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Monogamy in animals - Wikipedia
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<span class="vector-toc-numb">2</span> <span>Social monogamy</span> </div> </a> <ul id="toc-Social_monogamy-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genetic_monogamy" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Genetic_monogamy"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Genetic monogamy</span> </div> </a> <ul id="toc-Genetic_monogamy-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolution_of_monogamy_in_animals" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Evolution_of_monogamy_in_animals"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Evolution of monogamy in animals</span> </div> </a> <button aria-controls="toc-Evolution_of_monogamy_in_animals-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Evolution of monogamy in animals subsection</span> </button> <ul id="toc-Evolution_of_monogamy_in_animals-sublist" class="vector-toc-list"> <li id="toc-Anisogamy" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Anisogamy"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Anisogamy</span> </div> </a> <ul id="toc-Anisogamy-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Facultative_monogamy" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Facultative_monogamy"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Facultative monogamy</span> </div> </a> <ul id="toc-Facultative_monogamy-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Obligate_monogamy" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Obligate_monogamy"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Obligate monogamy</span> </div> </a> <ul id="toc-Obligate_monogamy-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Enforcement" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Enforcement"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Enforcement</span> </div> </a> <ul id="toc-Enforcement-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mate_assistance_and_territory_defense" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Mate_assistance_and_territory_defense"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Mate assistance and territory defense</span> </div> </a> <ul id="toc-Mate_assistance_and_territory_defense-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Consequences_of_monogamous_mating" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Consequences_of_monogamous_mating"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Consequences of monogamous mating</span> </div> </a> <ul id="toc-Consequences_of_monogamous_mating-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Sexual_dimorphism" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Sexual_dimorphism"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Sexual dimorphism</span> </div> </a> <ul id="toc-Sexual_dimorphism-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Sperm_quality" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Sperm_quality"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Sperm quality</span> </div> </a> <ul id="toc-Sperm_quality-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Animals" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Animals"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>Animals</span> </div> </a> <button aria-controls="toc-Animals-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 Animals subsection</span> </button> <ul id="toc-Animals-sublist" class="vector-toc-list"> <li id="toc-Sexual_dimorphism_2" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Sexual_dimorphism_2"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.1</span> <span>Sexual dimorphism</span> </div> </a> <ul id="toc-Sexual_dimorphism_2-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Testis_size" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Testis_size"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.2</span> <span>Testis size</span> </div> </a> <ul id="toc-Testis_size-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Monogamy_as_a_best_response" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Monogamy_as_a_best_response"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.3</span> <span>Monogamy as a best response</span> </div> </a> <ul id="toc-Monogamy_as_a_best_response-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Monogamous_species" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Monogamous_species"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.4</span> <span>Monogamous species</span> </div> </a> <ul id="toc-Monogamous_species-sublist" class="vector-toc-list"> </ul> </li> </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">11</span> <span>See also</span> </div> </a> <button aria-controls="toc-See_also-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 See also subsection</span> </button> <ul id="toc-See_also-sublist" class="vector-toc-list"> <li id="toc-Monogamy_topics" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Monogamy_topics"> <div class="vector-toc-text"> <span class="vector-toc-numb">11.1</span> <span>Monogamy topics</span> </div> </a> <ul id="toc-Monogamy_topics-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolution_topics" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Evolution_topics"> <div 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<h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Monogamy in animals</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 8 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-8" 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">8 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%A7%D9%84%D8%B2%D9%88%D8%A7%D8%AC_%D8%A7%D9%84%D8%A3%D8%AD%D8%A7%D8%AF%D9%8A_%D9%81%D9%8A_%D8%A7%D9%84%D8%AD%D9%8A%D9%88%D8%A7%D9%86%D8%A7%D8%AA" title="الزواج الأحادي في الحيوانات – Arabic" lang="ar" hreflang="ar" data-title="الزواج الأحادي في الحيوانات" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Heyvanlarda_monoqamiya" title="Heyvanlarda monoqamiya – Azerbaijani" lang="az" hreflang="az" data-title="Heyvanlarda monoqamiya" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%AA%DA%A9%E2%80%8C%D9%87%D9%85%D8%B3%D8%B1%DB%8C_%D8%AF%D8%B1_%D8%AC%D8%A7%D9%86%D9%88%D8%B1%D8%A7%D9%86" title="تکهمسری در جانوران – Persian" lang="fa" hreflang="fa" data-title="تکهمسری در جانوران" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D4%BF%D5%A5%D5%B6%D5%A4%D5%A1%D5%B6%D5%AB%D5%B6%D5%A5%D6%80%D5%AB_%D5%B4%D5%A5%D5%B6%D5%A1%D5%B4%D5%B8%D6%82%D5%BD%D5%B6%D5%B8%D6%82%D5%A9%D5%B5%D5%B8%D6%82%D5%B6" title="Կենդանիների մենամուսնություն – Armenian" lang="hy" hreflang="hy" data-title="Կենդանիների մենամուսնություն" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%9E%D7%95%D7%A0%D7%95%D7%92%D7%9E%D7%99%D7%94_%D7%91%D7%91%D7%A2%D7%9C%D7%99_%D7%97%D7%99%D7%99%D7%9D" title="מונוגמיה בבעלי חיים – Hebrew" lang="he" hreflang="he" data-title="מונוגמיה בבעלי חיים" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Monogamiya_(biologiya)" title="Monogamiya (biologiya) – Uzbek" lang="uz" hreflang="uz" data-title="Monogamiya (biologiya)" 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-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Monogamia_(biologia)" title="Monogamia (biologia) – Polish" lang="pl" hreflang="pl" data-title="Monogamia (biologia)" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Chung_th%E1%BB%A7y_%E1%BB%9F_%C4%91%E1%BB%99ng_v%E1%BA%ADt" title="Chung thủy ở động vật – Vietnamese" lang="vi" hreflang="vi" data-title="Chung thủy ở động vật" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li> </ul> <div 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id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Natural history of mating systems in which species pair bond to raise offspring</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">This article is about pairing for animals in non-human species. For monogamy in humans, see <a href="/wiki/Monogamy" title="Monogamy">monogamy</a>.</div> <p><b>Monogamous pairing in animals</b> refers to the natural history of <a href="/wiki/Mating_system" title="Mating system">mating systems</a> in which species <a href="/wiki/Pair_bond" title="Pair bond">pair bond</a> to raise offspring. This is associated, usually implicitly, with <a href="/wiki/Sexual_monogamy" class="mw-redirect" title="Sexual monogamy">sexual monogamy</a>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2010)">citation needed</span></a></i>]</sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Monogamous_mating">Monogamous mating</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=1" title="Edit section: Monogamous mating"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Monogamy is defined as a pair bond between two adult animals of the same species. This pair may cohabitate in an area or territory for some duration of time, and in some cases may <a href="/wiki/Copulation_(zoology)" title="Copulation (zoology)">copulate</a> and reproduce with only each other. Monogamy may either be short-term, lasting one to a few seasons or long-term, lasting many seasons and in extreme cases, life-long. Monogamy can be partitioned into two categories, social monogamy and genetic monogamy which may occur together in some combination, or completely independently of one another.<sup id="cite_ref-Ophir-2008_1-0" class="reference"><a href="#cite_note-Ophir-2008-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> As an example, in the cichlid species <i><a href="/wiki/Variabilichromis_moorii" title="Variabilichromis moorii">Variabilichromis moorii</a></i>, a monogamous pair will care for eggs and young together, but the eggs may not all be fertilized by the male giving the care.<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> Monogamy in mammals is rather rare, only occurring in 3–9% of these species.<sup id="cite_ref-Kleiman-1977_3-0" class="reference"><a href="#cite_note-Kleiman-1977-3"><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-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> A larger percentage of avian species are known to have monogamous relationships (about 90%),<sup id="cite_ref-Lack-1968_6-0" class="reference"><a href="#cite_note-Lack-1968-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> but most avian species practice social but not genetic monogamy in contrast to what was previously assumed by researchers.<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> Monogamy is quite rare in fish and amphibians, but not unheard of, appearing in a select few species.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Social_monogamy">Social monogamy</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=2" title="Edit section: Social monogamy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Social monogamy refers to the cohabitation of one male and one female. The two individuals may cooperate in search of resources such as food and shelter and/or in caring for young.<sup id="cite_ref-Ophir-2008_1-1" class="reference"><a href="#cite_note-Ophir-2008-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><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> Paternal care in monogamous species is commonly displayed through carrying, feeding, defending, and socializing offspring.<sup id="cite_ref-Kleiman-1977_3-1" class="reference"><a href="#cite_note-Kleiman-1977-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><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> With social monogamy there may not be an expected sexual fidelity between the males and the females.<sup id="cite_ref-Ophir-2008_1-2" class="reference"><a href="#cite_note-Ophir-2008-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Mainwaring-2013_13-0" class="reference"><a href="#cite_note-Mainwaring-2013-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> The existence of purely social monogamy is a polygamous or polyandrous social pair with <a href="/wiki/Extra-pair_copulation" title="Extra-pair copulation">extra pair coupling</a>.<sup id="cite_ref-Sundin-2009_14-0" class="reference"><a href="#cite_note-Sundin-2009-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Social monogamy has been shown to increase fitness in prairie voles. It has been shown that female prairie voles live longer when paired with males in a social monogamous relationship. This could be because of the shared energy expenditure by the males and females lower each individual's input.<sup id="cite_ref-Ophir-2008_1-3" class="reference"><a href="#cite_note-Ophir-2008-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> In largemouth bass, females are sometimes seen to exhibit <a href="/wiki/Cuckoo" title="Cuckoo">cuckoo</a> behavior by laying some of their eggs in another female's nest, thus "stealing" fertilizations from other females.<sup id="cite_ref-Dewoody-2000_15-0" class="reference"><a href="#cite_note-Dewoody-2000-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> Sexual conflicts that have been proposed to arise from social monogamy include infidelity and parental investment. The proposed conflict is derived from the conflict-centric differential allocation hypothesis, which states that there is a tradeoff between investment and attractiveness.<sup id="cite_ref-Mainwaring-2013_13-1" class="reference"><a href="#cite_note-Mainwaring-2013-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Genetic_monogamy">Genetic monogamy</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=3" title="Edit section: Genetic monogamy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Genetic monogamy refers to a mating system in which fidelity of the bonding pair is exhibited.<sup id="cite_ref-Ophir-2008_1-4" class="reference"><a href="#cite_note-Ophir-2008-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Though individual pairs may be genetically monogamous, no one species has been identified as fully genetically monogamous. </p><p>In some species, genetic monogamy has been enforced.<sup id="cite_ref-Schacht-2016_16-0" class="reference"><a href="#cite_note-Schacht-2016-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Female voles have shown no difference in fecundity with genetic monogamy, but it may be enforced by males in some instances.<sup id="cite_ref-Ophir-2008_1-5" class="reference"><a href="#cite_note-Ophir-2008-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Mate guarding is a typical tactic in monogamous species.<sup id="cite_ref-Mainwaring-2013_13-2" class="reference"><a href="#cite_note-Mainwaring-2013-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Dewoody-2000_15-1" class="reference"><a href="#cite_note-Dewoody-2000-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kokko-2006_17-0" class="reference"><a href="#cite_note-Kokko-2006-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> It is present in many animal species and can sometimes be expressed in lieu of parental care by males. This may be for many reasons, including paternity assurance.<sup id="cite_ref-Schacht-2016_16-1" class="reference"><a href="#cite_note-Schacht-2016-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kokko-2006_17-1" class="reference"><a href="#cite_note-Kokko-2006-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Evolution_of_monogamy_in_animals">Evolution of monogamy in animals</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=4" title="Edit section: Evolution of monogamy in animals"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>While the evolution of monogamy in animals cannot be broadly ascertained, there are several theories as to how monogamy may have evolved. </p> <div class="mw-heading mw-heading3"><h3 id="Anisogamy">Anisogamy</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=5" title="Edit section: Anisogamy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Anisogamy" title="Anisogamy">Anisogamy</a> is a form of sexual reproduction which involves the fusion of two unequally-sized gametes. In many animals, there are two sexes: the male, in which the gamete is small, motile, usually plentiful, and less energetically expensive, and the female, in which the gamete is larger, more energetically expensive, made at a lower rate, and largely immobile. Anisogamy is thought to have evolved from <a href="/wiki/Isogamy" title="Isogamy">isogamy</a>, the fusion of similar gametes, multiple times in many different species.<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>The introduction of anisogamy has caused males and females to tend to have different optimal mating strategies.<sup id="cite_ref-Sundin-2009_14-1" class="reference"><a href="#cite_note-Sundin-2009-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> This is because males may increase their fitness by mating with many females, whereas females are limited by their own fecundity. Females are therefore typically more likely to be selective in choosing mates.<sup id="cite_ref-Avise-2010_19-0" class="reference"><a href="#cite_note-Avise-2010-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> Monogamy is suggested to limit fitness differences, as males and females will mate in pairs.<sup id="cite_ref-Dewoody-2000_15-2" class="reference"><a href="#cite_note-Dewoody-2000-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> This would seem to be non-beneficial to males, but may not be in all cases. Several behaviors and ecological concerns may have led to the evolution of monogamy as a relevant mating strategy. Partner and resource availability, enforcement, mate assistance, and territory defense may be some of the most prevalent factors affecting animal behavior. </p> <div class="mw-heading mw-heading3"><h3 id="Facultative_monogamy">Facultative monogamy</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=6" title="Edit section: Facultative monogamy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>First introduced by Kleiman,<sup id="cite_ref-Kleiman-1977_3-2" class="reference"><a href="#cite_note-Kleiman-1977-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> facultative monogamy occurs when females are widely dispersed. This can either occur because females in a species tend to be solitary or because the distribution of resources available cause females to thrive when separated into distinct territories. In these instances, there is less of a chance for a given male to find multiple females to mate with. In such a case, it becomes more advantageous for a male to remain with a female, rather than seeking out another and risking (a) not finding another female and or (b) not being able to fight off another male from interfering with his offspring by mating with the female or through <a href="/wiki/Infanticide_(zoology)" title="Infanticide (zoology)">infanticide</a>. In these situations, male-to-male competition is reduced and female choice is limited. The end result is that the <a href="/wiki/Mate_choice" title="Mate choice">mate choice</a> is more random than in a more dense population, which has a number of effects including limiting dimorphism and <a href="/wiki/Sexual_selection" title="Sexual selection">sexual selection</a>.<sup id="cite_ref-Schacht-2016_16-2" class="reference"><a href="#cite_note-Schacht-2016-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kokko-2006_17-2" class="reference"><a href="#cite_note-Kokko-2006-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p><p>With resource availability limited, mating with multiple mates may be harder because the density of individuals is lowered. The habitat cannot sustain multiple mates, so monogamy may be more prevalent. This is because resources may be found more easily for the pair than for the individual. The argument for resource availability has been shown in many species, but in several species, once resource availability increases, monogamy is still apparent.<sup id="cite_ref-Sundin-2009_14-2" class="reference"><a href="#cite_note-Sundin-2009-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Schacht-2016_16-3" class="reference"><a href="#cite_note-Schacht-2016-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kokko-2006_17-3" class="reference"><a href="#cite_note-Kokko-2006-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p><p>With increased resource availability, males may be offsetting the restriction of their fitness through several means. In instances of social monogamy, males may offset any lowered fitness through <a href="/wiki/Extra-pair_copulation" title="Extra-pair copulation">extra pair coupling</a>. Extra pair coupling refers to male and females mating with several mates but only raising offspring with one mate. The male may not be related to all of the offspring of his main mate, but some offspring are being raised in other broods by other males and females, thereby offsetting any limitation of monogamy.<sup id="cite_ref-Sundin-2009_14-3" class="reference"><a href="#cite_note-Sundin-2009-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Males are cuckolds, but because they have other female sexual partners, they cuckold other males and increase their own fitness. Males exhibit parental care habits in order to be an acceptable mate to the female. Any males that do not exhibit parental care would not be accepted as a sexual partner for socially monogamous females in an enforcement pattern.<sup id="cite_ref-Sundin-2009_14-4" class="reference"><a href="#cite_note-Sundin-2009-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Obligate_monogamy">Obligate monogamy</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=7" title="Edit section: Obligate monogamy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Kleiman also offered a second theory. In obligate monogamy, the driving force behind monogamy is a greater need for paternal investment. This theory assumes that without biparental care fitness level of offspring would be greatly reduced. This paternal care may or may not be equal to that of the maternal care. </p><p>Related to paternal care, some researchers have argued that infanticide is the true cause of monogamy.<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> This theory has not garnered much support, however, critiqued by several authors including Lukas and Clutton-Brock and Dixson.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Enforcement">Enforcement</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=8" title="Edit section: Enforcement"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Monogamous mating may also be caused simply by enforcement through tactics such as mate guarding.<sup id="cite_ref-Mainwaring-2013_13-3" class="reference"><a href="#cite_note-Mainwaring-2013-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Dewoody-2000_15-3" class="reference"><a href="#cite_note-Dewoody-2000-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> In these species, the males will prevent other males from copulating with their chosen female or vice versa.<sup id="cite_ref-Schacht-2016_16-4" class="reference"><a href="#cite_note-Schacht-2016-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Males will help to fend off other aggressive males, and keep their mate for themselves. This is not seen in all species, such as some primates, in which the female may be more dominant than the male and may not need help to avoid unwanted mating; the pair may still benefit from some form of mate assistance, however, and therefore monogamy may be enforced to ensure the assistance of males. Bi-parental care is not seen in all monogamous species, however, so this may not be the only cause of female enforcement.<sup id="cite_ref-Sundin-2009_14-5" class="reference"><a href="#cite_note-Sundin-2009-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Mate_assistance_and_territory_defense">Mate assistance and territory defense</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=9" title="Edit section: Mate assistance and territory defense"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In species where mate guarding is not needed, there may still be a need for the pair to protect each other. An example of this would be sentinel behavior in avian species.<sup id="cite_ref-Mainwaring-2013_13-4" class="reference"><a href="#cite_note-Mainwaring-2013-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> The main advantage of sentinel behavior is that many survival tactics are improved. As stated, the male or female will act as a sentinel and signal to their mate if a predator is present. This can lead to an increase in survivorship, foraging, and incubation of eggs.<sup id="cite_ref-Mainwaring-2013_13-5" class="reference"><a href="#cite_note-Mainwaring-2013-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>Male care for offspring is rather rare in some taxa of species. This is because males may increase their fitness by searching for multiple mates.<sup id="cite_ref-Schacht-2016_16-5" class="reference"><a href="#cite_note-Schacht-2016-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Gubernick-2000_23-0" class="reference"><a href="#cite_note-Gubernick-2000-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Females are limited in fitness by their fecundity, so multiple mating does not affect their fitness to the same extent.<sup id="cite_ref-Avise-2010_19-1" class="reference"><a href="#cite_note-Avise-2010-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> Males have the opportunity to find a new mate earlier than females when there is internal fertilization or the females exhibit the majority of the care for the offspring.<sup id="cite_ref-Gubernick-2000_23-1" class="reference"><a href="#cite_note-Gubernick-2000-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> When males are shown to care for offspring as well as females, it is referred to as bi-parental care. </p><p>Bi-parental care may occur when there is a lower chance of survival of the offspring without male care. The evolution of this care has been associated with energetically expensive offspring.<sup id="cite_ref-Schacht-2016_16-6" class="reference"><a href="#cite_note-Schacht-2016-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Bi-parental care is exhibited in many avian species.<sup id="cite_ref-Mainwaring-2013_13-6" class="reference"><a href="#cite_note-Mainwaring-2013-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> In these cases, the male has a greater chance to increase his own fitness by seeing that his offspring live long enough to reproduce. If the male is not present in these populations, the survivorship of the offspring is drastically lowered and there is a lowering in male fitness.<sup id="cite_ref-Mainwaring-2013_13-7" class="reference"><a href="#cite_note-Mainwaring-2013-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Gubernick-2000_23-2" class="reference"><a href="#cite_note-Gubernick-2000-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Without monogamy, bi-parental care is less common and there is an increased chance of <a href="/wiki/Infanticide" title="Infanticide">infanticide</a>.<sup id="cite_ref-Ophir-2008_1-6" class="reference"><a href="#cite_note-Ophir-2008-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Infanticide with monogamous pairing would lead to a lowered fitness for socially monogamous males and is not seen to a wide extent.<sup id="cite_ref-Schacht-2016_16-7" class="reference"><a href="#cite_note-Schacht-2016-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Consequences_of_monogamous_mating">Consequences of monogamous mating</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=10" title="Edit section: Consequences of monogamous mating"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Monogamy as a mating system in animals has been thought to lower levels of some pre and post copulatory competition methods.<sup id="cite_ref-Dewoody-2000_15-4" class="reference"><a href="#cite_note-Dewoody-2000-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kokko-2006_17-4" class="reference"><a href="#cite_note-Kokko-2006-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-vanderHorst-2014_24-0" class="reference"><a href="#cite_note-vanderHorst-2014-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> Because of this reduction in competition in some instances the regulation of certain morphological characteristics may be lowered. This would result in a vast variety of morphological and physiological differences such as sexual dimorphism and sperm quality. </p> <div class="mw-heading mw-heading2"><h2 id="Sexual_dimorphism">Sexual dimorphism</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=11" title="Edit section: Sexual dimorphism"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Sexual dimorphism denotes the differences in males and females of the same species. Even in animals with seemingly no morphological sexual dimorphism visible there is still dimorphism in the gametes. Among mammals, males have the smaller gametes and females have the larger gametes. As soon as the two sexes emerge the dimorphism in the gamete structures and sizes may lead to further dimorphism in the species.<sup id="cite_ref-Lehtonen-2016_25-0" class="reference"><a href="#cite_note-Lehtonen-2016-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Sexual dimorphism is often caused through evolution in response to <a href="/wiki/Male_male_competition" class="mw-redirect" title="Male male competition">male male competition</a> and female choice.<sup id="cite_ref-Dewoody-2000_15-5" class="reference"><a href="#cite_note-Dewoody-2000-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> In polygamous species there is a noted sexual dimorphism. The sexual dimorphism is seen typically in sexual signaling aspects of morphology. Males typically exhibit these dimorphic traits and they are typically traits which help in signaling to females or male male competition.<sup id="cite_ref-Kokko-2006_17-5" class="reference"><a href="#cite_note-Kokko-2006-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> In monogamous species sexual conflict is thought to be lessened, and typically little to no sexual dimorphism is noted as there is less ornamentation and armor. This is because there is a relaxation of sexual selection.<sup id="cite_ref-Kokko-2006_17-6" class="reference"><a href="#cite_note-Kokko-2006-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> This may have something to do with a feedback loop caused by a low population density. If sexual selection is too strenuous in a population where there is a low density the population will shrink. In the continuing generations sexual selection will become less and less relevant as mating becomes more random.<sup id="cite_ref-Kokko-2006_17-7" class="reference"><a href="#cite_note-Kokko-2006-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> A similar feedback loop is thought to occur for the sperm quality in genetically monogamous pairs. </p> <div class="mw-heading mw-heading2"><h2 id="Sperm_quality">Sperm quality</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=12" title="Edit section: Sperm quality"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Once anisogamy has emerged in a species due to gamete dimorphism there is an inherent level of competition. This could be seen as <a href="/wiki/Sperm_competition" title="Sperm competition">sperm competition</a> in the very least. Sperm competition is defined as a post copulatory mode of sexual selection which causes the diversity of sperm across species.<sup id="cite_ref-vanderHorst-2014_24-1" class="reference"><a href="#cite_note-vanderHorst-2014-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> As soon as sperm and egg are the predominant mating types there is an increase in the need for the male gametes. This is because there will be a large number of unsuccessful sperm which will cost a certain level of expenditure on energy without a benefit from the individual sperm. Sperm in polygamous sexual encounters have evolved for size, speed, structure, and quantity.<sup id="cite_ref-vanderHorst-2014_24-2" class="reference"><a href="#cite_note-vanderHorst-2014-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> This competition causes selection for competitive traits which can be pre or post copulatory.<sup id="cite_ref-Lehtonen-2016_25-1" class="reference"><a href="#cite_note-Lehtonen-2016-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> In species where <a href="/wiki/Cryptic_female_choice" title="Cryptic female choice">cryptic female choice</a> is one of the main sources of competition females are able to choose sperm from among various male suitors.<sup id="cite_ref-vanderHorst-2014_24-3" class="reference"><a href="#cite_note-vanderHorst-2014-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><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><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> Typically the sperm of the highest quality are selected.<sup id="cite_ref-vanderHorst-2014_24-4" class="reference"><a href="#cite_note-vanderHorst-2014-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>In genetically monogamous species it can be expected that sperm competition is absent or otherwise severely limited. There is no selection for the highest quality sperm amongst the sperm of multiple males, and copulation is more random than it is in polygamous situations. Therefore, sperm quality for monogamous species has a higher variation and lower quality sperm have been noted in several species. The lack of sperm competition is not advantageous for sperm quality. An example of this is in the Eurasian bullfinch which exhibits relaxed selection and sperm competition. The sperm of these males have a lower velocity than other closely related but polygamous passerine bird species and the amount of abnormalities in sperm structure, length, and count when compared to similar bird families is increased.<sup id="cite_ref-vanderHorst-2014_24-5" class="reference"><a href="#cite_note-vanderHorst-2014-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Animals">Animals</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=13" title="Edit section: Animals"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Evolution" title="Evolution">evolution</a> of mating systems in animals has received an enormous amount of attention from biologists. This section briefly reviews three main findings about the evolution of monogamy in animals. </p><p>The amount of social monogamy in animals varies across taxa, with over 90% of birds engaging in social monogamy while only 3–9% of mammals are known to do the same.<sup id="cite_ref-Lack-1968_6-1" class="reference"><a href="#cite_note-Lack-1968-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Reichard-2002_29-0" class="reference"><a href="#cite_note-Reichard-2002-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Barash-2001_30-0" class="reference"><a href="#cite_note-Barash-2001-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p><p>This list is not complete. Other factors may also contribute to the evolution of social monogamy. Moreover, different sets of factors may explain the evolution of social monogamy in different species. There is no one-size-fits-all explanation of why different species evolved monogamous mating systems. </p> <div class="mw-heading mw-heading3"><h3 id="Sexual_dimorphism_2">Sexual dimorphism</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=14" title="Edit section: Sexual dimorphism"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a> refers to differences in body characteristics between females and males. A frequently studied type of sexual dimorphism is body size. For example, among mammals, males typically have larger bodies than females. In other orders, however, females have larger bodies than males. Sexual dimorphism in body size has been linked to mating behavior.<sup id="cite_ref-Owens-1998_31-0" class="reference"><a href="#cite_note-Owens-1998-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Frayer-1985_32-0" class="reference"><a href="#cite_note-Frayer-1985-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Geary-2001_33-0" class="reference"><a href="#cite_note-Geary-2001-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Dunn-2001_34-0" class="reference"><a href="#cite_note-Dunn-2001-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p><p>In <a href="/wiki/Polygyny" title="Polygyny">polygynous</a> species, males compete for control over sexual access to females. Large males have an advantage in the competition for access to females, and they consequently pass their genes along to a greater number of offspring. This eventually leads to large differences in body size between females and males. Polygynous males are often 1.5 to 2.0 times larger in size than females. In monogamous species, on the other hand, females and males have more equal access to mates, so there is little or no sexual dimorphism in body size. From a new biological point of view, monogamy could result from <a href="/wiki/Mate_guarding" title="Mate guarding">mate guarding</a> and is engaged as a result of <a href="/wiki/Sexual_conflict" title="Sexual conflict">sexual conflict</a>.<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><p>Some researchers have attempted to infer the evolution of human mating systems from the evolution of sexual dimorphism. Several studies have reported a large amount of sexual dimorphism in <i><a href="/wiki/Australopithecus" title="Australopithecus">Australopithecus</a></i>, an evolutionary ancestor of human beings that lived between 2 and 5 million years ago.<sup id="cite_ref-Frayer-1985_32-1" class="reference"><a href="#cite_note-Frayer-1985-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Geary-2001_33-1" class="reference"><a href="#cite_note-Geary-2001-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Flinn-2004_36-0" class="reference"><a href="#cite_note-Flinn-2004-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Lockwood-1996_37-0" class="reference"><a href="#cite_note-Lockwood-1996-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p><p>These studies raise the possibility that <i>Australopithecus</i> had a polygamous mating system. Sexual dimorphism then began to decrease. Studies suggest sexual dimorphism reached modern human levels around the time of <i><a href="/wiki/Homo_erectus" title="Homo erectus">Homo erectus</a></i> 0.5 to 2 million years ago.<sup id="cite_ref-Frayer-1985_32-2" class="reference"><a href="#cite_note-Frayer-1985-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Geary-2001_33-2" class="reference"><a href="#cite_note-Geary-2001-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Flinn-2004_36-1" class="reference"><a href="#cite_note-Flinn-2004-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Arsuaga-1997_38-0" class="reference"><a href="#cite_note-Arsuaga-1997-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> This line of reasoning suggests human ancestors started out polygamous and began the transition to monogamy somewhere between 0.5 million and 2 million years ago. </p><p>Attempts to infer the evolution of monogamy based on sexual dimorphism remain controversial for three reasons: </p> <blockquote> <ul><li>The skeletal remains of <i>Australopithecus</i> are quite fragmentary. This makes it difficult to identify the sex of the fossils. Researchers sometimes identify the sex of the fossils by their size, which, of course, can exaggerate findings of sexual dimorphism.</li> <li>Recent studies using new methods of measurement suggest <i>Australopithecus</i> had the same amount of sexual dimorphism as modern humans.<sup id="cite_ref-Reno-2003_39-0" class="reference"><a href="#cite_note-Reno-2003-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Larsen-2003_40-0" class="reference"><a href="#cite_note-Larsen-2003-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> This raises questions about the amount of sexual dimorphism in <i>Australopithecus</i>.</li> <li>Humans may have been partially unique in that selection pressures for sexual dimorphism might have been related to the new niches that humans were entering at the time, and how that might have interacted with potential early cultures and tool use. If these early humans had a differentiation of gender roles, with men hunting and women gathering, selection pressures in favor of increased size may have been distributed unequally between the sexes.</li> <li>Even if future studies clearly establish sexual dimorphism in <i>Australopithecus</i>, other studies have shown the relationship between sexual dimorphism and mating system is unreliable.<sup id="cite_ref-Owens-1998_31-1" class="reference"><a href="#cite_note-Owens-1998-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Frayer-1985_32-3" class="reference"><a href="#cite_note-Frayer-1985-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> Some polygamous species show little or no sexual dimorphism. Some monogamous species show a large amount of sexual dimorphism.</li></ul> </blockquote> <p>Studies of sexual dimorphism raise the possibility that early human ancestors were polygamous rather than monogamous. But this line of research remains highly controversial. It may be that early human ancestors showed little sexual dimorphism, and it may be that sexual dimorphism in early human ancestors had no relationship to their mating systems. </p> <div class="mw-heading mw-heading3"><h3 id="Testis_size">Testis size</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=15" title="Edit section: Testis size"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Schimpanse_Zoo_Leipzig.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/62/Schimpanse_Zoo_Leipzig.jpg/240px-Schimpanse_Zoo_Leipzig.jpg" decoding="async" width="240" height="180" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/62/Schimpanse_Zoo_Leipzig.jpg/360px-Schimpanse_Zoo_Leipzig.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/62/Schimpanse_Zoo_Leipzig.jpg/480px-Schimpanse_Zoo_Leipzig.jpg 2x" data-file-width="2560" data-file-height="1920" /></a><figcaption><a href="/wiki/Chimpanzee" title="Chimpanzee">Chimpanzee</a></figcaption></figure> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Silverback.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f1/Silverback.JPG/240px-Silverback.JPG" decoding="async" width="240" height="180" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f1/Silverback.JPG/360px-Silverback.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f1/Silverback.JPG/480px-Silverback.JPG 2x" data-file-width="1280" data-file-height="960" /></a><figcaption>Male and female <a href="/wiki/Gorilla" title="Gorilla">gorilla</a></figcaption></figure> <p>The relative sizes of male <a href="/wiki/Testes" class="mw-redirect" title="Testes">testes</a> often reflect mating systems.<sup id="cite_ref-Pitcher-2005_41-0" class="reference"><a href="#cite_note-Pitcher-2005-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Simmons-2004_42-0" class="reference"><a href="#cite_note-Simmons-2004-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Dixson-2001_43-0" class="reference"><a href="#cite_note-Dixson-2001-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Harcourt-1981_44-0" class="reference"><a href="#cite_note-Harcourt-1981-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> In species with promiscuous mating systems, where many males mate with many females, the testes tend to be relatively large. This appears to be the result of sperm competition. Males with large testes produce more sperm and thereby gain an advantage impregnating females. In polygynous species, where one male controls sexual access to females, the testes tend to be small. One male defends exclusive sexual access to a group of females and thereby eliminates sperm competition. </p><p>Studies of <a href="/wiki/Primate" title="Primate">primates</a> support the relationship between testis size and mating system.<sup id="cite_ref-Dixson-2001_43-1" class="reference"><a href="#cite_note-Dixson-2001-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Harcourt-1981_44-1" class="reference"><a href="#cite_note-Harcourt-1981-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Birkhead-2000_45-0" class="reference"><a href="#cite_note-Birkhead-2000-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> <sup class="noprint Inline-Template noprint Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:AGE_MATTERS" class="mw-redirect" title="Wikipedia:AGE MATTERS"><span title="These sources are 20-40yo, newer more detailed studies exists. (April 2019)">obsolete source</span></a></i>]</sup> <a href="/wiki/Chimpanzee" title="Chimpanzee">Chimpanzees</a>, which have a promiscuous mating system, have large testes compared to other primates. <a href="/wiki/Gorilla" title="Gorilla">Gorillas</a>, which have a polygynous mating system, have smaller testes than other primates. <a href="/wiki/Human" title="Human">Humans</a>, which have a <a href="/wiki/Incidence_of_monogamy" class="mw-redirect" title="Incidence of monogamy">socially monogamous mating system</a>, have moderately sized testes. <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. (April 2019)">citation needed</span></a></i>]</sup> The moderate amounts of sexual non-monogamy in humans may result in a low to moderate amount of sperm competition. <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. (April 2019)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="Monogamy_as_a_best_response">Monogamy as a best response</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=16" title="Edit section: Monogamy as a best response"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In species where the young are particularly vulnerable and may benefit from protection by both parents, monogamy may be an optimal strategy. Monogamy tends to also occur when populations are small and dispersed. This is not conductive to polygamous behavior as the male would spend far more time searching for another mate. The monogamous behavior allows the male to have a mate consistently, without having to waste energy searching for other females. Furthermore, there is an apparent connection between the time a male invests in their offspring and their monogamous behavior. A male which is required to care for the offspring to ensure their survival is much more likely to exhibit monogamous behavior over one that does not. </p><p>The selection factors in favor of different mating strategies for a species of animal, however, may potentially operate on a large number of factors throughout that animal's life cycle. For instance, with many species of bear, the female will often drive a male off soon after mating, and will later guard her cubs from him. It is thought that this may be due to the fact that too many bears close to one another may deplete the food available to the relatively small but growing cubs. Monogamy may be social but rarely genetic.<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. (May 2012)">citation needed</span></a></i>]</sup> For example, in the cichlid species <i><a href="/wiki/Variabilichromis_moorii" title="Variabilichromis moorii">Variabilichromis moorii</a></i>, a monogamous pair will care for their eggs and young but the eggs are not all fertilized by the same male.<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> <a href="/wiki/Thierry_Lod%C3%A9" title="Thierry Lodé">Thierry Lodé</a><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> argued that monogamy should result from conflict of interest between the sexes called <a href="/wiki/Sexual_conflict" title="Sexual conflict">sexual conflict</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Monogamous_species">Monogamous species</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=17" title="Edit section: Monogamous species"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are species which have adopted monogamy with great success. For instance, the male <a href="/wiki/Prairie_vole" title="Prairie vole">prairie vole</a> will mate exclusively with the first female he ever mates with. The vole is extremely loyal and will go as far as to even attack other females that may approach him. This type of behavior has been linked to the hormone <a href="/wiki/Vasopressin" title="Vasopressin">vasopressin</a>. This hormone is released when a male mates and cares for young. Due to this hormone's rewarding effects, the male experiences a positive feeling when they maintain a monogamous relationship. To further test this theory, the receptors that control vasopressin were placed into another species of vole that is promiscuous. After this addition, the originally unfaithful voles became monogamous with their selected partner. These very same receptors can be found in human brain, and have been found to vary at the individual level—which could explain why some human males tend to be more loyal than others.<sup id="cite_ref-NSF-2013_48-0" class="reference"><a href="#cite_note-NSF-2013-48"><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><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><p><a href="/wiki/Black_vulture" title="Black vulture">Black vultures</a> stay together as it is more beneficial for their young to be taken care of by both parents. They take turns incubating the eggs, and then supplying their fledglings with food. Black vultures will also attack other vultures that are participating in <a href="/wiki/Extra_pair_copulation" class="mw-redirect" title="Extra pair copulation">extra pair copulation</a>, this is an attempt to increase monogamy and decrease promiscuous behavior.<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. (May 2020)">citation needed</span></a></i>]</sup> Similarly, <a href="/wiki/Emperor_penguin" title="Emperor penguin">emperor penguins</a> also stay together to care for their young. This is due to the harshness of the Antarctic weather, predators and the scarcity of food. One parent will protect the chick, while the other finds food. However, these penguins only remain monogamous until the chick is able to go off on their own. After the chick no longer needs their care, approximately 85% of parents will part ways and typically find a new partner every breeding season. </p><p><a href="/wiki/Hornbill" title="Hornbill">Hornbills</a> are a socially monogamous bird species that usually only have one mate throughout their lives, much like the prairie vole. The female will close herself up in a nest cavity, sealed with a nest plug, for two months. At this time, she will lay eggs and will be cared for by her mate. The males are willing to work to support himself, his mate, and his offspring in order for survival; however, unlike the emperor penguin, the hornbills do not find new partners each season.<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> </p><p>It is relatively uncommon to find monogamous relationships in fish, amphibians and reptiles; however, the <a href="/wiki/Red-backed_salamander" title="Red-backed salamander">red-backed salamander</a> as well as the Caribbean cleaner <a href="/wiki/Goby" title="Goby">goby</a> practice monogamy as well. However, the male Caribbean cleaner goby fish has been found to separate from the female suddenly, leaving her abandoned. In a study conducted by Oregon State University, it was found that this fish practices not true monogamy, but serial monogamy. This essentially means that the goby will have multiple monogamous relationships throughout its life – but only be in one relationship at a time.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> The red-backed salamander exhibited signs of social monogamy, which is the idea that animals form pairs to mate and raise offspring, but still will partake in extra pair copulation with various males or females in order to increase their biological fitness. This is a relatively new concept in salamanders, and has not been seen frequently – it is also concerning that the act of monogamy may inhibit the salamanders reproductive rates and biological success. However, the study which was conducted in cooperation by the University of Louisiana, Lafayette, and the University of Virginia showed that the salamanders are not inhibited by this monogamy if they show alternative strategies with other mates.<sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Azara%27s_night_monkey" title="Azara's night monkey">Azara's night monkeys</a> are another species that proved to be monogamous. In an 18-year study conducted by the University of Pennsylvania, these monkeys proved to be entirely monogamous, exhibiting no genetic information or visual information that could lead to the assumption that extra pair copulation was occurring. This explained the question as to why the male owl monkey invested so much time in protecting and raising their own offspring. Because monogamy is often referred to as "placing all your eggs in one basket" the male wants to ensure his young survive, and thus pass on his genes.<sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> </p><p>The desert grass spider, <i><a href="/wiki/Agelenopsis_aperta" title="Agelenopsis aperta">Agelenopsis aperta</a>,</i> is mostly monogamous as well. Male size is the determining factor in fights over a female, with the larger male emerging as the winner since their size signifies success in future offspring.<sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> </p><p>Other monogamous species include <a href="/wiki/Gray_wolf" class="mw-redirect" title="Gray wolf">wolves</a>,<sup id="cite_ref-56" class="reference"><a href="#cite_note-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> certain species of <a href="/wiki/Fox" title="Fox">fox</a>,<sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Otter" title="Otter">otters</a>, a few hooved animals, some <a href="/wiki/Bat" title="Bat">bats</a>, and the <a href="/wiki/Eurasian_beaver" title="Eurasian beaver">Eurasian beaver</a>. This beaver is particularly interesting, as it is practicing monogamy in its reintroduction to certain parts of Europe; however, its <a href="/wiki/North_American_beaver" title="North American beaver">American counterpart</a> is not monogamous at all and often partakes in promiscuous behavior. The two species are quite similar in ecology, but American beavers tend to be less aggressive than European beavers. In this instance, the scarcity of the European beavers' population could drive its monogamous behavior; moreover, it lowers the risk of parasite transmission which is correlated with biological fitness. Monogamy is proving to be very efficient for this beaver, as their population is climbing.<sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">[</span>59<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=Monogamy_in_animals&action=edit&section=18" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Monogamy_topics">Monogamy topics</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=19" title="Edit section: Monogamy topics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Animal_sexual_behaviour#Monogamy" title="Animal sexual behaviour">Animal sexual behaviour#Monogamy</a></li> <li><a href="/wiki/Monogamy" title="Monogamy">Monogamy</a></li> <li><a href="/wiki/Social_monogamy_in_mammalian_species" title="Social monogamy in mammalian species">Social monogamy in mammalian species</a></li> <li><a href="/wiki/Varieties_of_monogamy" class="mw-redirect" title="Varieties of monogamy">Varieties of monogamy</a></li></ul> <div class="mw-heading mw-heading3"><h3 id="Evolution_topics">Evolution topics</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=20" title="Edit section: Evolution topics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Animal_sexuality" class="mw-redirect" title="Animal sexuality">Animal sexuality</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Evolution of sexual reproduction</a></li> <li><a href="/wiki/History_of_human_sexuality" title="History of human sexuality">History of human sexuality</a></li> <li><a href="/wiki/Human_evolution" title="Human evolution">Human evolution</a></li> <li><a href="/wiki/R/K_selection_theory" title="R/K selection theory">r/K selection theory</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Monogamy_in_animals&action=edit&section=21" 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-Ophir-2008-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Ophir-2008_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Ophir-2008_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Ophir-2008_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Ophir-2008_1-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Ophir-2008_1-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Ophir-2008_1-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Ophir-2008_1-6"><sup><i><b>g</b></i></sup></a></span> <span class="reference-text">Ophir, Alexander G., Phelps, Steven M., Sorin, Anna Bess & O. Wolff, J. (2008). <a rel="nofollow" class="external text" href="https://www.academia.edu/download/40573878/Social_but_not_genetic_monogamy_is_assoc20151202-28200-1o7cfoe.pdf">Social but not genetic monogamy is associated with greater breeding success in prairie voles</a><sup class="noprint Inline-Template"><span style="white-space: nowrap;">[<i><a href="/wiki/Wikipedia:Link_rot" title="Wikipedia:Link rot"><span title=" Dead link tagged July 2022">dead link</span></a></i><span style="visibility:hidden; color:transparent; padding-left:2px">‍</span>]</span></sup></span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFSefcMattersdorferSturmbauerKoblmüller2008" class="citation journal cs1">Sefc, Kristina M.; Mattersdorfer, Karin; Sturmbauer, Christian; Koblmüller, Stephan (2008). 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Lewiston, New York: Edwin Mellen Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-7734-6310-3" title="Special:BookSources/978-0-7734-6310-3"><bdi>978-0-7734-6310-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=World+Religions+and+Social+Evolution+of+the+Old+World+Oikumene+Civilizations%3A+A+Cross-cultural+Perspective&rft.place=Lewiston%2C+New+York&rft.edition=First&rft.pub=Edwin+Mellen+Press&rft.date=2004&rft.isbn=978-0-7734-6310-3&rft.aulast=Korotayev&rft.aufirst=Andrey&rft_id=https%3A%2F%2Fwww.academia.edu%2F29329039&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMonogamy+in+animals" class="Z3988"></span></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist 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href="/wiki/Template:Evolution" title="Template:Evolution"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolution" title="Template talk:Evolution"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolution" title="Special:EditPage/Template:Evolution"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_biology" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li> <li><a href="/wiki/Outline_of_evolution" title="Outline of evolution">Outline</a></li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li> <li><a href="/wiki/History_of_life" title="History of life">History of life</a></li> <li><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution" title="Evolution">Evolution</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></li> <li><a href="/wiki/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a href="/wiki/Cheating_(biology)" title="Cheating (biology)">Cheating</a></li> <li><a href="/wiki/Reciprocal_altruism" title="Reciprocal altruism">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a> <ul><li><a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li></ul></li> <li><a href="/wiki/Common_descent" title="Common descent">Common descent</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergence</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergence</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li><a href="/wiki/Evidence_of_common_descent" title="Evidence of common descent">Evidence of evolution</a></li> <li><a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">Evolutionary arms race</a></li> <li><a href="/wiki/Evolutionary_pressure" title="Evolutionary pressure">Evolutionary pressure</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Extinction" title="Extinction">Extinction</a> <ul><li><a href="/wiki/Extinction_event" title="Extinction event">Event</a></li></ul></li> <li><a href="/wiki/Homology_(biology)" title="Homology (biology)">Homology</a></li> <li><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Macroevolution" title="Macroevolution">Macroevolution</a></li> <li><a href="/wiki/Microevolution" title="Microevolution">Microevolution</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Nonadaptive_radiation" title="Nonadaptive radiation">Non-adaptive radiation</a></li> <li><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></li> <li><a href="/wiki/Panspermia" title="Panspermia">Panspermia</a></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a> <ul><li><a href="/wiki/Handicap_principle" title="Handicap principle">Handicap principle</a></li></ul></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a> <ul><li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li></ul></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Taxonomy</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centered view of evolution</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Population_genetics" title="Population genetics">Population<br />genetics</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Selective_breeding" title="Selective breeding">Artificial selection</a></li> <li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness" title="Inclusive fitness">Inclusive</a></li></ul></li> <li><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a></li> <li><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a> <ul><li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a></li> <li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a> <ul><li><a href="/wiki/Sexual_selection_in_flowering_plants" title="Sexual selection in flowering plants">Flowering plants</a></li> <li><a href="/wiki/Sexual_selection_in_fungi" title="Sexual selection in fungi">Fungi</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Variation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Development</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Inversion_(evolutionary_biology)" title="Inversion (evolutionary biology)">Inversion</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Phenotypic plasticity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Taxon" title="Taxon">taxa</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_bacteria" title="Evolution of bacteria">Bacteria</a></li> <li><a href="/wiki/Evolution_of_birds" title="Evolution of birds">Birds</a> <ul><li><a href="/wiki/Origin_of_birds" title="Origin of birds">origin</a></li></ul></li> <li><a href="/wiki/Evolution_of_brachiopods" title="Evolution of brachiopods">Brachiopods</a></li> <li><a href="/wiki/Evolution_of_molluscs" title="Evolution of molluscs">Molluscs</a> <ul><li><a href="/wiki/Evolution_of_cephalopods" title="Evolution of cephalopods">Cephalopods</a></li></ul></li> <li><a href="/wiki/Dinosaur#Evolutionary_history" title="Dinosaur">Dinosaurs</a></li> <li><a href="/wiki/Evolution_of_fish" title="Evolution of fish">Fish</a></li> <li><a href="/wiki/Evolution_of_fungi" title="Evolution of fungi">Fungi</a></li> <li><a href="/wiki/Evolution_of_insects" title="Evolution of insects">Insects</a> <ul><li><a href="/wiki/Evolution_of_butterflies" title="Evolution of butterflies">butterflies</a></li></ul></li> <li><a href="/wiki/History_of_life" title="History of life">Life</a></li> <li><a href="/wiki/Evolution_of_mammals" title="Evolution of mammals">Mammals</a> <ul><li><a href="/wiki/Cat_gap" title="Cat gap">cats</a></li> <li><a href="/wiki/Canidae#Evolution" title="Canidae">canids</a> <ul><li><a href="/wiki/Evolution_of_the_wolf" title="Evolution of the wolf">wolves</a></li> <li><a href="/wiki/Domestication_of_the_dog" title="Domestication of the dog">dogs</a></li></ul></li> <li><a href="/wiki/Hyena#Evolution" title="Hyena">hyenas</a></li> <li><a href="/wiki/Evolution_of_cetaceans" title="Evolution of cetaceans">dolphins and whales</a></li> <li><a href="/wiki/Evolution_of_the_horse" title="Evolution of the horse">horses</a></li> <li><a href="/wiki/Evolution_of_Macropodidae" title="Evolution of Macropodidae">Kangaroos</a></li> <li><a href="/wiki/Evolution_of_primates" title="Evolution of primates">primates</a> <ul><li><a href="/wiki/Human_evolution" title="Human evolution">humans</a></li> <li><a href="/wiki/Evolution_of_lemurs" title="Evolution of lemurs">lemurs</a></li></ul></li> <li><a href="/wiki/Evolution_of_sirenians" title="Evolution of sirenians">sea cows</a></li></ul></li> <li><a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">Plants</a> <ul><li><a href="/wiki/Pollinator-mediated_selection" title="Pollinator-mediated selection">pollinator-mediated</a></li></ul></li> <li><a href="/wiki/Evolution_of_reptiles" title="Evolution of reptiles">Reptiles</a></li> <li><a href="/wiki/Evolution_of_spiders" title="Evolution of spiders">Spiders</a></li> <li><a href="/wiki/Evolution_of_tetrapods" title="Evolution of tetrapods">Tetrapods</a></li> <li><a href="/wiki/Viral_evolution" title="Viral evolution">Viruses</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">organs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_cells" title="Evolution of cells">Cell</a></li> <li><a href="/wiki/Models_of_DNA_evolution" title="Models of DNA evolution">DNA</a></li> <li><a href="/wiki/Evolution_of_flagella" title="Evolution of flagella">Flagella</a></li> <li><a href="/wiki/Eukaryote#Origin_of_eukaryotes" title="Eukaryote">Eukaryotes</a> <ul><li><a href="/wiki/Symbiogenesis" title="Symbiogenesis">symbiogenesis</a></li> <li><a href="/wiki/Chromosome#Eukaryotes" title="Chromosome">chromosome</a></li> <li><a href="/wiki/Endomembrane_system#Evolution" title="Endomembrane system">endomembrane system</a></li> <li><a href="/wiki/Mitochondrion#Origin_and_evolution" title="Mitochondrion">mitochondria</a></li> <li><a href="/wiki/Cell_nucleus#Evolution" title="Cell nucleus">nucleus</a></li> <li><a href="/wiki/Plastid#Origin" title="Plastid">plastids</a></li></ul></li> <li>In animals <ul><li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">eye</a></li> <li><a href="/wiki/Hair#Evolution" title="Hair">hair</a></li> <li><a href="/wiki/Evolution_of_mammalian_auditory_ossicles" title="Evolution of mammalian auditory ossicles">auditory ossicle</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">nervous system</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">brain</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Biological_process" title="Biological process">processes</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_ageing" title="Evolution of ageing">Aging</a> <ul><li><a href="/wiki/Death#Evolution_of_aging_and_mortality" title="Death">Death</a></li> <li><a href="/wiki/Programmed_cell_death#Evolutionary_origin_of_mitochondrial_apoptosis" title="Programmed cell death">Programmed cell death</a></li></ul></li> <li><a href="/wiki/Origin_of_avian_flight" title="Origin of avian flight">Avian flight</a></li> <li><a href="/wiki/Evolution_of_biological_complexity" title="Evolution of biological complexity">Biological complexity</a></li> <li><a href="/wiki/Cooperation_(evolution)" title="Cooperation (evolution)">Cooperation</a></li> <li><a href="/wiki/Evolution_of_color_vision" title="Evolution of color vision">Color vision</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">in primates</a></li></ul></li> <li><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></li> <li><a href="/wiki/Empathy#Evolution_across_species" title="Empathy">Empathy</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Ethics</a></li> <li><a href="/wiki/Evolution_of_eusociality" title="Evolution of eusociality">Eusociality</a></li> <li><a href="/wiki/Immune_system#Evolution_and_other_mechanisms" title="Immune system">Immune system</a></li> <li><a href="/wiki/Metabolism#Evolution" title="Metabolism">Metabolism</a></li> <li><a class="mw-selflink selflink">Monogamy</a></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a></li> <li><a href="/wiki/Mosaic_evolution" title="Mosaic evolution">Mosaic evolution</a></li> <li><a href="/wiki/Multicellular_organism#Evolutionary_history" title="Multicellular organism">Multicellularity</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Sexual reproduction</a> <ul><li><a href="/wiki/Anisogamy#Evolution" title="Anisogamy">Gamete differentiation/sexes</a></li> <li><a href="/wiki/Biological_life_cycle" title="Biological life cycle">Life cycles/nuclear phases</a></li> <li><a href="/wiki/Mating_type" title="Mating type">Mating types</a></li> <li><a href="/wiki/Origin_and_function_of_meiosis" title="Origin and function of meiosis">Meiosis</a></li> <li><a href="/wiki/Sex-determination_system#Evolution" title="Sex-determination system">Sex-determination</a></li></ul></li> <li><a href="/wiki/Evolution_of_snake_venom" title="Evolution of snake venom">Snake venom</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tempo_and_Mode_in_Evolution" title="Tempo and Mode in Evolution">Tempo and modes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Phyletic_gradualism" title="Phyletic gradualism">Gradualism</a>/<a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a>/<a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Point_mutation" title="Point mutation">Micromutation</a>/<a href="/wiki/Mutationism#Later_mutationist_theories" title="Mutationism">Macromutation</a></li> <li><a href="/wiki/Uniformitarianism" title="Uniformitarianism">Uniformitarianism</a>/<a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Speciation" title="Speciation">Speciation</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allopatric_speciation" title="Allopatric speciation">Allopatric</a></li> <li><a href="/wiki/Anagenesis" title="Anagenesis">Anagenesis</a></li> <li><a href="/wiki/Catagenesis_(biology)" title="Catagenesis (biology)">Catagenesis</a></li> <li><a href="/wiki/Cladogenesis" title="Cladogenesis">Cladogenesis</a></li> <li><a href="/wiki/Cospeciation" title="Cospeciation">Cospeciation</a></li> <li><a href="/wiki/Ecological_speciation" title="Ecological speciation">Ecological</a></li> <li><a href="/wiki/Hybrid_speciation" title="Hybrid speciation">Hybrid</a></li> <li><a href="/wiki/Nonecological_speciation" title="Nonecological speciation">Non-ecological</a></li> <li><a href="/wiki/Parapatric_speciation" title="Parapatric speciation">Parapatric</a></li> <li><a href="/wiki/Peripatric_speciation" title="Peripatric speciation">Peripatric</a></li> <li><a href="/wiki/Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a></li> <li><a href="/wiki/Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance and Enlightenment</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Transmutation of species</a></li> <li><a href="/wiki/David_Hume" title="David Hume">David Hume</a> <ul><li><i><a href="/wiki/Dialogues_Concerning_Natural_Religion" title="Dialogues Concerning Natural Religion">Dialogues Concerning Natural Religion</a></i></li></ul></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> <ul><li><i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a></i></li></ul></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">History of paleontology</a></li> <li><a href="/wiki/Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li> <li><a href="/wiki/Blending_inheritance" title="Blending inheritance">Blending inheritance</a></li> <li><a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">The eclipse of Darwinism</a></li> <li><a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">History of molecular evolution</a></li> <li><a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">Extended evolutionary synthesis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Philosophy</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Darwinism" title="Darwinism">Darwinism</a></li> <li><a href="/wiki/Alternatives_to_Darwinian_evolution" title="Alternatives to Darwinian evolution">Alternatives</a> <ul><li><a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Orthogenesis" title="Orthogenesis">Orthogenesis</a></li> <li><a href="/wiki/Mutationism" title="Mutationism">Mutationism</a></li> <li><a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Structuralism_(biology)" title="Structuralism (biology)">Structuralism</a> <ul><li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">Spandrel</a></li></ul></li> <li><a href="/wiki/Theistic_evolution" title="Theistic evolution">Theistic</a></li> <li><a href="/wiki/Vitalism" title="Vitalism">Vitalism</a></li></ul></li> <li><a href="/wiki/Teleology_in_biology" title="Teleology in biology">Teleology in biology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a> <ul><li><a href="/wiki/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li></ul></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory of evolution</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Astrobiology" title="Astrobiology">Astrobiology</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a> <ul><li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Tree</a></li></ul></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Evolutionary_biology" title="Category:Evolutionary biology">Category</a></li> <li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Symbol_portal_class.svg" class="mw-file-description" title="Portal"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/16px-Symbol_portal_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/23px-Symbol_portal_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/31px-Symbol_portal_class.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span> <a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Portal</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Animal_sexual_behaviour" 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_sexual_behavior" title="Template:Animal sexual behavior"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Animal_sexual_behavior" title="Template talk:Animal sexual behavior"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Animal_sexual_behavior" title="Special:EditPage/Template:Animal sexual behavior"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Animal_sexual_behaviour" style="font-size:114%;margin:0 4em"><a href="/wiki/Animal_sexual_behaviour" title="Animal sexual behaviour">Animal sexual behaviour</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Animal_sexual_behaviour" title="Animal sexual behaviour">General</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a></li> <li><a href="/wiki/Sexual_reproduction" title="Sexual reproduction">Sexual reproduction</a> <ul><li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">evolution</a></li> <li><a href="/wiki/Reproductive_system" title="Reproductive system">reproductive system</a></li> <li><a href="/wiki/Effects_of_hormones_on_sexual_motivation" title="Effects of hormones on sexual motivation">hormonal motivation</a></li></ul></li> <li><a href="/wiki/Courtship_display" title="Courtship display">Courtship display</a> <ul><li><a href="/wiki/Biological_ornament" title="Biological ornament">sexual ornamentation</a></li> <li><a href="/wiki/Handicap_principle" title="Handicap principle">handicap principle</a></li> <li><a href="/wiki/Sexy_son_hypothesis" title="Sexy son hypothesis">sexy son hypothesis</a></li> <li><a href="/wiki/Fisherian_runaway" title="Fisherian runaway">Fisherian runaway</a></li></ul></li> <li><a href="/wiki/Mating_system" title="Mating system">Mating systems</a> <ul><li><a href="/wiki/Mate_choice" title="Mate choice">mate choice</a></li> <li><a href="/wiki/Mating_call" title="Mating call">mating call</a></li> <li><a href="/wiki/Mate_guarding" title="Mate guarding">mate guarding</a></li> <li><a href="/wiki/Mating_plug" title="Mating plug">mating plug</a></li> <li><a href="/wiki/Lek_mating" title="Lek mating">lek mating</a></li></ul></li> <li><a href="/wiki/Copulation_(zoology)" title="Copulation (zoology)">Copulation</a> <ul><li><a href="/wiki/Cloacal_kiss" class="mw-redirect" title="Cloacal kiss">cloacal kiss</a></li> <li><a href="/wiki/Sexual_intercourse" title="Sexual intercourse">sexual intercourse</a></li> <li><a href="/wiki/Pelvic_thrust" title="Pelvic thrust">Pelvic thrust</a></li> <li><a href="/wiki/Pseudocopulation" title="Pseudocopulation">pseudocopulation</a></li></ul></li> <li><a href="/wiki/Fertilisation#Fertilisation_in_animals" title="Fertilisation">Fertilisation</a> <ul><li><a href="/wiki/Internal_fertilization" title="Internal fertilization">internal</a></li> <li><a href="/wiki/External_fertilization" title="External fertilization">external</a></li> <li><a href="/wiki/Sperm_competition" title="Sperm competition">sperm competition</a></li> <li><a href="/wiki/Traumatic_insemination" title="Traumatic insemination">traumatic insemination</a></li> <li><a href="/wiki/Penile_spines" title="Penile spines">penile spines</a></li></ul></li> <li><a href="/wiki/Modes_of_reproduction" title="Modes of reproduction">Modes</a> <ul><li><a class="mw-selflink selflink">monogamy</a></li> <li><a href="/wiki/Promiscuity" title="Promiscuity">promiscuity</a></li> <li><a href="/wiki/Polyandry_in_animals" title="Polyandry in animals">polyandry</a></li> <li><a href="/wiki/Polygyny_in_animals" title="Polygyny in animals">polygyny</a></li> <li><a href="/wiki/Polygynandry" title="Polygynandry">polygynandry</a></li> <li><a href="/wiki/Semelparity_and_iteroparity" title="Semelparity and iteroparity">semelparity and iteroparity</a></li> <li><a href="/wiki/Opportunistic_breeder" title="Opportunistic breeder">opportunistic</a></li> <li><a href="/wiki/Hermaphrodite" title="Hermaphrodite">hermaphroditism</a></li> <li><a href="/wiki/Cuckold" title="Cuckold">cuckoldry</a></li> <li><a href="/wiki/Seasonal_breeder" title="Seasonal breeder">seasonal</a></li> <li><a href="/wiki/Reproductive_synchrony" title="Reproductive synchrony">synchrony</a></li></ul></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a> <ul><li><a href="/wiki/Anisogamy" title="Anisogamy">anisogamy</a></li> <li><a href="/wiki/Oogamy" title="Oogamy">oogamy</a></li> <li><a href="/wiki/Bateman%27s_principle" title="Bateman's principle">Bateman's principle</a></li> <li><a href="/wiki/Sexual_bimaturism" title="Sexual bimaturism">bimaturism</a></li> <li><a href="/wiki/Sexual_cannibalism" title="Sexual cannibalism">cannibalism</a></li> <li><a href="/wiki/Sexual_coercion_among_animals" title="Sexual coercion among animals">coercion</a></li></ul></li> <li><a href="/wiki/Sexual_conflict" title="Sexual conflict">Sexual conflict</a> <ul><li><a href="/wiki/Interlocus_sexual_conflict" title="Interlocus sexual conflict">interlocus</a></li> <li><a href="/wiki/Intralocus_sexual_conflict" title="Intralocus sexual conflict">intralocus</a></li></ul></li> <li><a href="/wiki/Hybrid_(biology)" title="Hybrid (biology)">Interspecies breeding</a></li> <li><a href="/wiki/Non-reproductive_sexual_behavior_in_animals" title="Non-reproductive sexual behavior in animals">Non-reproductive behavior</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Animal_sexual_behaviour#Invertebrates" title="Animal sexual behaviour">Invertebrates</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Arthropod#Reproduction_and_development" title="Arthropod">Arthropods</a> <ul><li><a href="/wiki/Crab#Reproduction_and_lifecycle" title="Crab">crab</a></li> <li><a href="/wiki/Spider#Reproduction_and_life_cycle" title="Spider">spider</a></li> <li><a href="/wiki/Scorpion#Reproduction" title="Scorpion">scorpion</a></li> <li><a href="/wiki/Beetle#Mating" title="Beetle">beetle</a></li> <li><a href="/wiki/Insect#Reproduction_and_development" title="Insect">insect</a> <ul><li><a href="/wiki/Butterfly#Life_cycle" title="Butterfly">butterfly</a></li></ul></li></ul></li> <li><a href="/wiki/Cephalopod#Reproduction_and_life_cycle" title="Cephalopod">Cephalopods</a> <ul><li><a href="/wiki/Octopus#Reproduction" title="Octopus">octopus</a></li></ul></li> <li><a href="/wiki/Cnidaria#Sexual" title="Cnidaria">Cnidaria</a> <ul><li><a href="/wiki/Sea_anemone#Lifecycle" title="Sea anemone">sea anemone</a></li> <li><a href="/wiki/Jellyfish#Reproduction" title="Jellyfish">jellyfish</a></li> <li><a href="/wiki/Coral#Sexual" title="Coral">coral</a></li></ul></li> <li><a href="/wiki/Echinoderm#Reproduction" title="Echinoderm">Echinoderms</a></li> <li><a href="/wiki/Mating_of_gastropods" title="Mating of gastropods">Gastropods</a> <ul><li><a href="/wiki/Apophallation" title="Apophallation">apophallation</a></li> <li><a href="/wiki/Love_dart" title="Love dart">love dart</a></li></ul></li> <li><a href="/wiki/Sponge#Reproduction" title="Sponge">Sponge</a></li> <li>Worms <ul><li><a href="/wiki/Earthworm#Reproduction" title="Earthworm">earthworm</a></li> <li><a href="/wiki/Epitoky" title="Epitoky">epitoky</a></li> <li><a href="/wiki/Penis_fencing" title="Penis fencing">penis fencing</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Fish_reproduction" title="Fish reproduction">Fish</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Spawn_(biology)#Spawning_strategies" class="mw-redirect" title="Spawn (biology)">Spawning strategies</a></li> <li><a href="/wiki/Polyandry_in_fish" title="Polyandry in fish">Polyandry in fish</a></li> <li><a href="/wiki/Eel_life_history" title="Eel life history">Eels</a></li> <li><a href="/wiki/Salmon_run" title="Salmon run">Salmon run</a></li> <li><a href="/wiki/Seahorse#Reproduction" title="Seahorse">Seahorse</a></li> <li><a href="/wiki/Shark#Life_history" title="Shark">Sharks</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Amphibian#Reproduction" title="Amphibian">Amphibians</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Sexual_selection_in_amphibians" title="Sexual selection in amphibians">Sexual selection</a> <ul><li><a href="/wiki/Sexual_selection_in_amphibians" title="Sexual selection in amphibians">frogs</a></li></ul></li> <li><a href="/wiki/Frog#Reproduction" title="Frog">Frog reproduction</a></li> <li><a href="/wiki/Salamander#Reproduction_and_development" title="Salamander">Salamanders</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Reptile#Reproduction" title="Reptile">Reptiles</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Sexual_selection_in_scaled_reptiles" title="Sexual selection in scaled reptiles">Sexual selection in scaled reptiles</a> <ul><li><a href="/wiki/Sexual_selection_in_scaled_reptiles" title="Sexual selection in scaled reptiles">lizards</a></li> <li><a href="/wiki/Sexual_selection_in_scaled_reptiles" title="Sexual selection in scaled reptiles">snakes</a></li> <li><a href="/wiki/Common_side-blotched_lizard" title="Common side-blotched lizard">side-blotched lizard</a></li></ul></li> <li><a href="/wiki/Crocodilia#Reproduction_and_parenting" title="Crocodilia">Crocodilians</a></li> <li><a href="/wiki/Tuatara#Reproduction" title="Tuatara">Tuatara</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Bird#Breeding" title="Bird">Birds</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Sexual_selection_in_birds" title="Sexual selection in birds">Sexual selection</a></li> <li><a href="/wiki/Bird#Breeding" title="Bird">Breeding behaviour</a> <ul><li><a href="/wiki/Reproduction_and_life_cycle_of_the_golden_eagle" title="Reproduction and life cycle of the golden eagle">golden eagle</a></li> <li><a href="/wiki/Seabird_breeding_behavior" title="Seabird breeding behavior">seabirds</a></li></ul></li> <li><a href="/wiki/List_of_birds_displaying_homosexual_behavior" title="List of birds displaying homosexual behavior">Homosexual behavior</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Mammalian_reproduction" title="Mammalian reproduction">Mammals</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Sexual_selection_in_mammals" title="Sexual selection in mammals">Sexual selection</a> <ul><li><a href="/wiki/Rut_(mammalian_reproduction)" title="Rut (mammalian reproduction)">rut</a></li></ul></li> <li><a href="/wiki/Lordosis_behavior" title="Lordosis behavior">Lordosis behavior</a></li> <li><a href="/wiki/List_of_mammals_displaying_homosexual_behavior" title="List of mammals displaying homosexual behavior">Homosexual behavior</a></li> <li><a href="/wiki/Canidae#Life_history" title="Canidae">Canid</a> <ul><li><a href="/wiki/African_wild_dog#Social_and_reproductive_behaviour" title="African wild dog">African wild dog</a></li> <li><a href="/wiki/Coyote#Social_and_reproductive_behaviors" title="Coyote">coyote</a></li> <li><a href="/wiki/Dingo#Reproduction" title="Dingo">dingo</a></li> <li><a href="/wiki/Canine_reproduction#Copulation" title="Canine reproduction">domestic dog</a></li> <li><a href="/wiki/Wolf#Reproduction_and_development" title="Wolf">gray wolf</a></li> <li><a href="/wiki/Red_fox#Reproduction_and_development" title="Red fox">red fox</a></li></ul></li> <li><a href="/wiki/Dolphin#Reproduction_and_sexuality" title="Dolphin">Dolphin</a></li> <li><a href="/wiki/Elephant#Sexual_behaviour" title="Elephant">Elephant</a></li> <li><a href="/wiki/European_badger#Reproduction_and_development" title="European badger">European badger</a></li> <li><a href="/wiki/Felidae" title="Felidae">Felidae</a> <ul><li><a href="/wiki/Lion#Reproduction_and_life_cycle" title="Lion">lion</a></li> <li><a href="/wiki/Tiger#Reproduction" title="Tiger">tiger</a></li> <li><a href="/wiki/Cheetah#Reproduction" title="Cheetah">cheetah</a></li> <li><a href="/wiki/Cat#Reproduction" title="Cat">domestic cat</a></li></ul></li> <li><a href="/wiki/Fossa_(animal)#Breeding" title="Fossa (animal)">Fossa</a></li> <li><a href="/wiki/Hippopotamus#Reproduction" title="Hippopotamus">Hippopotamus</a></li> <li><a href="/wiki/Spotted_hyena#Reproduction_and_development" title="Spotted hyena">Spotted hyena</a></li> <li><a href="/wiki/Marsupial#Reproductive_system" title="Marsupial">Marsupial</a> <ul><li><a href="/wiki/Kangaroo#Sexual_behavior" title="Kangaroo">kangaroo</a></li></ul></li> <li><a href="/wiki/Pinniped#Reproductive_behavior" title="Pinniped">Pinnipeds</a> <ul><li><a href="/wiki/Walrus#Reproduction" title="Walrus">walrus</a></li></ul></li> <li>Primates <ul><li><a href="/wiki/Human_sexual_activity" title="Human sexual activity">human</a></li> <li><a href="/wiki/Bonobo#Sociosexual_behaviour" title="Bonobo">bonobo</a></li> <li><a href="/wiki/Gorilla#Reproduction_and_parenting" title="Gorilla">gorilla</a></li> <li><a href="/wiki/Olive_baboon#Reproduction_and_parenting" title="Olive baboon">olive baboon</a></li> <li><a href="/wiki/Mandrill#Mating" title="Mandrill">mandrill</a></li> <li><a href="/wiki/Ring-tailed_lemur#Breeding_and_reproduction" title="Ring-tailed lemur">ringtailed lemur</a></li> <li><a href="/wiki/Sexual_swelling" title="Sexual swelling">sexual swelling</a></li></ul></li> <li><a href="/wiki/Raccoon#Reproduction" title="Raccoon">Raccoon</a></li> <li><a href="/wiki/Rodent#Mating_strategies" title="Rodent">Rodent</a></li> <li><a href="/wiki/Short-beaked_echidna#Reproduction" title="Short-beaked echidna">Short-beaked echidna</a></li></ul> </div></td></tr></tbody></table></div> <!-- NewPP limit report Parsed by mw‐web.codfw.main‐f69cdc8f6‐rhwrx Cached time: 20241122142443 Cache expiry: 2592000 Reduced expiry: false Complications: [vary‐revision‐sha1, show‐toc] CPU time usage: 0.968 seconds Real time usage: 1.137 seconds Preprocessor visited node count: 5416/1000000 Post‐expand include size: 201284/2097152 bytes Template argument size: 5783/2097152 bytes Highest expansion depth: 17/100 Expensive parser function count: 8/500 Unstrip recursion depth: 1/20 Unstrip post‐expand size: 237796/5000000 bytes Lua time usage: 0.564/10.000 seconds Lua memory usage: 6043953/52428800 bytes Number of Wikibase entities loaded: 0/400 --> <!-- Transclusion expansion time report (%,ms,calls,template) 100.00% 953.173 1 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