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href="#Total_evidence_dating"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.4</span> <span>Total evidence dating</span> </div> </a> <ul id="toc-Total_evidence_dating-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Methods" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Methods"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.5</span> <span>Methods</span> </div> </a> <ul id="toc-Methods-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Non-constant_rate_of_molecular_clock" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Non-constant_rate_of_molecular_clock"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Non-constant rate of molecular clock</span> </div> </a> <ul id="toc-Non-constant_rate_of_molecular_clock-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Uses" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Uses"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Uses</span> </div> </a> <ul id="toc-Uses-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" 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href="https://bn.wikipedia.org/wiki/%E0%A6%86%E0%A6%A3%E0%A6%AC%E0%A6%BF%E0%A6%95_%E0%A6%98%E0%A6%A1%E0%A6%BC%E0%A6%BF" title="আণবিক ঘড়ি – Bangla" lang="bn" hreflang="bn" data-title="আণবিক ঘড়ি" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9C%D0%BE%D0%BB%D0%B5%D0%BA%D1%83%D0%BB%D1%8F%D1%80%D0%B5%D0%BD_%D1%87%D0%B0%D1%81%D0%BE%D0%B2%D0%BD%D0%B8%D0%BA" title="Молекулярен часовник – Bulgarian" lang="bg" hreflang="bg" data-title="Молекулярен часовник" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Molekularni_sat" title="Molekularni sat – Bosnian" lang="bs" hreflang="bs" data-title="Molekularni sat" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Rellotge_molecular" title="Rellotge molecular – Catalan" lang="ca" hreflang="ca" data-title="Rellotge molecular" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Molekul%C3%A1rn%C3%AD_hodiny" title="Molekulární hodiny – Czech" lang="cs" hreflang="cs" data-title="Molekulární hodiny" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Det_molekyl%C3%A6re_ur" title="Det molekylære ur – Danish" lang="da" hreflang="da" data-title="Det molekylære ur" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Molekulare_Uhr" title="Molekulare Uhr – German" lang="de" hreflang="de" data-title="Molekulare Uhr" 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/Molekulaarne_kell" title="Molekulaarne kell – Estonian" lang="et" hreflang="et" data-title="Molekulaarne kell" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%9C%CE%BF%CF%81%CE%B9%CE%B1%CE%BA%CF%8C_%CF%81%CE%BF%CE%BB%CF%8C%CE%B9" title="Μοριακό ρολόι – Greek" lang="el" hreflang="el" data-title="Μοριακό ρολόι" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Reloj_molecular" title="Reloj molecular – Spanish" lang="es" hreflang="es" data-title="Reloj molecular" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Molekula_horlo%C4%9Do" title="Molekula horloĝo – Esperanto" lang="eo" hreflang="eo" data-title="Molekula horloĝo" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Erloju_molekular" title="Erloju molekular – Basque" lang="eu" hreflang="eu" data-title="Erloju molekular" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%B3%D8%A7%D8%B9%D8%AA_%D9%85%D9%88%D9%84%DA%A9%D9%88%D9%84%DB%8C" title="ساعت مولکولی – Persian" lang="fa" hreflang="fa" data-title="ساعت مولکولی" 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/Horloge_mol%C3%A9culaire" title="Horloge moléculaire – French" lang="fr" hreflang="fr" data-title="Horloge moléculaire" 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/Reloxo_molecular" title="Reloxo molecular – Galician" lang="gl" hreflang="gl" data-title="Reloxo molecular" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EB%B6%84%EC%9E%90_%EC%8B%9C%EA%B3%84" title="분자 시계 – Korean" lang="ko" hreflang="ko" data-title="분자 시계" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Jam_molekuler" title="Jam molekuler – Indonesian" lang="id" hreflang="id" data-title="Jam molekuler" 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/Orologio_molecolare" title="Orologio molecolare – Italian" lang="it" hreflang="it" data-title="Orologio molecolare" 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%A9%D7%A2%D7%95%D7%9F_%D7%9E%D7%95%D7%9C%D7%A7%D7%95%D7%9C%D7%A8%D7%99" title="שעון מולקולרי – Hebrew" lang="he" hreflang="he" data-title="שעון מולקולרי" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%A4%E0%B5%BB%E0%B4%AE%E0%B4%BE%E0%B4%A4%E0%B5%8D%E0%B4%B0%E0%B4%BE%E0%B4%98%E0%B4%9F%E0%B4%BF%E0%B4%95%E0%B4%BE%E0%B4%B0%E0%B4%82" title="തൻമാത്രാഘടികാരം – Malayalam" lang="ml" hreflang="ml" data-title="തൻമാത്രാഘടികാരം" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Moleculaire_klok" title="Moleculaire klok – Dutch" lang="nl" hreflang="nl" data-title="Moleculaire klok" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E5%88%86%E5%AD%90%E6%99%82%E8%A8%88" title="分子時計 – Japanese" lang="ja" hreflang="ja" data-title="分子時計" 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/Det_molekyl%C3%A6re_ur" title="Det molekylære ur – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Det molekylære ur" 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-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Zegar_molekularny" title="Zegar molekularny – Polish" lang="pl" hreflang="pl" data-title="Zegar molekularny" 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/Rel%C3%B3gio_molecular" title="Relógio molecular – Portuguese" lang="pt" hreflang="pt" data-title="Relógio molecular" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9C%D0%BE%D0%BB%D0%B5%D0%BA%D1%83%D0%BB%D1%8F%D1%80%D0%BD%D1%8B%D0%B5_%D1%87%D0%B0%D1%81%D1%8B" title="Молекулярные часы – Russian" lang="ru" hreflang="ru" data-title="Молекулярные часы" 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/Molecular_clock" title="Molecular clock – Simple English" lang="en-simple" hreflang="en-simple" data-title="Molecular clock" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Molekyylikello" title="Molekyylikello – Finnish" lang="fi" hreflang="fi" data-title="Molekyylikello" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Molekyl%C3%A4r_klocka" title="Molekylär klocka – Swedish" lang="sv" hreflang="sv" data-title="Molekylär klocka" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%AE%E0%AF%82%E0%AE%B2%E0%AE%95%E0%AF%8D%E0%AE%95%E0%AF%82%E0%AE%B1%E0%AF%81_%E0%AE%95%E0%AE%BE%E0%AE%B2%E0%AE%99%E0%AF%8D%E0%AE%95%E0%AE%BE%E0%AE%9F%E0%AF%8D%E0%AE%9F%E0%AE%BF" title="மூலக்கூறு காலங்காட்டி – Tamil" lang="ta" hreflang="ta" data-title="மூலக்கூறு காலங்காட்டி" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Molek%C3%BCler_saat" title="Moleküler saat – Turkish" lang="tr" hreflang="tr" data-title="Moleküler saat" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9C%D0%BE%D0%BB%D0%B5%D0%BA%D1%83%D0%BB%D1%8F%D1%80%D0%BD%D0%B8%D0%B9_%D0%B3%D0%BE%D0%B4%D0%B8%D0%BD%D0%BD%D0%B8%D0%BA" title="Молекулярний годинник – Ukrainian" lang="uk" hreflang="uk" data-title="Молекулярний годинник" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D8%B3%D8%A7%D9%84%D9%85%D8%A7%D8%AA%DB%8C_%DA%AF%DA%BE%DA%91%DB%8C" title="سالماتی گھڑی – Urdu" lang="ur" hreflang="ur" data-title="سالماتی گھڑی" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target"><span>اردو</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/%C4%90%E1%BB%93ng_h%E1%BB%93_ph%C3%A2n_t%E1%BB%AD" title="Đồng hồ phân tử – Vietnamese" lang="vi" hreflang="vi" data-title="Đồng hồ phân tử" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E5%88%86%E5%AD%90%E9%90%98" title="分子鐘 – Chinese" lang="zh" hreflang="zh" data-title="分子鐘" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q827541#sitelinks-wikipedia" title="Edit interlanguage links" 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dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Technique to deduce the time in prehistory when two or more life forms diverged</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Not to be confused with <a href="/wiki/Chemical_clock" title="Chemical clock">Chemical clock</a> or <a href="/wiki/Biological_clock_(disambiguation)" class="mw-redirect mw-disambig" title="Biological clock (disambiguation)">Biological clock</a>.</div> <p class="mw-empty-elt"> </p> <style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist 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style="background:#90ee90;padding:0.2em;font-size:175%;font-weight:bold"><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></th></tr><tr><td class="sidebar-image"><span class="notpageimage" typeof="mw:File"><a href="/wiki/File:Darwin%27s_finches_by_Gould.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Darwin%27s_finches_by_Gould.jpg/180px-Darwin%27s_finches_by_Gould.jpg" decoding="async" width="180" height="136" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Darwin%27s_finches_by_Gould.jpg/270px-Darwin%27s_finches_by_Gould.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Darwin%27s_finches_by_Gould.jpg/360px-Darwin%27s_finches_by_Gould.jpg 2x" data-file-width="1748" data-file-height="1319" /></a></span><div class="sidebar-caption"><a href="/wiki/Darwin%27s_finches" title="Darwin's finches">Darwin's finches</a> by <a href="/wiki/John_Gould" title="John Gould">John Gould</a></div></td></tr><tr><td class="sidebar-above" style="display:block; margin-bottom:0.55em; background-color: transparent;"> <div class="hlist"> <ul><li><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li> <li><a href="/wiki/Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li> <li><a href="/wiki/Evolution" title="Evolution">Main</a></li> <li><a href="/wiki/Outline_of_evolution" title="Outline of evolution">Outline</a></li></ul> <div class="hlist"> </div> <ul><li><a href="/wiki/Glossary_of_evolutionary_biology" class="mw-redirect" title="Glossary of evolutionary biology">Glossary</a></li> <li><a href="/wiki/Evidence_of_common_descent" title="Evidence of common descent">Evidence</a></li> <li><a href="/wiki/History_of_life" title="History of life">History</a></li></ul> </div></td></tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Processes and outcomes</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist"> <ul><li><a href="/wiki/Population_genetics" title="Population genetics">Population genetics</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Variation</a></li> <li><a href="/wiki/Genetic_diversity" title="Genetic diversity">Diversity</a></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></li> <li><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a></li> <li><a href="/wiki/Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li> <li><a href="/wiki/Cooperation_(evolution)" title="Cooperation (evolution)">Co-operation</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a></li> <li><a href="/wiki/Coextinction" title="Coextinction">Coextinction</a></li> <li><a href="/wiki/Contingency_(evolutionary_biology)" title="Contingency (evolutionary biology)">Contingency</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergence</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergence</a></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Extinction" title="Extinction">Extinction</a></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Natural history</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist"> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></li> <li><a href="/wiki/Common_descent" title="Common descent">Common descent</a></li> <li><a href="/wiki/History_of_life" title="History of life">History of life</a></li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li> <li><a href="/wiki/Human_evolution" title="Human evolution">Human evolution</a> <ul><li><a href="/wiki/Recent_human_evolution" title="Recent human evolution">Recent human evolution</a></li></ul></li> <li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Phylogeny</a></li> <li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Classification</a></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Evolutionary taxonomy</a></li> <li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li> <li><a href="/wiki/Extinction_event" title="Extinction event">Extinction event</a></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">History of evolutionary theory</div><div class="sidebar-list-content 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controversy</a></li></ul></div></div></td> </tr><tr><td class="sidebar-below hlist"> <ul><li><span class="nowrap"><span class="nowrap"><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Tree_of_life.svg" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/16px-Tree_of_life.svg.png" decoding="async" width="16" height="13" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/24px-Tree_of_life.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/32px-Tree_of_life.svg.png 2x" data-file-width="915" data-file-height="760" /></a></span> </span><a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Evolutionary biology portal</a></span></li> <li><span class="nowrap"><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Evolutionary_biology" title="Category:Evolutionary biology">Category</a></span></li></ul></td></tr><tr><td class="sidebar-navbar"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evolutionary_biology" title="Template:Evolutionary biology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolutionary_biology" title="Template talk:Evolutionary biology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolutionary_biology" title="Special:EditPage/Template:Evolutionary biology"><abbr title="Edit this template">e</abbr></a></li></ul></div></td></tr></tbody></table> <p>The <b>molecular clock </b> is a figurative term for a technique that uses the <a href="/wiki/Mutation_rate" title="Mutation rate">mutation rate</a> of <a href="/wiki/Biomolecule" title="Biomolecule">biomolecules</a> to <a href="/wiki/Chronological_dating" title="Chronological dating">deduce the time</a> in <a href="/wiki/Prehistory" title="Prehistory">prehistory</a> when two or more <a href="/wiki/Life_form" class="mw-redirect" title="Life form">life forms</a> <a href="/wiki/Genetic_divergence" title="Genetic divergence">diverged</a>. The biomolecular data used for such calculations are usually <a href="/wiki/Nucleotide" title="Nucleotide">nucleotide</a> <a href="/wiki/DNA_sequence" class="mw-redirect" title="DNA sequence">sequences</a> for <a href="/wiki/DNA" title="DNA">DNA</a>, <a href="/wiki/RNA" title="RNA">RNA</a>, or <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a> sequences for <a href="/wiki/Protein" title="Protein">proteins</a>. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Early_discovery_and_genetic_equidistance">Early discovery and genetic equidistance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Molecular_clock&action=edit&section=1" title="Edit section: Early discovery and genetic equidistance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The notion of the existence of a so-called "molecular clock" was first attributed to <a href="/wiki/Emile_Zuckerkandl" title="Emile Zuckerkandl">Émile Zuckerkandl</a> and <a href="/wiki/Linus_Pauling" title="Linus Pauling">Linus Pauling</a> who, in 1962, noticed that the number of <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a> differences in <a href="/wiki/Hemoglobin" title="Hemoglobin">hemoglobin</a> between different lineages changes roughly <a href="/wiki/Linear_function" title="Linear function">linearly</a> with time, as estimated from fossil evidence.<sup id="cite_ref-Zuckerkand62_1-0" class="reference"><a href="#cite_note-Zuckerkand62-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> They generalized this observation to assert that the rate of <a href="/wiki/Evolution" title="Evolution">evolutionary</a> change of any specified <a href="/wiki/Protein" title="Protein">protein</a> was approximately constant over time and over different lineages (known as the <b>molecular clock hypothesis</b>). </p><p>The <b>genetic equidistance</b> phenomenon was first noted in 1963 by <a href="/wiki/Emanuel_Margoliash" title="Emanuel Margoliash">Emanuel Margoliash</a>, who wrote: "It appears that the number of residue differences between <a href="/wiki/Cytochrome_c" title="Cytochrome c">cytochrome c</a> of any two species is mostly conditioned by the time elapsed since the lines of evolution leading to these two species originally diverged. If this is correct, the cytochrome c of all mammals should be equally different from the cytochrome c of all birds. Since fish diverges from the main stem of vertebrate evolution earlier than either birds or mammals, the cytochrome c of both mammals and birds should be equally different from the cytochrome c of fish. Similarly, all vertebrate cytochrome c should be equally different from the yeast protein."<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> For example, the difference between the cytochrome c of a carp and a frog, turtle, chicken, rabbit, and horse is a very constant 13% to 14%. Similarly, the difference between the cytochrome c of a bacterium and yeast, wheat, moth, tuna, pigeon, and horse ranges from 64% to 69%. Together with the work of Emile Zuckerkandl and Linus Pauling, the genetic equidistance result led directly to the formal postulation of the molecular clock hypothesis in the early 1960s.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>Similarly, <a href="/wiki/Vincent_Sarich" title="Vincent Sarich">Vincent Sarich</a> and <a href="/wiki/Allan_Wilson_(biologist)" title="Allan Wilson (biologist)">Allan Wilson</a> in 1967 demonstrated that molecular differences among modern <a href="/wiki/Primate" title="Primate">Primates</a> in <a href="/wiki/Albumin" title="Albumin">albumin</a> proteins showed that approximately constant rates of change had occurred in all the lineages they assessed.<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> The basic logic of their analysis involved recognizing that if one species lineage had evolved more quickly than a sister species lineage since their common ancestor, then the molecular differences between an outgroup (more distantly related) species and the faster-evolving species should be larger (since more molecular changes would have accumulated on that lineage) than the molecular differences between the outgroup species and the slower-evolving species. This method is known as the <a href="/wiki/Relative_rate_test" title="Relative rate test">relative rate test</a>. Sarich and Wilson's paper reported, for example, that human (<i><a href="/wiki/Homo_sapiens" class="mw-redirect" title="Homo sapiens">Homo sapiens</a></i>) and chimpanzee (<i><a href="/wiki/Common_chimpanzee" class="mw-redirect" title="Common chimpanzee">Pan troglodytes</a></i>) albumin immunological cross-reactions suggested they were about equally different from <a href="/wiki/New_World_monkey" title="New World monkey">Ceboidea</a> (New World Monkey) species (within experimental error). This meant that they had both accumulated approximately equal changes in albumin since their shared common ancestor. This pattern was also found for all the primate comparisons they tested. When calibrated with the few well-documented fossil branch points (such as no Primate fossils of modern aspect found before the <a href="/wiki/Cretaceous%E2%80%93Paleogene_boundary" title="Cretaceous–Paleogene boundary">K-T boundary</a>), this led Sarich and Wilson to argue that the human-chimp divergence probably occurred only ~4–6 million years ago.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Relationship_with_neutral_theory">Relationship with neutral theory</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Molecular_clock&action=edit&section=2" title="Edit section: Relationship with neutral theory"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The observation of a clock-like rate of molecular change was originally purely <a href="/wiki/Phenomenology_(philosophy)" title="Phenomenology (philosophy)">phenomenological</a>. Later, the work of <a href="/wiki/Motoo_Kimura" title="Motoo Kimura">Motoo Kimura</a><sup id="cite_ref-Kimura68_6-0" class="reference"><a href="#cite_note-Kimura68-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> developed the <a href="/wiki/Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">neutral theory of molecular evolution</a>, which predicted a molecular clock. Let there be N individuals, and to keep this calculation simple, let the individuals be <a href="/wiki/Ploidy" title="Ploidy">haploid</a> (i.e. have one copy of each gene). Let the rate of neutral <a href="/wiki/Mutation" title="Mutation">mutations</a> (i.e. mutations with no effect on <a href="/wiki/Fitness_(biology)" title="Fitness (biology)">fitness</a>) in a new individual be <span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \mu }"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>μ</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \mu }</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/9fd47b2a39f7a7856952afec1f1db72c67af6161" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:1.402ex; height:2.176ex;" alt="{\displaystyle \mu }"></span>. The probability that this new mutation will become <a href="/wiki/Fixation_(population_genetics)" title="Fixation (population genetics)">fixed</a> in the population is then 1/N, since each copy of the gene is as good as any other. Every generation, each individual can have new mutations, so there are <span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \mu }"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>μ</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \mu }</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/9fd47b2a39f7a7856952afec1f1db72c67af6161" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:1.402ex; height:2.176ex;" alt="{\displaystyle \mu }"></span>N new neutral mutations in the population as a whole. That means that each generation, <span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \mu }"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>μ</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \mu }</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/9fd47b2a39f7a7856952afec1f1db72c67af6161" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:1.402ex; height:2.176ex;" alt="{\displaystyle \mu }"></span> new neutral mutations will become fixed. If most changes seen during <a href="/wiki/Molecular_evolution" title="Molecular evolution">molecular evolution</a> are neutral, then <a href="/wiki/Fixation_(population_genetics)" title="Fixation (population genetics)">fixations</a> in a population will accumulate at a clock-rate that is equal to the rate of neutral <a href="/wiki/Mutation" title="Mutation">mutations</a> in an individual. </p> <div class="mw-heading mw-heading2"><h2 id="Calibration">Calibration</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Molecular_clock&action=edit&section=3" title="Edit section: Calibration"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>To use molecular clocks to estimate divergence times, molecular clocks need to be "calibrated". This is because molecular data alone does not contain any information on absolute times. For viral phylogenetics and <a href="/wiki/Ancient_DNA" title="Ancient DNA">ancient DNA</a> studies—two areas of evolutionary biology where it is possible to sample sequences over an evolutionary timescale—the dates of the intermediate samples can be used to calibrate the molecular clock. However, most phylogenies require that the molecular clock be <a href="/wiki/Calibration" title="Calibration">calibrated</a> using independent evidence about dates, such as the <a href="/wiki/Fossil" title="Fossil">fossil</a> record.<sup id="cite_ref-Benton01_7-0" class="reference"><a href="#cite_note-Benton01-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> There are two general methods for calibrating the molecular clock using fossils: node calibration and tip calibration.<sup id="cite_ref-Donoghue02_8-0" class="reference"><a href="#cite_note-Donoghue02-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Node_calibration">Node calibration</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Molecular_clock&action=edit&section=4" title="Edit section: Node calibration"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Sometimes referred to as node dating, node calibration is a method for time-scaling <a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">phylogenetic trees</a> by specifying time constraints for one or more nodes in the tree. Early methods of clock calibration only used a single fossil constraint (e.g. non-parametric rate smoothing),<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> but newer methods (BEAST<sup id="cite_ref-ReferenceA_10-0" class="reference"><a href="#cite_note-ReferenceA-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> and <a href="https://sourceforge.net/projects/r8s/" class="extiw" title="sourceforge:projects/r8s/">r8s</a><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>) allow for the use of multiple fossils to calibrate molecular clocks. The oldest fossil of a <a href="/wiki/Clade" title="Clade">clade</a> is used to constrain the minimum possible age for the node representing the most recent common ancestor of the clade. However, due to incomplete fossil preservation and other factors, clades are typically older than their oldest fossils.<sup id="cite_ref-Donoghue02_8-1" class="reference"><a href="#cite_note-Donoghue02-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> In order to account for this, nodes are allowed to be older than the minimum constraint in node calibration analyses. However, determining how much older the node is allowed to be is challenging. There are a number of strategies for deriving the maximum bound for the age of a clade including those based on birth-death models, fossil <a href="/wiki/Stratigraphy" title="Stratigraphy">stratigraphic</a> distribution analyses, or <a href="/wiki/Taphonomy" title="Taphonomy">taphonomic</a> controls.<sup id="cite_ref-O'Reilly03_12-0" class="reference"><a href="#cite_note-O'Reilly03-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Alternatively, instead of a maximum and a minimum, a <a href="/wiki/Probability_density" class="mw-redirect" title="Probability density">probability density</a> can be used to represent the uncertainty about the age of the clade. These calibration densities can take the shape of standard probability densities (e.g. <a href="/wiki/Normal_distribution" title="Normal distribution">normal</a>, <a href="/wiki/Log-normal_distribution" title="Log-normal distribution">lognormal</a>, <a href="/wiki/Exponential_distribution" title="Exponential distribution">exponential</a>, <a href="/wiki/Gamma_distribution" title="Gamma distribution">gamma</a>) that can be used to express the uncertainty associated with divergence time estimates. <sup id="cite_ref-ReferenceA_10-1" class="reference"><a href="#cite_note-ReferenceA-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Determining the shape and parameters of the probability distribution is not trivial, but there are methods that use not only the oldest fossil but a larger sample of the fossil record of clades to estimate calibration densities empirically.<sup id="cite_ref-Claramunt2022_13-0" class="reference"><a href="#cite_note-Claramunt2022-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Studies have shown that increasing the number of fossil constraints increases the accuracy of divergence time estimation.<sup id="cite_ref-Zheng04_14-0" class="reference"><a href="#cite_note-Zheng04-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Tip_calibration">Tip calibration</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Molecular_clock&action=edit&section=5" title="Edit section: Tip calibration"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Sometimes referred to as <a href="/wiki/Tip_dating" title="Tip dating">tip dating</a>, tip calibration is a method of molecular clock calibration in which fossils are treated as <a href="/wiki/Taxon" title="Taxon">taxa</a> and placed on the tips of the tree. This is achieved by creating a matrix that includes a <a href="/wiki/Molecular_phylogenetics" title="Molecular phylogenetics">molecular</a> dataset for the <a href="/wiki/Neontology" title="Neontology">extant taxa</a> along with a <a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphological</a> dataset for both the extinct and the extant taxa.<sup id="cite_ref-O'Reilly03_12-1" class="reference"><a href="#cite_note-O'Reilly03-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Unlike node calibration, this method reconstructs the tree topology and places the fossils simultaneously. Molecular and morphological models work together simultaneously, allowing morphology to inform the placement of fossils.<sup id="cite_ref-Donoghue02_8-2" class="reference"><a href="#cite_note-Donoghue02-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Tip calibration makes use of all relevant fossil taxa during clock calibration, rather than relying on only the oldest fossil of each clade. This method does not rely on the interpretation of negative evidence to infer maximum clade ages.<sup id="cite_ref-O'Reilly03_12-2" class="reference"><a href="#cite_note-O'Reilly03-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Expansion_calibration">Expansion calibration</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Molecular_clock&action=edit&section=6" title="Edit section: Expansion calibration"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Demographic changes in populations can be detected as fluctuations in historical coalescent <a href="/wiki/Effective_population_size" title="Effective population size">effective population size</a> from a sample of extant genetic variation in the population using coalescent theory.<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><sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Ancient population expansions that are well documented and dated in the geological record can be used to calibrate a rate of molecular evolution in a manner similar to node calibration. However, instead of calibrating from the known age of a node, expansion calibration uses a two-epoch model of constant population size followed by population growth, with the time of transition between epochs being the parameter of interest for calibration.<sup id="cite_ref-:0_18-0" class="reference"><a href="#cite_note-:0-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> Expansion calibration works at shorter, intraspecific timescales in comparison to node calibration, because expansions can only be detected after the <a href="/wiki/Most_recent_common_ancestor" title="Most recent common ancestor">most recent common ancestor</a> of the species in question. Expansion dating has been used to show that molecular clock rates can be inflated at short timescales<sup id="cite_ref-:0_18-1" class="reference"><a href="#cite_note-:0-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> (< 1 MY) due to incomplete fixation of alleles, as discussed below<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_21-0" class="reference"><a href="#cite_note-:1-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Total_evidence_dating">Total evidence dating</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Molecular_clock&action=edit&section=7" title="Edit section: Total evidence dating"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>This approach to tip calibration goes a step further by simultaneously estimating fossil placement, topology, and the evolutionary timescale. In this method, the age of a fossil can inform its phylogenetic position in addition to morphology. By allowing all aspects of tree reconstruction to occur simultaneously, the risk of biased results is decreased.<sup id="cite_ref-Donoghue02_8-3" class="reference"><a href="#cite_note-Donoghue02-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> This approach has been improved upon by pairing it with different models. One current method of molecular clock calibration is total evidence dating paired with the fossilized birth-death (FBD) model and a model of morphological evolution.<sup id="cite_ref-Heath05_22-0" class="reference"><a href="#cite_note-Heath05-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> The FBD model is novel in that it allows for "sampled ancestors", which are fossil taxa that are the direct ancestor of a living taxon or <a href="/wiki/Lineage_(evolution)" title="Lineage (evolution)">lineage</a>. This allows fossils to be placed on a branch above an extant organism, rather than being confined to the tips.<sup id="cite_ref-Gavryushkina06_23-0" class="reference"><a href="#cite_note-Gavryushkina06-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Methods">Methods</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Molecular_clock&action=edit&section=8" title="Edit section: Methods"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Bayesian methods can provide more appropriate estimates of divergence times, especially if large datasets—such as those yielded by <a href="/wiki/Phylogenomics" title="Phylogenomics">phylogenomics</a>—are employed.<sup id="cite_ref-Dos_Reis2012_24-0" class="reference"><a href="#cite_note-Dos_Reis2012-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Non-constant_rate_of_molecular_clock">Non-constant rate of molecular clock</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Molecular_clock&action=edit&section=9" title="Edit section: Non-constant rate of molecular clock"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Sometimes only a single divergence date can be estimated from fossils, with all other dates inferred from that. Other sets of species have abundant fossils available, allowing the hypothesis of constant divergence rates to be tested. DNA sequences experiencing low levels of <a href="/wiki/Negative_selection_(natural_selection)" title="Negative selection (natural selection)">negative selection</a> showed divergence rates of 0.7–0.8% per <a href="/wiki/Myr" class="mw-redirect" title="Myr">Myr</a> in bacteria, mammals, invertebrates, and plants.<sup id="cite_ref-Ochman87_25-0" class="reference"><a href="#cite_note-Ochman87-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> In the same study, genomic regions experiencing very high negative or purifying selection (encoding rRNA) were considerably slower (1% per 50 Myr). </p><p>In addition to such variation in rate with genomic position, since the early 1990s variation among taxa has proven fertile ground for research too,<sup id="cite_ref-Douzery03_26-0" class="reference"><a href="#cite_note-Douzery03-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> even over comparatively short periods of evolutionary time (for example <a href="/wiki/Mockingbird" title="Mockingbird">mockingbirds</a><sup id="cite_ref-Hunt01_27-0" class="reference"><a href="#cite_note-Hunt01-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup>). <a href="/wiki/Procellariiformes" title="Procellariiformes">Tube-nosed seabirds</a> have molecular clocks that on average run at half speed of many other birds,<sup id="cite_ref-Rheindt05_28-0" class="reference"><a href="#cite_note-Rheindt05-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> possibly due to long <a href="/wiki/Generation" title="Generation">generation</a> times, and many turtles have a molecular clock running at one-eighth the speed it does in small mammals, or even slower.<sup id="cite_ref-Avise92_29-0" class="reference"><a href="#cite_note-Avise92-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> Effects of <a href="/wiki/Small_population_size" title="Small population size">small population size</a> are also likely to confound molecular clock analyses. Researchers such as <a href="/wiki/Francisco_J._Ayala" title="Francisco J. Ayala">Francisco J. Ayala</a> have more fundamentally challenged the molecular clock hypothesis.<sup id="cite_ref-Ayala99_30-0" class="reference"><a href="#cite_note-Ayala99-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Schwartz06_31-0" class="reference"><a href="#cite_note-Schwartz06-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Pascual2019_32-0" class="reference"><a href="#cite_note-Pascual2019-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> According to Ayala's 1999 study, five factors combine to limit the application of molecular clock models: </p> <ul><li>Changing generation times (If the rate of new mutations depends at least partly on the number of generations rather than the number of years)</li> <li>Population size (<a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a> is stronger in small populations, and so more mutations are effectively neutral)</li> <li>Species-specific differences (due to differing metabolism, ecology, evolutionary history, ...)</li> <li>Change in function of the protein studied (can be avoided in closely related species by utilizing <a href="/wiki/Non-coding_DNA" title="Non-coding DNA">non-coding DNA</a> sequences or emphasizing <a href="/wiki/Silent_mutation" title="Silent mutation">silent mutations</a>)</li> <li>Changes in the intensity of natural selection.</li></ul> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Molecular_evolution_bamboos.svg" class="mw-file-description"><img alt="Phylogram showing three groups, one of which has strikingly longer branches than the two others" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3f/Molecular_evolution_bamboos.svg/300px-Molecular_evolution_bamboos.svg.png" decoding="async" width="300" height="300" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3f/Molecular_evolution_bamboos.svg/450px-Molecular_evolution_bamboos.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3f/Molecular_evolution_bamboos.svg/600px-Molecular_evolution_bamboos.svg.png 2x" data-file-width="360" data-file-height="360" /></a><figcaption>Woody bamboos (tribes <a href="/wiki/Arundinarieae" title="Arundinarieae">Arundinarieae</a> and <a href="/wiki/Bambuseae" title="Bambuseae">Bambuseae</a>) have long generation times and lower mutation rates, as expressed by short branches in the <a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">phylogenetic tree</a>, than the fast-evolving herbaceous bamboos (<a href="/wiki/Olyreae" title="Olyreae">Olyreae</a>).</figcaption></figure> <p>Molecular clock users have developed workaround solutions using a number of statistical approaches including <a href="/wiki/Maximum_likelihood" class="mw-redirect" title="Maximum likelihood">maximum likelihood</a> techniques and later <a href="/wiki/Bayesian_statistics" title="Bayesian statistics">Bayesian modeling</a>. In particular, models that take into account rate variation across lineages have been proposed in order to obtain better estimates of divergence times. These models are called <b>relaxed molecular clocks</b><sup id="cite_ref-Drummond06_33-0" class="reference"><a href="#cite_note-Drummond06-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> because they represent an intermediate position between the 'strict' molecular clock hypothesis and <a href="/wiki/Joseph_Felsenstein" title="Joseph Felsenstein">Joseph Felsenstein</a>'s many-rates model<sup id="cite_ref-Felsenstein01_34-0" class="reference"><a href="#cite_note-Felsenstein01-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> and are made possible through <a href="/wiki/Markov_chain_Monte_Carlo" title="Markov chain Monte Carlo">MCMC</a> techniques that explore a weighted range of tree topologies and simultaneously estimate parameters of the chosen substitution model. It must be remembered that divergence dates inferred using a molecular clock are based on statistical <a href="/wiki/Inference" title="Inference">inference</a> and not on direct <a href="/wiki/Evidence" title="Evidence">evidence</a>. </p><p>The molecular clock runs into particular challenges at very short and very long timescales. At long timescales, the problem is <a href="/wiki/Saturation_(genetic)" class="mw-redirect" title="Saturation (genetic)">saturation</a>. When enough time has passed, many sites have undergone more than one change, but it is impossible to detect more than one. This means that the observed number of changes is no longer <a href="/wiki/Linear_function" title="Linear function">linear</a> with time, but instead flattens out. Even at intermediate genetic distances, with phylogenetic data still sufficient to estimate topology, signal for the overall scale of the tree can be weak under complex likelihood models, leading to highly uncertain molecular clock estimates.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p><p>At very short time scales, many differences between samples do not represent <a href="/wiki/Fixation_(population_genetics)" title="Fixation (population genetics)">fixation</a> of different sequences in the different populations. Instead, they represent alternative <a href="/wiki/Alleles" class="mw-redirect" title="Alleles">alleles</a> that were both present as part of a polymorphism in the common ancestor. The inclusion of differences that have not yet become <a href="/wiki/Fixation_(population_genetics)" title="Fixation (population genetics)">fixed</a> leads to a potentially dramatic inflation of the apparent rate of the molecular clock at very short timescales.<sup id="cite_ref-:1_21-1" class="reference"><a href="#cite_note-:1-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><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-heading2"><h2 id="Uses">Uses</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Molecular_clock&action=edit&section=10" title="Edit section: Uses"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The molecular clock technique is an important tool in <a href="/wiki/Molecular_systematics" class="mw-redirect" title="Molecular systematics">molecular systematics</a>, <a href="/wiki/Macroevolution" title="Macroevolution">macroevolution</a>, and <a href="/wiki/Phylogenetic_comparative_methods" title="Phylogenetic comparative methods">phylogenetic comparative methods</a>. Estimation of the dates of <a href="/wiki/Phylogeny" class="mw-redirect" title="Phylogeny">phylogenetic</a> events, including those not documented by <a href="/wiki/Fossils" class="mw-redirect" title="Fossils">fossils</a>, such as the divergences between living <a href="/wiki/Taxon" title="Taxon">taxa</a> has allowed the study of macroevolutionary processes in organisms that had limited fossil records. Phylogenetic comparative methods rely heavily on calibrated phylogenies. </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=Molecular_clock&action=edit&section=11" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a></li> <li><a href="/wiki/Gene_orders" title="Gene orders">Gene orders</a></li> <li><a href="/wiki/Human_mitochondrial_molecular_clock" title="Human mitochondrial molecular clock">Human mitochondrial molecular clock</a></li> <li><a href="/wiki/Mitochondrial_Eve" title="Mitochondrial Eve">Mitochondrial Eve</a> and <a href="/wiki/Y-chromosomal_Adam" title="Y-chromosomal Adam">Y-chromosomal Adam</a></li> <li><a href="/wiki/Models_of_DNA_evolution" title="Models of DNA evolution">Models of DNA evolution</a></li> <li><a href="/wiki/Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">Neutral theory of molecular evolution</a></li> <li><a href="/wiki/Glottochronology" title="Glottochronology">Glottochronology</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Molecular_clock&action=edit&section=12" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width reflist-columns-2"> <ol class="references"> <li id="cite_note-Zuckerkand62-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-Zuckerkand62_1-0">^</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 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C., et al. 2016. Inflation of molecular clock rates and dates: molecular phylogenetics, biogeography, and diversification of a global cicada radiation from Australasia (Hemiptera: Cicadidae: Cicadettini). <a rel="nofollow" class="external text" href="https://academic.oup.com/sysbio/article/65/1/16/2461540/Inflation-of-Molecular-Clock-Rates-and-Dates">Systematic Biology 65(1):16–34</a>.</span> </li> <li id="cite_note-36"><span class="mw-cite-backlink"><b><a href="#cite_ref-36">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPetersonMasel2009" class="citation journal cs1">Peterson GI, Masel J (November 2009). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2912466">"Quantitative prediction of molecular clock and ka/ks at short timescales"</a>. <i>Molecular Biology and Evolution</i>. <b>26</b> (11): 2595–2603. <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%2Fmolbev%2Fmsp175">10.1093/molbev/msp175</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2912466">2912466</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19661199">19661199</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Molecular+Biology+and+Evolution&rft.atitle=Quantitative+prediction+of+molecular+clock+and+ka%2Fks+at+short+timescales&rft.volume=26&rft.issue=11&rft.pages=2595-2603&rft.date=2009-11&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2912466%23id-name%3DPMC&rft_id=info%3Apmid%2F19661199&rft_id=info%3Adoi%2F10.1093%2Fmolbev%2Fmsp175&rft.aulast=Peterson&rft.aufirst=GI&rft.au=Masel%2C+J&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2912466&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMolecular+clock" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Molecular_clock&action=edit&section=13" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHo,_S.Y.W.2020" class="citation book cs1">Ho, S.Y.W., ed. (2020). <i>The Molecular Evolutionary Clock: Theory and Practice</i>. Springer, Cham. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2F978-3-030-60181-2">10.1007/978-3-030-60181-2</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-3-030-60180-5" title="Special:BookSources/978-3-030-60180-5"><bdi>978-3-030-60180-5</bdi></a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:231672167">231672167</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Molecular+Evolutionary+Clock%3A+Theory+and+Practice&rft.pub=Springer%2C+Cham&rft.date=2020&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A231672167%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1007%2F978-3-030-60181-2&rft.isbn=978-3-030-60180-5&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMolecular+clock" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKumar2005" class="citation journal cs1">Kumar S (August 2005). "Molecular clocks: four decades of evolution". <i>Nature Reviews. Genetics</i>. <b>6</b> (8): 654–662. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnrg1659">10.1038/nrg1659</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/16136655">16136655</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:14261833">14261833</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Reviews.+Genetics&rft.atitle=Molecular+clocks%3A+four+decades+of+evolution&rft.volume=6&rft.issue=8&rft.pages=654-662&rft.date=2005-08&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A14261833%23id-name%3DS2CID&rft_id=info%3Apmid%2F16136655&rft_id=info%3Adoi%2F10.1038%2Fnrg1659&rft.aulast=Kumar&rft.aufirst=S&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMolecular+clock" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMorgan1998" class="citation journal cs1">Morgan GJ (1998). "Emile Zuckerkandl, Linus Pauling, and the molecular evolutionary clock, 1959-1965". <i>Journal of the History of Biology</i>. <b>31</b> (2): 155–178. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1023%2FA%3A1004394418084">10.1023/A:1004394418084</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11620303">11620303</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:5660841">5660841</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+the+History+of+Biology&rft.atitle=Emile+Zuckerkandl%2C+Linus+Pauling%2C+and+the+molecular+evolutionary+clock%2C+1959-1965&rft.volume=31&rft.issue=2&rft.pages=155-178&rft.date=1998&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A5660841%23id-name%3DS2CID&rft_id=info%3Apmid%2F11620303&rft_id=info%3Adoi%2F10.1023%2FA%3A1004394418084&rft.aulast=Morgan&rft.aufirst=GJ&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMolecular+clock" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFZuckerkandlPauling1965" class="citation book cs1"><a href="/wiki/Emile_Zuckerkandl" title="Emile Zuckerkandl">Zuckerkandl E</a>, <a href="/wiki/Linus_Pauling" title="Linus Pauling">Pauling LB</a> (1965). "Evolutionary divergence and convergence in proteins". In Bryson V, Vogel HJ (eds.). <i>Evolving Genes and Proteins</i>. Academic Press, New York. pp. 97–166.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Evolutionary+divergence+and+convergence+in+proteins&rft.btitle=Evolving+Genes+and+Proteins&rft.pages=97-166&rft.pub=Academic+Press%2C+New+York&rft.date=1965&rft.aulast=Zuckerkandl&rft.aufirst=E&rft.au=Pauling%2C+LB&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMolecular+clock" 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=Molecular_clock&action=edit&section=14" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20100525101859/http://awcmee.massey.ac.nz/aw.htm">Allan Wilson and the molecular clock</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20090213150149/http://rtis.com/nat/user/elsberry/evobio/evc/argresp/sequence.html">Molecular clock explanation of the molecular equidistance phenomenon</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20061107013958/http://www.fossilrecord.net/dateaclade/index.html">Date-a-Clade service for the molecular tree of life</a></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 .navbox-inner,.mw-parser-output .navbox-subgroup{width:100%}.mw-parser-output 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class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight:normal;background:#cc9;">Pre-Julian / Julian</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Roman_calendar" title="Roman calendar">Pre-Julian Roman</a></li> <li><a href="/wiki/Julian_calendar" title="Julian calendar">Original Julian</a></li> <li><a href="/wiki/Proleptic_Julian_calendar" title="Proleptic Julian calendar">Proleptic Julian</a></li> <li><a href="/wiki/Revised_Julian_calendar" title="Revised Julian calendar">Revised Julian</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight:normal;background:#cc9;">Gregorian</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Gregorian_calendar" title="Gregorian calendar">Gregorian</a></li> <li><a href="/wiki/Proleptic_Gregorian_calendar" title="Proleptic Gregorian calendar">Proleptic Gregorian</a></li> <li><a href="/wiki/Old_Style_and_New_Style_dates" title="Old Style and New Style dates">Old Style and New Style dates</a></li> <li><a href="/wiki/Adoption_of_the_Gregorian_calendar" title="Adoption of the Gregorian calendar">Adoption of the Gregorian calendar</a></li> <li><a href="/wiki/Dual_dating" title="Dual dating">Dual dating</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight:normal;background:#cc9;">Astronomical</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Lunisolar_calendar" title="Lunisolar calendar">Lunisolar</a> (<a href="/wiki/Hebrew_calendar" title="Hebrew calendar">Hebrew</a>, <a href="/wiki/Hindu_calendar" title="Hindu calendar">Hindu</a>)</li> <li><a href="/wiki/Solar_calendar" title="Solar calendar">Solar</a></li> <li><a href="/wiki/Lunar_calendar" title="Lunar calendar">Lunar</a> (<a href="/wiki/Islamic_calendar" title="Islamic calendar">Islamic</a>)</li> <li><a href="/wiki/Astronomical_year_numbering" title="Astronomical year numbering">Astronomical year numbering</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight:normal;background:#cc9;">Others</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Sexagenary_cycle" title="Sexagenary cycle">Chinese sexagenary cycle</a></li> <li><a href="/wiki/Geologic_Calendar" title="Geologic Calendar">Geologic Calendar</a></li> <li><a href="/wiki/Iranian_calendars" title="Iranian calendars">Iranian</a></li> <li><a href="/wiki/ISO_week_date" title="ISO week date">ISO week date</a></li> <li><a href="/wiki/Mesoamerican_calendars" title="Mesoamerican calendars">Mesoamerican</a> <ul><li><a href="/wiki/Maya_calendar" title="Maya calendar">Maya</a></li> <li><a href="/wiki/Aztec_calendar" title="Aztec calendar">Aztec</a></li></ul></li> <li><a href="/wiki/Winter_count" title="Winter count">Winter count</a></li> <li><a href="/wiki/New_Earth_Time" title="New Earth Time">New Earth Time</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;line-height:1.2em;background:#cc9;"><a href="/wiki/Astronomical_chronology" title="Astronomical chronology">Astronomic time</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0;background:#fff;"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cosmic_Calendar" title="Cosmic Calendar">Cosmic Calendar</a></li> <li><a href="/wiki/Ephemeris" title="Ephemeris">Ephemeris</a></li> <li><a href="/wiki/Galactic_year" title="Galactic year">Galactic year</a></li> <li><a href="/wiki/Metonic_cycle" title="Metonic cycle">Metonic cycle</a></li> <li><a href="/wiki/Milankovitch_cycles" title="Milankovitch cycles">Milankovitch cycles</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;line-height:1.2em;background:#cc9;"><a href="/wiki/Geologic_time_scale" title="Geologic time scale">Geologic time</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" 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:8.0em;font-weight:normal;background:#cc9;">Concepts</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Deep_time" title="Deep time">Deep time</a></li> <li><a href="/wiki/Geological_history_of_Earth" title="Geological history of Earth">Geological history of Earth</a></li> <li><a href="/wiki/Geologic_time_scale#Terminology" title="Geologic time scale">Geological time units</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight:normal;background:#cc9;">Standards</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Global_Standard_Stratigraphic_Age" title="Global Standard Stratigraphic Age">Global Standard Stratigraphic Age (GSSA)</a></li> <li><a href="/wiki/Global_Boundary_Stratotype_Section_and_Point" title="Global Boundary Stratotype Section and Point">Global Boundary Stratotype Section and Point (GSSP)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight:normal;background:#cc9;">Methods</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Chronostratigraphy" title="Chronostratigraphy">Chronostratigraphy</a></li> <li><a href="/wiki/Geochronology" title="Geochronology">Geochronology</a></li> <li><a href="/wiki/Isotope_geochemistry" title="Isotope geochemistry">Isotope geochemistry</a></li> <li><a href="/wiki/Law_of_superposition" title="Law of superposition">Law of superposition</a></li> <li><a href="/wiki/Luminescence_dating" title="Luminescence dating">Luminescence dating</a></li> <li><a href="/wiki/Samarium%E2%80%93neodymium_dating" title="Samarium–neodymium dating">Samarium–neodymium dating</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;line-height:1.2em;background:#cc9;"><a href="/wiki/Chronological_dating" title="Chronological dating">Chronological<br />dating</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" 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:8.0em;font-weight:normal;background:#cc9;"><a href="/wiki/Absolute_dating" title="Absolute dating">Absolute dating</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Amino_acid_dating" title="Amino acid dating">Amino acid racemisation</a></li> <li><a href="/wiki/Archaeomagnetic_dating" title="Archaeomagnetic dating">Archaeomagnetic dating</a></li> <li><a href="/wiki/Dendrochronology" title="Dendrochronology">Dendrochronology</a></li> <li><a href="/wiki/Ice_core" title="Ice core">Ice core</a></li> <li><a href="/wiki/Incremental_dating" title="Incremental dating">Incremental dating</a></li> <li><a href="/wiki/Lichenometry" title="Lichenometry">Lichenometry</a></li> <li><a href="/wiki/Paleomagnetism" title="Paleomagnetism">Paleomagnetism</a></li> <li><a href="/wiki/Radiometric_dating" title="Radiometric dating">Radiometric dating</a> <ul><li><a href="/wiki/Lead%E2%80%93lead_dating" title="Lead–lead dating">Lead–lead</a></li> <li><a href="/wiki/K%E2%80%93Ar_dating" title="K–Ar dating">Potassium–argon</a></li> <li><a href="/wiki/Radiocarbon_dating" title="Radiocarbon dating">Radiocarbon</a></li> <li><a href="/wiki/Uranium%E2%80%93lead_dating" title="Uranium–lead dating">Uranium–lead</a></li></ul></li> <li><a href="/wiki/Tephrochronology" title="Tephrochronology">Tephrochronology</a></li> <li><a href="/wiki/Luminescence_dating" title="Luminescence dating">Luminescence dating</a></li> <li><a href="/wiki/Thermoluminescence_dating" title="Thermoluminescence dating">Thermoluminescence dating</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight:normal;background:#cc9;"><a href="/wiki/Relative_dating" title="Relative dating">Relative dating</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Fluorine_absorption_dating" title="Fluorine absorption dating">Fluorine absorption</a></li> <li><a href="/wiki/Nitrogen_dating" title="Nitrogen dating">Nitrogen dating</a></li> <li><a href="/wiki/Obsidian_hydration_dating" title="Obsidian hydration dating">Obsidian hydration</a></li> <li><a href="/wiki/Seriation_(archaeology)" title="Seriation (archaeology)">Seriation</a></li> <li><a href="/wiki/Stratigraphy_(archaeology)" title="Stratigraphy (archaeology)">Stratigraphy</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;line-height:1.2em;background:#cc9;">Genetic methods</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a class="mw-selflink selflink">Molecular clock</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;line-height:1.2em;background:#cc9;">Linguistic methods</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Glottochronology" title="Glottochronology">Glottochronology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;line-height:1.2em;background:#cc9;">Related topics</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Chronicle" title="Chronicle">Chronicle</a></li> <li><a href="/wiki/New_Chronology_(Fomenko)" class="mw-redirect" title="New Chronology (Fomenko)">New Chronology</a></li> <li><a href="/wiki/Synchronoptic_view" title="Synchronoptic view">Synchronoptic view</a></li> <li><a href="/wiki/Timeline" title="Timeline">Timeline</a></li> <li><a href="/wiki/Year_zero" title="Year zero">Year zero</a></li> <li><a href="/wiki/Floruit" title="Floruit">Floruit</a></li> <li><a href="/wiki/Terminus_post_quem" title="Terminus post quem">Terminus post quem</a></li> <li><a href="/wiki/ASPRO_chronology" title="ASPRO chronology">ASPRO chronology</a></li></ul> </div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" 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=Molecular_clock&oldid=1252817864">https://en.wikipedia.org/w/index.php?title=Molecular_clock&oldid=1252817864</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:Evolutionary_biology_concepts" title="Category:Evolutionary biology concepts">Evolutionary biology concepts</a></li><li><a href="/wiki/Category:Molecular_evolution" title="Category:Molecular evolution">Molecular 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