (evolutionary history), and develop early within an organism's lifespan (development). It is easy to confuse such questions—for example, to argue that people eat because they are hungry and not to acquire nutrients—without realizing that the reason people experience hunger is because it causes them to acquire nutrients.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<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=Ethology&action=edit&section=24" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1266661725">.mw-parser-output .portalbox{padding:0;margin:0.5em 0;display:table;box-sizing:border-box;max-width:175px;list-style:none}.mw-parser-output .portalborder{border:1px solid 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<li><a href="/wiki/Cognitive_ethology" title="Cognitive ethology">Cognitive ethology</a></li> <li><a href="/wiki/Deception_in_animals" title="Deception in animals">Deception in animals</a></li> <li><a href="/wiki/Human_ethology" title="Human ethology">Human ethology</a></li> <li><a href="/wiki/List_of_abnormal_behaviours_in_animals" title="List of abnormal behaviours in animals">List of abnormal behaviours in animals</a></li> <li><a href="/wiki/Tool_use_by_non-human_animals" class="mw-redirect" title="Tool use by non-human animals">Tool use by non-human animals</a></li></ul> </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=Ethology&action=edit&section=25" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-insectbehavior-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-insectbehavior_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-insectbehavior_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-insectbehavior_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-insectbehavior_1-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFMatthewsMatthews,_Robert_W.2009" class="citation book cs1">Matthews, Janice R.; Matthews, Robert W. 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Retrieved <span class="nowrap">21 September</span> 2022</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=lib.uchicago.edu&rft.atitle=Guide+to+the+Charles+Otis+Whitman+Collection+ca.+1911&rft_id=https%3A%2F%2Fwww.lib.uchicago.edu%2Fe%2Fscrc%2Ffindingaids%2Fview.php%3Feadid%3DICU.SPCL.WHITMANCO%23idp149584456&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEthology" class="Z3988"></span></span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSchulze-HagenBirkhead2015" class="citation journal cs1">Schulze-Hagen, Karl; Birkhead, Timothy R. 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Retrieved <span class="nowrap">9 September</span> 2016</span>. <q>The Nobel Prize in Physiology or Medicine 1973 was awarded jointly to Karl von Frisch, Konrad Lorenz and Nikolaas Tinbergen 'for their discoveries concerning organization and elicitation of individual and social behaviour patterns'.</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Nobelprize.org.&rft.atitle=The+Nobel+Prize+in+Physiology+or+Medicine+1973&rft_id=https%3A%2F%2Fwww.nobelprize.org%2Fnobel_prizes%2Fmedicine%2Flaureates%2F1973%2Findex.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEthology" class="Z3988"></span></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"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://ishe.org/goals/">"Goals"</a>. <a href="/wiki/International_Society_for_Human_Ethology" title="International Society for Human Ethology">International Society for Human Ethology</a><span class="reference-accessdate">. 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John Wiley & Sons. pp. <span class="nowrap">79–</span>80. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-470-26215-3" title="Special:BookSources/978-0-470-26215-3"><bdi>978-0-470-26215-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Beyond+Revenge%3A+The+Evolution+of+the+Forgiveness+Instinct&rft.pages=%3Cspan+class%3D%22nowrap%22%3E79-%3C%2Fspan%3E80&rft.pub=John+Wiley+%26+Sons&rft.date=2008&rft.isbn=978-0-470-26215-3&rft.aulast=McCullough&rft.aufirst=Michael+E.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DdaomTGYZuW4C%26pg%3DPA79&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEthology" class="Z3988"></span></span> </li> <li id="cite_note-Waal2001-41"><span class="mw-cite-backlink"><b><a href="#cite_ref-Waal2001_41-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDe_Waal2001" class="citation book cs1"><a href="/wiki/Frans_de_Waal" title="Frans de Waal">De Waal, Frans</a> (2001). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/apesushimastercu0000waal"><i>The Ape and the Sushi Master: Cultural Reflections by a Primatologist</i></a></span>. 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(2002) <i>Human Evolutionary Psychology</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/9780691096223" title="Special:BookSources/9780691096223">9780691096223</a><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. (June 2023)">page needed</span></a></i>]</sup></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=Ethology&action=edit&section=26" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="http://dmr.bsu.edu/utils/getfile/collection/ConspectusH/id/603/filename/604.pdf">Burkhardt, Richard W. Jr. "On the Emergence of Ethology as a Scientific Discipline." Conspectus of History 1.7 (1981).</a></li></ul> <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=Ethology&action=edit&section=27" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Commons-logo.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/12px-Commons-logo.svg.png" decoding="async" width="12" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/18px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/24px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></a></span> Media related to <a 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.navbox-abovebelow{background-color:#e6e6ff}.mw-parser-output .navbox-even{background-color:#f7f7f7}.mw-parser-output .navbox-odd{background-color:transparent}.mw-parser-output .navbox .hlist td dl,.mw-parser-output .navbox .hlist td ol,.mw-parser-output .navbox .hlist td ul,.mw-parser-output .navbox td.hlist dl,.mw-parser-output .navbox td.hlist ol,.mw-parser-output .navbox td.hlist ul{padding:0.125em 0}.mw-parser-output .navbox .navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Ethology159" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" 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navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Animal_cognition" title="Animal cognition">Animal cognition</a></li> <li><a href="/wiki/Animal_communication" title="Animal communication">Animal communication</a></li> <li><a href="/wiki/Animal_consciousness" title="Animal consciousness">Animal consciousness</a></li> <li><a href="/wiki/Animal_culture" title="Animal culture">Animal culture</a></li> <li><a href="/wiki/Animal_sexual_behaviour" title="Animal sexual behaviour">Animal sexual behaviour</a></li> <li><a href="/wiki/Animal_welfare_science" title="Animal welfare science">Animal welfare science</a></li> <li><a href="/wiki/Anthrozoology" title="Anthrozoology">Anthrozoology</a></li> <li><a href="/wiki/Bee_learning_and_communication" title="Bee learning and communication">Bee learning and communication</a></li> <li><a href="/wiki/Behavioral_ecology" title="Behavioral ecology">Behavioural ecology</a></li> <li><a href="/wiki/Behavioral_endocrinology" title="Behavioral endocrinology">Behavioral endocrinology</a></li> <li><a href="/wiki/Behavioural_genetics" title="Behavioural genetics">Behavioural genetics</a></li> <li><a href="/wiki/Breed" title="Breed">Breed</a></li> <li><a href="/wiki/Cognitive_ethology" title="Cognitive ethology">Cognitive ethology</a></li> <li><a href="/wiki/Comfort_behaviour_in_animals" title="Comfort behaviour in animals">Comfort behaviour</a> <ul><li><a href="/wiki/Personal_grooming" title="Personal grooming">Grooming</a></li></ul></li> <li><a href="/wiki/Comparative_psychology" title="Comparative psychology">Comparative psychology</a></li> <li><a href="/wiki/Emotion_in_animals" title="Emotion in animals">Emotion in animals</a></li> <li><a href="/wiki/Ethogram" title="Ethogram">Ethogram</a></li> <li><a href="/wiki/Evolutionary_neuroscience" title="Evolutionary neuroscience">Evolutionary neuroscience</a></li> <li><a href="/wiki/List_of_feeding_behaviours" title="List of feeding behaviours">Feeding</a></li> <li><a href="/wiki/Hover_(behaviour)" title="Hover (behaviour)">Hover</a></li> <li><a href="/wiki/Human_ethology" title="Human ethology">Human ethology</a></li> <li><a href="/wiki/Instinct" title="Instinct">Instinct</a></li> <li><a href="/wiki/Learning" title="Learning">Learning</a></li> <li><a href="/wiki/Neuroethology" title="Neuroethology">Neuroethology</a></li> <li><a href="/wiki/Pain_in_animals" title="Pain in animals">Pain in animals</a></li> <li><a href="/wiki/Philosophical_ethology" title="Philosophical ethology">Philosophical ethology</a></li> <li><a href="/wiki/Sociobiology" title="Sociobiology">Sociobiology</a></li> <li><a href="/wiki/Stereotypy_(non-human)" title="Stereotypy (non-human)">Stereotypy</a></li> <li><a href="/wiki/Structures_built_by_animals" title="Structures built by animals">Structures</a> <ul><li><a href="/wiki/Beehive" title="Beehive">Hive</a></li> <li><a href="/wiki/Honeycomb_structure" title="Honeycomb structure">Honeycomb</a></li> <li><a href="/wiki/Nest" title="Nest">Nest</a></li> <li><a href="/wiki/Nesting_instinct" title="Nesting instinct">Instinct</a></li></ul></li> <li><a href="/wiki/Swarm_behaviour" title="Swarm behaviour">Swarm</a></li> <li><a href="/wiki/Tool_use_by_non-humans" title="Tool use by non-humans">Tool use by non-humans</a></li> <li><a href="/wiki/Zoosemiotics" title="Zoosemiotics">Zoosemiotics</a></li> <li><a href="/wiki/Zoomusicology" title="Zoomusicology">Zoomusicology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Ethologists</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/Patrick_Bateson" title="Patrick Bateson">Patrick Bateson</a></li> <li><a href="/wiki/Marc_Bekoff" title="Marc Bekoff">Marc Bekoff</a></li> <li><a href="/wiki/Donald_Broom" title="Donald Broom">Donald Broom</a></li> <li><a href="/wiki/John_B._Calhoun" title="John B. Calhoun">John B. Calhoun</a></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a></li> <li><a href="/wiki/Marian_Dawkins" title="Marian Dawkins">Marian Dawkins</a></li> <li><a href="/wiki/Richard_Dawkins" title="Richard Dawkins">Richard Dawkins</a></li> <li><a href="/wiki/Iren%C3%A4us_Eibl-Eibesfeldt" title="Irenäus Eibl-Eibesfeldt">Irenäus Eibl-Eibesfeldt</a></li> <li><a href="/wiki/Dian_Fossey" title="Dian Fossey">Dian Fossey</a></li> <li><a href="/wiki/Karl_von_Frisch" title="Karl von Frisch">Karl von Frisch</a></li> <li><a href="/wiki/Jane_Goodall" title="Jane Goodall">Jane Goodall</a></li> <li><a href="/wiki/Heini_Hediger" title="Heini Hediger">Heini Hediger</a></li> <li><a href="/wiki/Julian_Huxley" title="Julian Huxley">Julian Huxley</a></li> <li><a href="/wiki/Konrad_Lorenz" title="Konrad Lorenz">Konrad Lorenz</a></li> <li><a href="/wiki/Desmond_Morris" title="Desmond Morris">Desmond Morris</a></li> <li><a href="/wiki/Thomas_Sebeok" title="Thomas Sebeok">Thomas Sebeok</a></li> <li><a href="/wiki/William_Homan_Thorpe" title="William Homan Thorpe">William Homan Thorpe</a></li> <li><a href="/wiki/Nikolaas_Tinbergen" title="Nikolaas Tinbergen">Nikolaas Tinbergen</a></li> <li><a href="/wiki/Jakob_von_Uexk%C3%BCll" class="mw-redirect" title="Jakob von Uexküll">Jakob von Uexküll</a></li> <li><a href="/wiki/Wolfgang_Wickler" title="Wolfgang Wickler">Wolfgang Wickler</a></li> <li><a href="/wiki/E._O._Wilson" title="E. O. Wilson">E. O. Wilson</a></li> <li><a href="/wiki/Solly_Zuckerman" class="mw-redirect" title="Solly Zuckerman">Solly Zuckerman</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Societies</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/Association_for_the_Study_of_Animal_Behaviour" title="Association for the Study of Animal Behaviour">Association for the Study of Animal Behaviour</a></li> <li><a href="/wiki/International_Society_for_Applied_Ethology" title="International Society for Applied Ethology">International Society for Applied Ethology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Journals</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Animal_Behaviour_(journal)" title="Animal Behaviour (journal)">Animal Behaviour</a></i></li> <li><i><a href="/wiki/Animal_Cognition" title="Animal Cognition">Animal Cognition</a></i></li> <li><i><a href="/wiki/Animal_Welfare_(journal)" title="Animal Welfare (journal)">Animal Welfare</a></i></li> <li><i><a href="/wiki/Behavioral_Ecology_(journal)" title="Behavioral Ecology (journal)">Behavioral Ecology</a></i></li> <li><i><a href="/wiki/Behaviour_(journal)" title="Behaviour (journal)">Behaviour</a></i></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Ethology" title="Category:Ethology">Category</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Biological_rhythms67" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Biological_rhythm" title="Template:Biological rhythm"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Biological_rhythm" title="Template talk:Biological rhythm"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Biological_rhythm" title="Special:EditPage/Template:Biological rhythm"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Biological_rhythms67" style="font-size:114%;margin:0 4em"><a href="/wiki/Biological_rhythm" title="Biological rhythm">Biological rhythms</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Internal rhythms</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/Circadian_rhythm" title="Circadian rhythm">Circadian rhythm</a></li> <li><a href="/wiki/Infradian_rhythm" title="Infradian rhythm">Infradian rhythm</a></li> <li><a href="/wiki/Ultradian_rhythm" title="Ultradian rhythm">Ultradian rhythm</a></li> <li><a href="/wiki/Lunar_effect" title="Lunar effect">Lunar rhythm</a></li> <li><a href="/wiki/Season" title="Season">Annual rhythm</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="4" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Light_Ethology.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Light_Ethology.svg/150px-Light_Ethology.svg.png" decoding="async" width="150" height="60" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Light_Ethology.svg/225px-Light_Ethology.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Light_Ethology.svg/300px-Light_Ethology.svg.png 2x" data-file-width="512" data-file-height="205" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">External cycles</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/Diurnality" title="Diurnality">Diurnality</a> <ul><li><a href="/wiki/Diurnal_cycle" title="Diurnal cycle">Diurnal cycle</a></li></ul></li> <li><a href="/wiki/Nocturnality" title="Nocturnality">Nocturnality</a></li> <li><a href="/wiki/Cathemerality" title="Cathemerality">Cathemerality</a></li> <li><a href="/wiki/Crepuscular_animal" title="Crepuscular animal">Crepuscular</a> <ul><li><a href="/wiki/Matutinal" title="Matutinal">Matutinal</a></li> <li><a href="/wiki/Vespertine_(biology)" title="Vespertine (biology)">Vespertine</a></li></ul></li> <li><a href="/wiki/Season" title="Season">Seasonality</a></li> <li><a href="/wiki/Photoperiodism" title="Photoperiodism">Photoperiodism</a></li> <li><a href="/wiki/Vernalization" title="Vernalization">Vernalization</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Fields</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/Behavioral_ecology" title="Behavioral ecology">Behavioural ecology</a></li> <li><a href="/wiki/Chronobiology" title="Chronobiology">Chronobiology</a></li> <li><a class="mw-selflink selflink">Ethology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">See also</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/Chronotype" title="Chronotype">Chronotype</a></li> <li><a href="/wiki/Diel_vertical_migration" title="Diel vertical migration">Diel vertical migration</a></li> <li><a href="/wiki/Light_pollution" title="Light pollution">Light pollution</a></li> <li><a href="/wiki/Solunar_theory" title="Solunar theory">Solunar theory</a></li> <li><a href="/wiki/Template:Parts_of_a_day" title="Template:Parts of a day">Parts of a day</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="Sociobiology421" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Sociobiology" title="Template:Sociobiology"><abbr title="View this 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theory</a></li> <li><a class="mw-selflink selflink">Ethology</a></li> <li><a href="/wiki/Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Evolution of morality</a></li> <li><a href="/wiki/Evolutionary_models_of_food_sharing" title="Evolutionary models of food sharing">Evolutionary models of food sharing</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a></li> <li><a href="/wiki/Kin_recognition" title="Kin recognition">Kin recognition</a></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a></li> <li><a href="/wiki/Male_warrior_hypothesis" title="Male warrior hypothesis">Male warrior hypothesis</a></li> <li><a href="/wiki/Reciprocal_altruism" title="Reciprocal altruism">Reciprocal altruism</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a> <ul><li><a href="/wiki/Sexual_selection_in_humans" title="Sexual selection in humans">in humans</a></li></ul></li> <li><a href="/wiki/Sex_and_psychology" class="mw-redirect" title="Sex and psychology">Sex and psychology</a></li> <li><a href="/wiki/Sociality" title="Sociality">Sociality</a> <ul><li><a href="/wiki/Eusociality" title="Eusociality">eusociality</a> <ul><li><a href="/wiki/Evolution_of_eusociality" title="Evolution of eusociality">evolution</a></li></ul></li> <li><a href="/wiki/Presociality" class="mw-redirect" title="Presociality">presociality</a></li></ul></li> <li><a href="/wiki/Dunbar%27s_number" title="Dunbar's number">Dunbar's number</a></li> <li><a href="/wiki/Polyethism" title="Polyethism">Polyethism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Supporters</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/Anne_Campbell_(academic)" title="Anne Campbell (academic)">Anne Campbell</a></li> <li><a href="/wiki/Noam_Chomsky" title="Noam Chomsky">Noam Chomsky</a></li> <li><a href="/wiki/Richard_Dawkins" title="Richard Dawkins">Richard Dawkins</a></li> <li><a href="/wiki/Daniel_Dennett" title="Daniel Dennett">Daniel Dennett</a></li> <li><a href="/wiki/Sarah_Blaffer_Hrdy" title="Sarah Blaffer Hrdy">Sarah Blaffer Hrdy</a></li> <li><a href="/wiki/Steven_Pinker" title="Steven Pinker">Steven Pinker</a></li> <li><a href="/wiki/Frans_de_Waal" title="Frans de Waal">Frans de Waal</a></li> <li><a href="/wiki/E._O._Wilson" title="E. O. Wilson">E. O. Wilson</a> <ul><li><i><a href="/wiki/Sociobiology:_The_New_Synthesis" title="Sociobiology: The New Synthesis">Sociobiology: The New Synthesis</a></i></li></ul></li> <li><a href="/wiki/Robert_Wright_(journalist)" title="Robert Wright (journalist)">Robert Wright</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Opponents</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/Stephen_Jay_Gould" title="Stephen Jay Gould">Stephen Jay Gould</a></li> <li><a href="/wiki/Leon_Kamin" title="Leon Kamin">Leon Kamin</a></li> <li><a href="/wiki/Richard_Lewontin" title="Richard Lewontin">Richard Lewontin</a></li> <li><a href="/wiki/Steven_Rose" title="Steven Rose">Steven Rose</a> <ul><li><i><a href="/wiki/Not_in_Our_Genes" title="Not in Our Genes">Not in Our Genes</a></i></li></ul></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <b><a href="/wiki/Category:Sociobiology" title="Category:Sociobiology">Category</a></b></li> <li><span class="noviewer" typeof="mw:File"><span title="Commons page"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/12px-Commons-logo.svg.png" decoding="async" width="12" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/18px-Commons-logo.svg.png 1.5x, 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biology portal</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="Animal_communication54" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Animal_communication" title="Template:Animal communication"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a 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href="/wiki/Anecdotal_cognitivism" title="Anecdotal cognitivism">Anecdotal cognitivism</a></li> <li><a href="/wiki/Bioacoustics" title="Bioacoustics">Bioacoustics</a></li> <li><a href="/wiki/Deception_in_animals" title="Deception in animals">Deception in animals</a> <ul><li><a href="/wiki/Tactical_deception_in_animals" title="Tactical deception in animals">Tactical</a></li></ul></li> <li><a class="mw-selflink selflink">Ethology</a></li> <li><a href="/wiki/Evolutionary_linguistics" title="Evolutionary linguistics">Evolutionary linguistics</a></li> <li><a href="/wiki/FOXP2" title="FOXP2">FOXP2</a></li> <li><a href="/wiki/FOXP2_and_human_evolution" class="mw-redirect" title="FOXP2 and human evolution">FOXP2 and human evolution</a></li> <li><a href="/wiki/Human%E2%80%93animal_communication" title="Human–animal communication">Human–animal communication</a></li> <li><a href="/wiki/Mating_call" title="Mating call">Mating call</a></li> <li><a href="/wiki/Origin_of_language" title="Origin of language">Origin of language</a></li> <li><a href="/wiki/Seismic_communication" title="Seismic communication">Seismic communication</a></li> <li><a href="/wiki/Self-anointing_in_animals" title="Self-anointing in animals">Self-anointing in animals</a></li> <li><a href="/wiki/Talking_animal" title="Talking animal">Talking animals</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Animal-specific</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/Communication_in_aquatic_animals" title="Communication in aquatic animals">Aquatic animals</a> <ul><li><a href="/wiki/Whale_vocalization" title="Whale vocalization">Whale vocalization</a></li></ul></li> <li><a href="/wiki/Bee_learning_and_communication" title="Bee learning and communication">Bees</a> <ul><li><a href="/wiki/Bumblebee_communication" title="Bumblebee communication">Bumblebees</a></li></ul></li> <li><a href="/wiki/Bird_vocalization" title="Bird vocalization">Birds</a> <ul><li><a href="/wiki/Talking_bird" title="Talking bird">Talking birds</a></li></ul></li> <li><a href="/wiki/Cat_communication" title="Cat communication">Cats</a></li> <li><a href="/wiki/Dog_communication" title="Dog communication">Dogs</a></li> <li><a href="/wiki/Elephant_communication" title="Elephant communication">Elephants</a></li> <li><a href="/wiki/Frog_hearing_and_communication" title="Frog hearing and communication">Frogs</a></li> <li><a href="/wiki/Great_ape_language" title="Great ape language">Great ape language</a> <ul><li><a href="/wiki/Yerkish" title="Yerkish">Yerkish</a></li></ul></li> <li><a href="/wiki/Chemical_communication_in_insects" title="Chemical communication in insects">Insects</a></li> <li><a href="/wiki/Lizard_communication" title="Lizard communication">Lizards</a></li> <li><a href="/wiki/Wolf_communication" title="Wolf communication">Wolves</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Roar" title="Roar">Roar</a></li> <li><a href="/wiki/Category:Talking_birds" title="Category:Talking birds">Category:Talking birds</a></li> <li><a href="/wiki/Category:Individual_apes_involved_in_language_studies" title="Category:Individual apes involved in language studies">Category:Individual apes involved in language studies</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="Zoology332" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2" style="background:"><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:Zoology" title="Template:Zoology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Zoology" title="Template talk:Zoology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Zoology" title="Special:EditPage/Template:Zoology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Zoology332" style="font-size:114%;margin:0 4em"><a href="/wiki/Zoology" title="Zoology">Zoology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2" style="background:"><div> <ul><li><a href="/wiki/Outline_of_zoology" title="Outline of zoology">Outline</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background:;width:1%">Branches</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/Anthrozoology" title="Anthrozoology">Anthrozoology</a></li> <li><a href="/wiki/Arthropodology" title="Arthropodology">Arthropodology</a> (<a href="/wiki/Arachnology" title="Arachnology">Arachnology</a> (<a href="/wiki/Acarology" title="Acarology">Acarology</a>)</li> <li><a href="/wiki/Carcinology" title="Carcinology">Carcinology</a></li> <li><a href="/wiki/Entomology" title="Entomology">Entomology</a> (<a href="/wiki/Coleopterology" title="Coleopterology">Coleopterology</a></li> <li><a href="/wiki/Lepidopterology" title="Lepidopterology">Lepidopterology</a></li> <li><a href="/wiki/Melittology" title="Melittology">Melittology</a></li> <li><a href="/wiki/Myrmecology" title="Myrmecology">Myrmecology</a></li> <li><a href="/wiki/Orthopterology" title="Orthopterology">Orthopterology</a>)</li> <li><a href="/wiki/Myriapodology" title="Myriapodology">Myriapodology</a>)</li> <li><a href="/wiki/Biological_anthropology" title="Biological anthropology">Biological anthropology</a></li> <li><a href="/wiki/Bryozoology" title="Bryozoology">Bryozoology</a></li> <li><a href="/wiki/Cnidariology" class="mw-redirect" title="Cnidariology">Cnidariology</a></li> <li><a href="/wiki/Ethnozoology" title="Ethnozoology">Ethnozoology</a></li> <li><a class="mw-selflink selflink">Ethology</a></li> <li><a href="/wiki/Helminthology" title="Helminthology">Helminthology</a></li> <li><a href="/wiki/Herpetology" title="Herpetology">Herpetology</a> (<a href="/wiki/Batrachology" title="Batrachology">Batrachology</a></li> <li><a href="/wiki/Ophiology" class="mw-redirect" title="Ophiology">Ophiology</a></li> <li><a href="/wiki/Testudinology" class="mw-redirect" title="Testudinology">Testudinology</a>)</li> <li><a href="/wiki/Ichthyology" title="Ichthyology">Ichthyology</a></li> <li><a href="/wiki/Malacology" title="Malacology">Malacology</a> (<a href="/wiki/Conchology" title="Conchology">Conchology</a></li> <li><a href="/wiki/Teuthology" title="Teuthology">Teuthology</a>)</li> <li><a href="/wiki/Mammalogy" title="Mammalogy">Mammalogy</a> (<a href="/wiki/Cetology" title="Cetology">Cetology</a></li> <li><a href="/wiki/Cynology" title="Cynology">Cynology</a></li> <li><a href="/wiki/Felinology" title="Felinology">Felinology</a></li> <li><a href="/wiki/Hippology" title="Hippology">Hippology</a></li> <li><a href="/wiki/Primatology" title="Primatology">Primatology</a>)</li> <li><a href="/wiki/Nematology" title="Nematology">Nematology</a></li> <li><a href="/wiki/Neuroethology" title="Neuroethology">Neuroethology</a></li> <li><a href="/wiki/Ornithology" title="Ornithology">Ornithology</a></li> <li><a href="/wiki/Parasitology" title="Parasitology">Parasitology</a></li> <li><a href="/wiki/Paleozoology" title="Paleozoology">Paleozoology</a></li> <li><a href="/wiki/Planktology" title="Planktology">Planktology</a></li> <li><a href="/wiki/Zooarchaeology" title="Zooarchaeology">Zooarchaeology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background:;width:1%"><a href="/wiki/Animal" title="Animal">Animal</a> groups</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/Sponge" title="Sponge">Porifera</a></li> <li><a href="/wiki/Ctenophora" title="Ctenophora">Ctenophora</a></li> <li><a href="/wiki/Placozoa" title="Placozoa">Placozoa</a></li> <li><a href="/wiki/Cnidaria" title="Cnidaria">Cnidaria</a></li> <li><a href="/wiki/Xenacoelomorpha" title="Xenacoelomorpha">Xenacoelomorpha</a></li> <li><a href="/wiki/Ambulacraria" title="Ambulacraria">Ambulacraria</a></li> <li><a href="/wiki/Chordate" title="Chordate">Chordata</a></li> <li><a href="/wiki/Aschelminthes" class="mw-redirect" title="Aschelminthes">Aschelminthes</a></li> <li><a href="/wiki/Arthropod" title="Arthropod">Arthropoda</a></li> <li><a href="/wiki/Flatworm" title="Flatworm">Platyhelminthes</a></li> <li><a href="/wiki/Mollusca" title="Mollusca">Mollusca</a></li> <li><a href="/wiki/Annelid" title="Annelid">Annelida</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background:;width:1%"><a href="/wiki/Anatomy" title="Anatomy">Animal anatomy</a><br /><a href="/wiki/Morphology_(biology)" title="Morphology (biology)">Animal morphology</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/Histology" title="Histology">Histology</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/Tissue_(biology)#Animal_tissue" title="Tissue (biology)">Tissues</a> <ul><li><a href="/wiki/Epithelium" title="Epithelium">Epithelial tissue</a></li> <li><a href="/wiki/Muscle_tissue" class="mw-redirect" title="Muscle tissue">Muscular tissue</a></li> <li><a href="/wiki/Nervous_tissue" title="Nervous tissue">Nervous tissue</a></li> <li><a href="/wiki/Connective_tissue" title="Connective tissue">Connective tissues</a></li> <li><a href="/wiki/Mineralized_tissues" title="Mineralized tissues">Mineralized tissues</a></li></ul></li> <li><a href="/wiki/Molecular_anatomy" title="Molecular anatomy">Molecular anatomy</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Comparative_anatomy" title="Comparative anatomy">Anatomy <br /> and morphology</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/Mammal#Anatomy" title="Mammal">Mammalian anatomy and morphology</a> <ul><li><a href="/wiki/Human_anatomy" title="Human anatomy">Human</a></li> <li><a href="/wiki/Dog_anatomy" title="Dog anatomy">Dog</a></li> <li><a href="/wiki/Elephant#Anatomy_and_morphology" title="Elephant">Elephant</a></li> <li><a href="/wiki/Cat_anatomy" title="Cat anatomy">Cat</a></li></ul></li> <li><a href="/wiki/Bird_anatomy" title="Bird anatomy">Bird anatomy</a></li> <li><a href="/wiki/Fish_anatomy" title="Fish anatomy">Fish anatomy</a> <ul><li><a href="/wiki/Shark_anatomy" title="Shark anatomy">Shark anatomy</a></li></ul></li> <li><a href="/wiki/Spider_anatomy" title="Spider anatomy">Spider anatomy</a></li> <li><a href="/wiki/Insect_morphology" title="Insect morphology">Insect morphology</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="background:;width:1%"><a href="/wiki/Physiology#Animals" title="Physiology">Animal physiology</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%">General physiology</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/Respiration_(physiology)" title="Respiration (physiology)">Respiration</a> <ul><li><a href="/wiki/Respiratory_system" title="Respiratory system">Respiratory system</a></li> <li><a href="/wiki/Breathing" title="Breathing">Breathing</a></li> <li><a href="/wiki/Gas_exchange" title="Gas exchange">Gas exchange</a></li> <li><a href="/wiki/Respiratory_pigment" title="Respiratory pigment">Respiratory pigment</a></li> <li><a href="/wiki/Cellular_respiration" title="Cellular respiration">Cellular respiration</a></li></ul></li> <li><a href="/wiki/Vascular_system" class="mw-redirect" title="Vascular system">Vascular system</a> <ul><li><a href="/wiki/Blood" title="Blood">Blood</a></li> <li><a href="/wiki/Lymph" title="Lymph">Lymph</a></li> <li><a href="/wiki/Blood_vessel" title="Blood vessel">Blood vessels</a> <ul><li><a href="/wiki/Arteries" class="mw-redirect" title="Arteries">Arteries</a></li> <li><a href="/wiki/Vein" title="Vein">Veins</a></li> <li><a href="/wiki/Capillaries" class="mw-redirect" title="Capillaries">Capillaries</a></li></ul></li> <li><a href="/wiki/Heart" title="Heart">Heart</a></li> <li><a href="/wiki/Water_vascular_system" title="Water vascular system">Water vascular system</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">By species</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/Insect_physiology" title="Insect physiology">Insect physiology</a></li> <li><a href="/wiki/Fish_physiology" title="Fish physiology">Fish physiology</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="background:;width:1%"><a href="/wiki/List_of_zoologists" class="mw-redirect" title="List of zoologists">Zoologists</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/Karl_Ernst_von_Baer" title="Karl Ernst von Baer">Karl Ernst von Baer</a></li> <li><a href="/wiki/Georges_Cuvier" title="Georges Cuvier">Georges Cuvier</a></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a></li> <li><a href="/wiki/Jean-Henri_Fabre" title="Jean-Henri Fabre">Jean-Henri Fabre</a></li> <li><a href="/wiki/William_Kirby_(entomologist)" title="William Kirby (entomologist)">William Kirby</a></li> <li><a href="/wiki/Carl_Linnaeus" title="Carl Linnaeus">Carl Linnaeus</a></li> <li><a href="/wiki/Konrad_Lorenz" title="Konrad Lorenz">Konrad Lorenz</a></li> <li><a href="/wiki/Thomas_Say" title="Thomas Say">Thomas Say</a></li> <li><a href="/wiki/Jakob_von_Uexk%C3%BCll" class="mw-redirect" title="Jakob von Uexküll">Jakob von Uexküll</a></li> <li><a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background:;width:1%">History</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/History_of_zoology_through_1859" title="History of zoology through 1859">Pre-Darwin</a></li> <li><a href="/wiki/History_of_zoology_(1859%E2%80%93present)" title="History of zoology (1859–present)">Post-Darwin</a></li> <li><a href="/wiki/Timeline_of_zoology" title="Timeline of zoology">Timeline of zoology</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="background:"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Zoology" title="Category:Zoology">Category</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_psychology1033" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evolutionary_psychology" title="Template:Evolutionary psychology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolutionary_psychology" title="Template talk:Evolutionary psychology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolutionary_psychology" title="Special:EditPage/Template:Evolutionary psychology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_psychology1033" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/History_of_evolutionary_psychology" title="History of evolutionary psychology">History</a> <ul><li><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">Evolutionary thought</a></li></ul></li> <li><a href="/wiki/Theoretical_foundations_of_evolutionary_psychology" title="Theoretical foundations of evolutionary psychology">Theoretical foundations</a> <ul><li><a href="/wiki/Adaptationism" title="Adaptationism">Adaptationism</a></li> <li><a href="/wiki/Cognitive_revolution" title="Cognitive revolution">Cognitive revolution</a></li> <li><a href="/wiki/Cognitivism_(psychology)" title="Cognitivism (psychology)">Cognitivism</a></li> <li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene selection theory</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li></ul></li> <li><a href="/wiki/Criticism_of_evolutionary_psychology" title="Criticism of evolutionary psychology">Criticism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;"><a href="/wiki/Human_evolution" title="Human evolution">Evolutionary<br />processes</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/Psychological_adaptation" title="Psychological adaptation">Adaptations</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a href="/wiki/Cheating_(biology)" title="Cheating (biology)">Cheating</a></li> <li><a href="/wiki/Hamiltonian_spite" title="Hamiltonian spite">Hamiltonian spite</a></li> <li><a href="/wiki/Reciprocal_altruism_in_humans" title="Reciprocal altruism in humans">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">By-products</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness_in_humans" title="Inclusive fitness in humans">Inclusive</a></li></ul></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a> <ul><li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Sexual_selection_in_humans" title="Sexual selection in humans">Sexual selection</a> <ul><li><a href="/wiki/Costly_signaling_theory_in_evolutionary_psychology" title="Costly signaling theory in evolutionary psychology">Costly signaling</a></li> <li><a href="/wiki/Male_intrasexual_competition" title="Male intrasexual competition">Male</a>/<a href="/wiki/Female_intrasexual_competition" title="Female intrasexual competition">female intrasexual competition</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;">Areas</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/Evolution_of_cognition" title="Evolution of cognition">Cognition</a> /<br /><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</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/Affect_(psychology)" title="Affect (psychology)">Affect</a> <ul><li><a href="/wiki/Affect_display" title="Affect display">Display</a></li> <li><a href="/wiki/Display_rules" title="Display rules">Display rules</a></li> <li><a href="/wiki/Facial_expression" title="Facial expression">Facial expression</a></li></ul></li> <li><a href="/wiki/Behavioral_modernity" title="Behavioral modernity">Behavioral modernity</a></li> <li><a href="/wiki/Cognitive_module" title="Cognitive module">Cognitive module</a>/<a href="/wiki/Modularity_of_mind" title="Modularity of mind">modularity of mind</a> <ul><li><a href="/wiki/Automatic_and_controlled_processes" title="Automatic and controlled processes">Automatic and controlled processes</a></li> <li><a href="/wiki/Computational_theory_of_mind" title="Computational theory of mind">Computational theory of mind</a></li> <li><a href="/wiki/Domain-general_learning" title="Domain-general learning">Domain generality</a></li> <li><a href="/wiki/Domain_specificity" title="Domain specificity">Domain specificity</a></li> <li><a href="/wiki/Dual_process_theory" title="Dual process theory">Dual process theory</a></li></ul></li> <li><a href="/wiki/Cognitive_tradeoff_hypothesis" title="Cognitive tradeoff hypothesis">Cognitive tradeoff hypothesis</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">Evolution of the brain</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">Evolution of nervous systems</a></li> <li><a href="/wiki/Fight-or-flight_response" title="Fight-or-flight response">Fight-or-flight response</a> <ul><li><a href="/wiki/Arachnophobia" title="Arachnophobia">Arachnophobia</a></li> <li><a href="/wiki/Fear_of_falling" title="Fear of falling">Basophobia</a></li> <li><a href="/wiki/Ophidiophobia" title="Ophidiophobia">Ophidiophobia</a></li></ul></li> <li><a href="/wiki/Folk_biology" title="Folk biology">Folk biology</a>/<a href="/wiki/Folk_taxonomy" title="Folk taxonomy">taxonomy</a></li> <li><a href="/wiki/Folk_psychology" title="Folk psychology">Folk psychology</a>/<a href="/wiki/Theory_of_mind" title="Theory of mind">theory of mind</a></li> <li><a href="/wiki/Evolution_of_human_intelligence" title="Evolution of human intelligence">Intelligence</a> <ul><li><a href="/wiki/Flynn_effect" title="Flynn effect">Flynn effect</a></li> <li><a href="/wiki/Wason_selection_task" title="Wason selection task">Wason selection task</a></li></ul></li> <li><a href="/wiki/Motor_control" title="Motor control">Motor control</a>/<a href="/wiki/Motor_skill" title="Motor skill">skill</a></li> <li><a href="/wiki/Human_multitasking" title="Human multitasking">Multitasking</a></li> <li><a href="/wiki/Neuroscience_of_sleep" title="Neuroscience of sleep">Sleep</a></li> <li><a href="/wiki/Visual_perception" title="Visual perception">Visual perception</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">Color vision</a></li> <li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">Eye</a></li> <li><a href="/wiki/Na%C3%AFve_physics" title="Naïve physics">Naïve physics</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolutionary_psychology_and_culture" title="Evolutionary psychology and culture">Culture</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolutionary_aesthetics" title="Evolutionary aesthetics">Aesthetics</a> <ul><li><a href="/wiki/Darwinian_literary_studies" title="Darwinian literary studies">Literary criticism</a></li> <li><a href="/wiki/Evolutionary_musicology" title="Evolutionary musicology">Musicology</a></li></ul></li> <li><a href="/wiki/Evolutionary_anthropology" title="Evolutionary anthropology">Anthropology</a> <ul><li><a href="/wiki/Biological_anthropology" title="Biological anthropology">Biological</a></li></ul></li> <li><a href="/wiki/Biosocial_criminology" title="Biosocial criminology">Crime</a></li> <li><a href="/wiki/Evolutionary_linguistics" title="Evolutionary linguistics">Language</a> <ul><li><a href="/wiki/Origin_of_language" title="Origin of language">Origin</a></li> <li><a href="/wiki/Evolutionary_psychology_of_language" title="Evolutionary psychology of language">Psychology</a></li> <li><a href="/wiki/Origin_of_speech" title="Origin of speech">Speech</a></li></ul></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a> <ul><li><a href="/wiki/Moral_foundations_theory" title="Moral foundations theory">Moral foundations</a></li></ul></li> <li><a href="/wiki/Evolutionary_psychology_of_religion" title="Evolutionary psychology of religion">Religion</a> <ul><li><a href="/wiki/Evolutionary_origin_of_religions" class="mw-redirect" title="Evolutionary origin of religions">Origin</a></li></ul></li> <li><a href="/wiki/Cultural_universal" title="Cultural universal">Universals</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolutionary_developmental_psychology" title="Evolutionary developmental psychology">Development</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/Attachment_theory" title="Attachment theory">Attachment</a></li> <li><a href="/wiki/Human_bonding" title="Human bonding">Bonding</a></li> <li><a href="/wiki/Affectional_bond" title="Affectional bond">Affectional</a>/<a href="/wiki/Maternal_bond" title="Maternal bond">maternal</a>/<a href="/wiki/Paternal_bond" title="Paternal bond">paternal bond</a></li> <li><a href="/wiki/Maternal_deprivation" title="Maternal deprivation">Caregiver deprivation</a></li> <li><a href="/wiki/Attachment_in_children" title="Attachment in children">Childhood attachment</a></li> <li><a href="/wiki/Cinderella_effect" title="Cinderella effect">Cinderella effect</a></li> <li><a href="/wiki/Cognitive_development" title="Cognitive development">Cognitive development</a></li> <li><a href="/wiki/Evolutionary_educational_psychology" title="Evolutionary educational psychology">Education</a></li> <li><a href="/wiki/Language_acquisition" title="Language acquisition">Language acquisition</a></li> <li><a href="/wiki/Personality_development" title="Personality development">Personality development</a></li> <li><a href="/wiki/Socialization" title="Socialization">Socialization</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ergonomics" title="Ergonomics">Human factors</a> /<br /><a href="/wiki/Evolutionary_psychiatry" title="Evolutionary psychiatry">Mental health</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/Cognitive_ergonomics" title="Cognitive ergonomics">Cognitive ergonomics</a> <ul><li><a href="/wiki/Computer-mediated_communication" title="Computer-mediated communication">Computer-mediated communication</a></li> <li><a href="/wiki/Engineering_psychology" title="Engineering psychology">Engineering psychology</a></li> <li><a href="/wiki/Human%E2%80%93computer_interaction" title="Human–computer interaction">Human–computer interaction</a></li> <li><a href="/wiki/Media_naturalness_theory" title="Media naturalness theory">Media naturalness theory</a></li> <li><a href="/wiki/Neuroergonomics" title="Neuroergonomics">Neuroergonomics</a></li></ul></li> <li><a href="/wiki/Evolutionary_approaches_to_depression" title="Evolutionary approaches to depression">Depression</a></li> <li><a href="/wiki/Digital_media_use_and_mental_health" title="Digital media use and mental health">Digital media use and mental health</a></li> <li><a href="/wiki/Accident-proneness" title="Accident-proneness">Hypophobia</a></li> <li><a href="/wiki/Imprinted_brain_hypothesis" title="Imprinted brain hypothesis">Imprinted brain hypothesis</a></li> <li><a href="/wiki/Mind-blindness" title="Mind-blindness">Mind-blindness</a></li> <li><a href="/wiki/Psychological_effects_of_Internet_use" title="Psychological effects of Internet use">Psychological effects of Internet use</a></li> <li><a href="/wiki/Rank_theory_of_depression" title="Rank theory of depression">Rank theory of depression</a></li> <li><a href="/wiki/Evolution_of_schizophrenia" title="Evolution of schizophrenia">Schizophrenia</a></li> <li><a href="/wiki/Screen_time" title="Screen time">Screen time</a></li> <li><a href="/wiki/Smartphones_and_pedestrian_safety" title="Smartphones and pedestrian safety">Smartphones and pedestrian safety</a></li> <li><a href="/wiki/Social_aspects_of_television" title="Social aspects of television">Social aspects of television</a></li> <li><a href="/wiki/Societal_effects_of_cars" title="Societal effects of cars">Societal effects of cars</a> <ul><li><a href="/wiki/Distracted_driving" title="Distracted driving">Distracted driving</a></li> <li><a href="/wiki/Lead%E2%80%93crime_hypothesis" title="Lead–crime hypothesis">Lead–crime hypothesis</a></li> <li><a href="/wiki/Mobile_phones_and_driving_safety" title="Mobile phones and driving safety">Mobile phones and driving safety</a></li> <li><a href="/wiki/Texting_while_driving" title="Texting while driving">Texting while driving</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sexology" title="Sexology">Sex</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/Human_sexual_activity" title="Human sexual activity">Activity</a></li> <li><a href="/wiki/Attachment_in_adults" title="Attachment in adults">Adult attachment</a></li> <li><a href="/wiki/Age_disparity_in_sexual_relationships" title="Age disparity in sexual relationships">Age disparity</a></li> <li><a href="/wiki/Sexual_arousal" title="Sexual arousal">Arousal</a></li> <li><a href="/wiki/Concealed_ovulation" title="Concealed ovulation">Concealed ovulation</a></li> <li><a href="/wiki/Coolidge_effect" title="Coolidge effect">Coolidge effect</a></li> <li><a href="/wiki/Sexual_desire" title="Sexual desire">Desire</a></li> <li><a href="/wiki/Sexual_fantasy" title="Sexual fantasy">Fantasy</a></li> <li><a href="/wiki/Effects_of_hormones_on_sexual_motivation" title="Effects of hormones on sexual motivation">Hormonal motivation</a></li> <li><a href="/wiki/Sexual_jealousy" title="Sexual jealousy">Jealousy</a></li> <li><a href="/wiki/Mate_guarding_in_humans" title="Mate guarding in humans">Mate guarding</a></li> <li><a href="/wiki/Mating_preferences" title="Mating preferences">Mating preferences</a></li> <li><a href="/wiki/Human_mating_strategies" title="Human mating strategies">Mating strategies</a></li> <li><a href="/wiki/Biology_and_sexual_orientation" title="Biology and sexual orientation">Orientation</a></li> <li><a href="/wiki/Ovulatory_shift_hypothesis" title="Ovulatory shift hypothesis">Ovulatory shift hypothesis</a></li> <li><a href="/wiki/Pair_bond" title="Pair bond">Pair bond</a></li> <li><a href="/wiki/Physical_attractiveness" title="Physical attractiveness">Physical</a>/<a href="/wiki/Sexual_attraction" title="Sexual attraction">Sexual attraction</a></li> <li><a href="/wiki/Human_sexuality" title="Human sexuality">Sexuality</a>/<a href="/wiki/Human_male_sexuality" title="Human male sexuality">male</a>/<a href="/wiki/Human_female_sexuality" title="Human female sexuality">female</a></li> <li><a href="/wiki/Sexy_son_hypothesis" title="Sexy son hypothesis">Sexy son hypothesis</a></li> <li><a href="/wiki/Westermarck_effect" title="Westermarck effect">Westermarck effect</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sex_differences_in_psychology" title="Sex differences in psychology">Sex differences</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/Aggression" title="Aggression">Aggression</a></li> <li><a href="/wiki/Sex_and_gender_differences_in_autism" title="Sex and gender differences in autism">Autism</a></li> <li><a href="/wiki/Sex_differences_in_cognition" title="Sex differences in cognition">Cognition</a></li> <li><a href="/wiki/Sex_differences_in_crime" title="Sex differences in crime">Crime</a></li> <li><a href="/wiki/Sexual_division_of_labour" title="Sexual division of labour">Division of labour</a></li> <li><a href="/wiki/Sex_differences_in_emotional_intelligence" title="Sex differences in emotional intelligence">Emotional intelligence</a></li> <li><a href="/wiki/Empathising%E2%80%93systemising_theory" title="Empathising–systemising theory">Empathising–systemising theory</a></li> <li><a href="/wiki/Gender_role" title="Gender role">Gender role</a></li> <li><a href="/wiki/Sex_differences_in_intelligence" title="Sex differences in intelligence">Intelligence</a></li> <li><a href="/wiki/Sex_differences_in_memory" title="Sex differences in memory">Memory</a></li> <li><a href="/wiki/Mental_disorders_and_gender" title="Mental disorders and gender">Mental health</a></li> <li><a href="/wiki/Sex_differences_in_narcissism" title="Sex differences in narcissism">Narcissism</a></li> <li><a href="/wiki/Neuroscience_of_sex_differences" title="Neuroscience of sex differences">Neuroscience</a></li> <li><a href="/wiki/Sex_differences_in_schizophrenia" title="Sex differences in schizophrenia">Schizophrenia</a></li> <li><a href="/wiki/Substance_abuse" title="Substance abuse">Substance abuse</a></li> <li><a href="/wiki/Gender_differences_in_suicide" title="Gender differences in suicide">Suicide</a></li> <li><a href="/wiki/Variability_hypothesis" title="Variability hypothesis">Variability hypothesis</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;">Related subjects</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%">Academic disciplines</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/Behavioral_economics" title="Behavioral economics">Behavioral</a>/<a href="/wiki/Evolutionary_economics" title="Evolutionary economics">evolutionary economics</a></li> <li><a href="/wiki/Behavioral_epigenetics" title="Behavioral epigenetics">Behavioral epigenetics</a>/<a href="/wiki/Behavioural_genetics" title="Behavioural genetics">genetics</a></li> <li><a href="/wiki/Affective_neuroscience" title="Affective neuroscience">Affective</a>/<a href="/wiki/Behavioral_neuroscience" title="Behavioral neuroscience">behavioral</a>/<a href="/wiki/Cognitive_neuroscience" title="Cognitive neuroscience">cognitive</a>/<a href="/wiki/Evolutionary_neuroscience" title="Evolutionary neuroscience">evolutionary neuroscience</a></li> <li><a href="/wiki/Biocultural_anthropology" title="Biocultural anthropology">Biocultural anthropology</a></li> <li><a href="/wiki/Biological_psychiatry" title="Biological psychiatry">Biological psychiatry</a></li> <li><a href="/wiki/Cognitive_psychology" title="Cognitive psychology">Cognitive psychology</a></li> <li><a href="/wiki/Cognitive_science" title="Cognitive science">Cognitive science</a></li> <li><a href="/wiki/Cross-cultural_psychology" title="Cross-cultural psychology">Cross-cultural psychology</a></li> <li><a class="mw-selflink selflink">Ethology</a></li> <li><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Functional_psychology" title="Functional psychology">Functional psychology</a></li> <li><a href="/wiki/Neuropsychology" title="Neuropsychology">Neuropsychology</a></li> <li><a href="/wiki/Philosophy_of_mind" title="Philosophy of mind">Philosophy of mind</a></li> <li><a href="/wiki/Population_genetics" title="Population genetics">Population genetics</a></li> <li><a href="/wiki/Primatology" title="Primatology">Primatology</a></li> <li><a href="/wiki/Sociobiology" title="Sociobiology">Sociobiology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Research topics</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Evolutionary_epistemology" title="Evolutionary epistemology">Evolutionary epistemology</a></li> <li><a href="/wiki/Great_ape_language" title="Great ape language">Great ape language</a></li> <li><a href="/wiki/Human%E2%80%93animal_communication" title="Human–animal communication">Human–animal communication</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Primate_cognition" title="Primate cognition">Primate cognition</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a></li> <li><a href="/wiki/Recent_human_evolution" title="Recent human evolution">Recent human evolution</a></li> <li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Theoretical positions</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/Cultural_selection_theory" title="Cultural selection theory">Cultural selection theory</a></li> <li><a href="/wiki/Determinism" title="Determinism">Determinism</a>/<a href="/wiki/Indeterminism" title="Indeterminism">indeterminism</a> <ul><li><a href="/wiki/Biological_determinism" title="Biological determinism">Biological determinism</a></li> <li><a href="/wiki/Connectionism" title="Connectionism">Connectionism</a></li> <li><a href="/wiki/Cultural_determinism" title="Cultural determinism">Cultural determinism</a></li> <li><a href="/wiki/Environmental_determinism" title="Environmental determinism">Environmental determinism</a></li> <li><a href="/wiki/Nature_versus_nurture" title="Nature versus nurture">Nature versus nurture</a></li> <li><a href="/wiki/Psychological_nativism" title="Psychological nativism">Psychological nativism</a></li> <li><a href="/wiki/Social_constructionism" title="Social constructionism">Social constructionism</a></li> <li><a href="/wiki/Social_determinism" title="Social determinism">Social determinism</a></li> <li><a href="/wiki/Standard_social_science_model" title="Standard social science model">Standard social science model</a></li></ul></li> <li><a href="/wiki/Functionalism_(philosophy_of_mind)" title="Functionalism (philosophy of mind)">Functionalism</a></li> <li><a href="/wiki/Memetics" title="Memetics">Memetics</a></li> <li><a href="/wiki/Multilineal_evolution" title="Multilineal evolution">Multilineal evolution</a></li> <li><a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li> <li><a href="/wiki/Neoevolutionism" title="Neoevolutionism">Neoevolutionism</a></li> <li><a href="/wiki/Sociocultural_evolution" title="Sociocultural evolution">Sociocultural evolution</a></li> <li><a href="/wiki/Unilineal_evolution" title="Unilineal evolution">Unilineal evolution</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><b><a href="/wiki/List_of_evolutionary_psychologists" title="List of evolutionary psychologists">Evolutionary psychologists</a></b></li> 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