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id="toc-Research_history" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Research_history"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Research history</span> </div> </a> <ul id="toc-Research_history-sublist" class="vector-toc-list"> <li id="toc-Discovery" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Discovery"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1.1</span> <span>Discovery</span> </div> </a> <ul id="toc-Discovery-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-"P._crassidens"" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#"P._crassidens""> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1.2</span> <span>"<i>P. crassidens</i>"</span> </div> </a> <ul id="toc-"P._crassidens"-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Gigantopithecus" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Gigantopithecus"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1.3</span> <span><i>Gigantopithecus</i></span> </div> </a> <ul id="toc-Gigantopithecus-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-P._boisei" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#P._boisei"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1.4</span> <span><i>P. boisei</i></span> </div> </a> <ul id="toc-P._boisei-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Classification" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Classification"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Classification</span> </div> </a> <ul id="toc-Classification-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Anatomy" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Anatomy"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Anatomy</span> </div> </a> <button aria-controls="toc-Anatomy-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 Anatomy subsection</span> </button> <ul id="toc-Anatomy-sublist" class="vector-toc-list"> <li id="toc-Head" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Head"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Head</span> </div> </a> <ul id="toc-Head-sublist" class="vector-toc-list"> <li id="toc-Skull" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Skull"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1.1</span> <span>Skull</span> </div> </a> <ul id="toc-Skull-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Brain" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Brain"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1.2</span> <span>Brain</span> </div> </a> <ul id="toc-Brain-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Blood_vessels" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Blood_vessels"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1.3</span> <span>Blood vessels</span> </div> </a> <ul id="toc-Blood_vessels-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Torso" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Torso"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Torso</span> </div> </a> <ul id="toc-Torso-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Limbs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Limbs"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Limbs</span> </div> </a> <ul id="toc-Limbs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Size" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Size"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>Size</span> </div> </a> <ul id="toc-Size-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Palaeobiology" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Palaeobiology"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Palaeobiology</span> </div> </a> <button aria-controls="toc-Palaeobiology-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 Palaeobiology subsection</span> </button> <ul id="toc-Palaeobiology-sublist" class="vector-toc-list"> <li id="toc-Diet" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Diet"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Diet</span> </div> </a> <ul id="toc-Diet-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Social_structure" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Social_structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Social structure</span> </div> </a> <ul id="toc-Social_structure-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Technology" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Technology"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Technology</span> </div> </a> <ul id="toc-Technology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Development" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Development"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.4</span> <span>Development</span> </div> </a> <ul id="toc-Development-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Life_history" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Life_history"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.5</span> <span>Life history</span> </div> </a> <ul id="toc-Life_history-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Pathology" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Pathology"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.6</span> <span>Pathology</span> </div> </a> <ul id="toc-Pathology-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Palaeoecology" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Palaeoecology"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Palaeoecology</span> </div> </a> <button aria-controls="toc-Palaeoecology-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 Palaeoecology subsection</span> </button> <ul id="toc-Palaeoecology-sublist" class="vector-toc-list"> <li id="toc-Fossil-bearing_deposits" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Fossil-bearing_deposits"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Fossil-bearing deposits</span> </div> </a> <ul id="toc-Fossil-bearing_deposits-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Predation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Predation"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Predation</span> </div> </a> <ul id="toc-Predation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Extinction" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Extinction"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Extinction</span> </div> </a> <ul id="toc-Extinction-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><i>Paranthropus robustus</i></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 40 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-40" 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">40 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Afrikaans" lang="af" hreflang="af" data-title="Paranthropus robustus" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target"><span>Afrikaans</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%AA%E0%A7%8D%E0%A6%AF%E0%A6%BE%E0%A6%B0%E0%A6%BE%E0%A6%A8%E0%A6%A5%E0%A7%8D%E0%A6%B0%E0%A7%8B%E0%A6%AA%E0%A6%BE%E0%A6%B8_%E0%A6%B0%E0%A7%8B%E0%A6%AC%E0%A6%BE%E0%A6%B8%E0%A7%8D%E0%A6%9F%E0%A6%BE%E0%A6%B8" title="প্যারানথ্রোপাস রোবাস্টাস – Bangla" lang="bn" hreflang="bn" data-title="প্যারানথ্রোপাস রোবাস্টাস" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Belarusian" lang="be" hreflang="be" data-title="Paranthropus robustus" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target"><span>Беларуская</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Catalan" lang="ca" hreflang="ca" data-title="Paranthropus robustus" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-ceb mw-list-item"><a href="https://ceb.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Cebuano" lang="ceb" hreflang="ceb" data-title="Paranthropus robustus" data-language-autonym="Cebuano" data-language-local-name="Cebuano" class="interlanguage-link-target"><span>Cebuano</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Czech" lang="cs" hreflang="cs" data-title="Paranthropus robustus" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – German" lang="de" hreflang="de" data-title="Paranthropus robustus" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Estonian" lang="et" hreflang="et" data-title="Paranthropus robustus" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Spanish" lang="es" hreflang="es" data-title="Paranthropus robustus" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Basque" lang="eu" hreflang="eu" data-title="Paranthropus robustus" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%BE%D8%B1%D8%A7%D9%85%D8%B1%D8%AF%D9%85_%D8%AA%D9%86%D8%A7%D9%88%D8%B1" title="پرامردم تناور – Persian" lang="fa" hreflang="fa" data-title="پرامردم تناور" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – French" lang="fr" hreflang="fr" data-title="Paranthropus robustus" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Galician" lang="gl" hreflang="gl" data-title="Paranthropus robustus" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%ED%8C%8C%EB%9E%80%ED%8A%B8%EB%A1%9C%ED%91%B8%EC%8A%A4_%EB%A1%9C%EB%B6%80%EC%8A%A4%ED%88%AC%EC%8A%A4" title="파란트로푸스 로부스투스 – Korean" lang="ko" hreflang="ko" data-title="파란트로푸스 로부스투스" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Croatian" lang="hr" hreflang="hr" data-title="Paranthropus robustus" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Indonesian" lang="id" hreflang="id" data-title="Paranthropus robustus" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Italian" lang="it" hreflang="it" data-title="Paranthropus robustus" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%A4%D7%A8%D7%A0%D7%AA%D7%A8%D7%95%D7%A4%D7%95%D7%A1_%D7%A8%D7%95%D7%91%D7%95%D7%A1%D7%98%D7%95%D7%A1" title="פרנתרופוס רובוסטוס – Hebrew" lang="he" hreflang="he" data-title="פרנתרופוס רובוסטוס" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Georgian" lang="ka" hreflang="ka" data-title="Paranthropus robustus" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-lb mw-list-item"><a href="https://lb.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Luxembourgish" lang="lb" hreflang="lb" data-title="Paranthropus robustus" data-language-autonym="Lëtzebuergesch" data-language-local-name="Luxembourgish" class="interlanguage-link-target"><span>Lëtzebuergesch</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Hungarian" lang="hu" hreflang="hu" data-title="Paranthropus robustus" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Dutch" lang="nl" hreflang="nl" data-title="Paranthropus robustus" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%83%91%E3%83%A9%E3%83%B3%E3%83%88%E3%83%AD%E3%83%97%E3%82%B9%E3%83%BB%E3%83%AD%E3%83%96%E3%82%B9%E3%83%88%E3%82%B9" title="パラントロプス・ロブストス – Japanese" lang="ja" hreflang="ja" data-title="パラントロプス・ロブストス" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Australopithecus_robustus" title="Australopithecus robustus – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Australopithecus robustus" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Australopithecus_robustus" title="Australopithecus robustus – Occitan" lang="oc" hreflang="oc" data-title="Australopithecus robustus" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Polish" lang="pl" hreflang="pl" data-title="Paranthropus robustus" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Portuguese" lang="pt" hreflang="pt" data-title="Paranthropus robustus" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Romanian" lang="ro" hreflang="ro" data-title="Paranthropus robustus" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Russian" lang="ru" hreflang="ru" data-title="Paranthropus robustus" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Simple English" lang="en-simple" hreflang="en-simple" data-title="Paranthropus robustus" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a 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class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Swedish" lang="sv" hreflang="sv" data-title="Paranthropus robustus" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Tagalog" lang="tl" hreflang="tl" data-title="Paranthropus robustus" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target"><span>Tagalog</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Paranthropus_robustus" title="Paranthropus robustus – Turkish" lang="tr" hreflang="tr" data-title="Paranthropus robustus" 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font-size: 100%"> <tbody><tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(235,235,210)"><i>Paranthropus robustus</i><br /><div style="font-size: 85%;">Temporal range: <a href="/wiki/Pleistocene" title="Pleistocene">Pleistocene</a>, <span class="noprint"><span style="display:inline-block;"></span><span style="display:inline-block;">2.27/2–1/0.87 <a href="/wiki/Megaannum" class="mw-redirect" title="Megaannum">Ma</a></span> <span style="display:inline-block;"></span><div id="Timeline-row" style="margin: 4px auto 0; clear:both; width:220px; padding:0px; height:18px; overflow:visible; white-space:nowrap; border:1px #666; border-style:solid none; position:relative; z-index:0; font-size:97%;"> <div style="position:absolute; height:100%; left:0px; width:207.23076923077px; padding-left:5px; text-align:left; background-color:rgb(254,217,106); background-image: linear-gradient(to right, rgba(255,255,255,1), rgba(254,217,106,1) 15%, rgba(254,217,106,1));"><a href="/wiki/Precambrian" title="Precambrian">PreꞒ</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(127,160,86); left:37.636923076923px; width:18.073846153846px;"><a href="/wiki/Cambrian" title="Cambrian">Ꞓ</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(0,146,112); left:55.710769230769px; width:14.08px;"><a href="/wiki/Ordovician" title="Ordovician">O</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(179,225,182); left:69.790769230769px; width:8.3261538461539px;"><a href="/wiki/Silurian" title="Silurian">S</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(203,140,55); left:78.116923076923px; width:20.409230769231px;"><a href="/wiki/Devonian" title="Devonian">D</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(103,165,153); left:98.526153846154px; width:20.307692307692px;"><a href="/wiki/Carboniferous" title="Carboniferous">C</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(240,64,40); left:118.83384615385px; width:15.907015384615px;"><a href="/wiki/Permian" title="Permian">P</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(129,43,146); left:134.74086153846px; width:17.092984615385px;"><a href="/wiki/Triassic" title="Triassic">T</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(52,178,201); left:151.83384615385px; width:19.089230769231px;"><a href="/wiki/Jurassic" title="Jurassic">J</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(127,198,78); left:170.92307692308px; width:26.738461538462px;"><a href="/wiki/Cretaceous" title="Cretaceous">K</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(253,154,82); left:197.66153846154px; width:14.543692307692px;"><a href="/wiki/Paleogene" title="Paleogene">Pg</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(255,230,25); left:212.20523076923px; width:6.9215384615385px;"><a href="/wiki/Neogene" title="Neogene">N</a></div> <div id="end-border" style="position:absolute; height:100%; background-color:#666; width:1px; left:219px"></div><div style="margin:0 auto; line-height:0; clear:both; width:220px; padding:0px; height:8px; overflow:visible; background-color:transparent; position:relative; top:-4px; z-index:100;"> <div style="position:absolute; left:219.61584615385px; font-size:50%"><div style="position:relative; left:-0.42em">↓</div></div> </div> </div></span></div> </th></tr> <tr> <td colspan="2" style="text-align: center"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Original_of_Paranthropus_robustus_Face.jpg" class="mw-file-description"><img alt="a skull in face-view, lacking a forehead, and having large and rectangular eye sockets, an even larger triangular opening for the nose, flaring cheeks strongly angled inward, and missing the front teeth" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a1/Original_of_Paranthropus_robustus_Face.jpg/220px-Original_of_Paranthropus_robustus_Face.jpg" decoding="async" width="220" height="178" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a1/Original_of_Paranthropus_robustus_Face.jpg/330px-Original_of_Paranthropus_robustus_Face.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a1/Original_of_Paranthropus_robustus_Face.jpg/440px-Original_of_Paranthropus_robustus_Face.jpg 2x" data-file-width="3195" data-file-height="2592" /></a></span> </td></tr> <tr> <td colspan="2" style="text-align: center; font-size: 88%">Cast of the presumed-male <a href="/wiki/SK_48" title="SK 48">SK 48</a> skull </td></tr> <tr> <th colspan="2" style="color:inherit; min-width:15em; text-align: center; background-color: rgb(235,235,210)"><a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Scientific classification</a> <span class="plainlinks taxobox-edit-taxonomy skin-invert" style="font-size:smaller; float:right; padding-right:0.4em; margin-left:-3em;"><span typeof="mw:File"><a href="/wiki/Template:Taxonomy/Paranthropus" title="Edit this classification"><img alt="Edit this classification" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/15px-OOjs_UI_icon_edit-ltr.svg.png" decoding="async" width="15" height="15" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/23px-OOjs_UI_icon_edit-ltr.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/30px-OOjs_UI_icon_edit-ltr.svg.png 2x" data-file-width="20" data-file-height="20" /></a></span></span> </th></tr> <tr> <td>Domain: </td> <td><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a> </td></tr> <tr> <td>Kingdom: </td> <td><a href="/wiki/Animal" title="Animal">Animalia</a> </td></tr> <tr> <td>Phylum: </td> <td><a href="/wiki/Chordate" title="Chordate">Chordata</a> </td></tr> <tr> <td>Class: </td> <td><a href="/wiki/Mammal" title="Mammal">Mammalia</a> </td></tr> <tr> <td>Order: </td> <td><a href="/wiki/Primate" title="Primate">Primates</a> </td></tr> <tr> <td>Suborder: </td> <td><a href="/wiki/Haplorhini" title="Haplorhini">Haplorhini</a> </td></tr> <tr> <td>Infraorder: </td> <td><a href="/wiki/Simian" title="Simian">Simiiformes</a> </td></tr> <tr> <td>Family: </td> <td><a href="/wiki/Hominidae" title="Hominidae">Hominidae</a> </td></tr> <tr> <td>Subfamily: </td> <td><a href="/wiki/Homininae" title="Homininae">Homininae</a> </td></tr> <tr> <td>Tribe: </td> <td><a href="/wiki/Hominini" title="Hominini">Hominini</a> </td></tr> <tr> <td>Genus: </td> <td><span style="font-style:normal;font-weight:normal;">†</span><a href="/wiki/Paranthropus" title="Paranthropus"><i>Paranthropus</i></a> </td></tr> <tr> <td>Species: </td> <td><div style="display:inline" class="species">†<i><b>P. robustus</b></i></div> </td></tr> <tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(235,235,210)"><a href="/wiki/Binomial_nomenclature" title="Binomial nomenclature">Binomial name</a> </th></tr> <tr> <td colspan="2" style="text-align: center"><b><span class="binomial"><span style="font-weight:normal;">†</span><i>Paranthropus robustus</i></span></b><br /><div style="font-size: 85%;"><a href="/wiki/Robert_Broom" title="Robert Broom">Broom</a>, 1938</div> </td></tr> <tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(235,235,210)"><a href="/wiki/Synonym_(taxonomy)" title="Synonym (taxonomy)">Synonyms</a> </th></tr> <tr> <td colspan="2" style="text-align: left"> <ul><li><i>P. crassidens</i><br /><small><a href="/wiki/Robert_Broom" title="Robert Broom">Broom</a> and <a href="/wiki/John_Talbot_Robinson" title="John Talbot Robinson">Robinson</a>, 1948</small></li> <li><i>Australopithecus robustus</i><br /><small><a href="/wiki/Sherwood_Washburn" title="Sherwood Washburn">Washburn</a> and Patterson, 1951</small></li></ul> </td></tr> </tbody></table><style data-mw-deduplicate="TemplateStyles:r1238732961">@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox.biota tr{background:transparent!important}html.skin-theme-clientpref-night .mw-parser-output .infobox.biota img{background:transparent}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox.biota tr{background:transparent!important}html.skin-theme-clientpref-os .mw-parser-output .infobox.biota img{background:white}}.mw-parser-output .infobox.biota .taxobox-edit-taxonomy img{background:transparent!important}body.skin-vector .mw-parser-output table.biota.infobox{margin-top:0.5em}body.skin--responsive .mw-parser-output table.biota.infobox tr.taxonrow td{padding:2px 10px}</style> <p><i><b>Paranthropus robustus</b></i> is a <a href="/wiki/Species" title="Species">species</a> of robust <a href="/wiki/Australopithecine" title="Australopithecine">australopithecine</a> from the <a href="/wiki/Early_Pleistocene" title="Early Pleistocene">Early</a> and possibly <a href="/wiki/Middle_Pleistocene" class="mw-redirect" title="Middle Pleistocene">Middle</a> Pleistocene of the <a href="/wiki/Cradle_of_Humankind" title="Cradle of Humankind">Cradle of Humankind</a>, <a href="/wiki/South_Africa" title="South Africa">South Africa</a>, about 2.27 to 0.87 (or, more conservatively, 2 to 1) million years ago.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> It has been identified in <a href="/wiki/Kromdraai" class="mw-redirect" title="Kromdraai">Kromdraai</a>, <a href="/wiki/Swartkrans" title="Swartkrans">Swartkrans</a>, <a href="/wiki/Sterkfontein" title="Sterkfontein">Sterkfontein</a>, <a href="/wiki/Gondolin_Cave" title="Gondolin Cave">Gondolin</a>, <a href="/wiki/Cooper%27s_Cave" title="Cooper's Cave">Cooper's</a>, and <a href="/wiki/Drimolen" title="Drimolen">Drimolen</a> Caves. Discovered in 1938, it was among the first early <a href="/wiki/Hominin" class="mw-redirect" title="Hominin">hominins</a> described, and became the <a href="/wiki/Type_species" title="Type species">type species</a> for the <a href="/wiki/Genus" title="Genus">genus</a> <i><a href="/wiki/Paranthropus" title="Paranthropus">Paranthropus</a></i>. However, it has been argued by some that <i>Paranthropus</i> is an invalid grouping and <a href="/wiki/Synonym_(taxonomy)" title="Synonym (taxonomy)">synonymous</a> with <i><a href="/wiki/Australopithecus" title="Australopithecus">Australopithecus</a></i>, so the species is also often classified as <i><b>Australopithecus robustus</b></i>. </p><p>Robust australopithecines—as opposed to <a href="/wiki/Gracility" title="Gracility">gracile</a> australopithecines—are characterised by heavily built skulls capable of producing high <a href="/wiki/Stress_(physics)" class="mw-redirect" title="Stress (physics)">stresses</a> and <a href="/wiki/Bite_force" class="mw-redirect" title="Bite force">bite forces</a>, as well as inflated <a href="/wiki/Cheek_teeth" title="Cheek teeth">cheek teeth</a> (<a href="/wiki/Molar_(tooth)" title="Molar (tooth)">molars</a> and <a href="/wiki/Premolar" title="Premolar">premolars</a>). Males had more heavily built skulls than females. <i>P. robustus</i> may have had a <a href="/wiki/Genetic_susceptibility" class="mw-redirect" title="Genetic susceptibility">genetic susceptibility</a> for <a href="/wiki/Pitting_enamel_hypoplasia" title="Pitting enamel hypoplasia">pitting enamel hypoplasia</a> on the teeth, and seems to have had a dental <a href="/wiki/Tooth_decay" title="Tooth decay">cavity</a> rate similar to non-agricultural modern humans. The species is thought to have exhibited marked <a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">sexual dimorphism</a>, with males substantially larger and more robust than females. Based on 3 specimens, males may have been 132 cm (4 ft 4 in) tall and females 110 cm (3 ft 7 in). Based on 4 specimens, males averaged 40 kg (88 lb) in weight and females 30 kg (66 lb). The brain volume of the specimen SK 1585 is estimated to have been 476<i> </i>cc, and of DNH 155 about 450<i> </i>cc (for comparison, the brain volume of contemporary <i><a href="/wiki/Homo" title="Homo">Homo</a></i> varied from 500 to 900<i> </i>cc). <i>P. robustus</i> limb anatomy is similar to that of other australopithecines, which may indicate a less efficient walking ability than modern humans, and perhaps some degree of <a href="/wiki/Arboreality" class="mw-redirect" title="Arboreality">arboreality</a> (movement in the trees). </p><p><i>P. robustus</i> seems to have consumed a high proportion of <a href="/wiki/C4_carbon_fixation" title="C4 carbon fixation">C<sub>4</sub> savanna plants</a>. In addition, it may have also eaten fruits, underground <a href="/wiki/Storage_organ" title="Storage organ">storage organs</a> (such as roots and tubers), and perhaps <a href="/wiki/Honey" title="Honey">honey</a> and <a href="/wiki/Termite" title="Termite">termites</a>. <i>P. robustus</i> may have used bones as tools to extract and process food. It is unclear if <i>P. robustus</i> lived in a <a href="/wiki/Harem_(zoology)" title="Harem (zoology)">harem</a> society like <a href="/wiki/Gorilla" title="Gorilla">gorillas</a> or a multi-male society like <a href="/wiki/Baboon" title="Baboon">baboons</a>. <i>P. robustus</i> society may have been <a href="/wiki/Patrilocal" class="mw-redirect" title="Patrilocal">patrilocal</a>, with adult females more likely to leave the group than males, but males may have been more likely to be evicted as indicated by higher male mortality rates and assumed increased risk of predation to solitary individuals. <i>P. robustus</i> contended with <a href="/wiki/Sabertooth_cat" class="mw-redirect" title="Sabertooth cat">sabertooth cats</a>, <a href="/wiki/Leopard" title="Leopard">leopards</a>, and <a href="/wiki/Hyena" title="Hyena">hyenas</a> on the mixed, open-to-closed landscape, and <i>P. robustus</i> bones probably accumulated in caves due to big cat predation. It is typically found in what were mixed open and wooded environments, and may have gone extinct in the <a href="/wiki/Mid-Pleistocene_Transition" title="Mid-Pleistocene Transition">Mid-Pleistocene Transition</a> characterised by the continual prolonging of dry cycles and subsequent retreat of such habitat. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Taxonomy">Taxonomy</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=1" title="Edit section: Taxonomy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <table id="Container" role="presentation" class="nomobile toccolours searchaux" style="text-align:left;padding:0 0.5em;border-style:solid;float:right;clear:right;margin:0.3em 0 0.8em 1.4em;overflow:hidden;min-width:29.95em"><tbody><tr><td colspan="3" id="Title" style="padding:1em 1em 0 1em"><div 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none;position:absolute;text-align:center;margin:0;padding:0;top:26.665em;height:23.335em;left:0.000em;width:1.300em"><div style="font-size:100%;position:relative;top:0em;left:-0.1em"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-11.667em;transform:translateY(-50%);z-index:5"><b><style data-mw-deduplicate="TemplateStyles:r1231500821">@supports(writing-mode:vertical-lr){.mw-parser-output .ts-vertical-text{letter-spacing:-0.12em;line-height:1em;text-orientation:upright;writing-mode:vertical-lr;width:1em}}</style><span class="ts-vertical-text" style=""><a href="/wiki/Miocene" title="Miocene"><span style="color:#ffffff;">Miocene</span></a></span></b></div></div></div><div class="notheme" style="font-size:100%;background-color:#f1c309;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:12.900em;height:13.765em;left:0.000em;width:1.300em"><div style="font-size:100%;position:relative;top:0.5em;left:-0.1em"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-6.883em;transform:translateY(-50%);z-index:5"><b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1231500821"><span class="ts-vertical-text" style=""><a href="/wiki/Pliocene" title="Pliocene">Pliocene</a></span></b></div></div></div><div class="notheme" style="font-size:100%;background-color:#c1e0c1;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.058em;height:12.841em;left:0.000em;width:1.300em"><div style="font-size:100%;position:relative;top:0.75em;left:-0.1em"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-6.421em;transform:translateY(-50%);z-index:5"><b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1231500821"><span class="ts-vertical-text" style=""><a href="/wiki/Pleistocene" title="Pleistocene">Pleistocene</a></span></b></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:14.000em;height:36.000em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:-6.7em;left:-0.3em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-18.000em;transform:translateY(-50%);z-index:5"><span style="font-size:120%"><b><a href="/wiki/Hominini" title="Hominini">Hominini</a></b></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:48.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0em;left:0.1em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5"><i><span style="font-size:90%;"><a href="/wiki/Nakalipithecus" title="Nakalipithecus">Nakalipithecus</a></span></i></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:46.500em;height:1.000em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0.5em;left:0.6em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.500em;transform:translateY(-50%);z-index:5"><i><span style="font-size:90%;"><a href="/wiki/Samburupithecus" title="Samburupithecus">Samburupithecus</a></span></i></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:44.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0em;left:0.2em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5"><i><span style="font-size:90%;"><a href="/wiki/Ouranopithecus" title="Ouranopithecus">Ouranopithecus</a></span></i><br /><span style="font-size:85%;"><i>(<a href="/wiki/Ouranopithecus_turkae" title="Ouranopithecus turkae">Ou. turkae</a>)</i><br /><i>(<a href="/wiki/Ouranopithecus_macedoniensis" title="Ouranopithecus macedoniensis">Ou. macedoniensis</a>)</i></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:39.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0.1em;left:0.4em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5"><i><span style="font-size:90%;"><a href="/wiki/Chororapithecus" title="Chororapithecus">Chororapithecus</a></span></i></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:35.000em;height:10.000em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:-1.75em;left:-0.2em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-5.000em;transform:translateY(-50%);z-index:5"><i><span style="font-size:90%;"><a href="/wiki/Oreopithecus" title="Oreopithecus">Oreopithecus</a></span></i></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:39.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:1.75em;left:-0.4em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5"><i><span style="font-size:90%;"><a href="/wiki/Sivapithecus" title="Sivapithecus">Sivapithecus</a></span></i></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:34.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0.5em;left:0.3em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5"><i><span style="font-size:90%;"><a href="/wiki/Sahelanthropus" title="Sahelanthropus">Sahelanthropus</a></span></i></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:34.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:2em;left:0.3em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5"><i><span style="font-size:90%;"><a href="/wiki/Graecopithecus" title="Graecopithecus">Graecopithecus</a></span></i></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:29.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0.5em;left:-1.5em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5"><i><span style="font-size:90%;"><a href="/wiki/Orrorin" title="Orrorin">Orrorin</a></span></i></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:29.500em;height:0.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:2.5em;left:0.15em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.250em;transform:translateY(-50%);z-index:5"><span style="font-size:85%;"><i>(<a href="/wiki/Orrorin" title="Orrorin">O. praegens</a>)</i><br /><i>(<a href="/wiki/Orrorin" title="Orrorin">O. tugenensis</a>)</i></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:22.000em;height:6.850em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:-3.75em;left:-0.3em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-3.425em;transform:translateY(-50%);z-index:5"><i><b><span style="font-size:90%;"><a href="/wiki/Ardipithecus" title="Ardipithecus">Ardipithecus</a></span></b></i></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:27.700em;height:1.150em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0.75em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.575em;transform:translateY(-50%);z-index:5"><span style="font-size:85%;"><i>(<a href="/wiki/Ardipithecus_kadabba" title="Ardipithecus kadabba">Ar. kadabba</a>)</i></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:21.500em;height:1.000em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:1em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.500em;transform:translateY(-50%);z-index:5"><span style="font-size:85%;"><i>(<a href="/wiki/Ardipithecus_ramidus" title="Ardipithecus ramidus">Ar. ramidus</a>)</i></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:6.000em;height:16.500em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:3em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-8.250em;transform:translateY(-50%);z-index:5"><i><b><span style="font-size:90%;"><a href="/wiki/Australopithecus" title="Australopithecus">Australopithecus</a></span></b></i><br /><span style="font-size:85%;"><i>(<a href="/wiki/Australopithecus_africanus" title="Australopithecus africanus">Au. africanus</a>)</i><br /><i>(<a href="/wiki/Australopithecus_afarensis" title="Australopithecus afarensis">Au. afarensis</a>)</i><br /><i>(<a href="/wiki/Australopithecus_anamensis" title="Australopithecus anamensis">Au. anamensis</a>)</i></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#DCEEFF;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:8.250em;height:5.750em;left:2.600em;width:10.400em"><div style="font-size:100%;position:relative;top:0.9em;left:-0.5em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.875em;transform:translateY(-50%);z-index:5"><i><b><span style="font-size:90%;"><a href="/wiki/Homo_habilis" title="Homo habilis">H. habilis</a></span></b></i><br /><span style="font-size:85%;"><i>(<a href="/wiki/Homo_rudolfensis" title="Homo rudolfensis">H. rudolfensis</a>)</i><br /><i>(<a href="/wiki/Australopithecus_garhi" title="Australopithecus garhi">Au. garhi</a>)</i></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#FFE4F1;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.570em;height:9.430em;left:3.900em;width:9.100em"><div style="font-size:100%;position:relative;top:1.5em;left:-1em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-4.715em;transform:translateY(-50%);z-index:5"><i><b><span style="font-size:90%;"><a href="/wiki/Homo_erectus" title="Homo erectus">H. erectus</a></span></b></i><br /><span style="font-size:85%;"><i>(<a href="/wiki/Homo_antecessor" title="Homo antecessor">H. antecessor</a>)</i><br /><i>(<a href="/wiki/Homo_ergaster" title="Homo ergaster">H. ergaster</a>)</i><br /><i>(<a href="/wiki/Australopithecus_sediba" title="Australopithecus sediba">Au. sediba</a>)</i></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#FFE4F1;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:6.000em;height:2.500em;left:2.600em;width:10.400em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-1.250em;transform:translateY(-50%);z-index:5"></div></div></div><div class="notheme" style="font-size:100%;background-color:#7DFFFF;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:1.000em;height:2.500em;left:2.600em;width:10.400em"><div style="font-size:100%;position:relative;top:1em;left:-0.2em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-1.250em;transform:translateY(-50%);z-index:5"><i><span style="font-size:90%;"><a href="/wiki/Homo_heidelbergensis" title="Homo heidelbergensis">H. heidelbergensis</a></span></i></div></div></div><div class="notheme" style="font-size:100%;background-color:#F3F300;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:1.500em;left:3.900em;width:9.100em"><div style="font-size:100%;position:relative;top:0em;left:-0.7em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.750em;transform:translateY(-50%);z-index:5"><i><b><span style="font-size:90%;"><a href="/wiki/Anatomically_modern_human" class="mw-redirect" title="Anatomically modern human">Homo sapiens</a></span></b></i></div></div></div><div class="notheme" style="font-size:100%;background-color:#F3F300;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:0.200em;left:1.300em;width:3.900em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.100em;transform:translateY(-50%);z-index:5"></div></div></div><div class="notheme" style="font-size:100%;background-color:#7DFFFF;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.200em;height:1.050em;left:2.600em;width:1.300em"><div style="font-size:100%;position:relative;top:1.8em;left:2.6em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.525em;transform:translateY(-50%);z-index:5"><span class="nowrap"><span style="font-size:90%;"><a href="/wiki/Neanderthal" title="Neanderthal">Neanderthals</a></span></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#F3F300;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.200em;height:0.370em;left:1.300em;width:11.700em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.185em;transform:translateY(-50%);z-index:5"></div></div></div><div class="notheme" style="font-size:100%;background-color:#7DFFFF;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.200em;height:0.370em;left:2.600em;width:1.300em"><div style="font-size:100%;position:relative;top:1.4em;left:2.8em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.185em;transform:translateY(-50%);z-index:5"><span class="nowrap"><span style="font-size:90%;"><a href="/wiki/Denisovan" title="Denisovan">Denisovans</a></span></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#ABFFAB;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:6.000em;height:8.500em;left:11.700em;width:1.300em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-4.250em;transform:translateY(-50%);z-index:5"> </div></div></div></td><td id="Annotations" style="padding:0;margin:0.7em 0 0.7em 0;float:left;position:relative;width:11.05em;height:50em"><table role="presentation" style="position:absolute;z-index:15;top:50.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Timeline_of_human_evolution#Hominidae" title="Timeline of human evolution">Earlier apes</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:45.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Gorilla%E2%80%93human_last_common_ancestor" title="Gorilla–human last common ancestor">Gorilla split</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:42.500em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Chimpanzee%E2%80%93human_last_common_ancestor" title="Chimpanzee–human last common ancestor">Chimpanzee split</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:35.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Human_skeletal_changes_due_to_bipedalism" title="Human skeletal changes due to bipedalism">Earliest bipedal</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:29.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Ardipithecus_kadabba" title="Ardipithecus kadabba">Earliest sign of <i>Ardipithecus</i></a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:22.500em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Australopithecus_anamensis" title="Australopithecus anamensis">Earliest sign of <i>Australopithecus</i></a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:17.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Stone_Age#Beginning_of_the_Stone_Age" title="Stone Age">Earliest stone tools</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:14.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.25em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/LD_350-1" title="LD 350-1">Earliest sign of<br /> <i>Homo</i></a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:10.600em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.25em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Early_expansions_of_hominins_out_of_Africa" title="Early expansions of hominins out of Africa">Dispersal beyond Africa</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:9.550em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Origin_of_language" title="Origin of language">Earliest language</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:8.500em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Control_of_fire_by_early_humans" title="Control of fire by early humans">Earliest fire</a> / <a href="/wiki/Cooking#History" title="Cooking">cooking</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:1.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.6em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><span style="font-size:95%;"><a href="/wiki/Rock_art#East_Asia" title="Rock art">Earliest rock art</a></span></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:0.625em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.1em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><span style="font-size:95%;"><a href="/wiki/Clothing#Origin_and_history" title="Clothing">Earliest clothes</a></span></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:0.250em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.4em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><span style="font-size:95%;"><a href="/wiki/Behavioral_modernity" title="Behavioral modernity">Modern humans</a></span></div></div></td></tr></tbody></table><div class="annot-bar" style="width:auto;font-size:100%;position:absolute;text-align:center;margin-top:21.000em"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;text-align:left;top:0.25em;left:-5em;width:8.8em"><div style="position:relative;width:auto;z-index:10;font-size:90%;color:var( --color-base, #000);vertical-align:middle;line-height:105%;bottom:0"><span style="display:block; text-align:center; font-size:110%"><b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1231500821"><span class="ts-vertical-text" style=""><a href="/wiki/Hominidae" title="Hominidae">H o m i n i d s</a></span></b></span></div></div></div><div class="annot-bar" style="width:auto;font-size:100%;position:absolute;text-align:center;margin-top:3.000em"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;text-align:left;top:3.25em;left:-5em;width:8.8em"><div style="position:relative;width:auto;z-index:10;font-size:90%;color:var( --color-base, #000);vertical-align:middle;line-height:105%;bottom:0"><span style="display:block; text-align:center; font-size:85%;"><b><span style="font-size:85%;"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1231500821"><span class="ts-vertical-text" style=""><a href="/wiki/Paranthropus" title="Paranthropus">P a r a n t h r o p u s</a></span></span></b></span></div></div></div></td></tr><tr><td id="Caption" colspan="3" style="padding:0;margin:0 0.2em 0.7em 0.2em"><div id="Caption" class="toccolours" style="margin:0 0.5em;border-style:none;clear:both;text-align:center;width:26.8em"><div style="float:left;font-size:85%;">(<a href="/wiki/Myr" class="mw-redirect" title="Myr">million years ago</a>)</div></div></td></tr></tbody></table> <div class="mw-heading mw-heading3"><h3 id="Research_history">Research history</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=2" title="Edit section: Research history"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading4"><h4 id="Discovery">Discovery</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=3" title="Edit section: Discovery"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:TM_1517.jpg" class="mw-file-description"><img alt="Preserved left half of the skull and right half of the jaw mounted on a wall" src="//upload.wikimedia.org/wikipedia/commons/thumb/0/06/TM_1517.jpg/220px-TM_1517.jpg" decoding="async" width="220" height="169" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/06/TM_1517.jpg/330px-TM_1517.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/06/TM_1517.jpg/440px-TM_1517.jpg 2x" data-file-width="2932" data-file-height="2248" /></a><figcaption>Cast of the face of <a href="/wiki/TM_1517" title="TM 1517">TM 1517</a>, the <a href="/wiki/Holotype_specimen" class="mw-redirect" title="Holotype specimen">holotype specimen</a> of <i>P. robustus</i></figcaption></figure> <p>The first remains, a partial skull including a part of the jawbone (<a href="/wiki/TM_1517" title="TM 1517">TM 1517</a>), were discovered in June 1938 at the <a href="/wiki/Kromdraai_fossil_site" title="Kromdraai fossil site">Kromdraai cave site</a>, South Africa, by local schoolboy Gert Terblanche. He gave the remains to South African conservationist Charles Sydney Barlow, who then relayed them to South African palaeontologist <a href="/wiki/Robert_Broom" title="Robert Broom">Robert Broom</a>.<sup id="cite_ref-Broom1938_2-0" class="reference"><a href="#cite_note-Broom1938-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Broom began investigating the site, and, a few weeks later, recovered a right distal <a href="/wiki/Humerus" title="Humerus">humerus</a> (the lower part of the upper arm bone), a proximal right <a href="/wiki/Ulna" title="Ulna">ulna</a> (upper part of a lower arm bone) and a <a href="/wiki/Distal_phalanx" class="mw-redirect" title="Distal phalanx">distal phalanx</a> bone of the big toe, all of which he assigned to TM 1517. He also identified a distal toe phalanx which he believed belonged to a <a href="/wiki/Baboon" title="Baboon">baboon</a>, but has since been associated with TM 1517.<sup id="cite_ref-Cazenave2019_3-0" class="reference"><a href="#cite_note-Cazenave2019-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Broom noted the Kromdraai remains were especially robust compared to other <a href="/wiki/Hominin" class="mw-redirect" title="Hominin">hominins</a>.<sup id="cite_ref-Broom1938_2-1" class="reference"><a href="#cite_note-Broom1938-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> In August 1938, Broom classified the robust Kromdraai remains into a new <a href="/wiki/Genus" title="Genus">genus</a>, as <i>Paranthropus robustus</i>.<sup id="cite_ref-Broom1938_2-2" class="reference"><a href="#cite_note-Broom1938-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> "<i><a href="/wiki/Paranthropus" title="Paranthropus">Paranthropus</a></i>" derives from the <a href="/wiki/Ancient_Greek" title="Ancient Greek">Ancient Greek</a> <span title="Ancient Greek (to 1453)-language text"><span lang="grc">παρα</span></span> <span title="Ancient Greek (to 1453)-language romanization"><i lang="grc-Latn">para</i></span>, beside or alongside; and <span title="Ancient Greek (to 1453)-language text"><span lang="grc">άνθρωπος</span></span> <span title="Ancient Greek (to 1453)-language romanization"><i lang="grc-Latn">ánthropos</i></span>, man.<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> </p><p>At this point in time, Australian anthropologist <a href="/wiki/Raymond_Dart" title="Raymond Dart">Raymond Dart</a> had made the very first claim (quite controversially at the time) of an early <a href="/wiki/Ape" title="Ape">ape</a>-like human ancestor in 1924 from South Africa, <i><a href="/wiki/Australopithecus_africanus" title="Australopithecus africanus">Australopithecus africanus</a></i>, based on the <a href="/wiki/Taung_child" class="mw-redirect" title="Taung child">Taung child</a>.<sup id="cite_ref-Tobias1998_5-0" class="reference"><a href="#cite_note-Tobias1998-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 285–288">: 285–288 </span></sup> In 1936, Broom had described "<i>Plesianthropus transvaalensis</i>" (now <a href="/wiki/Synonym_(taxonomy)" title="Synonym (taxonomy)">synonymised</a> with <i>A. africanus</i>) from the <a href="/wiki/Sterkfontein" title="Sterkfontein">Sterkfontein</a> Caves only 2 km (1.2 mi) west from Kromdraai. All these species dated to the <a href="/wiki/Pleistocene" title="Pleistocene">Pleistocene</a> and were found in the same general vicinity (now called the "<a href="/wiki/Cradle_of_Humankind" title="Cradle of Humankind">Cradle of Humankind</a>"). Broom considered them evidence of a greater diversity of hominins in the <a href="/wiki/Pliocene" title="Pliocene">Pliocene</a> from which they and modern humans descended, and consistent with several hominin taxa existing alongside human ancestors.<sup id="cite_ref-Broom1938_2-3" class="reference"><a href="#cite_note-Broom1938-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>The Kromdraai taxon, classified as <i>Paranthropus robustus</i>, was later discovered at the nearby <a href="/wiki/Swartkrans" title="Swartkrans">Swartkrans</a> Cave in 1948. <i>P. robustus</i> was only definitively identified at Kromdraai and Swartkrans until around the turn of the century when the species was reported elsewhere in the Cradle of Humankind at Sterkfontein, <a href="/wiki/Gondolin_Cave" title="Gondolin Cave">Gondolin</a>, <a href="/wiki/Cooper%27s_Cave" title="Cooper's Cave">Cooper's</a>, and <a href="/wiki/Drimolen" title="Drimolen">Drimolen</a> Caves. The species has not been found outside this small area.<sup id="cite_ref-Constantino2004_6-0" class="reference"><a href="#cite_note-Constantino2004-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id=""P._crassidens""><span id=".22P._crassidens.22"></span>"<i>P. crassidens</i>"</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=4" title="Edit section: "P. crassidens""><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1948, at the nearby Swartkrans Cave, Broom described "<i>P. crassidens</i>" (distinct from <i>P. robustus</i>) based on a subadult jaw, SK 6,<sup id="cite_ref-Constantino2004_6-1" class="reference"><a href="#cite_note-Constantino2004-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> because Swartkrans and Kromdraai clearly dated to different time intervals based on the diverging animal assemblages in these caves.<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> At this point in time, humans and allies were classified into the <a href="/wiki/Family_(biology)" title="Family (biology)">family</a> <a href="/wiki/Hominidae" title="Hominidae">Hominidae</a>, and non-human <a href="/wiki/Great_ape" class="mw-redirect" title="Great ape">great apes</a> into "<a href="/wiki/Pongidae" title="Pongidae">Pongidae</a>"; in 1950, Broom suggested separating early hominins into the <a href="/wiki/Subfamilies" class="mw-redirect" title="Subfamilies">subfamilies</a> Australopithecinae (<i>Au. africanus</i> and "<i>Pl. transvaalensis</i>"), "Paranthropinae" (<i>Pa. robustus</i> and "<i>Pa. crassidens</i>"), and "Archanthropinae" ("<i>Au. prometheus</i>").<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> This scheme was widely criticised for being too liberal in demarcating species.<sup id="cite_ref-Robinson1954_9-0" class="reference"><a href="#cite_note-Robinson1954-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Further, the remains were not firmly dated, and it was debated if there were indeed multiple hominin lineages or if there was only a single one leading to humans. Most prominently, Broom and South African palaeontologist <a href="/wiki/John_Talbot_Robinson" title="John Talbot Robinson">John Talbot Robinson</a> continued arguing for the validity of <i>Paranthropus</i>.<sup id="cite_ref-Johanson_10-0" class="reference"><a href="#cite_note-Johanson-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p><p>Anthropologists <a href="/wiki/Sherwood_Washburn" title="Sherwood Washburn">Sherwood Washburn</a> and Bruce D. Patterson were the first to recommend synonymising <i>Paranthropus</i> with <i><a href="/wiki/Australopithecus" title="Australopithecus">Australopithecus</a></i> in 1951, wanting to limit hominin genera to only that and <i><a href="/wiki/Homo" title="Homo">Homo</a></i>,<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> and it has since been debated whether or not <i>Paranthropus</i> is a <a href="/wiki/Junior_synonym" class="mw-redirect" title="Junior synonym">junior synonym</a> of <i>Australopithecus</i>.<sup id="cite_ref-Constantino2004_6-2" class="reference"><a href="#cite_note-Constantino2004-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> In the spirit of tightening <a href="/wiki/Lumping_and_splitting" class="mw-redirect" title="Lumping and splitting">splitting</a> criteria for hominin taxa, in 1954, Robinson suggested demoting "<i>P. crassidens</i>" to subspecies level as "<i>P. r. crassidens</i>", and also moved the Indonesian <i><a href="/wiki/Meganthropus" title="Meganthropus">Meganthropus</a></i> into the genus as "<i>P. palaeojavanicus</i>".<sup id="cite_ref-Robinson1954_9-1" class="reference"><a href="#cite_note-Robinson1954-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> <i>Meganthropus</i> has since been variously reclassified as a synonym of the Asian <i><a href="/wiki/Homo_erectus" title="Homo erectus">Homo erectus</a></i>, "<i><a href="/wiki/Anthropopithecus" title="Anthropopithecus">Pithecanthropus dubius</a></i>", <i>Pongo</i> (<a href="/wiki/Orangutan" title="Orangutan">orangutans</a>), and so on, and in 2019 it was again argued to be a valid genus.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p><p>In 1949, also in Swartkrans Cave, Broom and Robinson found a mandible which they preliminary described as "intermediate between one of the ape-men and true man," classifying it as a new genus and species "<i>Telanthropus capensis</i>". Most immediate reactions favoured synonymising "<i>T. capensis</i>" with "<i>P. crassidens</i>", whose remains were already abundantly found in the cave.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> In 1957, though, Italian biologist Alberto Simonetta moved it to the genus "<i>Pithecanthropus</i>", and Robinson (without a specific reason why) decided to synonymise it with <i>H. erectus</i> (African <i>H. erectus</i> are sometimes called <i><a href="/wiki/H._ergaster" class="mw-redirect" title="H. ergaster">H. ergaster</a></i> today). In 1965, South African palaeoanthropologist <a href="/wiki/Phillip_V._Tobias" class="mw-redirect" title="Phillip V. Tobias">Phillip V. Tobias</a> questioned whether this classification is completely sound or not.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p><p>By the 21st century, "<i>P. crassidens</i>" had more or less fallen out of use in favour of <i>P. robustus</i>. American palaeoanthropologist <a href="/wiki/Frederick_E._Grine" title="Frederick E. Grine">Frederick E. Grine</a> is the primary opponent of synonymisation of the two species.<sup id="cite_ref-Constantino2004_6-3" class="reference"><a href="#cite_note-Constantino2004-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Gigantopithecus"><i>Gigantopithecus</i></h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=5" title="Edit section: Gigantopithecus"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1939, Broom hypothesised that <i>P. robustus</i> was closely related to the similarly large-toothed ape <i><a href="/wiki/Gigantopithecus" title="Gigantopithecus">Gigantopithecus</a></i> from Asia (extinct apes were primarily known from Asia at the time) believing <i>Gigantopithecus</i> to have been a hominin.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> Primarily influenced by the mid-century opinions of Jewish German anthropologist <a href="/wiki/Franz_Weidenreich" title="Franz Weidenreich">Franz Weidenreich</a> and German-Dutch palaeontologist <a href="/wiki/Gustav_Heinrich_Ralph_von_Koenigswald" title="Gustav Heinrich Ralph von Koenigswald">Ralph von Koenigswald</a> that <i>Gigantopithecus</i> was, respectively, the direct ancestor of the Asian <i>H. erectus</i> or closely related, much debate followed over whether <i>Gigantopithecus</i> was a hominin or a non-human ape.<sup id="cite_ref-ZhangHarrison2017_16-0" class="reference"><a href="#cite_note-ZhangHarrison2017-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p><p>In 1972, Robinson suggested including <i>Gigantopithecus</i> in "Paranthropinae", with the <a href="/wiki/Miocene" title="Miocene">Miocene</a> Pakistani "<i>G. bilaspurensis</i>" (now <i><a href="/wiki/Indopithecus" title="Indopithecus">Indopithecus</a></i>) as the ancestor of <i>Paranthropus</i> and the Chinese <i>G. blacki</i>. He also believed that they both had a massive build. In contrast, he reported a very small build for <i>A. africanus</i> (which he referred to as "<i>Homo</i>" <i>africanus</i>) and speculated it had some cultural and hunting abilities, being a member of the human lineage, which "paranthropines" lacked.<sup id="cite_ref-Wolpoff1974_17-0" class="reference"><a href="#cite_note-Wolpoff1974-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> With the popularisation of <a href="/wiki/Cladistic" class="mw-redirect" title="Cladistic">cladistics</a> by the late 1970s to 1980s, and better resolution on how Miocene apes relate to later apes, <i>Gigantopithecus</i> was entirely removed from <a href="/wiki/Homininae" title="Homininae">Homininae</a>, and is now placed in the subfamily <a href="/wiki/Ponginae" title="Ponginae">Ponginae</a> with orangutans.<sup id="cite_ref-ZhangHarrison2017_16-1" class="reference"><a href="#cite_note-ZhangHarrison2017-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="P._boisei"><i>P. boisei</i></h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=6" title="Edit section: P. boisei"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Australophithecus_boisei_(cast),_Olduvai_Gorge_-_Springfield_Science_Museum_-_Springfield,_MA_-_DSC03368.JPG" class="mw-file-description"><img alt="Large, white, complete skull in three-quarter view facing left" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Australophithecus_boisei_%28cast%29%2C_Olduvai_Gorge_-_Springfield_Science_Museum_-_Springfield%2C_MA_-_DSC03368.JPG/220px-Australophithecus_boisei_%28cast%29%2C_Olduvai_Gorge_-_Springfield_Science_Museum_-_Springfield%2C_MA_-_DSC03368.JPG" decoding="async" width="220" height="203" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Australophithecus_boisei_%28cast%29%2C_Olduvai_Gorge_-_Springfield_Science_Museum_-_Springfield%2C_MA_-_DSC03368.JPG/330px-Australophithecus_boisei_%28cast%29%2C_Olduvai_Gorge_-_Springfield_Science_Museum_-_Springfield%2C_MA_-_DSC03368.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Australophithecus_boisei_%28cast%29%2C_Olduvai_Gorge_-_Springfield_Science_Museum_-_Springfield%2C_MA_-_DSC03368.JPG/440px-Australophithecus_boisei_%28cast%29%2C_Olduvai_Gorge_-_Springfield_Science_Museum_-_Springfield%2C_MA_-_DSC03368.JPG 2x" data-file-width="3495" data-file-height="3227" /></a><figcaption><a href="/wiki/OH_5" class="mw-redirect" title="OH 5">OH 5</a> (reconstructed skull above) was made the <a href="/wiki/Holotype_specimen" class="mw-redirect" title="Holotype specimen">holotype specimen</a> of the second <i><a href="/wiki/Paranthropus" title="Paranthropus">Paranthropus</a></i> species, <i><a href="/wiki/P._boisei" class="mw-redirect" title="P. boisei">P. boisei</a></i>.</figcaption></figure> <p>In 1959, another and much more robust australopithecine was discovered in East Africa, <i><a href="/wiki/P._boisei" class="mw-redirect" title="P. boisei">P. boisei</a></i>, and in 1975, the <i>P. boisei</i> skull <a href="/wiki/KNM-ER_406" title="KNM-ER 406">KNM-ER 406</a> was demonstrated to have been contemporaneous with the <i>H. ergaster/H. erectus</i> skull <a href="/wiki/KNM_ER_3733" title="KNM ER 3733">KNM ER 3733</a> (which is considered a human ancestor). This is generally taken to show that <i>Paranthropus</i> was a <a href="/wiki/Sister_taxon" class="mw-redirect" title="Sister taxon">sister taxon</a> to <i>Homo</i>, both developing from some <i>Australopithecus</i> species, which at the time only included <i>A. africanus</i>.<sup id="cite_ref-Johanson_10-1" class="reference"><a href="#cite_note-Johanson-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p><p>In 1979, a year after describing <i><a href="/wiki/A._afarensis" class="mw-redirect" title="A. afarensis">A. afarensis</a></i> from East Africa, anthropologists <a href="/wiki/Donald_Johanson" title="Donald Johanson">Donald Johanson</a> and <a href="/wiki/Tim_D._White" title="Tim D. White">Tim D. White</a> suggested that <i>A. afarensis</i> was instead the last common ancestor between <i>Homo</i> and <i>Paranthropus</i>, and <i>A. africanus</i> was the earliest member of the <i>Paranthropus</i> lineage or at least was ancestral to <i>P. robustus</i>, because <i>A. africanus</i> inhabited South Africa before <i>P. robustus</i>, and <i>A. afarensis</i> was at the time the oldest known hominin species at roughly 3.5 million years old.<sup id="cite_ref-Johanson_10-2" class="reference"><a href="#cite_note-Johanson-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Now, the earliest-known South African australopithecine ("<a href="/wiki/Little_Foot" title="Little Foot">Little Foot</a>") dates to 3.67 million years ago, contemporaneous with <i>A. afarensis</i>.<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> The matter is still debated.<sup id="cite_ref-McNulty2016_19-0" class="reference"><a href="#cite_note-McNulty2016-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p><p>It was long assumed that if <i>Paranthropus</i> is a valid genus then <i>P. robustus</i> was the ancestor of <i>P. boisei</i>, but in 1985, anthropologists <a href="/wiki/Alan_Walker_(anthropologist)" title="Alan Walker (anthropologist)">Alan Walker</a> and <a href="/wiki/Richard_Leakey" title="Richard Leakey">Richard Leakey</a> found that the 2.5-million-year-old East African skull <a href="/wiki/KNM_WT_17000" title="KNM WT 17000">KNM WT 17000</a>—which they assigned to a new species <i><a href="/wiki/Australopithecus_aethiopicus" class="mw-redirect" title="Australopithecus aethiopicus">A. aethiopicus|A. aethiopicus</a></i>—was ancestral to <i>A. boisei</i> (they considered <i>Paranthropus</i> synonymous with <i>Australopithecus</i>), thus establishing the <i>boisei</i> lineage as beginning long before <i>robustus</i> had existed.<sup id="cite_ref-Walker1986_20-0" class="reference"><a href="#cite_note-Walker1986-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Classification">Classification</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=7" title="Edit section: Classification"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The genus <i>Paranthropus</i> (otherwise known as "robust australopithecines", in contrast to the "<a href="/wiki/Gracile" class="mw-redirect" title="Gracile">gracile</a> australopithecines") now also includes the East African <i>P. boisei</i> and <i>P. aethiopicus</i>. It is still debated if this is a valid natural grouping (<a href="/wiki/Monophyletic" class="mw-redirect" title="Monophyletic">monophyletic</a>) or an invalid grouping of similar-looking hominins (<a href="/wiki/Paraphyletic" class="mw-redirect" title="Paraphyletic">paraphyletic</a>). Because skeletal elements are so limited in these species, their <a href="/wiki/Affinity_(taxonomy)" title="Affinity (taxonomy)">affinities</a> with each other and with other <a href="/wiki/Australopithecine" title="Australopithecine">australopithecines</a> are difficult to gauge with accuracy. The jaws are the main argument for monophyly, but jaw anatomy is strongly influenced by diet and environment, and could have evolved independently in <i>P. robustus</i> and <i>P. boisei</i>. Proponents of monophyly consider <i>P. aethiopicus</i> to be ancestral to the other two species, or closely related to the ancestor. Proponents of paraphyly allocate these three species to the genus <i>Australopithecus</i> as <i>A. boisei</i>, <i>A. aethiopicus</i>, and <i>A. robustus</i>.<sup id="cite_ref-wood2_21-0" class="reference"><a href="#cite_note-wood2-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> In 2020, palaeoanthropologist Jesse M. Martin and colleagues' <a href="/wiki/Phylogenetic_analyses" class="mw-redirect" title="Phylogenetic analyses">phylogenetic analyses</a> reported the monophyly of <i>Paranthropus</i>, but also that <i>P. robustus</i> had branched off before <i>P. aethiopicus</i> (that <i>P. aethiopicus</i> was ancestral to only <i>P. boisei</i>).<sup id="cite_ref-Martin2020_22-0" class="reference"><a href="#cite_note-Martin2020-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> The exact classification of <i>Australopithecus</i> species with each other is quite contentious.<sup id="cite_ref-McNulty2016_19-1" class="reference"><a href="#cite_note-McNulty2016-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p> <style data-mw-deduplicate="TemplateStyles:r1216315873">.mw-parser-output div.clade-gallery{display:table;margin:0 auto}.mw-parser-output div.clade-gallery>div{line-height:1.5em;padding:0.2em 0.4em;border:1px solid #a2a9b1}.mw-parser-output div.clade-gallery div.main-caption{text-align:center;font-weight:bold;border:none}.mw-parser-output div.clade-gallery div.main-footer{text-align:center}.mw-parser-output div.clade-gallery div.main-content{display:table}.mw-parser-output div.clade-gallery table.gallery-element{float:left!important;border:1px none;border:1px solid #a2a9b1;padding:5px;margin:1px;border-collapse:collapse}.mw-parser-output div.clade-gallery div.gallery-element{float:left;border:1px none;border:1px solid #a2a9b1;padding:5px;margin:1px;border-collapse:collapse}.mw-parser-output div.clade-gallery .no-border{border:none}.mw-parser-output div.clade-gallery table.gallery-element.vertical{float:none}.mw-parser-output div.clade-gallery table.gallery-element caption.caption{text-align:center;font-weight:bold;border:none}.mw-parser-output div.clade-gallery table.gallery-element th.header{text-align:center;font-weight:bold}.mw-parser-output div.clade-gallery table.gallery-element td.cladogram{padding:10px;margin:-1px 0}.mw-parser-output div.clade-gallery table.gallery-element td.footer{border:1px solid #a2a9b1;margin:0}.mw-parser-output div.clade-gallery div.header.shading{background-color:#eaecf0}.mw-parser-output div.clade-gallery div.cladogram.shading{background-color:#f8f9fa}.mw-parser-output div.clade-gallery div.footer.shading{background-color:#f8f9fa}</style><div class="clade-gallery" style="float:centre"><div class="main-content"><table class="gallery-element" style="width:200px;height:225px;"><caption class="caption">Monophyly<sup id="cite_ref-wood2_21-1" class="reference"><a href="#cite_note-wood2-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 119">: 119 </span></sup></caption><tbody><tr><td class="cladogram" style="height:225px"> <div class="clade"><style data-mw-deduplicate="TemplateStyles:r1258728058">body.skin-vector-2022 .mw-parser-output div.clade,body.skin-minerva .mw-parser-output div.clade{overflow-x:auto;overflow-y:hidden}body.skin-minerva .mw-parser-output div.clade p{font-size:inherit}.mw-parser-output table.clade{border-spacing:0;margin:0;font-size:100%;line-height:100%;border-collapse:separate;width:auto}.mw-parser-output table.clade table.clade{width:100%;line-height:inherit}.mw-parser-output table.clade td.clade-label{min-width:0.2em;width:0.2em;padding:0.1em 0.25em;vertical-align:bottom;text-align:center;border-left:1px solid;border-bottom:1px solid;white-space:nowrap}.mw-parser-output table.clade td.clade-label::before,.mw-parser-output table.clade td.clade-slabel::before{content:"\2060 "}.mw-parser-output table.clade td.clade-fixed-width{overflow:hidden;text-overflow:ellipsis}.mw-parser-output table.clade td.clade-fixed-width:hover{overflow:visible}.mw-parser-output table.clade td.clade-label.first{border-left:none;border-right:none}.mw-parser-output table.clade td.clade-label.reverse{border-left:none;border-right:1px solid}.mw-parser-output table.clade td.clade-slabel{padding:0.1em 0.25em;vertical-align:top;text-align:center;border-left:1px solid;white-space:nowrap}.mw-parser-output table.clade td.clade-slabel:hover{overflow:visible}.mw-parser-output table.clade td.clade-slabel.last{border-left:none;border-right:none}.mw-parser-output table.clade td.clade-slabel.reverse{border-left:none;border-right:1px solid}.mw-parser-output table.clade td.clade-bar{vertical-align:middle;text-align:left;padding:0 0.5em;position:relative}.mw-parser-output table.clade td.clade-bar.reverse{text-align:right;position:relative}.mw-parser-output table.clade td.clade-leaf{border:0;padding:0;text-align:left}.mw-parser-output table.clade td.clade-leafR{border:0;padding:0;text-align:right}.mw-parser-output table.clade td.clade-leaf.reverse{text-align:right}.mw-parser-output table.clade:hover span.linkA{background-color:yellow}.mw-parser-output table.clade:hover span.linkB{background-color:green}</style> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><i><a href="/wiki/Australopithecus_africanus" title="Australopithecus africanus">A. africanus</a></i> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><i><a href="/wiki/P._aethiopicus" class="mw-redirect" title="P. aethiopicus">P. aethiopicus</a></i> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><i><a href="/wiki/P._boisei" class="mw-redirect" title="P. boisei">P. boisei</a></i> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <p><i><b>P. robustus</b></i> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div></td></tr></tbody></table><table class="gallery-element" style="width:200px;height:225px;"><caption class="caption">Paraphyly<sup id="cite_ref-wood2_21-2" class="reference"><a href="#cite_note-wood2-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 119">: 119 </span></sup></caption><tbody><tr><td class="cladogram" style="height:225px"> <div class="clade"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><i><a href="/wiki/Australopithecus_africanus" title="Australopithecus africanus">A. africanus</a></i> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <p><i><b>P. robustus</b></i> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><i><a href="/wiki/P._aethiopicus" class="mw-redirect" title="P. aethiopicus">P. aethiopicus</a></i> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <p><i><a href="/wiki/P._boisei" class="mw-redirect" title="P. boisei">P. boisei</a></i> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div></td></tr></tbody></table><table class="gallery-element" style="width:200px;height:225px;"><caption class="caption">Monophyly<sup id="cite_ref-Martin2020_22-1" class="reference"><a href="#cite_note-Martin2020-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup></caption><tbody><tr><td class="cladogram" style="height:225px"> <div class="clade"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><i><a href="/wiki/Australopithecus_africanus" title="Australopithecus africanus">A. africanus</a></i> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><i><a href="/wiki/Homo" title="Homo">Homo</a></i> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <p><i><a href="/wiki/Kenyanthropus" title="Kenyanthropus">Kenyanthropus</a></i> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><i><b>P. robustus</b></i> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><i><a href="/wiki/P._aethiopicus" class="mw-redirect" title="P. aethiopicus">P. aethiopicus</a></i> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <p><i><a href="/wiki/P._boisei" class="mw-redirect" title="P. boisei">P. boisei</a></i> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div></td></tr></tbody></table></div><div class="main-footer">Three examples of family trees with <i>P. robustus</i> (note, they are not absolute)</div></div> <p>In 2023, fragmentary genetic material belonging to this species was reported from 2 million year-old teeth, being the oldest genetic evidence to be retrieved from a human.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Anatomy">Anatomy</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=8" title="Edit section: Anatomy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Head">Head</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=9" title="Edit section: Head"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading4"><h4 id="Skull">Skull</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=10" title="Edit section: Skull"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1237032888/mw-parser-output/.tmulti">.mw-parser-output .tmulti .multiimageinner{display:flex;flex-direction:column}.mw-parser-output .tmulti .trow{display:flex;flex-direction:row;clear:left;flex-wrap:wrap;width:100%;box-sizing:border-box}.mw-parser-output .tmulti .tsingle{margin:1px;float:left}.mw-parser-output .tmulti .theader{clear:both;font-weight:bold;text-align:center;align-self:center;background-color:transparent;width:100%}.mw-parser-output .tmulti .thumbcaption{background-color:transparent}.mw-parser-output .tmulti .text-align-left{text-align:left}.mw-parser-output .tmulti .text-align-right{text-align:right}.mw-parser-output .tmulti .text-align-center{text-align:center}@media all and (max-width:720px){.mw-parser-output .tmulti .thumbinner{width:100%!important;box-sizing:border-box;max-width:none!important;align-items:center}.mw-parser-output .tmulti .trow{justify-content:center}.mw-parser-output .tmulti .tsingle{float:none!important;max-width:100%!important;box-sizing:border-box;text-align:center}.mw-parser-output .tmulti .tsingle .thumbcaption{text-align:left}.mw-parser-output .tmulti .trow>.thumbcaption{text-align:center}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .tmulti .multiimageinner img{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .tmulti .multiimageinner img{background-color:white}}</style><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:408px;max-width:408px"><div class="trow"><div class="tsingle" style="width:202px;max-width:202px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:DNH7.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/DNH7.jpg/200px-DNH7.jpg" decoding="async" width="200" height="267" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/DNH7.jpg/300px-DNH7.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/09/DNH7.jpg/400px-DNH7.jpg 2x" data-file-width="3456" data-file-height="4608" /></a></span></div></div><div class="tsingle" style="width:202px;max-width:202px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Paranthropus_robustus_top_(University_of_Zurich).JPG" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8e/Paranthropus_robustus_top_%28University_of_Zurich%29.JPG/200px-Paranthropus_robustus_top_%28University_of_Zurich%29.JPG" decoding="async" width="200" height="267" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8e/Paranthropus_robustus_top_%28University_of_Zurich%29.JPG/300px-Paranthropus_robustus_top_%28University_of_Zurich%29.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8e/Paranthropus_robustus_top_%28University_of_Zurich%29.JPG/400px-Paranthropus_robustus_top_%28University_of_Zurich%29.JPG 2x" data-file-width="1944" data-file-height="2592" /></a></span></div></div></div><div class="trow" style="display:flex"><div class="thumbcaption">Female DNH 7 (left) and cast of male <a href="/wiki/SK_48" title="SK 48">SK 48</a> (right) skulls</div></div></div></div> <p>Typical of <i>Paranthropus</i>, <i>P. robustus</i> exhibits <a href="/wiki/Post-canine_megadontia" class="mw-redirect" title="Post-canine megadontia">post-canine megadontia</a> with enormous <a href="/wiki/Cheek_teeth" title="Cheek teeth">cheek teeth</a> but human-sized <a href="/wiki/Incisor" title="Incisor">incisors</a> and <a href="/wiki/Canine_tooth" title="Canine tooth">canines</a>. The <a href="/wiki/Premolar" title="Premolar">premolars</a> are shaped like <a href="/wiki/Molar_(tooth)" title="Molar (tooth)">molars</a>.<sup id="cite_ref-Cartmill2009_24-0" class="reference"><a href="#cite_note-Cartmill2009-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Tooth_enamel" title="Tooth enamel">enamel</a> thickness on the cheek teeth is relatively on par with that of modern humans, though australopithecine cheek tooth enamel thickens especially at the tips of the cusps, whereas in humans it thickens at the base of the cusps.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p><p><i>P. robustus</i> has a tall face with slight <a href="/wiki/Prognathism" title="Prognathism">prognathism</a> (the jaw jutted out somewhat). The skulls of males have a well-defined <a href="/wiki/Sagittal_crest" title="Sagittal crest">sagittal crest</a> on the midline of the skullcap and inflated <a href="/wiki/Cheek" title="Cheek">cheek</a> bones, which likely supported massive <a href="/wiki/Temporal_muscle" class="mw-redirect" title="Temporal muscle">temporal muscles</a> important in biting. The cheeks project so far from the face that, when in top-view, the nose appears to sit at the bottom of a concavity (a dished face). This displaced the <a href="/wiki/Eye_socket" class="mw-redirect" title="Eye socket">eye sockets</a> forward somewhat, causing a weak brow ridge and receding forehead. The inflated cheeks also would have pushed the <a href="/wiki/Masseter_muscle" title="Masseter muscle">masseter muscle</a> (important in biting down) forward and pushed the tooth rows back, which would have created a higher <a href="/wiki/Bite_force" class="mw-redirect" title="Bite force">bite force</a> on the premolars. The ramus of the jawbone, which connects the lower jaw to the upper jaw, is tall, which would have increased <a href="/wiki/Lever_arm" class="mw-redirect" title="Lever arm">lever arm</a> (and thereby, torque) of the masseter and <a href="/wiki/Medial_pterygoid_muscle" title="Medial pterygoid muscle">medial pterygoid</a> muscles (both important in biting down), further increasing bite force.<sup id="cite_ref-Cartmill2009_24-1" class="reference"><a href="#cite_note-Cartmill2009-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>The well-defined sagittal crest and inflated cheeks are absent in the presumed-female skull DNH-7, so Keyser suggested that male <i>P. robustus</i> may have been more heavily built than females (<i>P. robustus</i> was <a href="/wiki/Sexually_dimorphic" class="mw-redirect" title="Sexually dimorphic">sexually dimorphic</a>).<sup id="cite_ref-Keyser2000_26-0" class="reference"><a href="#cite_note-Keyser2000-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> The Drimolen material, being more basal, is comparatively more gracile and consequently probably had a smaller bite force than the younger Swartkrans and Kromdraii <i>P. robustus</i>. The brows of the former also are rounded off rather than squared, and the sagittal crest of the presumed-male DNH 155 is more posteriorly (towards the back of the head) positioned.<sup id="cite_ref-Martin2020_22-2" class="reference"><a href="#cite_note-Martin2020-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>The <a href="/wiki/Semicircular_canals#Posterior_semicircular_canal" title="Semicircular canals">posterior semicircular canals</a> in the <a href="/wiki/Inner_ear" title="Inner ear">inner ear</a> of <a href="/wiki/SK_46" title="SK 46">SK 46</a> and SK 47 are unlike those of the apelike <i>Australopithecus</i> or <i>Homo</i>, suggesting different locomotory and head movement patterns, since inner ear anatomy affects the <a href="/wiki/Vestibular_system" title="Vestibular system">vestibular system</a> (sense of balance). The posterior semicircular canals of modern humans are thought to aid in stabilisation while running, which could mean <i>P. robustus</i> was not an endurance runner.<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> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Paranthropus_robustus_female.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/49/Paranthropus_robustus_female.jpg/220px-Paranthropus_robustus_female.jpg" decoding="async" width="220" height="225" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/49/Paranthropus_robustus_female.jpg/330px-Paranthropus_robustus_female.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/49/Paranthropus_robustus_female.jpg/440px-Paranthropus_robustus_female.jpg 2x" data-file-width="2839" data-file-height="2909" /></a><figcaption>Reconstruction of a female <i>P. robustus</i></figcaption></figure> <div class="mw-heading mw-heading4"><h4 id="Brain">Brain</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=11" title="Edit section: Brain"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Upon describing the species, Broom estimated the fragmentary braincase of TM 1517 as 600<i> </i>cc,<sup id="cite_ref-Broom1938_2-4" class="reference"><a href="#cite_note-Broom1938-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> and he, along with South African anthropologist Gerrit Willem Hendrik Schepers, revised this to 575–680 cc in 1946.<sup id="cite_ref-Clark1947_28-0" class="reference"><a href="#cite_note-Clark1947-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> For comparison, the brain volume of contemporary <i>Homo</i> varied from 500 to 900<i> </i>cc.<sup id="cite_ref-Spoor2015_29-0" class="reference"><a href="#cite_note-Spoor2015-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> A year later, British primatologist <a href="/wiki/Wilfrid_Le_Gros_Clark" title="Wilfrid Le Gros Clark">Wilfrid Le Gros Clark</a> commented that, since only a part of the <a href="/wiki/Temporal_bone" title="Temporal bone">temporal bone</a> on one side is known, brain volume cannot be accurately measured for this specimen.<sup id="cite_ref-Clark1947_28-1" class="reference"><a href="#cite_note-Clark1947-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> In 2001, Polish anthropologist Katarzyna Kaszycka said that Broom quite often artificially inflated brain size in early hominins, and the true value was probably much lower.<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p><p>In 1972, American physical anthropologist <a href="/wiki/Ralph_Holloway" title="Ralph Holloway">Ralph Holloway</a> measured the skullcap SK 1585, which is missing part of the <a href="/wiki/Frontal_bone" title="Frontal bone">frontal bone</a>, and reported a volume of about 530<i> </i>cc. He also noted that, compared to other australopithecines, <i>Paranthropus</i> seems to have had an expanded <a href="/wiki/Cerebellum" title="Cerebellum">cerebellum</a> like <i>Homo</i>, echoing what Tobias said while studying <i>P. boisei</i> skulls in 1967.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> In 2000, American neuroanthropologist <a href="/wiki/Dean_Falk" title="Dean Falk">Dean Falk</a> and colleagues filled in frontal bone anatomy of SK 1585 using the <i>P. boisei</i> specimens KNM-ER 407, <a href="/wiki/OH_5" class="mw-redirect" title="OH 5">OH 5</a>, and KNM-ER 732, and recalculated the brain volume to about 476<i> </i>cc. They stated overall brain anatomy of <i>P. robustus</i> was more like that of non-human apes.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p><p>In 2020, the nearly complete skull DNH 155 was discovered and was measured to have had a brain volume of 450<i> </i>cc.<sup id="cite_ref-Martin2020_22-3" class="reference"><a href="#cite_note-Martin2020-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Blood_vessels">Blood vessels</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=12" title="Edit section: Blood vessels"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1983, while studying SK 1585 (<i>P. robustus</i>) and KNM-ER 407 (<i>P. boisei</i>, which he referred to as <i>robustus</i>), French anthropologist Roger Saban stated that the parietal branch of the <a href="/wiki/Middle_meningeal_artery" title="Middle meningeal artery">middle meningeal artery</a> originated from the posterior branch in <i>P. robustus</i> and <i>P. boisei</i> instead of the anterior branch as in earlier hominins, and considered this a derived characteristic due to increased brain capacity.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> It has since been demonstrated that, at least for <i>P. boisei</i>, the parietal branch could originate from either the anterior or posterior branches, sometimes both in a single specimen on opposite sides of the skull.<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p><p>Regarding the <a href="/wiki/Dural_venous_sinuses" title="Dural venous sinuses">dural venous sinuses</a>, in 1983, Falk and anthropologist Glenn Conroy suggested that, unlike <i>A. africanus</i> or modern humans, all <i>Paranthropus</i> (and <i>A. afarensis</i>) had expanded <a href="/wiki/Occipital_sinus" title="Occipital sinus">occipital</a> and marginal (around the <a href="/wiki/Foramen_magnum" title="Foramen magnum">foramen magnum</a>) sinuses, completely supplanting the <a href="/wiki/Transverse_sinus" class="mw-redirect" title="Transverse sinus">transverse</a> and <a href="/wiki/Sigmoid_sinus" title="Sigmoid sinus">sigmoid sinuses</a>. They suggested the setup would have increased blood flow to the <a href="/wiki/Internal_vertebral_venous_plexuses" title="Internal vertebral venous plexuses">internal vertebral venous plexuses</a> or <a href="/wiki/Internal_jugular_vein" title="Internal jugular vein">internal jugular vein</a>, and was thus related to the reorganisation of the blood vessels supplying the head as an immediate response to <a href="/wiki/Biped" class="mw-redirect" title="Biped">bipedalism</a>, which relaxed as bipedalism became more developed.<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> In 1988, Falk and Tobias demonstrated that early hominins (at least <i>A. africanus</i> and <i>P. boisei</i>) could have both an occipital/marginal and transverse/sigmoid systems concurrently or on opposite halves of the skull.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Torso">Torso</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=13" title="Edit section: Torso"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Few <a href="/wiki/Vertebra" title="Vertebra">vertebrae</a> are assigned to <i>P. robustus</i>. The only thoracolumbar series (<a href="/wiki/Thoracic_vertebrae" title="Thoracic vertebrae">thoracic</a> and <a href="/wiki/Lumbar_vertebrae" title="Lumbar vertebrae">lumbar</a> series) preserved belongs to the juvenile SKW 14002, and either represents the 1st to the 4th lumbar vertebrae, or the 2nd to the 5th. SK 3981 preserves a 12th thoracic vertebra (the last in the series), and a lower lumbar vertebra. The 12th thoracic vertebra is relatively elongated, and the articular surface (where it joins with another vertebra) is <a href="/wiki/Kidney" title="Kidney">kidney</a>-shaped. The T12 is more compressed in height than that of other australopithecines and modern apes.<sup id="cite_ref-Sanders1998_37-0" class="reference"><a href="#cite_note-Sanders1998-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> Modern humans who suffer from <a href="/wiki/Spinal_disc_herniation" title="Spinal disc herniation">spinal disc herniation</a> often have vertebrae that are more similar to those of <a href="/wiki/Chimpanzee" title="Chimpanzee">chimpanzees</a> than healthy humans. Early hominin vertebrae are similar to those of a pathological human, including the only other 12th thoracic vertebra known for <i>P. robustus</i>, the juvenile SK 853. Conversely, SK 3981 is more similar to those of healthy humans, which could be explained as: SK 3981 is abnormal, the vertebrae took on a more humanlike condition with maturity, or one of these specimens is assigned to the wrong species.<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> The shape of the lumbar vertebrae is much more similar to that of <a href="/wiki/Turkana_Boy" title="Turkana Boy">Turkana Boy</a> (<i>H. ergaster</i>/<i>H. erectus</i>) and humans than other australopithecines. The pedicles (which jut out diagonally from the vertebra) of the lower lumbar vertebra are much more robust than in other australopithecines and are within the range of humans, and the transverse processes (which jut out to the sides of the vertebra) indicate powerful <a href="/wiki/Iliolumbar_ligament" title="Iliolumbar ligament">iliolumbar ligaments</a>. These could have bearing on the amount of time spent upright compared to other australopithecines.<sup id="cite_ref-Sanders1998_37-1" class="reference"><a href="#cite_note-Sanders1998-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:DNH_43_ventral.jpg" class="mw-file-description"><img alt="Pelvis in face-view preserving the sacrum and the right ilium and the upper portion of the ischium" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ad/DNH_43_ventral.jpg/220px-DNH_43_ventral.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ad/DNH_43_ventral.jpg/330px-DNH_43_ventral.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ad/DNH_43_ventral.jpg/440px-DNH_43_ventral.jpg 2x" data-file-width="1985" data-file-height="1329" /></a><figcaption>Illustration of the pelvis of DNH 43 (front view)</figcaption></figure> <p>The <a href="/wiki/Pelvis" title="Pelvis">pelvis</a> is similar to the pelvises of <i>A. africanus</i> and <i>A. afarensis</i>, but it has a wider <a href="/wiki/Iliac_blade" class="mw-redirect" title="Iliac blade">iliac blade</a> and smaller <a href="/wiki/Acetabulum" title="Acetabulum">acetabulum</a> and hip joint.<sup id="cite_ref-Wood2000_39-0" class="reference"><a href="#cite_note-Wood2000-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> Like modern humans, the ilium of <i>P. robustus</i> features development of the surface and thickening of the <a href="/wiki/Posterior_superior_iliac_spine" title="Posterior superior iliac spine">posterior superior iliac spine</a>, which are important in stabilising the <a href="/wiki/Sacrum" title="Sacrum">sacrum</a>, and indicates <a href="/wiki/Lumbar_lordosis" class="mw-redirect" title="Lumbar lordosis">lumbar lordosis</a> (curvature of the lumbar vertebrae) and thus bipedalism. The anatomy of the sacrum and the first lumbar vertebra (at least the vertebral arch), preserved in DNH 43, are similar to those of other australopithecines.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> The pelvis seems to indicate a more-or-less humanlike hip joint consistent with bipedalism, though differences in overall pelvic anatomy may indicate <i>P. robustus</i> used different muscles to generate force and perhaps had a different mechanism to direct force up the spine. This is similar to the condition seen in <i>A. africanus</i>. This could potentially indicate the lower limbs had a wider range of motion than those of modern humans.<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Limbs">Limbs</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=14" title="Edit section: Limbs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The distal (lower) <a href="/wiki/Humerus" title="Humerus">humerus</a> of <i>P. robustus</i> falls within the variation of both modern humans and chimps, as the distal humerus is quite similar between humans and chimpanzees.<sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Radius_(bone)" title="Radius (bone)">radius</a> of <i>P. robustus</i> is comparable in form to <i>Australopithecus</i> species. The <a href="/wiki/Wrist" title="Wrist">wrist</a> joint had the same maneuverability as that of modern humans rather than the greater <a href="/wiki/Flexion" class="mw-redirect" title="Flexion">flexion</a> achieved by non-human apes, but the <a href="/wiki/Head_of_radius" title="Head of radius">head of radius</a> (the <a href="/wiki/Elbow" title="Elbow">elbow</a>) seems to have been quite capable of maintaining stability when the forearm was flexed like non-human apes. It is possible this reflects some <a href="/wiki/Arboreal" class="mw-redirect" title="Arboreal">arboreal</a> activity (movement in the trees) as is controversially postulated in other australopithecines.<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> SKX 3602 exhibits robust <a href="/wiki/Radial_styloid_process" title="Radial styloid process">radial styloid processes</a> near the hand which indicate strong <a href="/wiki/Brachioradialis_muscle" class="mw-redirect" title="Brachioradialis muscle">brachioradialis muscles</a> and <a href="/wiki/Extensor_retinaculum_of_the_hand" title="Extensor retinaculum of the hand">extensor retinaculae</a>. Like humans, the finger bones are uncurved and have weaker muscle attachment than non-human apes, though the <a href="/wiki/Proximal_phalanges" class="mw-redirect" title="Proximal phalanges">proximal phalanges</a> are smaller than in humans. The intermediate phalanges are stout and straight like humans, but have stouter bases and better developed flexor impressions. The distal phalanges seem to be essentially humanlike. These could indicate a decreased climbing capacity compared to non-human apes<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> and <i>P. boisei</i>.<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> The <i>P. robustus</i> hand is consistent with a humanlike <a href="/wiki/Precision_grip" class="mw-redirect" title="Precision grip">precision grip</a> which would have made possible the production or usage of tools requiring greater motor functions than <a href="/wiki/Tool_use_by_animals#Primates" class="mw-redirect" title="Tool use by animals">non-human primate tools</a>.<sup id="cite_ref-Susman1988_46-0" class="reference"><a href="#cite_note-Susman1988-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> </p><p>The femur, as in <i>P. boisei</i> and <i>H. habilis</i>, is flattened anteroposteriorly (on the front and back side). This may indicate a walking gait more similar to early hominins than to modern humans (less efficient gait).<sup id="cite_ref-Wood2000_39-1" class="reference"><a href="#cite_note-Wood2000-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> Four femora assigned to <i>P. robustus</i>—SK 19, SK 82, SK 97, and SK 3121—exhibit an apparently high <a href="/wiki/Anisotropy" title="Anisotropy">anisotropic</a> <a href="/wiki/Trabecular_bone" class="mw-redirect" title="Trabecular bone">trabecular bone</a> (at the hip joint) structure, which could indicate reduced mobility of the hip joint compared to non-human apes, and the ability to produce forces consistent with humanlike bipedalism.<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> The <a href="/wiki/Femoral_head" title="Femoral head">femoral head</a> StW 311, which either belongs to <i>P. robustus</i> or early <i>Homo</i>, seems to have habitually been placed in highly flexed positions based on the wearing patterns, which would be consistent with frequent climbing activity. It is unclear if frequent squatting could be a valid alternative interpretation.<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> The textural complexity of the <a href="/wiki/Patella" title="Patella">kneecap</a> SKX 1084, which reflects <a href="/wiki/Cartilage" title="Cartilage">cartilage</a> thickness and thus usage of the knee joint and bipedality, is midway between modern humans and chimps.<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> The <a href="/wiki/First_metatarsal_bone" title="First metatarsal bone">big toe bone</a> of <i>P. robustus</i> is not dextrous, which indicates a humanlike foot posture and range of motion, but the more distal ankle joint would have inhibited the modern human toe-off <a href="/wiki/Bipedal_gait_cycle" title="Bipedal gait cycle">gait cycle</a>. <i>P. robustus</i> and <i>H. habilis</i> may have achieved about the same grade of bipedality.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Size">Size</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=15" title="Edit section: Size"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Broom had noted that the ankle bone and humerus of the holotype TM 1517 were about the same dimensions as that of a modern <a href="/wiki/San_people" title="San people">San</a> woman, and so assumed humanlike proportions in <i>P. robustus</i>. In 1972, Robinson estimated <i>Paranthropus</i> as having been massive. He calculated the humerus-to-femur ratio of <i>P. robustus</i> by using the presumed female humerus of STS 7 and comparing it with the presumed male femur of STS 14. He also had to estimate the length of the humerus using the femur assuming a similar degree of sexual dimorphism between <i>P. robustus</i> and humans. Comparing the ratio to humans, he concluded that <i>P. robustus</i> was a heavily built species with a height of 140–150 cm (4 ft 7 in – 4 ft 11 in) and a weight of 68–91 kg (150–201 lb). Consequently, Robinson had described its locomotory habits as, "a compromise between erectness and facility for quadrupedal climbing." In contrast, he estimated <i>A. africanus</i> (which he called "<i>H.</i>" <i>africanus</i>) to have been 1.2–1.4 m (4–4.5 ft) tall and 18–27 kg (40–60 lb) in weight, and to have also been completely bipedal.<sup id="cite_ref-Wolpoff1974_17-1" class="reference"><a href="#cite_note-Wolpoff1974-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p><p>Robinson's estimation of <i>P. robustus</i> size was soon challenged in 1974 by American palaeontologist <a href="/wiki/Stephen_Jay_Gould" title="Stephen Jay Gould">Stephen Jay Gould</a> and English palaeoanthropologist <a href="/wiki/David_Pilbeam" title="David Pilbeam">David Pilbeam</a>, who guessed from the available skeletal elements a weight of about 40.5 kg (89 lb).<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> Similarly, in 1988, American anthropologist <a href="/wiki/Henry_McHenry_(anthropologist)" title="Henry McHenry (anthropologist)">Henry McHenry</a> reported much lighter weights as well as notable sexual dimorphism for <i>Paranthropus</i>. McHenry plotted body size vs. the cross sectional area of the femoral head for a sample of just humans and a sample with all great apes including humans, and calculated <a href="/wiki/Linear_regression" title="Linear regression">linear regressions</a> for each one. Based on the average of these two regressions, he reported an average weight of 47.1 kg (104 lb) for <i>P. robustus</i> using the specimens SK 82 and SK 97.<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> In 1991, McHenry expanded his sample size, and also estimated the living size of Swartkrans specimens by scaling down the dimensions of an average modern human to meet a preserved leg or foot element (he considered the arm measurements too variable among hominins to give accurate estimates). At Members 1 and 2, about 35% of the <i>P. robustus</i> leg or foot specimens were the same size as those in a 28 kg (62 lb) human, 22% in a 43 kg (95 lb) human, and the remaining 43% bigger than the former but less than a 54 kg (119 lb) human except for KNM‐ER 1464 (an <a href="/wiki/Ankle_bone" class="mw-redirect" title="Ankle bone">ankle bone</a>). At Member 3, all individuals were consistent with a 45 kg (99 lb) human.<sup id="cite_ref-McHenry1991a_53-0" class="reference"><a href="#cite_note-McHenry1991a-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> Smaller adults thus seem to have been more common.<sup id="cite_ref-Braga2017_54-0" class="reference"><a href="#cite_note-Braga2017-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> McHenry also estimated the living height of three <i>P. robustus</i> specimens (male SK 82, male SK 97, and female or subadult SK 3155), by scaling down an average human to meet the estimated size of the preserved femur, as 126 cm (4 ft 2 in), 137 cm (4 ft 6 in), and 110 cm (3 ft 7 in), respectively. Based on just these three, he reported an average height of 132 cm (4 ft 4 in) for <i>P. robustus</i> males and 110 cm (3 ft 7 in) for females.<sup id="cite_ref-McHenry1991b_55-0" class="reference"><a href="#cite_note-McHenry1991b-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> </p><p>In 2001, palaeoanthropologist Randall L. Susman and colleagues, using two recently discovered proximal femoral fragments from Swartkrans, estimated an average of 42 kg (93 lb) for males and 30 kg (66 lb) for females. If these four proximal femur specimens—SK 82, SK 97, SKW 19, and SK 3121—are representative of the entire species, they said that this degree of sexual dimorphism is greater than what is exhibited in <a href="/wiki/Human" title="Human">humans</a> and <a href="/wiki/Chimpanzee" title="Chimpanzee">chimpanzees</a>, but less than <a href="/wiki/Orangutan" title="Orangutan">orangutans</a> and <a href="/wiki/Gorilla" title="Gorilla">gorillas</a>. Female <i>P. robustus</i> were about the same estimated weight as female <i>H. ergaster/H. erectus</i> in Swartkrans, but they estimated male <i>H. ergaster/H. erectus</i> as much bigger at 55 kg (121 lb).<sup id="cite_ref-Susman2001_56-0" class="reference"><a href="#cite_note-Susman2001-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> In 2012, American anthropologist Trenton Holliday, using the same equation as McHenry on three specimens, reported an average of 37 kg (82 lb) with a range of 30–43 kg (66–95 lb).<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> In 2015, biological anthropologist Mark Grabowski and colleagues, using nine specimens, estimated an average of 32.3 kg (71 lb) for males and 24 kg (53 lb) for females.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Palaeobiology">Palaeobiology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=16" title="Edit section: Palaeobiology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Diet">Diet</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=17" title="Edit section: Diet"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:SK_46.jpg" class="mw-file-description"><img alt="Skull in three-quarter view facing left, with the right side of the face missing (though the right dental row is still there)" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/9b/SK_46.jpg/220px-SK_46.jpg" decoding="async" width="220" height="184" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/9b/SK_46.jpg/330px-SK_46.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/9b/SK_46.jpg/440px-SK_46.jpg 2x" data-file-width="2692" data-file-height="2248" /></a><figcaption>Cast of SK 46 skull</figcaption></figure> <p>In 1954, Robinson suggested that the heavily built skull of <i>P. robustus</i> and resultantly exorbitant bite force was indicative of a <a href="/wiki/Generalist_and_specialist_species" title="Generalist and specialist species">specialist</a> diet adapted for frequently cracking hard foods such as nuts. Because of this, the predominant model of <i>Paranthropus</i> extinction for the latter half of the 20th century was that they were unable to adapt to the volatile climate of the Pleistocene, unlike the much more adaptable <i>Homo</i>. Subsequent researchers reinforced this model studying the musculature of the face, dental wearing patterns, and primate ecology.<sup id="cite_ref-Wood2004_59-0" class="reference"><a href="#cite_note-Wood2004-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> In 1981, English anthropologist Alan Walker, while studying the <i>P. boisei</i> skulls KNM-ER 406 and 729, pointed out that bite force is a measure of not only the total pressure exerted but also the surface area of the tooth over which the pressure is being exerted, and <i>Paranthropus</i> teeth are 4–5 times the size of modern human teeth. Because the chewing muscles are arranged the same way, Walker postulated that the heavy build was instead an adaptation to chew a large quantity of food at the same time. He also found that microwearing on 20 <i>P. boisei</i> molar specimens were indistinguishable from patterning recorded in <a href="/wiki/Mandrill" title="Mandrill">mandrills</a>, chimps, and orangutans.<sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> Despite subsequent arguments that <i>Paranthropus</i> were not specialist feeders, the predominant consensus in favour of Robinson's initial model did not change for the remainder of the 20th century.<sup id="cite_ref-Wood2004_59-1" class="reference"><a href="#cite_note-Wood2004-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> </p><p>In 2004, in their review of <i>Paranthropus</i> dietary literature, anthropologists Bernard Wood and David Strait concluded that <i>Paranthropus</i> were most definitely generalist feeders, and that <i>P. robustus</i> was an <a href="/wiki/Omnivore" title="Omnivore">omnivore</a>. They found that the microwear patterns in <i>P. robustus</i> suggest hard food was infrequently consumed, and therefore the heavy build of the skull was only relevant when eating less desirable fallback foods.<sup id="cite_ref-Wood2004_59-2" class="reference"><a href="#cite_note-Wood2004-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> Such a strategy is similar to that used by modern gorillas, which can sustain themselves entirely on lower quality fallback foods year-round, as opposed to lighter built chimpanzees (and presumably gracile australopithecines) which require steady access to high quality foods.<sup id="cite_ref-Laden2005_61-0" class="reference"><a href="#cite_note-Laden2005-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> In 1980, anthropologists Tom Hatley and John Kappelman suggested that early hominins (<a href="/wiki/Convergent_evolution" title="Convergent evolution">convergently</a> with <a href="/wiki/Bear" title="Bear">bears</a> and <a href="/wiki/Pig" title="Pig">pigs</a>) adapted to eating abrasive and calorie-rich underground <a href="/wiki/Storage_organ" title="Storage organ">storage organs</a> (USOs), such as roots and tubers.<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> Since then, hominin exploitation of USOs has gained more support. In 2005, biological anthropologists <a href="/wiki/Greg_Laden" title="Greg Laden">Greg Laden</a> and <a href="/wiki/Richard_Wrangham" title="Richard Wrangham">Richard Wrangham</a> proposed that <i>Paranthropus</i> relied on USOs as a fallback or possibly primary food source, and noted that there may be a correlation between high USO abundance and hominin occupation.<sup id="cite_ref-Laden2005_61-1" class="reference"><a href="#cite_note-Laden2005-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> </p><p>A 2006 <a href="/wiki/Isotope_analysis#Carbon-13" title="Isotope analysis">carbon isotope analysis</a> suggested that <i>P. robustus</i> subsisted on mainly <a href="/wiki/C4_carbon_fixation" title="C4 carbon fixation">C<sub>4</sub> savanna plants</a> or <a href="/wiki/C3_carbon_fixation" title="C3 carbon fixation">C<sub>3</sub> forest plants</a> depending on the season, which could indicate either seasonal shifts in diet or seasonal migration from forest to savanna.<sup id="cite_ref-Sponheimer2006_63-0" class="reference"><a href="#cite_note-Sponheimer2006-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> <i>H. ergaster/H. erectus</i> appears to have consumed about the same proportion of C<sub>3</sub> to C<sub>4</sub> based foods as <i>P. robustus</i>.<sup id="cite_ref-Lee2000_64-0" class="reference"><a href="#cite_note-Lee2000-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> <i>P. robustus</i> likely also commonly cracked hard foods such as seeds or nuts, as it had a moderate tooth-chipping rate (about 12% in a sample of 239 individuals, as opposed to little to none for <i>P. boisei</i>).<sup id="cite_ref-Sponheimer2006_63-1" class="reference"><a href="#cite_note-Sponheimer2006-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-65" class="reference"><a href="#cite_note-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> A high cavity rate could indicate <a href="/wiki/Honey" title="Honey">honey</a> consumption.<sup id="cite_ref-Towle2019_66-0" class="reference"><a href="#cite_note-Towle2019-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> Juvenile <i>P. robustus</i> may have relied more on tubers than adults, given the elevated levels of <a href="/wiki/Strontium" title="Strontium">strontium</a> compared to adults in teeth from Swartkrans Cave, which, in the area, was most likely sourced from tubers. <a href="/wiki/Dentin" title="Dentin">Dentin</a> exposure on juvenile teeth could indicate early weaning, or a more abrasive diet than adults which wore away the <a href="/wiki/Cementum" title="Cementum">cementum</a> and <a href="/wiki/Tooth_enamel" title="Tooth enamel">enamel</a> coatings, or both. It is also possible juveniles were instead less capable of removing grit from dug-up food rather than purposefully seeking out more abrasive foods.<sup id="cite_ref-Williams2015_67-0" class="reference"><a href="#cite_note-Williams2015-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Social_structure">Social structure</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=18" title="Edit section: Social structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Given the marked anatomical and physical differences with modern great apes, there may be no modern analogue for australopithecine societies, so comparisons drawn with modern primates are highly speculative.<sup id="cite_ref-Copeland2011_68-0" class="reference"><a href="#cite_note-Copeland2011-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kaszycka2016_69-0" class="reference"><a href="#cite_note-Kaszycka2016-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> </p><p>In 2007, anthropologist Charles Lockwood and colleagues pointed out that <i>P. robustus</i> appears to have had pronounced <a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">sexual dimorphism</a>, with males notably larger than females. This is commonly correlated with a male-dominated <a href="/wiki/Polygamous" class="mw-redirect" title="Polygamous">polygamous</a> society, such as the harem society of modern forest-dwelling <a href="/wiki/Silverback_gorilla" class="mw-redirect" title="Silverback gorilla">silverback gorillas</a> where one male has exclusive breeding rights to a group of females. Estimated male-female size disparity in <i>P. robustus</i> is comparable to gorillas (based on facial dimensions), and younger males were less robust than older males (delayed maturity is also exhibited in gorillas). Because the majority of sexed <i>P. robustus</i> specimens are male (or at least presumed male), males seem to have had a higher mortality rate than females. In a harem society, males are more likely to be evicted from the group given higher male–male competition over females, and lone males may have been put at a higher risk of predation. By this hypothesis, a female moving out of her birth group may have spent little time alone and transferred immediately to another established group.<sup id="cite_ref-Lockwood2007_70-0" class="reference"><a href="#cite_note-Lockwood2007-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:20070819_Dierenpark_Emmen_(20).JPG" class="mw-file-description"><img alt="A large troop of Hamadrayas baboons. The females are brown and shorthaired, whereas the males are white and longhaired." src="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/20070819_Dierenpark_Emmen_%2820%29.JPG/290px-20070819_Dierenpark_Emmen_%2820%29.JPG" decoding="async" width="290" height="218" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/20070819_Dierenpark_Emmen_%2820%29.JPG/435px-20070819_Dierenpark_Emmen_%2820%29.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/99/20070819_Dierenpark_Emmen_%2820%29.JPG/580px-20070819_Dierenpark_Emmen_%2820%29.JPG 2x" data-file-width="1600" data-file-height="1200" /></a><figcaption><i>P. robustus</i> could have lived in multi-male groups like <a href="/wiki/Hamadryas_baboon" title="Hamadryas baboon">Hamadryas baboons</a> (above troop at <a href="/wiki/Dierenpark_Emmen" title="Dierenpark Emmen">Dierenpark Emmen</a>).<sup id="cite_ref-Kaszycka2016_69-1" class="reference"><a href="#cite_note-Kaszycka2016-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>However, in 2011, palaeoanthropologist Sandi Copeland and colleagues studied the <a href="/wiki/Strontium_isotope" class="mw-redirect" title="Strontium isotope">strontium isotope</a> ratio of <i>P. robustus</i> teeth from the <a href="/wiki/Dolomite_(rock)" title="Dolomite (rock)">dolomite</a> Sterkfontein Valley, and found that like other hominins, but unlike other great apes, <i>P. robustus</i> females were more likely to leave their place of birth (<a href="/wiki/Patrilocal" class="mw-redirect" title="Patrilocal">patrilocal</a>). This discounts the plausibility of a harem society, which would have resulted in a <a href="/wiki/Matrilocal" class="mw-redirect" title="Matrilocal">matrilocal</a> society due to heightened male–male competition. Males did not seem to have ventured very far from the valley, which could either indicate small home ranges, or that they preferred dolomitic landscapes due to perhaps cave abundance or factors related to vegetation growth.<sup id="cite_ref-Copeland2011_68-1" class="reference"><a href="#cite_note-Copeland2011-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> Similarly, in 2016, Polish anthropologist Katarzyna Kaszycka rebutted that, among primates, delayed maturity is also exhibited in the <a href="/wiki/Rhesus_monkey" class="mw-redirect" title="Rhesus monkey">rhesus monkey</a> which has a multi-male society, and may not be an accurate indicator of social structure. If <i>P. robustus</i> preferred a savanna habitat, a multi-male society would have been more conducive in defending the troop from predators in the more exposed environment, much like baboons which live in the savanna. Even in a multi-male society, it is still possible that males were more likely to be evicted, explaining male-skewed mortality with the same mechanism.<sup id="cite_ref-Kaszycka2016_69-2" class="reference"><a href="#cite_note-Kaszycka2016-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> </p><p>In 2017, anthropologist Katharine Balolia and colleagues postulated that, because male non-human great apes have a larger sagittal crest than females (particularly gorillas and orangutans), the crest may be influenced by <a href="/wiki/Sexual_selection" title="Sexual selection">sexual selection</a> in addition to supporting chewing muscles. Further, the size of the sagittal crest (and the <a href="/wiki/Gluteus_muscle" class="mw-redirect" title="Gluteus muscle">gluteus muscles</a>) in male <a href="/wiki/Western_lowland_gorilla" title="Western lowland gorilla">western lowland gorillas</a> has been correlated with reproductive success. Balolia <i>et al.</i> extended their interpretation of the crest to the males of <i>Paranthropus</i> species, with the crest and resultantly larger head (at least in <i>P. boisei</i>) being used for some kind of <a href="/wiki/Display_(zoology)" title="Display (zoology)">display</a>. This contrasts with other primates which flash the typically enlarged canines in <a href="/wiki/Agonistic_display" class="mw-redirect" title="Agonistic display">agonistic display</a> (<i>Paranthropus</i> likely did not do this as the canines are comparatively small), though it is also possible that the crest is only so prominent in male gorillas and orangutans because they require larger temporalis muscles to achieve a wider gape to better display the canines.<sup id="cite_ref-71" class="reference"><a href="#cite_note-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Technology">Technology</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=19" title="Edit section: Technology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Cave sites in the Cradle of Humankind often have stone and bone tools, with the former attributed to early <i>Homo</i> and the latter generally to <i>P. robustus</i>, as bone tools are most abundant when <i>P. robustus</i> remains far outnumber <i>Homo</i> remains. Australopithecine bone technology was first proposed by Dart in the 1950s with what he termed the "<a href="/wiki/Osteodontokeratic_culture" title="Osteodontokeratic culture">osteodontokeratic culture</a>", which he attributed to <i>A. africanus</i> at <a href="/wiki/Makapansgat" title="Makapansgat">Makapansgat</a> dating to 3–2.6 million years ago. These bones are no longer considered to have been tools, and the existence of this culture is not supported. The first probable bone tool was reported by Robinson in 1959 at Sterkfontein Member 5. Excavations led by South African palaeontologist <a href="/wiki/Charles_Kimberlin_Brain" title="Charles Kimberlin Brain">Charles Kimberlin Brain</a> at Swartkrans in the late 1980s and early 1990s recovered 84 similar bone tools, and excavations led by Keyser at Drimolen recovered 23. These tools were all found alongside <a href="/wiki/Acheulean" title="Acheulean">Acheulean</a> <a href="/wiki/Stone_tool" title="Stone tool">stone tools</a>, except for those from Swartkrans Member 1 which bore <a href="/wiki/Oldowan" title="Oldowan">Oldowan</a> stone tools. Thus, there are 108 bone tool specimens from the region in total, and possibly an additional two from Kromdraai B. The two stone tools (either "Developed Oldowan" or "Early Acheulean") from Kromdraai B could possibly be attributed to <i>P. robustus</i>, as <i>Homo</i> has not been confidently identified in this layer, though it is possible that the stone tools were <a href="/wiki/Reworked_fossil" class="mw-redirect" title="Reworked fossil">reworked</a> (moved into the layer after the inhabitants had died). Bone tools may have been used to cut or process vegetation,<sup id="cite_ref-Stammers2018_72-0" class="reference"><a href="#cite_note-Stammers2018-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> process fruits (namely <a href="/wiki/Marula_fruit" class="mw-redirect" title="Marula fruit">marula fruit</a>), strip tree bark,<sup id="cite_ref-Errico2009_73-0" class="reference"><a href="#cite_note-Errico2009-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> or dig up tubers or termites.<sup id="cite_ref-Stammers2018_72-1" class="reference"><a href="#cite_note-Stammers2018-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Errico2009_73-1" class="reference"><a href="#cite_note-Errico2009-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Backwell2001_74-0" class="reference"><a href="#cite_note-Backwell2001-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> The form of <i>P. robustus</i> incisors appears to be intermediate between <i>H. erectus</i> and modern humans, which could possibly mean it did not have to regularly bite off mouthfuls of a large food item due to preparation with simple tools.<sup id="cite_ref-Williams2015_67-1" class="reference"><a href="#cite_note-Williams2015-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> The bone tools were typically sourced from the <a href="/wiki/Diaphysis" title="Diaphysis">shaft</a> of <a href="/wiki/Long_bone" title="Long bone">long bones</a> from medium- to large-sized mammals, but tools sourced from <a href="/wiki/Mandible" title="Mandible">mandibles</a>, <a href="/wiki/Rib" title="Rib">ribs</a>, and horn cores have also been found. They were not manufactured or purposefully shaped for a task, but since they display no weathering, and there is a preference displayed for certain bones, raw materials were likely specifically hand picked. This contrasts with East African bone tools which appear to have been modified and directly cut into specific shapes before using.<sup id="cite_ref-Stammers2018_72-2" class="reference"><a href="#cite_note-Stammers2018-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> </p><p>In 1988, Brain and South African archaeologist A. Sillent analysed the 59,488 bone fragments from Swartkrans Member 3, and found that 270 had been burnt, mainly belonging to medium-sized antelope, but also zebra, warthog, baboon, and <i>P. robustus</i>. They were found across the entire depth of Member 3, so fire was a regular event throughout its deposition. Based on colour and structural changes, they found that 46 were heated to below 300 °C (572 °F), 52 to 300–400 °C (572–752 °F), 45 to 400–500 °C (752–932 °F), and 127 above this. They concluded that these bones were, "the earliest direct evidence of fire use in the fossil record," and compared the temperatures with those achieved by experimental campfires burning <a href="/wiki/Celtis_africana" title="Celtis africana">white stinkwood</a> which commonly grows near the cave. Though some bones had cut marks consistent with butchery, they said it was also possible hominins were making fire to scare away predators or for warmth instead of cooking. Because both <i>P. robustus</i> and <i>H. ergaster</i>/<i>H. erectus</i> were found in the cave, they were unsure which species to attribute the fire to.<sup id="cite_ref-Brain1988_75-0" class="reference"><a href="#cite_note-Brain1988-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> As an alternative to hominin activity, because the bones were not burnt inside the cave, it is possible that they were naturally burnt in cyclically occurring <a href="/wiki/Wildfire" title="Wildfire">wildfires</a> (dry savanna grass as well as possible <a href="/wiki/Guano" title="Guano">guano</a> or plant accumulation in the cave may have left it susceptible to such a scenario), and then washed into what would become Member 3.<sup id="cite_ref-Pickering_76-0" class="reference"><a href="#cite_note-Pickering-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-77" class="reference"><a href="#cite_note-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> The now-earliest claim of fire usage is 1.7 million years ago at <a href="/wiki/Wonderwerk_Cave" title="Wonderwerk Cave">Wonderwerk Cave</a>, South Africa, made by South African archaeologist <a href="/wiki/Peter_Beaumont_(archaeologist)" title="Peter Beaumont (archaeologist)">Peter Beaumont</a> in 2011, which he attributed to <i>H. ergaster</i>/<i>H. erectus</i>.<sup id="cite_ref-Beaumont2011_78-0" class="reference"><a href="#cite_note-Beaumont2011-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Development">Development</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=20" title="Edit section: Development"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:SK11_Global.jpg" class="mw-file-description"><img alt="A tooth inside-view, top-view, and bottom-view" src="//upload.wikimedia.org/wikipedia/commons/thumb/5/54/SK11_Global.jpg/250px-SK11_Global.jpg" decoding="async" width="250" height="85" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/54/SK11_Global.jpg/375px-SK11_Global.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/54/SK11_Global.jpg/500px-SK11_Global.jpg 2x" data-file-width="9000" data-file-height="3056" /></a><figcaption>SKX 11 tooth</figcaption></figure> <p>Australopithecines are generally considered to have had a faster, apelike growth rate than modern humans largely due to dental development trends. Broadly speaking, the emergence of the first permanent molar in early hominins has been variously estimated anywhere from 2.5 to 4.5 years, which all contrast markedly with the modern human average of 5.8 years. The 1st permanent molar of SK 63, which may have died at 3.4–3.7 years of age, possibly erupted at 2.9–3.2 years. In modern apes (including humans), dental development trajectory is strongly correlated with life history and overall growth rate, but it is possible that early hominins simply had a faster dental trajectory but a slower life history due to environmental factors, such as early weaning age as is exemplified in modern <a href="/wiki/Indriidae" title="Indriidae">indriid</a> <a href="/wiki/Lemur" title="Lemur">lemurs</a>.<sup id="cite_ref-Kelley2012_79-0" class="reference"><a href="#cite_note-Kelley2012-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> In TM 1517, fusion of the elements of the distal humerus (at the elbow joint) occurred before the fusion of the elements in the distal big toe phalanx, much like in chimps and <a href="/wiki/Bonobo" title="Bonobo">bonobos</a>, but unlike humans, which could also indicate an apelike growth trajectory.<sup id="cite_ref-Cazenave2019_3-1" class="reference"><a href="#cite_note-Cazenave2019-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>While growing, the front part of the jaw in <i>P. robustus</i> is depository (so it grows) whereas the sides are resorptive (so they recede). For comparison, chimp jaws are generally depository reflecting prognathism, and modern humans resorptive reflecting a flat face. In <i>Paranthropus</i>, this may have functioned to thicken the <a href="/wiki/Palate" title="Palate">palate</a>. Unlike other apes and gracile australopithecines, but like humans, the <a href="/wiki/Premaxilla" title="Premaxilla">premaxillary</a> <a href="/wiki/Suture_(anatomy)" title="Suture (anatomy)">suture</a> between the premaxilla and the <a href="/wiki/Maxilla" title="Maxilla">maxilla</a> (on the palate) formed early in development. At early stages, the <i>P. robustus</i> jawbone was somewhat similar to that of modern humans, but the breadth grew in <i>P. robustus</i>, as to be expected from its incredible robustness in adulthood. By the time the first permanent molar erupts, the <a href="/wiki/Body_of_the_mandible" class="mw-redirect" title="Body of the mandible">body of the mandible</a> and the front jaw broadened, and the <a href="/wiki/Ramus_of_the_mandible" class="mw-redirect" title="Ramus of the mandible">ramus of the mandible</a> elongated, diverging from the modern human trajectory. Because the ramus was so tall, it is suggested that <i>P. robustus</i> experienced more anterior face rotation than modern humans and apes. Growth was most marked between the eruptions of the first and second permanent molars, most notably in terms of the distance from the back of the mouth to the front of the mouth, probably to make room for the massive postcanine teeth. Like humans, jaw robustness decreased with age, though it decreased slower in <i>P. robustus</i>.<sup id="cite_ref-Cofran2014_80-0" class="reference"><a href="#cite_note-Cofran2014-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> Regardless if <i>P. robustus</i> followed a human or non-human ape dental development timeframe, the premolars and molars would have had an accelerated growth rate to achieve their massive size. In contrast, the presence of <a href="/wiki/Perikyma" title="Perikyma">perikymata</a> on the incisors and canines (growth lines which typically are worn away after eruption) could indicate these teeth had a reduced growth rate.<sup id="cite_ref-81" class="reference"><a href="#cite_note-81"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup> The tooth roots of <i>P. robustus</i> molars may have grown at a faster rate than gracile australopithecines; the root length of SK 62's 1st molar, which was reaching emergence from the <a href="/wiki/Dental_alveolus" title="Dental alveolus">dental alveolus</a>, is about 6 mm (0.24 in). In contrast, those of other hominins reach 5–6 mm (0.20–0.24 in) after the tooth has emerged not only from the gums (a later stage of dental development). SK 62's growth trajectory is more similar to that of gorillas, whose roots typically measure 7 mm (0.28 in) when emerging from the gums.<sup id="cite_ref-Kelley2012_79-1" class="reference"><a href="#cite_note-Kelley2012-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> </p><p>Females may have reached skeletal maturity by the time the third molar erupted, but males appear to have continued growing after reaching dental maturity, during which time they become markedly more robust than females (<a href="/wiki/Sexual_bimaturism" title="Sexual bimaturism">sexual bimaturism</a>). Similarly, male gorillas complete dental development about the same time as females, but continue growing for up to 5 or 6 years; and male mandrills complete dental development before females, but continue growing for several years more.<sup id="cite_ref-Lockwood2007_70-1" class="reference"><a href="#cite_note-Lockwood2007-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> It is debated whether or not <i>P. robustus</i> had a defined growth spurt in terms of overall height during adolescence, an event unique to humans among modern apes.<sup id="cite_ref-Cofran2014_80-1" class="reference"><a href="#cite_note-Cofran2014-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Life_history">Life history</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=21" title="Edit section: Life history"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1968, American anthropologist Alan Mann, using dental maturity, stratified <i>P. robustus</i> specimens from Swartkrans into different ages, and found an average of 17.2 years at death (they did not necessarily die from old age), and the oldest specimen was 30–35 years old. He also reported an average of 22.2 years for <i>A. africanus</i>. Using these, he argued these hominins had a humanlike prolonged childhood.<sup id="cite_ref-82" class="reference"><a href="#cite_note-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> In response, in 1971, biologist Kelton McKinley repeated Mann's process with more specimens, and (including <i>P. boisei</i>) reported an average of 18 years. McKinley agreed with Mann that <i>P. robustus</i> may have had a prolonged childhood. McKinley also speculated that <a href="/wiki/Sexual_maturity" title="Sexual maturity">sexual maturity</a> was reached at approximately 11 years because it is about halfway between the averages for chimps (9 years) and humans (13). Based on this, he concluded babies were birthed at intervals of 3 to 4 years using a statistical test to maximise the number of children born.<sup id="cite_ref-83" class="reference"><a href="#cite_note-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> </p><p>In 1972, after estimating a foetal size of 1,230–1,390 g (2.7–3.1 lb) based on an adult female weight of 50 kg (110 lb), anthropologist Walter Leutenegger estimated foetal head size at about 110–160 cc (6.7–9.8 cu in), similar to a chimp.<sup id="cite_ref-84" class="reference"><a href="#cite_note-84"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> In 1973, using this and an equation between foetal head size and gestation (assuming foetal growth rate of 0.6 for all mammals), biologist John Frazer estimated a gestation of 300 days for <i>P. robustus</i>.<sup id="cite_ref-85" class="reference"><a href="#cite_note-85"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> In response, Leutenegger pointed out that apes have highly variable foetal growth rates, and "estimates on gestation periods based on this rate and birth weight are useless."<sup id="cite_ref-86" class="reference"><a href="#cite_note-86"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup> </p><p>In 1985, British biologists <a href="/wiki/Paul_H._Harvey" title="Paul H. Harvey">Paul H. Harvey</a> and <a href="/wiki/Tim_Clutton-Brock" title="Tim Clutton-Brock">Tim Clutton-Brock</a> came up with equations relating body size to life history events for primates, which McHenry applied to australopithecines in 1994. For <i>P. robustus</i>, he reported newborn brain size of 175<i> </i>cc and weight of 1.9 kg (4.2 lb), gestation 7.6 months, weaning after 30.1 months of age, maturation age 9.7 years, breeding age 11.4 years, birth interval 45 months, and lifespan 43.3 years. These roughly aligned with other australopithecines and chimps. However, for chimps, he got strongly inaccurate results when compared to actual data for newborn brain size, weaning age, and birth interval, and for humans all metrics except birth interval.<sup id="cite_ref-87" class="reference"><a href="#cite_note-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Pathology">Pathology</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=22" title="Edit section: Pathology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:SK_57.png" class="mw-file-description"><img alt="A tooth still in the jawbone which has numerous cracks all over, and the bottom left is chipped off. There is a white arrow pointing to a prominent black circle on the upper left tooth" src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cd/SK_57.png/220px-SK_57.png" decoding="async" width="220" height="235" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cd/SK_57.png/330px-SK_57.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cd/SK_57.png/440px-SK_57.png 2x" data-file-width="1757" data-file-height="1874" /></a><figcaption>The left upper 1st molar of SK 57 with <a href="/wiki/Tertiary_dentin" title="Tertiary dentin">tertiary dentin</a> (white arrow)</figcaption></figure> <p>Based on a sample of 402 teeth, <i>P. robustus</i> seems to have had a low incidence rate of about 12–16% for <a href="/wiki/Tertiary_dentin" title="Tertiary dentin">tertiary dentin</a>, which forms to repair tooth damage caused by excessive wearing or dental cavities. This is similar to what was found for <i>A. africanus</i> and <i><a href="/wiki/H._naledi" class="mw-redirect" title="H. naledi">H. naledi</a></i> (all three inhabited the Cradle of Humankind at different points in time). In contrast, chimpanzees have an incidence rate of 47%, and gorillas as much as 90%, probably due to a diet with a much higher content of tough plants.<sup id="cite_ref-88" class="reference"><a href="#cite_note-88"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup> </p><p><i>P. robustus</i> seems to have had notably high rates of <a href="/wiki/Pitting_enamel_hypoplasia" title="Pitting enamel hypoplasia">pitting enamel hypoplasia</a> (PEH), where tooth enamel formation is spotty instead of mostly uniform. In <i>P. robustus</i>, about 47% of <a href="/wiki/Baby_teeth" class="mw-redirect" title="Baby teeth">baby teeth</a> and 14% of adult teeth were affected, in comparison to about 6.7% and 4.3%, respectively, for the combined teeth of <i>A. africanus</i>, <i><a href="/wiki/A._sediba" class="mw-redirect" title="A. sediba">A. sediba</a></i>, early <i>Homo</i>, and <i>H. naledi</i>. The condition of these holes covering the entire tooth is consistent with the modern human ailment <a href="/wiki/Amelogenesis_imperfecta" title="Amelogenesis imperfecta">amelogenesis imperfecta</a>. Since circular holes in enamel coverage are uniform in size, only present on the <a href="/wiki/Molar_teeth" class="mw-redirect" title="Molar teeth">molar teeth</a>, and have the same severity across individuals, the PEH may have been a genetic condition. It is possible that the <a href="/wiki/Coding_region" title="Coding region">coding region</a> concerned with thickening enamel also increased the risk of developing PEH.<sup id="cite_ref-89" class="reference"><a href="#cite_note-89"><span class="cite-bracket">[</span>89<span class="cite-bracket">]</span></a></sup> </p><p>As many as four <i>P. robustus</i> individuals have been identified as having had dental <a href="/wiki/Tooth_decay" title="Tooth decay">cavities</a>, indicating a rate similar to non-agricultural modern humans (1–5%). This is odd as <i>P. robustus</i> is thought to have had a diet high in gritty foods, and gritty foods should decrease cavity incidence rate, so <i>P. robustus</i> may have often consumed high-sugar cavity-causing foods. PEH may have also increased susceptibility to cavities.<sup id="cite_ref-90" class="reference"><a href="#cite_note-90"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup> A molar from Drimolen showed a cavity on the tooth root, a rare occurrence in fossil great apes. In order for cavity-creating bacteria to reach this area, the individual would have also presented either <a href="/wiki/Alveolar_process#Alveolar_bone_loss" title="Alveolar process">alveolar resportion</a>, which is commonly associated with <a href="/wiki/Gum_disease" class="mw-redirect" title="Gum disease">gum disease</a>; or super-eruption of the tooth which occurs when it becomes worn down and has to erupt a bit more in order to maintain a proper bite, exposing the root in the process. The latter is most likely, and the exposed root seems to have caused <a href="/wiki/Hypercementosis" title="Hypercementosis">hypercementosis</a> to anchor the tooth in place. The cavity seems to have been healing, possibly due to a change in diet or <a href="/wiki/Oral_microbiology" title="Oral microbiology">mouth microbiome</a>, or the loss of the adjacent molar.<sup id="cite_ref-Towle2019_66-1" class="reference"><a href="#cite_note-Towle2019-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> </p><p>In a sample of 15 <i>P. robustus</i> specimens, all of them exhibited mild to moderate <a href="/wiki/Bone_destruction_patterns_in_periodontal_disease" title="Bone destruction patterns in periodontal disease">alveolar bone loss resulting from periodontal disease</a> (the wearing away of the bone which supports the teeth due to gum disease). In contrast, in a sample of 10 <i>A. africanus</i> specimens, three exhibited no pathologies of the alveolar bone. Measuring the distance between the alveolar bone and the <a href="/wiki/Cementoenamel_junction" title="Cementoenamel junction">cementoenamel junction</a>, <i>P. robustus</i> possibly suffered from a higher rate of <a href="/wiki/Tooth_mobility" title="Tooth mobility">tooth-attachment loss</a>, unless <i>P. robustus</i> had a higher cervical height (the slightly narrowed area where the crown meets the root) in which case these two species had the same rate of tooth-attachment loss. If the former is correct, then the difference may be due to different dietary habits, chewing strategies, more pathogenic mouth microflora in <i>P. robustus</i>, or some immunological difference which made <i>P. robustus</i> somewhat more susceptible to gum disease.<sup id="cite_ref-91" class="reference"><a href="#cite_note-91"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup> </p><p>While removing the <a href="/wiki/Matrix_(geology)" title="Matrix (geology)">matrix</a> encapsulating TM 1517, Schepers noted a large rock, which would have weighed 75 g (2.6 oz), which had driven itself into the braincase through the <a href="/wiki/Parietal_bone" title="Parietal bone">parietal bone</a>. He considered this evidence that another individual had killed TM 1517 by launching the rock as a projectile in either defense or attack, but the most parsimonious explanation is that the rock was deposited during the fossilisation process after TM 1517 had died. In 1961, science writer <a href="/wiki/Robert_Ardrey" title="Robert Ardrey">Robert Ardrey</a> noted two small holes about 2.5 cm (an inch) apart on the child skullcap SK 54, and believed this individual had been killed by being struck twice on the head in an assault; in 1970, Brain reinterpreted this as evidence of a <a href="/wiki/Leopard_attack" title="Leopard attack">leopard attack</a>.<sup id="cite_ref-92" class="reference"><a href="#cite_note-92"><span class="cite-bracket">[</span>92<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Palaeoecology">Palaeoecology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=23" title="Edit section: Palaeoecology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The Pleistocene Cradle of Humankind was mainly dominated by the <a href="/wiki/Springbok" title="Springbok">springbok</a> <i>Antidorcas recki</i>, but other antelope, <a href="/wiki/Giraffidae" title="Giraffidae">giraffes</a>, and <a href="/wiki/Elephantidae" title="Elephantidae">elephants</a> were also seemingly abundant megafauna. The carnivore assemblage comprises the <a href="/wiki/Sabertoothed_cat" class="mw-redirect" title="Sabertoothed cat">sabertoothed cats</a> <i><a href="/wiki/Dinofelis" title="Dinofelis">Dinofelis</a> spp.</i> and <i><a href="/wiki/Megantereon" title="Megantereon">Megantereon</a> spp.</i>, and the hyena <i><a href="/w/index.php?title=Lycyaenops_silberbergi&action=edit&redlink=1" class="new" title="Lycyaenops silberbergi (page does not exist)">Lycyaenops silberbergi</a></i>. Overall, the animal assemblage of the region broadly indicates a mixed, open-to-closed landscape featuring perhaps <a href="/wiki/Montane_grasslands_and_shrublands" title="Montane grasslands and shrublands">montane grasslands and shrublands</a>.<sup id="cite_ref-Adams_93-0" class="reference"><a href="#cite_note-Adams-93"><span class="cite-bracket">[</span>93<span class="cite-bracket">]</span></a></sup> Australopithecines and early <i>Homo</i> likely preferred cooler conditions than later <i>Homo</i>, as there are no australopithecine sites that were below 1,000 m (3,300 ft) in elevation at the time of deposition. This would mean that, like chimps, they often inhabited areas with an average diurnal temperature of 25 °C (77 °F), dropping to 10 or 5 °C (50 or 41 °F) at night.<sup id="cite_ref-David2016_94-0" class="reference"><a href="#cite_note-David2016-94"><span class="cite-bracket">[</span>94<span class="cite-bracket">]</span></a></sup> </p><p><i>P. robustus</i> also cohabited the Cradle of Humankind with <i>H. ergaster</i>/<i>H. erectus</i>.<sup id="cite_ref-Susman2001_56-1" class="reference"><a href="#cite_note-Susman2001-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Pickering_76-1" class="reference"><a href="#cite_note-Pickering-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Herries2020_95-0" class="reference"><a href="#cite_note-Herries2020-95"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup> In addition, these two species resided alongside <i>Australopithecus sediba</i> which is known from about 2 million years ago at <a href="/wiki/Malapa_Fossil_Site,_Cradle_of_Humankind" title="Malapa Fossil Site, Cradle of Humankind">Malapa</a>. The most recent <i>A. africanus</i> specimen, Sts 5, dates to about 2.07 million years ago, around the arrival of <i>P. robustus</i> and <i>H. erectus</i>.<sup id="cite_ref-Herries2020_95-1" class="reference"><a href="#cite_note-Herries2020-95"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup> It has been debated whether or not <i>P. robustus</i> would have had symbiotic, neutral, or antagonist relations with contemporary <i>Australopithecus</i> and <i>Homo</i>.<sup id="cite_ref-96" class="reference"><a href="#cite_note-96"><span class="cite-bracket">[</span>96<span class="cite-bracket">]</span></a></sup> It is possible that South Africa was a <a href="/wiki/Refuge_(ecology)" title="Refuge (ecology)">refugium</a> for <i>Australopithecus</i> until about 2 million years ago with the beginning of major climatic variability and volatility, and potentially competition with <i>Homo</i> and <i>Paranthropus</i>.<sup id="cite_ref-Herries2020_95-2" class="reference"><a href="#cite_note-Herries2020-95"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Fossil-bearing_deposits">Fossil-bearing deposits</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=24" title="Edit section: Fossil-bearing deposits"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1237032888/mw-parser-output/.tmulti"><div class="thumb tmulti tleft"><div class="thumbinner multiimageinner" style="width:358px;max-width:358px"><div class="trow"><div class="tsingle" style="width:162px;max-width:162px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Paranthropus_robustus_Africa.jpg" class="mw-file-description"><img alt="A map of the African continent with a red dot indicating the Cradle of Humankind in the northeast region of South Africa" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ac/Paranthropus_robustus_Africa.jpg/160px-Paranthropus_robustus_Africa.jpg" decoding="async" width="160" height="143" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ac/Paranthropus_robustus_Africa.jpg/240px-Paranthropus_robustus_Africa.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ac/Paranthropus_robustus_Africa.jpg/320px-Paranthropus_robustus_Africa.jpg 2x" data-file-width="2283" data-file-height="2035" /></a></span></div><div class="thumbcaption">Location of the <a href="/wiki/Cradle_of_Humankind" title="Cradle of Humankind">Cradle of Humankind</a></div></div><div class="tsingle" style="width:192px;max-width:192px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Cradle_of_Humankind_plan.jpg" class="mw-file-description"><img alt="A map of the Cradle of Humankind with 15 blue dots indicating various fossil-bearing caves. Paranthropus is known from Kromdraai, Swartkrans, Sterkfontein, Gondolin, Cooper's, and Drimolen Caves" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/20/Cradle_of_Humankind_plan.jpg/190px-Cradle_of_Humankind_plan.jpg" decoding="async" width="190" height="142" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/20/Cradle_of_Humankind_plan.jpg/285px-Cradle_of_Humankind_plan.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/20/Cradle_of_Humankind_plan.jpg/380px-Cradle_of_Humankind_plan.jpg 2x" data-file-width="1730" data-file-height="1291" /></a></span></div><div class="thumbcaption">Fossil-bearing caves (<i>P. robustus</i> is known from 1, 7, and 11–14)</div></div></div></div></div> <dl><dt>Swartkrans</dt></dl> <p>At Swartkrans, <i>P. robustus</i> has been identified from Members 1–3.<sup id="cite_ref-Herries2009_97-0" class="reference"><a href="#cite_note-Herries2009-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> <i>Homo</i> is also found in these deposits, but species identification in Members 1 and 2 is debated between <i>H. ergaster/H. erectus</i>, <i><a href="/wiki/H._habilis" class="mw-redirect" title="H. habilis">H. habilis</a></i>, <i><a href="/wiki/H._rudolfensis" class="mw-redirect" title="H. rudolfensis">H. rudolfensis</a></i>, or multiple species. In total, over 300 <i>P. robustus</i> specimens representing over 130 individuals,<sup id="cite_ref-98" class="reference"><a href="#cite_note-98"><span class="cite-bracket">[</span>98<span class="cite-bracket">]</span></a></sup> predominantly isolated teeth, have been recovered from Swartkrans.<sup id="cite_ref-Constantino2004_6-4" class="reference"><a href="#cite_note-Constantino2004-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>Member 1 and Member 3 have several mammal species in common, making dating by animal remains (<a href="/wiki/Biostratigraphy" title="Biostratigraphy">biostratigraphy</a>) yield overlapping time intervals. Like the East African <a href="/wiki/Olduvai_Gorge" title="Olduvai Gorge">Olduvai</a> Bed I (2.03–1.75 million years ago) and Lower Bed II (1.75–1.70 million years ago), Member 1 preserved the antelope <i><a href="/wiki/Parmularius" title="Parmularius">Parmularius angusticornis</a></i>, the <a href="/wiki/Wildebeest" title="Wildebeest">wildebeest</a>, and the <a href="/wiki/Cape_buffalo" class="mw-redirect" title="Cape buffalo">Cape buffalo</a>. The presence of the <a href="/wiki/Hamadryas_baboon" title="Hamadryas baboon">Hamadryas baboon</a> and <i><a href="/wiki/Dinopithecus" title="Dinopithecus">Dinopithecus</a></i> could mean Members 1–3 were deposited 1.9–1.65 million years ago, though the presence of <a href="/wiki/Warthog" title="Warthog">warthogs</a> suggests some sections of the deposits could date to after 1.5 million years ago. <a href="/wiki/Uranium%E2%80%93lead_dating" title="Uranium–lead dating">Uranium–lead dating</a> reports intervals of 3.21–0.45 million years ago for Member 1 (a very large error range), 1.65–1.07 million years ago for Member 2, and 1.04–0.62 million years ago for Member 3, though more likely the younger side of the estimate; this could mean <i>P. robustus</i> outlived <i>P. boisei</i>.<sup id="cite_ref-Herries2009_97-1" class="reference"><a href="#cite_note-Herries2009-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Cosmogenic_nuclide" title="Cosmogenic nuclide">Cosmogenic nuclide</a> geochronology has reported much more constrained dates of 2.2–1.8 million years ago for Member 1, and 0.96 million years ago for Member 3. No suitable section of Member 2 could be identified to date.<sup id="cite_ref-99" class="reference"><a href="#cite_note-99"><span class="cite-bracket">[</span>99<span class="cite-bracket">]</span></a></sup> </p> <dl><dt>Sterkfontein</dt></dl> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:The_cranium_of_an_Australopithecus_africanus_(3967774317).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/The_cranium_of_an_Australopithecus_africanus_%283967774317%29.jpg/220px-The_cranium_of_an_Australopithecus_africanus_%283967774317%29.jpg" decoding="async" width="220" height="130" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/The_cranium_of_an_Australopithecus_africanus_%283967774317%29.jpg/330px-The_cranium_of_an_Australopithecus_africanus_%283967774317%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fd/The_cranium_of_an_Australopithecus_africanus_%283967774317%29.jpg/440px-The_cranium_of_an_Australopithecus_africanus_%283967774317%29.jpg 2x" data-file-width="650" data-file-height="383" /></a><figcaption><a href="/wiki/Ronald_J._Clarke" title="Ronald J. Clarke">Ronald J. Clarke</a> suggested <a href="/wiki/StW_505" title="StW 505">StW 505</a> (above) is ancestral to <i>P. robustus.</i></figcaption></figure> <p>At Sterkfontein, only the specimens StW 566 and StW 569 are firmly assigned to <i>P. robustus</i>, coming from the "Oldowan infill" dating to 2–1.7 million years ago in a section of Member 5. Earlier members yielded <i>A. africanus</i>. In 1988, palaeoanthropologist <a href="/wiki/Ronald_J._Clarke" title="Ronald J. Clarke">Ronald J. Clarke</a> suggested <a href="/wiki/StW_505" title="StW 505">StW 505</a> from the earlier Member 4 was an ancestor to <i>P. robustus</i>. The specimen is still generally assigned to <i>A. africanus</i>, though the Sterkfontein hominins are known to have an exceedingly wide range of variation, and it is debated whether or not the materials represent multiple species instead of just <i>A. africanus</i>.<sup id="cite_ref-Constantino2004_6-5" class="reference"><a href="#cite_note-Constantino2004-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>The appearance of the baboon <i><a href="/wiki/Theropithecus_oswaldi" title="Theropithecus oswaldi">Theropithecus oswaldi</a></i>, <a href="/wiki/Zebra" title="Zebra">zebras</a>, <a href="/wiki/Lion" title="Lion">lions</a>, <a href="/wiki/Ostrich" title="Ostrich">ostriches</a>, <a href="/wiki/Springhare" class="mw-redirect" title="Springhare">springhares</a>, and several grazing antelope in Member 5 indicates the predominance of open grasslands, but sediment analysis indicates the cave opening was moist during deposition, which could point to a well-watered wooded grassland.<sup id="cite_ref-100" class="reference"><a href="#cite_note-100"><span class="cite-bracket">[</span>100<span class="cite-bracket">]</span></a></sup> </p> <dl><dt>Kromdraai</dt></dl> <p>At Kromdraai, <i>P. robustus</i> has been unearthed at Kromdraai B, and almost all <i>P. robustus</i> fossils discovered in the cave have been recovered from Member 3 (out of 5 members). A total of 31 specimens representing at least 17 individuals have been recovered. The only potential <i>Homo</i> specimen from Member 3 is KB 5223, but its classification is debated.<sup id="cite_ref-Stammers2018_72-3" class="reference"><a href="#cite_note-Stammers2018-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> The ear bones of the juvenile KB 6067 from Member 3 is consistent with that of <i>P. robustus</i>, but the dimensions of the <a href="/wiki/Cochlea" title="Cochlea">cochlea</a> and <a href="/wiki/Oval_window" title="Oval window">oval window</a> better align with the more ancient StW 53 from Sterkfontein Member 4 with undetermined species designation. KB 6067, therefore, may possibly be basal to (more ancient than) other <i>P. robustus</i> specimens, at least those for which ear morphology is known.<sup id="cite_ref-101" class="reference"><a href="#cite_note-101"><span class="cite-bracket">[</span>101<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Paleomagnetism" title="Paleomagnetism">Palaeomagnetism</a> suggests Member 3 may date to 1.78–1.6 million years ago, Member 2 to before 1.78 million years ago, and Member 1 to 2.11–1.95 million years ago.<sup id="cite_ref-Herries2009_97-2" class="reference"><a href="#cite_note-Herries2009-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> </p><p>The animal remains of Kromdraai A suggest deposition occurred anywhere between 1.89 and 1.63 million years ago, and the presence of Oldowan or Achulean tools indicates early <i>Homo</i> activity. The biostratigraphic dating of Kromdraai B is less clear as there are no animal species which are known to have existed in a narrow time interval, and many non-hominin specimens have not been assigned to a species (left at genus level).<sup id="cite_ref-Herries2009_97-3" class="reference"><a href="#cite_note-Herries2009-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> About 75% of mammalian remains other than <i>P. robustus</i> are monkeys, including leaf-eating <a href="/wiki/Colobine_monkey" class="mw-redirect" title="Colobine monkey">colobine monkeys</a>, possibly the earliest record of the Hamadryas baboon, <i><a href="/wiki/Gorgopithecus" title="Gorgopithecus">Gorgopithecus</a></i>, and <i><a href="/w/index.php?title=Papio_angusticeps&action=edit&redlink=1" class="new" title="Papio angusticeps (page does not exist)">Papio angusticeps</a></i> in South Africa. The absence of the baboons <i>T. oswaldi</i> and <i>Dinopithecus</i> could potentially mean Member 3 is older than Sterkfontein Member 5 and Swartkrans Member 1; which, if correct, would invalidate the results from palaeomagnetism, and make these specimens among the oldest representatives of the species.<sup id="cite_ref-Braga2017_54-1" class="reference"><a href="#cite_note-Braga2017-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> </p> <dl><dt>Gondolin Cave</dt></dl> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Sterkfontein_Caves_69.jpg" class="mw-file-description"><img alt="A tooth in three-quarter view, with the root facing the bottom right and the crown facing the upper left" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d8/Sterkfontein_Caves_69.jpg/220px-Sterkfontein_Caves_69.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d8/Sterkfontein_Caves_69.jpg/330px-Sterkfontein_Caves_69.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d8/Sterkfontein_Caves_69.jpg/440px-Sterkfontein_Caves_69.jpg 2x" data-file-width="3043" data-file-height="2027" /></a><figcaption>GDA-2 from <a href="/wiki/Gondolin_Cave" title="Gondolin Cave">Gondolin Cave</a></figcaption></figure> <p>Gondolin Cave has yielded 3 hominin specimens: a right third premolar assigned to early <i>Homo</i> (G14018), a partial left gracile australopithecine first or second molar (GDA-1), and a robust australopithecine second molar (GDA-2). The first hominin specimen (G14018) was found by German palaeontologist <a href="/wiki/Elisabeth_Vrba" title="Elisabeth Vrba">Elisabeth Vrba</a> in 1979, and the other two specimens were recovered in 1997 by, respectively, South African palaeoanthropologist <a href="/wiki/Andre_Keyser" title="Andre Keyser">Andre Keyser</a> and excavator L. Dihasu. GDA-2—measuring 18.8 mm × 18.1 mm (0.74 in × 0.71 in), an area of 340 mm<sup>2</sup> (0.53 sq in)—is exceptionally large for <i>P. robustus</i>, which has a recorded maximum of 290 mm<sup>2</sup> (0.45 sq in). This falls within the range of <i>P. boisei</i> 278–378 mm<sup>2</sup> (0.431–0.586 sq in), so the discoverers assigned it to an indeterminate species of <i>Paranthropus</i> rather than <i>P. robustus</i>.<sup id="cite_ref-102" class="reference"><a href="#cite_note-102"><span class="cite-bracket">[</span>102<span class="cite-bracket">]</span></a></sup> </p><p>GDA-2 was found alongside the pig <i><a href="/wiki/Metridiochoerus_andrewsi" class="mw-redirect" title="Metridiochoerus andrewsi">Metridiochoerus andrewsi</a></i>, which means the tooth must be 1.9–1.5 million years old.<sup id="cite_ref-Herries2009_97-4" class="reference"><a href="#cite_note-Herries2009-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> Using this and palaeomagnetism, it may date to roughly 1.8 million years ago.<sup id="cite_ref-Herries2009_97-5" class="reference"><a href="#cite_note-Herries2009-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> </p> <dl><dt>Cooper's Cave</dt></dl> <p>Cooper's Cave was first reported to yield <i>P. robustus</i> remains in 2000 by South African palaeoanthropologists Christine Steininger and <a href="/wiki/Lee_Rogers_Berger" class="mw-redirect" title="Lee Rogers Berger">Lee Rogers Berger</a>. Specimens include a crushed partial right face (COB 101), three isolated teeth, a juvenile jawbone, and several skull fragments.<sup id="cite_ref-Herries2009_97-6" class="reference"><a href="#cite_note-Herries2009-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> </p><p>The animal remains in the hominin-bearing deposit are similar to those of Swartkrans and Kromdraai A, so the Cooper's Cave deposits may date to 1.87–1.56 million years ago.<sup id="cite_ref-Herries2009_97-7" class="reference"><a href="#cite_note-Herries2009-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> </p> <dl><dt>Drimolen Cave</dt></dl> <p>Drimolen Cave was first discovered to have yielded hominin remains by Keyser in 1992, who, in eight years, oversaw the recovery of 79 <i>P. robustus</i> specimens.<sup id="cite_ref-Keyser2000_26-1" class="reference"><a href="#cite_note-Keyser2000-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Among these are the most complete <i>P. robustus</i> skulls: the presumed female DNH-7 (which also preserved articulated jawbone with almost all the teeth), and presumed male DNH 155.<sup id="cite_ref-Martin2020_22-4" class="reference"><a href="#cite_note-Martin2020-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> It was also associated with the <i>H. ergaster</i>/<i>H. erectus</i> skull DNH 134.<sup id="cite_ref-Herries2020_95-3" class="reference"><a href="#cite_note-Herries2020-95"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup> The Drimolen material preserves several basal characteristics relative to the Swartkrans and Kromdraai remains (meaning it may be older).<sup id="cite_ref-Martin2020_22-5" class="reference"><a href="#cite_note-Martin2020-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>The site is thought to be roughly 2–1.5 million years old based on animal remains which have also been recovered from Swartkrans Member 1.<sup id="cite_ref-Keyser2000_26-2" class="reference"><a href="#cite_note-Keyser2000-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> The animal assemblage is broadly similar to that of Cooper's Cave, meaning they probably are about the same age.<sup id="cite_ref-Herries2009_97-8" class="reference"><a href="#cite_note-Herries2009-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> In 2020, DNH 152 was palaeomagnetically dated to 2.04–1.95 million years ago, making it the oldest identified <i>P. robustus</i> specimen.<sup id="cite_ref-Herries2020_95-4" class="reference"><a href="#cite_note-Herries2020-95"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup> </p> <table border="0" cellspacing="2" cellpadding="0" align="center" style="font-size:smaller;"> <caption style="font-size:larger;"><b>African <a href="/wiki/Hominin" class="mw-redirect" title="Hominin">hominin</a> timeline</b> <small>(in mya)</small><br /><small><a href="/wiki/Template:African_hominin_timeline#References" title="Template:African hominin timeline">View references</a></small> </caption> <tbody><tr> <td> <div class="timeline-wrapper"><map name="timeline_bzjr3ohria7p49s0i73p1pez4j7lxmu"><area shape="rect" href="/wiki/H._sapiens" coords="510,309,601,330" title="H. sapiens" alt="H. sapiens" /><area shape="rect" href="/wiki/H._naledi" coords="521,293,607,314" title="H. naledi" alt="H. naledi" /><area shape="rect" href="/wiki/H._rhodesiensis" coords="389,278,508,298" title="H. rhodesiensis" alt="H. rhodesiensis" /><area shape="rect" href="/wiki/H._ergaster" coords="283,262,380,283" title="H. ergaster" alt="H. ergaster" /><area shape="rect" href="/wiki/Au._sediba" coords="298,246,390,267" title="Au. sediba" alt="Au. sediba" /><area shape="rect" href="/wiki/P._robustus" coords="283,230,380,251" title="P. robustus" alt="P. robustus" /><area shape="rect" href="/wiki/P._boisei" coords="279,215,365,235" title="P. boisei" alt="P. boisei" /><area shape="rect" href="/wiki/H._rudolfensis" coords="170,199,284,220" title="H. rudolfensis" alt="H. rudolfensis" /><area shape="rect" href="/wiki/H._habilis" coords="236,183,328,204" title="H. habilis" alt="H. habilis" /><area shape="rect" href="/wiki/Au._garhi" coords="254,167,340,188" title="Au. garhi" alt="Au. garhi" /><area shape="rect" href="/wiki/P._aethiopicus" coords="148,152,261,172" title="P. aethiopicus" alt="P. aethiopicus" /><area shape="rect" href="/wiki/LD_350%2D1" coords="235,136,315,157" title="LD 350-1" alt="LD 350-1" /><area shape="rect" href="/wiki/K._platyops" coords="122,120,219,141" title="K. platyops" alt="K. platyops" /><area shape="rect" href="/wiki/Au._bahrelghazali" coords="-4,104,127,125" title="Au. bahrelghazali" alt="Au. bahrelghazali" /><area shape="rect" href="/wiki/Au._deyiremeda" coords="47,89,161,109" title="Au. deyiremeda" alt="Au. deyiremeda" /><area shape="rect" href="/wiki/Au._africanus" coords="40,73,148,94" title="Au. africanus" alt="Au. africanus" /><area shape="rect" href="/wiki/Au._afarensis" coords="13,57,122,78" title="Au. afarensis" alt="Au. afarensis" /><area shape="rect" href="/wiki/Au._anamensis" coords="-22,41,86,62" title="Au. anamensis" alt="Au. anamensis" /><area shape="rect" href="/wiki/Ar._ramidus" coords="-26,26,71,46" title="Ar. ramidus" alt="Ar. ramidus" /><area shape="rect" href="/wiki/Ar._kadabba" coords="77,10,174,31" title="Ar. kadabba" alt="Ar. kadabba" /></map><img usemap="#timeline_bzjr3ohria7p49s0i73p1pez4j7lxmu" src="//upload.wikimedia.org/wikipedia/en/timeline/bzjr3ohria7p49s0i73p1pez4j7lxmu.png" /></div> </td></tr></tbody></table> <p><br /> </p> <div class="mw-heading mw-heading3"><h3 id="Predation">Predation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=25" title="Edit section: Predation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Sterkfontein_Caves_65.jpg" class="mw-file-description"><img alt="A skullcap with two prominent and closely spaced circles at the top" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/20/Sterkfontein_Caves_65.jpg/220px-Sterkfontein_Caves_65.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/20/Sterkfontein_Caves_65.jpg/330px-Sterkfontein_Caves_65.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/20/Sterkfontein_Caves_65.jpg/440px-Sterkfontein_Caves_65.jpg 2x" data-file-width="4566" data-file-height="3043" /></a><figcaption>SK 54 skullcap with two holes probably inflicted by a <a href="/wiki/Leopard_attack" title="Leopard attack">leopard attack</a></figcaption></figure> <p>Australopithecine bones may have accumulated in caves due to large carnivores dragging in carcasses, which was first explored in detail by Brain in his 1981 book <i><a href="/wiki/The_Hunters_or_the_Hunted%3F:_An_Introduction_to_African_Cave_Taphonomy" class="mw-redirect" title="The Hunters or the Hunted?: An Introduction to African Cave Taphonomy">The Hunters or the Hunted?: An Introduction to African Cave Taphonomy</a></i>. The juvenile <i>P. robustus</i> skullcap SK 54 has two puncture marks consistent with the lower canines of the <a href="/wiki/Leopard" title="Leopard">leopard</a> specimen SK 349 from the same deposits. Brain hypothesised that <i>Dinofelis</i> and perhaps also <a href="/wiki/Hunting_hyena" class="mw-redirect" title="Hunting hyena">hunting hyenas</a> specialised on killing australopithecines,<sup id="cite_ref-Brain1983_103-0" class="reference"><a href="#cite_note-Brain1983-103"><span class="cite-bracket">[</span>103<span class="cite-bracket">]</span></a></sup> but carbon isotope analysis indicates these species predominantly ate large grazers, while the leopard, the sabertoothed <i>Megantereon</i>, and the <a href="/wiki/Spotted_hyena" title="Spotted hyena">spotted hyena</a> were more likely to have regularly consumed <i>P. robustus</i>.<sup id="cite_ref-104" class="reference"><a href="#cite_note-104"><span class="cite-bracket">[</span>104<span class="cite-bracket">]</span></a></sup> Brain was unsure if these predators actively sought them out and brought them back to the cave den to eat, or inhabited deeper recesses of caves and ambushed them when they entered. Modern-day baboons in this region often shelter in <a href="/wiki/Sinkhole" title="Sinkhole">sinkholes</a> especially on cold winter nights, though Brain proposed that australopithecines seasonally migrated out of the <a href="/wiki/Highveld" title="Highveld">Highveld</a> and into the warmer <a href="/wiki/Bushveld" title="Bushveld">Bushveld</a>, only taking up cave shelters in spring and autumn.<sup id="cite_ref-Brain1983_103-1" class="reference"><a href="#cite_note-Brain1983-103"><span class="cite-bracket">[</span>103<span class="cite-bracket">]</span></a></sup> </p><p>As an antipredator behaviour, baboons often associate themselves with medium-to-large herbivores, most notably <a href="/wiki/Impala" title="Impala">impalas</a>, and it is possible that <i>P. robustus</i> as well as other early hominins which lived in open environments did so also, given they are typically associated with an abundance of medium-to-large bovid and horse remains.<sup id="cite_ref-105" class="reference"><a href="#cite_note-105"><span class="cite-bracket">[</span>105<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Extinction">Extinction</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=26" title="Edit section: Extinction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Though <i>P. robustus</i> was a rather hardy species with a tolerance for environmental variability, it seems to have preferred wooded environments, and similarly most <i>P. robustus</i> remains date to a wet period in South Africa 2–1.75 million years ago conducive to such biomes. The extinction of <i>P. robustus</i> coincided with the <a href="/wiki/Mid-Pleistocene_Transition" title="Mid-Pleistocene Transition">Mid-Pleistocene Transition</a>, and the doubling of glacial cycle duration. During glacial events, with more ice locked up at the poles, the <a href="/wiki/Tropical_rain_belt" title="Tropical rain belt">tropical rain belt</a> contracted towards the equator, subsequently causing the retreat of wetland and woodland environments. Before the transition, <i>P. robustus</i> populations possibly contracted to certain wooded refuge zones over 21,000-year cycles, becoming regionally extinct in certain areas until the wet cycle whereupon it would repopulate those zones. The continual prolonging of dry cycles may have caused its extinction, with the last occurrence in the fossil record 1–0.6 million years ago (though more likely 0.9 million years ago). <i>Homo</i> possibly was able to survive by inhabiting a much larger geographical range, more likely to find a suitable refuge area during unfavourable climate swings.<sup id="cite_ref-Caley2018_106-0" class="reference"><a href="#cite_note-Caley2018-106"><span class="cite-bracket">[</span>106<span class="cite-bracket">]</span></a></sup> </p><p>However, the geographical range of <i>P. robustus</i> in the fossil record is roughly 500 km<sup>2</sup> (190 sq mi), whereas the <a href="/wiki/Critically_endangered" class="mw-redirect" title="Critically endangered">critically endangered</a> <a href="/wiki/Eastern_gorilla" title="Eastern gorilla">eastern gorilla</a> (with the smallest range of any African ape) inhabits 70,000 km<sup>2</sup> (27,000 sq mi), the critically endangered <a href="/wiki/Western_gorilla" title="Western gorilla">western gorilla</a> 700,000 km<sup>2</sup> (270,000 sq mi), and the <a href="/wiki/Endangered" class="mw-redirect" title="Endangered">endangered</a> chimpanzee 2.6 million km<sup>2</sup> (1 million sq mi). Therefore, fossil distribution very unlikely represents the true range of the species; consequently, <i>P. robustus</i> possibly went extinct much more recently somewhere other than the Cradle of Humankind (<a href="/wiki/Signor%E2%80%93Lipps_effect" title="Signor–Lipps effect">Signor–Lipps effect</a>).<sup id="cite_ref-Caley2018_106-1" class="reference"><a href="#cite_note-Caley2018-106"><span class="cite-bracket">[</span>106<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=Paranthropus_robustus&action=edit&section=27" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 25em;"> <ul><li><a href="/wiki/African_archaeology" title="African archaeology">African archaeology</a></li> <li><a href="/wiki/Australopithecus_africanus" title="Australopithecus africanus"><i>Australopithecus africanus</i></a> – Extinct hominid from South Africa</li> <li><a href="/wiki/Australopithecus_sediba" title="Australopithecus sediba"><i>Australopithecus sediba</i></a> – Two-million-year-old hominin from the Cradle of Humankind</li> <li><a href="/wiki/Homo_ergaster" title="Homo ergaster"><i>Homo ergaster</i></a> – Extinct species or subspecies of archaic human</li> <li><a href="/wiki/Homo_habilis" title="Homo habilis"><i>Homo habilis</i></a> – Archaic human species from 2.8 to 1.65 mya</li> <li><a href="/wiki/Homo_naledi" title="Homo naledi"><i>Homo naledi</i></a> – South African archaic human species</li> <li><a href="/wiki/Homo_rudolfensis" title="Homo rudolfensis"><i>Homo rudolfensis</i></a> – Extinct hominin from the Early Pleistocene of East Africa</li> <li><a href="/wiki/Paranthropus_aethiopicus" title="Paranthropus aethiopicus"><i>Paranthropus aethiopicus</i></a> – Extinct species of hominin of East Africa</li> <li><a href="/wiki/Paranthropus_boisei" title="Paranthropus boisei"><i>Paranthropus boisei</i></a> – Extinct species of hominin of East Africa</li></ul></div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=28" 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-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFWoodDohertyBoyle2020" class="citation web cs1">Wood, Bernard; Doherty, Dandy; Boyle, Eve (2020-05-29). <a rel="nofollow" class="external text" href="https://oxfordre.com/anthropology/view/10.1093/acrefore/9780190854584.001.0001/acrefore-9780190854584-e-194">"Hominin Taxic Diversity"</a>. <i>Oxford Research Encyclopedia of Anthropology</i>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Facrefore%2F9780190854584.013.194">10.1093/acrefore/9780190854584.013.194</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a 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W. H. (1946). <i>The South African Fossil Ape-Men - The Australopithecinae</i>. The Transvaal Museum.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+South+African+Fossil+Ape-Men+-+The+Australopithecinae&rft.pub=The+Transvaal+Museum&rft.date=1946&rft.aulast=Broom&rft.aufirst=R.&rft.au=Schepers%2C+G.+W.+H.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AParanthropus+robustus" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRobinson1972" class="citation book cs1"><a href="/wiki/John_Talbot_Robinson" title="John Talbot Robinson">Robinson, J. T.</a> (1972). <i>Early Hominid Posture and Locomotion</i>. University of Chicago Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-226-72230-6" title="Special:BookSources/978-0-226-72230-6"><bdi>978-0-226-72230-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Early+Hominid+Posture+and+Locomotion&rft.pub=University+of+Chicago+Press&rft.date=1972&rft.isbn=978-0-226-72230-6&rft.aulast=Robinson&rft.aufirst=J.+T.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AParanthropus+robustus" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGrine1988" class="citation book cs1">Grine, F. E. (1988). <i>Evolutionary History of the "Robust" Australopithecines</i>. Aldine de Gruyter. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-202-02031-0" title="Special:BookSources/978-0-202-02031-0"><bdi>978-0-202-02031-0</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Evolutionary+History+of+the+%22Robust%22+Australopithecines&rft.pub=Aldine+de+Gruyter&rft.date=1988&rft.isbn=978-0-202-02031-0&rft.aulast=Grine&rft.aufirst=F.+E.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AParanthropus+robustus" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBragáThackeray2017" class="citation book cs1">Bragá, J.; Thackeray, J. F. (2017). <i>Kromdraai: A Birthplace of Paranthropus in the Cradle of Humankind</i>. African Sun Media. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-928355-06-9" title="Special:BookSources/978-1-928355-06-9"><bdi>978-1-928355-06-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Kromdraai%3A+A+Birthplace+of+Paranthropus+in+the+Cradle+of+Humankind&rft.pub=African+Sun+Media&rft.date=2017&rft.isbn=978-1-928355-06-9&rft.aulast=Brag%C3%A1&rft.aufirst=J.&rft.au=Thackeray%2C+J.+F.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AParanthropus+robustus" class="Z3988"></span></li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paranthropus_robustus&action=edit&section=30" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/45px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/59px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <span style="font-weight: bold; font-style: italic;"><a href="https://commons.wikimedia.org/wiki/Category:Paranthropus_robustus" class="extiw" title="commons:Category:Paranthropus robustus">Paranthropus robustus</a></span>.</div></div> </div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1235681985"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1237033735"><div class="side-box side-box-right plainlinks sistersitebox"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1126788409"> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/34px-Wikispecies-logo.svg.png" decoding="async" width="34" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/51px-Wikispecies-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/68px-Wikispecies-logo.svg.png 2x" data-file-width="941" data-file-height="1103" /></span></span></div> <div class="side-box-text plainlist"><a href="/wiki/Wikispecies" title="Wikispecies">Wikispecies</a> has information related to <i><b><a href="https://species.wikimedia.org/wiki/Special:Search/Paranthropus_robustus" class="extiw" title="wikispecies:Special:Search/Paranthropus robustus">Paranthropus robustus</a></b></i>.</div></div> </div> <ul><li><a rel="nofollow" class="external text" href="http://johnhawks.net/explainer/laboratory/meet-robustus.html">Meet <i>Australopithecus robustus</i></a> — <a href="/wiki/John_D._Hawks" title="John D. 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Hawks</a>' website</li> <li><a rel="nofollow" class="external text" href="http://humanorigins.si.edu/evidence/human-fossils/species/paranthropus-robustus"><i>Paranthropus robustus</i></a> - The <a href="/wiki/Smithsonian" class="mw-redirect" title="Smithsonian">Smithsonian</a>'s Human Origins Program</li> <li><a rel="nofollow" class="external text" href="http://humanorigins.si.edu/evidence/human-evolution-timeline-interactive">Human Timeline (Interactive)</a> – Smithsonian</li></ul> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1236075235">.mw-parser-output .navbox{box-sizing:border-box;border:1px solid #a2a9b1;width:100%;clear:both;font-size:88%;text-align:center;padding:1px;margin:1em auto 0}.mw-parser-output .navbox .navbox{margin-top:0}.mw-parser-output .navbox+.navbox,.mw-parser-output .navbox+.navbox-styles+.navbox{margin-top:-1px}.mw-parser-output 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.navbox-subgroup .navbox-abovebelow{background-color:#e6e6ff}.mw-parser-output .navbox-even{background-color:#f7f7f7}.mw-parser-output .navbox-odd{background-color:transparent}.mw-parser-output .navbox .hlist td dl,.mw-parser-output .navbox .hlist td ol,.mw-parser-output .navbox .hlist td ul,.mw-parser-output .navbox td.hlist dl,.mw-parser-output .navbox td.hlist ol,.mw-parser-output .navbox td.hlist ul{padding:0.125em 0}.mw-parser-output .navbox .navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Human_evolution" style="padding:3px"><table class="nowraplinks hlist mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2" style="text-align:center;"><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:Human_evolution" title="Template:Human evolution"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Human_evolution" title="Template talk:Human evolution"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Human_evolution" title="Special:EditPage/Template:Human evolution"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Human_evolution" style="font-size:114%;margin:0 4em"><a href="/wiki/Human_evolution" title="Human evolution">Human evolution</a></div></th></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%"><a href="/wiki/Human_taxonomy" title="Human taxonomy">Taxonomy</a><br />(<a href="/wiki/Hominini" title="Hominini">Hominins</a>)</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Most_recent_common_ancestor" title="Most recent common ancestor">Last common ancestors</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/Chimpanzee%E2%80%93human_last_common_ancestor" title="Chimpanzee–human last common ancestor">Chimpanzee–human</a></li> <li><a href="/wiki/Gorilla%E2%80%93human_last_common_ancestor" title="Gorilla–human last common ancestor">Gorilla–human</a></li> <li><a href="/wiki/Orangutan%E2%80%93human_last_common_ancestor" title="Orangutan–human last common ancestor">Orangutan–human</a></li> <li><a href="/wiki/Gibbon%E2%80%93human_last_common_ancestor" title="Gibbon–human last common ancestor">Gibbon–human</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Australopithecine" title="Australopithecine">Australopithecines</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Nakalipithecus" title="Nakalipithecus">Nakalipithecus</a></i></li> <li><i><a href="/wiki/Orrorin" title="Orrorin">Orrorin</a></i></li> <li><i><a href="/wiki/Sahelanthropus" title="Sahelanthropus">Sahelanthropus</a></i></li> <li><i><a href="/wiki/Kenyanthropus" title="Kenyanthropus">Kenyanthropus</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;font-weight:normal;"><i><a href="/wiki/Ardipithecus" title="Ardipithecus">Ardipithecus</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Ardipithecus_kadabba" title="Ardipithecus kadabba">A. kadabba</a></i></li> <li><i><a href="/wiki/Ardipithecus_ramidus" title="Ardipithecus ramidus">A. ramidus</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;font-weight:normal;"><i><a href="/wiki/Australopithecus" title="Australopithecus">Australopithecus</a></i></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Australopithecus_afarensis" title="Australopithecus afarensis">A. afarensis</a></i></li> <li><i><a href="/wiki/Australopithecus_africanus" title="Australopithecus africanus">A. africanus</a></i></li> <li><i><a href="/wiki/Australopithecus_anamensis" title="Australopithecus anamensis">A. anamensis</a></i></li> <li><i><a href="/wiki/Australopithecus_bahrelghazali" title="Australopithecus bahrelghazali">A. bahrelghazali</a></i></li> <li><i><a href="/wiki/Australopithecus_deyiremeda" title="Australopithecus deyiremeda">A. deyiremeda</a></i></li> <li><i><a href="/wiki/Australopithecus_garhi" title="Australopithecus garhi">A. garhi</a></i></li> <li><i><a href="/wiki/Australopithecus_sediba" title="Australopithecus sediba">A. sediba</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;font-weight:normal;"><i><a href="/wiki/Paranthropus" title="Paranthropus">Paranthropus</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Paranthropus_aethiopicus" title="Paranthropus aethiopicus">P. aethiopicus</a></i></li> <li><i><a href="/wiki/Paranthropus_boisei" title="Paranthropus boisei">P. boisei</a></i></li> <li><i><a class="mw-selflink selflink">P. robustus</a></i></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Homo" title="Homo">Humans and<br />proto-humans<br />(<i>Homo</i>)</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;font-weight:normal;">Proto-humans</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Homo_gautengensis" title="Homo gautengensis">H. gautengensis</a></i> (?)</li> <li><i><a href="/wiki/Homo_habilis" title="Homo habilis">H. habilis</a></i></li> <li><i><a href="/wiki/Homo_naledi" title="Homo naledi">H. naledi</a></i></li> <li><i><a href="/wiki/Homo_rudolfensis" title="Homo rudolfensis">H. rudolfensis</a></i> (?)</li> <li><i><a href="/wiki/Penghu_1" title="Penghu 1">H. tsaichangensis</a></i> (?)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;font-weight:normal;"><i><a href="/wiki/Homo_erectus" title="Homo erectus">Homo erectus</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Java_Man" title="Java Man">H. e. erectus</a></i></li> <li><i><a href="/wiki/Dmanisi_hominins" title="Dmanisi hominins">H. e. georgicus</a></i></li> <li><i><a href="/wiki/Lantian_Man" title="Lantian Man">H. e. lantianensis</a></i></li> <li><i><a href="/wiki/Nanjing_Man" title="Nanjing Man">H. e. nankinensis</a></i></li> <li><i><a href="/wiki/Peking_Man" title="Peking Man">H. e. pekinensis</a></i></li> <li><i><a href="/wiki/Solo_Man" title="Solo Man">H. e. soloensis</a></i></li> <li><i><a href="/wiki/Tautavel_Man" title="Tautavel Man">H. e. tautavelensis</a></i></li> <li><i><a href="/wiki/Yuanmou_Man" title="Yuanmou Man">H. e. yuanmouensis</a></i></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;font-weight:normal;"><a href="/wiki/Archaic_humans" title="Archaic humans">Archaic humans</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><i><a href="/wiki/Homo_antecessor" title="Homo antecessor">H. antecessor</a></i></li> <li><a href="/wiki/Denisovan" title="Denisovan">Denisovans</a></li> <li><i><a href="/wiki/Homo_ergaster" title="Homo ergaster">H. ergaster</a></i> (?)</li> <li><i><a href="/wiki/Homo_floresiensis" title="Homo floresiensis">H. floresiensis</a></i></li> <li><i><a href="/wiki/Homo_heidelbergensis" title="Homo heidelbergensis">H. heidelbergensis</a></i></li> <li><i><a href="/wiki/Homo_longi" title="Homo longi">H. longi</a></i> (?)</li> <li><i><a href="/wiki/Homo_luzonensis" title="Homo luzonensis">H. luzonensis</a></i></li> <li><i><a href="/wiki/Neanderthal" title="Neanderthal">H. neanderthalensis</a></i></li> <li><i><a href="/wiki/Homo_rhodesiensis" title="Homo rhodesiensis">H. rhodesiensis</a></i> (?)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;font-weight:normal;"><a href="/wiki/Human" title="Human">Modern humans</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Homo_sapiens" scope="row" class="navbox-group" style="width:1%;font-weight:normal;"><i><a href="/wiki/Human" title="Human">Homo sapiens</a></i></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/Early_modern_human" title="Early modern human"><i>H. s. sapiens</i> (archaic homo sapiens, anatomically modern humans)</a></li> <li><a href="/wiki/Jebel_Irhoud" title="Jebel Irhoud">Jebel Irhoud</a></li> <li><i><a href="/wiki/Herto_Man" title="Herto Man">H. s. idaltu</a></i></li> <li><a href="/wiki/Cro-Magnon" title="Cro-Magnon">Cro-Magnon</a></li> <li><a href="/wiki/Manot_1" title="Manot 1">Manot people</a></li> <li><a href="/wiki/Tam_Pa_Ling_Cave" title="Tam Pa Ling Cave">Tam Pa Ling</a></li> <li><a href="/wiki/Red_Deer_Cave_people" title="Red Deer Cave people">Red Deer Cave people</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%"><a href="/wiki/Human_evolution" title="Human evolution">Ancestors</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><i><a href="/wiki/Homo_habilis" title="Homo habilis">Homo habilis</a></i> → <i><a href="/wiki/Homo_ergaster" title="Homo ergaster">Homo ergaster</a></i> / <i><a href="/wiki/Homo_erectus" title="Homo erectus">Homo erectus</a></i> (→ <i><a href="/wiki/Homo_antecessor" title="Homo antecessor">Homo antecessor</a></i>)<sup>?</sup> → <i><a href="/wiki/Homo_heidelbergensis" title="Homo heidelbergensis">Homo heidelbergensis</a></i> → <i>archaic Homo sapiens</i> → <i><a href="/wiki/Human" title="Human">Homo sapiens</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%">Models</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">General models</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/Hunting_hypothesis" title="Hunting hypothesis">Hunting</a></li> <li><a href="/wiki/Gathering_hypothesis" title="Gathering hypothesis">Gathering</a></li> <li><a href="/wiki/Endurance_running_hypothesis" title="Endurance running hypothesis">Endurance running</a></li> <li><a href="/wiki/Aquatic_ape_hypothesis" title="Aquatic ape hypothesis">Aquatic ape</a></li> <li><a href="/wiki/Sexual_selection_in_humans" title="Sexual selection in humans">Sexual selection</a></li> <li><a href="/wiki/Self-domestication#In_humans" title="Self-domestication">Self-domestication</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Specific models</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>Diet <ul><li><a href="/wiki/Control_of_fire_by_early_humans#The_cooking_hypothesis" title="Control of fire by early humans">Cooking</a></li> <li><a href="/wiki/Expensive_tissue_hypothesis" title="Expensive tissue hypothesis">Expensive tissue</a></li> <li><a href="/wiki/Aquatic_ape_hypothesis#Diet" title="Aquatic ape hypothesis">Shore-based</a></li></ul></li> <li>Drugs <ul><li><a href="/wiki/Drunken_monkey_hypothesis" title="Drunken monkey hypothesis">Drunken monkey</a></li> <li><a href="/wiki/Evolutionary_models_of_human_drug_use" title="Evolutionary models of human drug use">Evolutionary models of human drug use</a></li> <li><a href="/wiki/Stoned_ape_theory" title="Stoned ape theory">Stoned ape theory</a></li></ul></li> <li>Behavior <ul><li><a href="/wiki/Killer_ape_theory" title="Killer ape theory">Killer ape</a></li> <li><a href="/wiki/Cooperative_eye_hypothesis" title="Cooperative eye hypothesis">Cooperative eye</a></li></ul></li> <li>Life history <ul><li><a href="/wiki/Grandmother_hypothesis" title="Grandmother hypothesis">Grandmother</a></li> <li><a href="/wiki/Patriarch_hypothesis" title="Patriarch hypothesis">Patriarch</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Topics<br /></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_bipedalism_in_humans" class="mw-redirect" title="Evolution of bipedalism in humans">Bipedalism</a></li> <li><a href="/wiki/Human_skeletal_changes_due_to_bipedalism" title="Human skeletal changes due to bipedalism">Skeleton</a></li> <li><a href="/wiki/Muscular_evolution_in_humans" title="Muscular evolution in humans">Muscles</a></li> <li><a href="/wiki/Evolution_of_skin_color" class="mw-redirect" title="Evolution of skin color">Skin color</a></li> <li><a href="/wiki/Evolution_of_hair" class="mw-redirect" title="Evolution of hair">Hair</a></li> <li><a href="/wiki/Cold_and_heat_adaptations_in_humans" title="Cold and heat adaptations in humans">Thermoregulation</a></li> <li><a href="/wiki/Origin_of_speech" title="Origin of speech">Speech</a></li> <li><a href="/wiki/Origin_of_language" title="Origin of language">Language</a></li> <li><a href="/wiki/Evolution_of_human_intelligence" title="Evolution of human intelligence">Intelligence</a></li> <li><a href="/wiki/Sexual_division_of_labour#Hypotheses_for_the_evolutionary_origins_of_SDL" title="Sexual division of labour">Gender roles</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Origin_of_modern_humans" class="mw-redirect" title="Origin of modern humans">Origin of modern humans</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/Recent_African_origin_of_modern_humans" title="Recent African origin of modern humans">Recent African origin</a></li> <li><a href="/wiki/Multiregional_origin_of_modern_humans" title="Multiregional origin of modern humans">Multiregional origin</a></li> <li><a href="/wiki/Interbreeding_between_archaic_and_modern_humans" title="Interbreeding between archaic and modern humans">Archaic admixture</a></li> <li><a href="/wiki/Behavioral_modernity" title="Behavioral modernity">Behavioral modernity</a></li> <li><a href="/wiki/Early_human_migrations" title="Early human migrations">Early migrations</a></li> <li><a href="/wiki/Recent_human_evolution" title="Recent human evolution">Recent evolution</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%">Timelines</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/Timeline_of_human_evolution" title="Timeline of human evolution">Human evolution</a></li> <li><a href="/wiki/Timeline_of_prehistory" title="Timeline of prehistory">Human prehistory</a></li> <li><a href="/wiki/Template:Human_timeline" title="Template:Human timeline">Human timeline</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%">Others</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/Category:Human_evolution_theorists" title="Category:Human evolution theorists">Theorists</a></li> <li><a href="/wiki/Category:Human_evolution_books" title="Category:Human evolution books">Books</a></li> <li><a href="/wiki/List_of_human_evolution_fossils" title="List of human evolution fossils">Fossils</a></li> <li><a href="/wiki/Evolutionary_anthropology" title="Evolutionary anthropology">Evolutionary anthropology</a> <ul><li><a href="/wiki/Paleoanthropology" title="Paleoanthropology">Paleoanthropology</a></li></ul></li> <li><a href="/wiki/Human_evolutionary_developmental_biology" title="Human evolutionary developmental biology">Human evolutionary developmental biology</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="text-align:center;"><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, 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style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Taxon_identifiers" style="font-size:114%;margin:0 4em"><a href="/wiki/Help:Taxon_identifiers" title="Help:Taxon identifiers">Taxon identifiers</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: left;"><i>Paranthropus robustus</i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><span style="white-space:nowrap;"><a href="/wiki/Wikidata" title="Wikidata">Wikidata</a>: <span class="uid"><span class="external"><a href="https://www.wikidata.org/wiki/Q310529" class="extiw" title="wikidata:Q310529">Q310529</a></span></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Wikispecies" title="Wikispecies">Wikispecies</a>: <span class="uid"><span class="external"><a href="https://species.wikimedia.org/wiki/Praeanthropus_robustus" class="extiw" title="wikispecies:Praeanthropus robustus">Praeanthropus robustus</a></span></span></span></li> <li><span style="white-space:nowrap;">BioLib: <span class="uid"><a rel="nofollow" class="external text" href="https://www.biolib.cz/en/taxon/id32376">32376</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Encyclopedia_of_Life" title="Encyclopedia of Life">EoL</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://eol.org/pages/4454196">4454196</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Global_Biodiversity_Information_Facility" title="Global Biodiversity Information Facility">GBIF</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.gbif.org/species/4827640">4827640</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Interim_Register_of_Marine_and_Nonmarine_Genera" title="Interim Register of Marine and Nonmarine Genera">IRMNG</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.irmng.org/aphia.php?p=taxdetails&id=11419625">11419625</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Open_Tree_of_Life" title="Open Tree of Life">Open Tree of Life</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://tree.opentreeoflife.org/taxonomy/browse?id=3607708">3607708</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Paleobiology_Database" title="Paleobiology Database">Paleobiology Database</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://paleobiodb.org/classic/basicTaxonInfo?taxon_no=385304">385304</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Paleobiology_Database" title="Paleobiology Database">Paleobiology Database</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://paleobiodb.org/classic/basicTaxonInfo?taxon_no=83069">83069</a></span></span></li></ul> </div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1&useformat=desktop" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" href="https://en.wikipedia.org/w/index.php?title=Paranthropus_robustus&oldid=1254950607">https://en.wikipedia.org/w/index.php?title=Paranthropus_robustus&oldid=1254950607</a>"</div></div> <div id="catlinks" class="catlinks" data-mw="interface"><div id="mw-normal-catlinks" class="mw-normal-catlinks"><a href="/wiki/Help:Category" title="Help:Category">Categories</a>: <ul><li><a href="/wiki/Category:Pleistocene_mammals_of_Africa" title="Category:Pleistocene mammals of Africa">Pleistocene mammals of Africa</a></li><li><a href="/wiki/Category:Paranthropus" 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